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COMPREHENSIVE INVENTORIES OF SELECTED BIOLOGICAL RESOURCES WITHIN TARGETED WATERSHEDS AND ECOLOGICAL CORRIDORS OF SOUTHWESTERN EL SALVADOR IMPROVED MANAGEMENT AND CONSERVATION OF CRITICAL WATERSHEDS PROJECT Biodiversity field inventories in selected watersheds 1 SEPTEMBER 2009 THIS PUBLICATION HAS BEEN PRODUCED FOR REVIEW BY THE UNITED STATES AGENCY FOR INTERNATIONAL DEVELOPMENT. IT WAS PREPARED BY DEVELOPMENT ALTERNATIVES INC. (DAI) AND SALVANATURA. COMPREHENSIVE INVENTORIES OF SELECTED BIOLOGICAL RESOURCES WITHIN TARGETED WATERSHEDS AND ECOLOGICAL CORRIDORS OF SOUTHWESTERN EL SALVADOR IMPROVED MANAGEMENT AND CONSERVATION OF CRITICAL WATERSHEDS (IMCW) PROJECT Contract No. EPP-I-00-04-00023-00 Strategic Objective No. 519-022 Contractor Name: DAI Date: September 2009 Oliver Komar, editor. Research Supervisor, Improved Management and Conservation of Critical Watersheds Project, and Director, Conservation Science Department, SalvaNATURA, San Salvador, El Salvador Chapter authors are Oliver Komar, José L. Linares, Francisco Chicas Batres, José Alberto González Leiva, Vladlen Henríquez, Xiomara Henríquez, Luis E. Girón, Melissa Rodríguez, and James G. Owen. Recommended citation: Komar, O. (Editor). 2009. Comprehensive Inventories of Selected Biological Resources within Targeted Watersheds and Ecological Corridors of Southwestern El Salvador. USAID El Salvador, Improved Management and Conservation of Critical Watersheds Project. Or for individual chapters, following this example: Linares, J. L. 2009. Flora inventory in Southwestern El Salvador. in Komar, O. (editor). Comprehensive Inventories of Selected Biological Resources within Targeted Watersheds and Ecological Corridors of Southwestern El Salvador. USAID El Salvador, Improved Management and Conservation of Critical Watersheds Project. This study is made possible by the support of the American People through the United States Agency for International Development (USAID.) The contents of this study are the sole responsibility of Development Alternatives, Inc. (DAI) and do not necessarily reflect the views of USAID or the United States Government. Cover: Cuscatlania vulcanicola, a species rediscovered in the Coatepeque watershed. Photo by Frank Sullyvan Cardoza. 2 Biodiversity field inventories in selected watersheds TABLE OF CONTENTS ACKNOWLEDGMENTS ............................................................................................................... 7 EXECUTIVE SUMMARY .............................................................................................................. 9 1. SYNTHESIS AND SUMMARY OF THE BIODIVERSITY FIELD INVENTORIES ................ 14 2. FLORA INVENTORY ............................................................................................................. 32 3. PRELIMINARY INVENTORY OF FISH SPECIES IN ELEVEN WATERSHEDS ................. 76 4. INVENTORY OF AMPHIBIANS AND REPTILES ............................................................... 107 5. INVENTORY OF BIRDS ...................................................................................................... 133 6. INVENTORY OF TERRESTRIAL MAMMALS ..................................................................... 203 REFERENCES ......................................................................................................................... 234 TABLES AND FIGURES TABLES 1. Geographic distribution of field days for the biodiversity surveys carried out during 2007 in the IMCW project area .................................................................................................. 18 2. Historical and new records of flora and fauna in the project area, through 2007 ................. 19 3. Species recorded in the IMCW project area, through 2007 .................................................. 21 4. Level of inventory completeness recorded by watershed ..................................................... 22 5. Level of inventory completeness in the municipalities ........................................................... 23 6. Level of inventory completeness recorded by ecosystem ..................................................... 24 7. Importance rankings by ecosystem ....................................................................................... 27 8. Summary of recommendations for conservation, organized by taxonomic group ................ 29 9. Sampling sites for trees and total botanical sampling effort .................................................. 34 Biodiversity field inventories in selected watersheds 3 10. Indicator species for inventory completeness of trees in the study area ............................ 38 11. Tree species recorded in the study area ............................................................................. 46 12. Flora of conservation importance in the project area .......................................................... 64 13. Inventory completeness for tree species and relative importance of ecosystems .............. 65 14. Plants endemic to the IMCW project area, with collection locations from the present study ........................................................................................................................................... 66 15. Botanical species of interest collected in the study area ..................................................... 67 16. Inventory completeness for tree species and relative importance of watersheds .............. 73 17. Inventory completeness for tree species and relative importance of municipalities ........... 74 18. Watersheds, municipalities, and ecosystems considered in the evaluation of the fish inventory ..................................................................................................................................... 80 19. List of fish species considered to be indicators of complete site inventories in southwestern El Salvador, but not yet recorded ........................................................................ 82 20. Fish recorded in southwestern El Salvador, by ecosystem, through 2008 ......................... 92 21. Level of fish inventory completeness by watershed in southwestern El Salvador .............. 99 22. Level of fish inventory completeness by municipality in southwestern El Salvador ......... 100 23. Level of fish inventory completeness by habitat in southwestern El Salvador.................. 101 24. Conservation-important fish in southwestern El Salvador ................................................. 102 25. Description of the sampling sites and effort for the herpetofauna study ........................... 109 26. Indicator species for inventory completeness .................................................................... 113 27. List of indicator species for conservation important sites .................................................. 116 28. Herpetofauna species recorded in the study area ............................................................. 118 29. Inventory completeness for amphibians and reptiles and relative importance of watersheds ............................................................................................................................... 125 30. Inventory completeness for amphibians and reptiles and relative importance of municipalities ............................................................................................................................ 126 31. Inventory completeness for amphibians and reptiles and relative importance of ecosystems ............................................................................................................................... 127 4 Biodiversity field inventories in selected watersheds 32. Inventory completeness for amphibians and reptiles and relative importance of protected natural areas ............................................................................................................ 128 33. Estimators of herpetofauna species richness in each protected natural area studied ...................................................................................................................................... 129 34. Estimators of herpetofauna species richness in each ecosystem studied ........................ 131 35. Localities where bird inventory field work was carried out by the IMCW project .............. 136 36. The 74 resident bird species, expected in any municipality or watershed of the study area, that serve as indicators of inventory completeness (―test‖ species) at the site level ..... 138 37. Bird species used as ―test species‖ to indicate relative levels of inventory completeness at the ecosystem level, in southwestern El Salvador ....................................... 142 38. Birds of national conservation concern (188 species), useful as indicators for site prioritization in the study area .................................................................................................. 152 39. Types of avian locality records available for the present analysis .................................... 163 40. Sampling effort for birds at each study site during 2007 field work ................................... 163 41. List of 431 bird species recorded in the project area ......................................................... 169 42. Inventory completeness for birds and relative importance of watersheds ........................ 198 43. Inventory completeness for birds and relative importance of municipalities ..................... 199 44. Inventory completeness for birds and relative importance of ecosystems........................ 200 45. Sites and sampling efforts for mammals ............................................................................ 205 46. Indicator species (mammals) of inventory completeness.................................................. 207 47. Mammals that are indicators of important sites for conservation ...................................... 209 48. Individual mammals counted during the study................................................................... 211 49. List of terrestrial mammal species recorded in southwestern El Salvador through 2007 . 221 50. Level of mammal inventory completeness in the protected natural areas ........................ 228 51. Inventory completeness for mammals and relative importance of watersheds ................ 228 52. Inventory completeness for mammals and relative importance of municipalities ............. 229 53. Inventory completeness for mammals and relative importance of ecosystems ................ 229 54. Estimators of terrestrial mammal species richness in the study area and in each ecosystem ................................................................................................................................. 231 Biodiversity field inventories in selected watersheds 5 FIGURES 1. The IMCW project area contains 11 watersheds and 25 municipalities, as well as several adjoining protected natural areas .................................................................................. 16 2. Ecoregions, natural ecosystems, and anthropogenic land uses in the study area ............... 17 3. improvements in the inventory of birds during 2007, for watersheds and municipalities ..... 25 4. Study area, watersheds, and sampling sites for flora ............................................................ 37 5. Selected photographs of flora taken during the study ..................................................... 60–63 6. Ichthyogeographic provinces of Central America .................................................................. 77 7. The Tropical Eastern Pacific biogeographic region ............................................................... 78 8. Study area in southwestern El Salvador, with watersheds and municipalities labeled ......... 81 9. Map of the sampling sites of the study of amphibians and reptiles during 2007 ................ 109 10. Selected photographs of amphibians and reptiles taken during the study ............... 120–123 11. Accumulation curve of herpetofauna species from each protected natural area studied ...................................................................................................................................... 129 12. Accumulation curve for herpetofauna species from southwestern El Salvador ................ 130 13. Accumulation curve for herpetofauna species from each ecosystem studied .................. 131 14. Map of bird inventory field sites during 2007 ..................................................................... 135 15. Selected photographs of birds taken during the field study....................................... 165–168 16. Increment in inventory completeness index for birds in watersheds, resulting from the field study ................................................................................................................... 197 17. Map of sites sampled for mammals in 2007 ...................................................................... 204 18. Mammal photographs taken during the study............................................................ 217–220 19. Species accumulation curve for terrestrial mammals in southwestern El Salvador ......... 230 20. Accumulation curves for terrestrial mammal species by ecosystem ................................. 231 6 Biodiversity field inventories in selected watersheds ACKNOWLEDGMENTS This study was funded by a contract from USAID to DAI, Inc. and a subcontract to SalvaNATURA. This study was made possible by the support of the land managers who gave their permissions to carry out field work, including managers of the Protected Natural Areas System at the Ministry of Environment and Natural Resources, in particular Zulma Ricord de Mendoza, Walter Rojas, Alfonso Sermeño, and Patricia Quintana. In the natural areas visited, we thank the following for their collaboration: Carlos Batres of Finca Monte Carlo Estate at Laguna de Las Ninfas; Nidia Lara of AGAPE at Plan de Amayo; the Instituto Salvadoreño de Turismo for work at Cerro Verde; Martha Huezo de Sandoval for lodging at Playa Los Cóbanos; FUNDARRECIFE, especially Luis Pineda, at Los Cóbanos; Walter Martínez of Asociación de Desarrollo Comunal Nueva Esperanza (ADESCONE) at Bosque Santa Rita; COEX and Finca El Pireo at Laguna de Las Ranas; Cooperativa San Rafael Los Naranjos at Cerro El Aguila; María Isabel Morales of Asociación Salvadoreña para la Conservación del Medio Ambiente (ASACMA) at Complejo San Marcelino; staff of the Natural Areas Department of SalvaNATURA (Francisco Soto and Enrique Fuentes in particular) at Los Volcanes National Park and at El Imposible National Park. Also personnel of CATIE and Cooperativa María Auxiliadora for assistance at Los Volcanes National Park. Personnel of the Santo Domingo de Guzmán mayor’s office assisted with visits to the municipality. The National Civil Police provided lodging at Barra Salada. Botanical field trips and reports were assisted by Frank Sullyvan Cardoza, Vladlen Henríquez, Xiomara Henríquez, and Manuel Méndez. Herpetology field trips were accompanied by student volunteers Diana Quijano, Eder Caceros, Esmeralda Martínez, Jorge Herrera, and Vanessa Amaya. Karla Lara collected herpetofaunal data at Los Cóbanos. Mammalogy field trips were assisted by Arnulfo Morán, Elias Delgado, Alexandro Molina, and volunteer students Karla Lara, Stefany Henríquez, Verónica Guzmán, Jonathan Hernández, Enrique Fajardo, and Jorge García. Ornithology field trips technicians included Jesse Fagan, Carlos Funes, Jorge Jiménez, Sofia Trujillo, Iselda Vega, and Carlos Zaldaña and several volunteers. All of the field trips were greatly assisted by numerous park wardens, for whom we are grateful, and apologize for not listing all of their names. Similarly, numerous officers of the National Civil Police, Environment division, provided security and assistance at several field sites. The data in the SalvaNATURA flora and fauna database comes from many sources, both published and unpublished. Of particular note are extensive unpublished observations of birds contributed by Néstor Herrera, Ricardo Ibarra, Tom Jenner, Oliver Komar, Alvaro Moises, and Walter Thurber. The data base contains thousands of banding and observation records from the SalvaNATURA bird monitoring program, collected mostly by Lety Andino, Vicky Galán, Roselvy Juárez, Jennifer Smith, and their assistants. Biodiversity field inventories in selected watersheds 7 Gerrit Davidse of the Missouri Botanical Garden graciously provided all the records from that institution that were requested. Frank Sullyvan Cardoza assisted with the flora database of the Escuela Agrícola Panamericana. Héctor Manuel Quezada, Karen Xiomara García, and German Bladimir Sánchez provided valuable collaboration in the compilation of información in the fish database. Authorities and technical staff of the Natural History Museum of El Salvador and Miriam Cortez de Galán, coordinator of the zoology collections at the School of Biology of the University of El Salvador, facilitated the review of preserved fish collections. Maps were prepared by Vladlen Henríquez, Juan Felipe Gutiérrez, and Luis Girón. For the many contributors, we extend sincere thanks and apologize for not listing everyone’s name. All chapters were reviewed by Oliver Komar. Selected chapters were reviewed by Néstor Herrera, Ricardo Ibarra Portillo, and Enrique Barraza. Marta Lilian Quezada and Melissa Rodríguez assisted with formatting. The director of the Improved Management and Conservation of Critical Watersheds Project during the implementation of this study was Steve Romanoff. 8 Biodiversity field inventories in selected watersheds EXECUTIVE SUMMARY Comprehensive biological inventories were carried out during the second half of 2007 as part of the Improved Management and Conservation of Critical Watersheds (IMCW) Project, in southwestern El Salvador. This field work focused on trees, herpetofauna (amphibians and reptiles), mammals, and birds in the principal natural areas, such as at Los Volcanes and El Imposible National Parks, and in targeted watersheds and ecological corridors. Specific field trips were designed to fill gaps in the biological inventory. Also, bibliographical and museum data on fishes were collected and reviewed. The objective of this work was to develop more complete information on the status of biological resources, and effectively complete baseline information on distribution of key indicator species. Widely-distributed species that are habitat generalists were used as indicators of inventory progress, permitting the estimation of relative levels of inventory completeness for the entire project area and also at the finer scales of watersheds and municipalities, and in some cases, protected natural areas. Globally and nationally threatened (red-listed) species were used as indicators for conservation importance of sites. Field surveys were carried out in six or more priority watersheds for each of the taxonomic groups studied, but not all taxonomic groups were studied in every watershed visited. In all, field studies of flora and/or fauna were carried out in 10 of the Project area’s 11 watersheds during the study period. The studies increased the completeness of the biological inventory in 10 different ecosystems, and in 11 municipalities. The study area contains the country’s two largest national parks, El Imposible and Los Volcanes, and other important natural areas such as the Los Cóbanos coral reef, with many species of conservation importance. The Barra de Santiago mangrove estuary is also well known for important natural resources, including some unique aquatic species found nowhere else in the country. Field work was carried out at Los Volcanes National Park (plants, herpetofauna, mammals, birds), El Imposible National Park (plants, herpetofauna, mammals), and numerous corridor sites, such as within the Barra de Santiago—Garita Palmera estuaries (reptiles, mammals, birds), the El Imposible—Los Volcanes corridor (all taxonomic groups), and Barra de Santiago—El Imposible (birds). Fish resources were evaluated, via a desk study, in all of these areas. The biodiversity inventory in the study area is probably the most complete of any area in El Salvador. We have identified 2719 species, including 584 trees and 1287 other plants, 220 fishes, 96 amphibians and reptiles, 101 mammals, and 431 birds, based on a cumulative database of nearly 40,000 modern and historical locality records. More than 1% of the species (37) are globally threatened (red-listed), and 15% (402) are nationally threatened. Some of the country’s rarest species exist only in the project area. Of special note are at least 22 vertebrate species (without considering fishes) that are restricted in El Salvador to the area’s natural habitats. One fish, described for science in 2007, is considered restricted (in the world) to Lake Coatepeque (Schmitter-Soto 2007). Even more impressive, at least 14 plant species Biodiversity field inventories in selected watersheds 9 (10 trees and four herbs) are unique to the study area, including 6 new plant species discovered for science during the field work; these are natural resources found nowhere else in the world. FLORA The field work was carried out during 65 days, between the months of June 2007 and January 2008, recording a total of 1164 herbarium samples coming from approximately 707 vascular plant species. The results found include up to 11 probable new species for science, plus two new records for Central America, 10 new records for El Salvador, and two rediscoveries. Of the 584 tree species recorded, 23 of these are on the IUCN list of globally threatened (redlisted) species and one more should be included due to its rarity. Fourteen plant species are restricted in the world to the area of the project, including 10 trees and 4 herbs. Six of these endemic plants have yet to be described for science. At the watershed level, the Cara Sucia watershed presents the most advanced tree inventory, with 82% of the expected tree species recorded. Before the execution of this study, only 64% had been recorded. At the municipal level, San Francisco Menéndez presents an 86% level of inventory completion, an increase from 68%. At the ecosystems level, Humid Forest and Savannas present 100% inventory completion, based on the presence of expected tree species, and the Cloud Forest 95%. Dry Forest presents an 86% level of completion. The Dry Forest’s ecosystem presents a greater amount of species that are recognized as important for conservation, 12 species; followed by Humid Forest with 9 and Cloud Forest with 4 species. FISH A comprehensive bibliographic revision was made on the fish studies in the study area up until 2007, and the collections at the School of Biology of the University of El Salvador and at the Museum of Natural History of El Salvador were also verified. Only 753 records of fish species were found, documenting 220 fish species, of which 19 are Freshwater, 16 Freshwater-Brackish, 57 Estuarine, 21 Marine, 18 Reef-Marine and 89 Reef. Consequently, the level of inventory completion per ecosystem was: Freshwater 63% of expected species recorded, Estuarine had 56%, Reef had 55% and Marine barely reached 18%. The estimated percentage progress of the inventory for the whole study area was 41%. The fish inventories within individual watersheds are very incomplete. The greatest inventory completion estimates at the watershed level were presented in Coatepeque with 32%, Los Cóbanos with 29%, Barra de Santiago with 24%, and Cara Sucia with 12%. The analysis by municipality shows records in just seven municipal districts, of which Acajutla has reached 27% inventory completion, Jujutla 24%, Tacuba and San Francisco Menéndez 19% and 12% respectively, while Sonsonate and Chalchuapa each register 8%. Fifty-one fish species have been recognized as important for conservation, including freshwater species and others that are strictly reef-associated. According to the IUCN Red List, three reef species and one marine species in the study area are globally threatened. The 10 Biodiversity field inventories in selected watersheds most important areas for conservation are the Los Cóbanos Reef System and Barra de Santiago estuary, as these are deemed highly productive areas and operate as integrated habitats for many coastal fish populations. Lake Coatepeque and the fluvial systems of the area are also included herein as conservation priorities, as these maintain the freshwater native species. AMPHIBIANS AND REPTILES Field work was carried out for a 130-day period, during the months of June and December 2007, registering a total of 96 species (22 amphibian species and 74 reptile species) with a total of 1279 records. The results obtained included 20 new records, five new reports for the project area and 17 new records at the departmental level. Of the 96 species that were registered, 50 are on the national or global threatened species list and of these eight are globally threatened. At the watershed levels, the Izalco watershed presents a more advanced level of inventory completion with 74% of the expected species (indicators of inventory completion). Before the execution of this Project, the level of completion for this watershed was 47%. At the municipal level, San Francisco Menéndez presents a high level of inventory completion at 92% (an 18% increase). At the ecosystems level, the Dry Forest presents a relatively complete inventory (100% of the indicator species were detected) and another four ecosystems have completed inventories in a similar or greater percentage to 80%. At the level of the six Protected Natural Areas, where this study was carried out, the El Imposible National Park presents an inventory completeness of 85%. The five remaining natural areas present a level of inventory completenesse between 65 and 80%. The Humid Forest ecosystem presents a greater amount of conservation importance species (28 species) followed by freshwater ecosystems (12 species) and the Dry Forest (10 species). BIRDS The ornithology team worked for 116 days in the field, at 8 sites, comprising 7 watersheds, 8 municipalities, and 6 ecosystems. The sites included 7 protected natural areas, and some non-protected areas. The team logged 10,724 mistnetting hours, and 296 hours of observations, generating 4,068 new locality records, supplementing the 21,724 locality records previously available from southwestern El Salvador. The bird list for the study area was increased from 412 to 424 bird species by the field work. Seven additional species (mostly marine species) have been discovered independently, thus the project area’s bird list is now 431 species (79% of the country’s avifauna). Ten of 11 watersheds now have reasonably complete inventories (on average, 86% of the expected inventory-indicator species have been recorded), compared to five watersheds with well-advanced inventories prior to this study. Only eight municipalities have sufficiently complete (>75%, mean 92%) inventories for analysis, compared with five prior to the study. Six of eight ecosystems now have complete inventories (>80%, mean of 90%). These 10 watersheds, 8 municipalities, and Biodiversity field inventories in selected watersheds 11 6 ecosystems can be compared for their importance for conservation of threatened bird species. The 188 nationally threatened bird species used as indicators for prioritizing sites of national conservation importance include 31 that are critically endangered, 69 endangered, and 88 vulnerable. The Cara Sucia and the Barra de Santiago watersheds (both include parts of El Imposible National Park and parts of Barra de Santiago estuary) have the highest conservation importance, with more than 70% of the area’s nationally-threatened bird species present. They are distantly followed by the Coatepeque and Izalco watersheds (which both include parts of the Apaneca-Ilamatepec Biosphere Reserve). At the municipal level, the most important cities are San Francisco Menéndez (El Imposible National Park and Santa Rita Forest) and Jujutla (Barra de Santiago Protected Natural Area), with 68% and 57% of the area’s threatened bird species, respectively. Estuaries have 39% of the threatened bird species (including many shorebirds and water birds). Although Humid Forest and Dry Forest have fewer threatened birds (33% and 26%, respectively), these are mostly forest specialists, which are at risk from fragmentation and climate change. MAMMALS Field work was carried out during 105 days, registering 1296 individuals of 72 mammal species, of which 49 were captured, 17 photographed with camera traps, 21 were recorded by observations and five were recorded only outside of the standardized sampling. Four species were recorded for the first time in the whole study area (including one that was new for the country, Van Gelder’s Bat Bauerus dubiaquercus), increasing the mammal list for the study area to 101 species. We estimated that an index of inventory completion for the study area went from 92% to 100% after this study, suggesting that the mammal inventory is virtually complete. Nonetheless, specific natural areas still have incomplete lists. The protected natural areas with the most advanced mammal inventories are: El Imposible (81%), Santa Rita (63%), and Los Volcanes (52%). Twenty-two mammal species recorded in the study area are nationally threatened, or are indicators of conservation-importance sites. Such species include the Margay Cat (Leopardus wiedii), the Ocelot (Leopardus pardalis), the River Otter (Lontra longicaudis), and Baird’s Tapir (Tapirus bairdi) (these last three species have been recorded in previous years, but not during this study). At the national level, 11 threatened species and seven endangered species are reported (MARN 2004), as well as one species considered globally endangered (IUCN 2008). Based on the presence of nationally and globally threatened and endangered terrestrial mammal species, the Dry Forest ecosystem is more important than the Humid Forest. However, the rest of the ecosystems, the majority of the watersheds, the municipalities, and the protected natural areas cannot be compared for conservation importance, because their mammal inventories are not yet reasonably complete. 12 Biodiversity field inventories in selected watersheds RECOMMENDED STRATEGIES FOR CONSERVATION AND SUSTAINABLE ECONOMIC DEVELOPMENT Conservation strategy recommendations for estuaries focus mostly on education and awareness-building, while for upland habitats, forests need to be expanded and organized to connect fragments via altitudinal corridors, ideally along rivers. More flora and faunal inventory work is needed (especially flora), as well as long-term population monitoring, in order to track progress with climate change adaptability projects that should be implemented in the region. Ten recommendations for sustainable economic development identify the need to increase or promote activities such as payments for environmental services, sustainable forestry, urban and rural planning, agricultural certification, wildlife farms for diverse markets and export opportunities, development of sustainable businesses (such as ecotourism services), and government involvement in facilitating access by ecotourists to natural areas. Scientific tourism presents great potential, as a sustainable use of natural resources, and could also generate greater demand among international tourists for visiting natural areas in the project area, such as the national parks, biosphere reserves, and coral reef. A reintroduction program for the locally-extinct Scarlet Macaw can help rally multilateral support for sustainable development in the Barra de Santiago–El Imposible biological corridor. Certain development ―opportunites‖ may cause more harm than good, however, and much care should be taken before promoting practices such as expansion of biofuels or other high-input agricultural crops, or factories that require high inputs of water or fuels. Biodiversity field inventories in selected watersheds 13 CHAPTER 1: SYNTHESIS AND SUMMARY OF THE BIODIVERSITY FIELD INVENTORIES Oliver Komar Conservation Science Department, SalvaNATURA San Salvador, El Salvador okomar@salvanatura.org 1. INTRODUCTION A key element to the sustainable economic development of southwestern El Salvador is conservation of biodiversity. This region of El Salvador contains several of the country’s most important wildlife reserves, including the two largest national parks and one of the country’s two biosphere reserves. It has substantial tourism and ecotourism potential, with resources such as ―la Ruta de las Flores‖, El Salvador’s largest volcano (Ilamatepec) and its youngest (Izalco), the country’s only coral reef, and extensive beaches. Despite a long history of biodiversity investigation in the area, the flora and fauna inventory was both taxonomically and geographically incomplete (Komar 2007). As a signatory to the Convention on Biological Diversity, for some time it has been a national priority in El Salvador to complete the inventories of all flora and fauna groups (MARN 2000a). Knowledge of the wildlife species present and their patterns of distribution and habitat use are essential for truly sustainable development to take place. A principal objective of the Improved Management and Conservation of Critical Watersheds (IMCW) Project, which was planned to run from the end of 2006 through the end of 2009, is the conservation of biodiversity in several watersheds of Ahuachapán and Sonsonate, southwestern El Salvador. One of the scheduled outputs of this project is the realization of a comprehensive, multi-taxon biodiversity field inventory in selected watersheds, which this report summarizes. The IMCW Project also commissioned a preliminary study of the baseline biodiversity information available, which was completed prior to the initiation of the present study (Komar 2007). That report summarized the state of knowledge about plants, butterflies, amphibians, reptiles, mammals, and birds. It determined that 2,457 species of flora, butterflies, amphibians, reptiles, mammals, and birds had been recorded through 2006. More than half (1716 species) were plants. They included 325 nationally threatened species and 24 globally threatened species of flora and fauna. This latter group includes mostly trees and just seven vertebrates (four sea turtles, two frogs, and tapir which is presumed to be extirpated). However, there are extensive areas where practically no inventory efforts have been 14 Biodiversity field inventories in selected watersheds undertaken, and the inventory is still far from complete for most taxonomic groups. Also reported in the baseline preliminary inventory was the presence of 48 plant species and 22 vertebrates (excluding fish) in the Project area that have not been recorded in other localities of El Salvador (Komar 2007). Reported separately was a unique fish in Lake Coatepeque, which thus became the only endemic vertebrate known to be unique to El Salvador (Schmitter-Soto 2007). The IMCW project was interested in determining if specific watersheds or sites within southwestern El Salvador were more important than others for conserving biodiversity. In order to conserve overall diversity, maintaining a large number of species in a region, a strategy must focus on assuring that habitat specialists (and their habitats) are preserved, even if some specialized habitats have relatively few (but unique) species. It is not necessary for every habitat conserved to be of high species richness or diversity. Thus in seeking conservation priorities across sites, one must look for unique and rare species at the sites, and not simply count species. Comparison of sites requires the existence of complete inventories at the site level, rather than standardized counts of species. The purpose of the present study was to advance the state of the flora and fauna inventories for a series of sites and ecosystems in southwestern El Salvador, such that relative value for biodiversity conservation could be assessed. The following chapters include the evaluation of the current state of inventories (at the beginning of 2008) for flora, fishes, herpetofauna (amphibians and reptiles), mammals, and birds in the entire IMCW project area. The present chapter includes a summary and comparison of the state of the inventories among these taxonomic groups. The main purpose is to establish conservation priorities and recommendations for future economic development. 2. STUDY AREA AND METHODS The study area includes 11 watersheds that drain into the Pacific Ocean (Fig. 1), defined following the preliminary analysis that served as a baseline to this study (Komar 2007), as follows. The relatively large Río Banderas watershed was divided into three watersheds for the purpose of biological analysis: the lower Río Banderas (coastal plain), the Izalco watershed which drains most of Los Volcanes National Park, and the San Julian watershed which drains the northwestern sector of the Sierra del Bálsamo. The Los Cóbanos area was treated as a single watershed, although technically it consists of seven minor watersheds, each draining a small segment of the Los Cóbanos plain (an ancient landslide originating from the Santa Ana volcano); the divides of these tiny watersheds are almost indistinguishable in the field. The large Barra de Santiago watershed was divided into two: ―Cara Sucia,‖ which drains the western two-thirds of El Imposible National Park, and ―Barra de Santiago‖ which drains the eastern third of the national park and several valleys to the east. Most of the rivers in the Cara Sucia watershed are drained or go underground before they reach the ocean. Data previously available (Komar 2007) was combined with new data generated from the field work for conservation analyses of 11 watersheds and 25 municipalities that form the IMCW Project area in southwestern El Salvador (Fig. 1). This area covers the southern region of the Biodiversity field inventories in selected watersheds 15 department of Ahuachapán, the entire department of Sonsonate, and marginal areas of the departments of Santa Ana and La Libertad. Three additional municipalities within the Project area (Ahuachapán, Atiquizaya, and El Congo) were excluded from the analysis because only insignificant areas of their territories are within the Project area. In total, the Project area covers 1545 km 2, or 7.4% of El Salvador. The Project area includes four terrestrial ecoregions, nine natural ecosystem types and a variety of anthropogenic land uses (Fig. 2). It also includes dozens of fragments of natural habitat areas, including 10 natural areas in various states of formal protection that were visited during the field surveys. Field inventories were conducted during 2007 (and January 2008) for selected taxonomic groups: trees (and opportunistically other types of flora), amphibians, reptiles, mammals, and birds. Other groups, such as insects and other invertebrates, were not included in the study plan because they are harder to identify, and relative importance for conservation has not been established at the species level (i.e, we don’t know which species are threatened with extinction). Butterflies were originally included in the study plan but removed when negotiations with a potential field team were unsuccessful. Fish were originally excluded from the study plan because El Salvador lacked a list of threatened fish species and because of lack of taxonomists in the country. Part of the savings from the canceled buttefly inventory were reinvested in a preliminary baseline evaluation of the state of knowledge of the fish inventory, which is included as a chapter in this report. FIGURE 1. THE IMCW PROJECT AREA CONTAINS 11 WATERSHEDS AND 25 MUNICIPALITIES, AS WELL AS SEVERAL ADJOINING PROTECTED NATURAL AREAS. 16 Biodiversity field inventories in selected watersheds FIGURE 2. ECOREGIONS, NATURAL ECOSYSTEMS, AND ANTHROPOGENIC LAND USES (TOP TO BOTTOM) IN THE STUDY AREA. Maps prepared by J. F. Gutiérrez. Biodiversity field inventories in selected watersheds 17 TABLE 1. GEOGRAPHIC DISTRIBUTION OF FIELD DAYS FOR THE BIODIVERSITY SURVEYS CARRIED OUT DURING 2007 IN THE IMCW PROJECT AREA. WaterMuniciAmphibians Location Flora Mammals Birds shed(s) palities & Reptiles Cara Sucia, San Franciso El Imposible 25 20 40 0 Barra de Menéndez, National Park Santiago Tacuba Los Volcanes Izalco, Santa Izalco 15 20 20 20 National Park Ana Plan de San Julian Caluco 15 20 10 12 Amayo PNA Complejo San Izalco, Izalco 5 20 5 0 Marcelino Coatepeque PNA Río Grande Cerro El de Juayúa 5 0 0 0 Águila PNA Sonsonate Santa Rita y San Franciso Cara Sucia 0 20 5 20 Zanjón El Menéndez Chino PNA Laguna de Río Grande Las Ninfas de Apaneca 0 20 5 11 PNA Sonsonate Los Cóbanos Los Sonsonate, 0 10 10 20 PNA Cóbanos Acajutla Laguna de Río Grande Juayúa, las Ranas de 0 0 10 10 Chalchuapa PNA Sonsonate Santo Santo Domingo de San Pedro Domingo de 0 0 0 4 Guzmán Guzmán Barra Salada Lower Río Sector (Los Sonsonate 0 0 0 19 Banderas Cóbanos PNA) Total days of field work 65 130 105 116 PNA = Protected Natural Area 18 Biodiversity field inventories in selected watersheds TABLE 2. HISTORICAL AND NEW RECORDS OF FLORA AND FAUNA IN THE IMCW PROJECT AREA, THROUGH 2007 (SOURCE: SALVANATURA’S FLORA AND FAUNA DATABASE*). Other records Taxon Specimens (observations, Totals photos, etc.) Plants 5,313 2,686 7,999 Fish 615 138 753 Amphibians & reptiles 1,500 500 2,000 Mammals 1,200 1,616 2,816 Birds 2,076 23,716 25,792 All combined 10,704 28,656 39,360 *Primary sources include museums, natural history collections (national and foreign), data published in scientific literature and gray literature (project technical reports, etc.), and unpublished data contributed by independent researchers. More details are included in the thematic chapters of this report and in a previous report (Komar 2007). Field collections or observations for the taxonomic groups were carried out in at least six watersheds, as required in the study’s terms of reference, but not necessarily in the same watersheds for each group. The locations for each field inventory during 2007 and early 2008 are given in Table 1. The various field methods used are detailed in the following chapters for each taxonomic group. In general, a team of botanists and zoologists sought to document the presence and relative abundance of as many species as possible, but with a focus on finding species that were not previously documented in the area. Thus field sites were selected with the aim of filling gaps in the inventory, rather than with the aim for standardized sampling (Komar 2007). This was justified by the objective of evaluating conservation importance, rather than comparing species diversity among sites. In order to ducument the presence of species, the biologists focused on collecting voucher specimens when possible, with one specimen collected for the El Salvador national museum of natural history, and a duplicate collected for a natural history museum or herbarium located outside of El Salvador. Additional specimens captured by various trapping methods were released unharmed. When possible, digital cameras were used to document the presence of species that could not be captured. The field teams used standard capture methods, which do not harm the individuals, such as mist nets for birds and harp traps for bats, under license from the Ministry of Environment and Natural Resources. Camera traps (movement-triggered digital and analog cameras) were used to document the presence of larger mammals. The new data collected from the field surveys was logged in flora and fauna databases managed in SalvaNATURA’s conservation science department. These data were added to the more than 29,000 locality records from the Study Area that had been assembled prior to the field surveys (Komar 2007; Table 2). After completing the field surveys, the updated data set was used to determine the relative level of inventory completeness, both for the whole Biodiversity field inventories in selected watersheds 19 project area as well as for individual watersheds, municipalities, and ecosystems. For some groups, completeness levels were evaluated for individual Protected Natural Areas. Completeness was evaluated using the same method used for the baseline evaluation (Komar 2007, following Gómez de Silva and Medellín 2001), which identifies the list of generalist and relatively common species that would be present in any site that has been subjected to a proper inventory effort. Because these are generalist species, they may exist in a variety of habitats, a variety of altitudes, and sites with or without disturbance. These species are used as a ―test‖ or indicator for the effectiveness of the efforts to collect data on all the species in an area, and the proportion of test species recorded represents an index of inventory completeness. When 100% of the test species have been recorded, the inventory is considered relatively complete, although some rare species may not yet have been recorded. The species that do not qualify as tests (indicators), such as species restricted to certain natural habitats (―habitat specialists‖), naturally rare species (predators at the top of the food chain), or species of cryptic behavior, may be present but not yet recorded. The method is nonetheless useful for identifying large gaps in the inventory, and to prioritize where to focus future inventory efforts. Threatened species (i.e., species that are considered endangered and require conservation efforts to prevent local or global extinction) were used as indicators of conservation importance. Threatened status was based on IUCN (2007), MARN (2004), Greenbaum and Komar (2005), and Komar et al. (2009). Conservation priorities among sites were determined by comparing the numbers of nationally and globally threatened species recorded from the field sites, with highest priority assigned to sites with the most threatened species. Since many threatened species are rare, sites with incomplete inventories may not be comparable simply because their threatened species have not yet been documented (Remsen 1994, Komar 2003, Greenbaum and Komar 2005, Komar 2007). Only sites with reasonably complete inventories were used to evaluate site conservation priorities. In the case of the bird inventory, statistics were used to demonstrate the level of completeness required to include a site in the analysis of conservation importance. The richness of threatened species was regressed against the inventory completeness index score to determine if inventory effort was a significant predictor of conservation importance. If the slope of the regression line was significantly different from zero (P<0.05), then sites with low completeness scores were removed from the analysis until the regression slope was not significantly different from zero (P>0.05), indicating that inventory completeness was not a good predictor for conservation importance. This occurred when all sites included in the analysis had reasonably complete inventories (see Chapter 6). 3. RESULTS More than 10,600 individual plants and animals were logged and mapped in the study area, permitting the documentation of numerous new species previously unknown from this area. As described in detail in the chapter on flora by José Linares, the 154 new tree species and many other new plant species documented included no fewer than 11 thought to be new species for science, as well as two first records for Central America, 10 additional first records for El Salvador, and two rediscoveries of species last reported by botanists in 1922, and known only from El Salvador. The project area appears to include all or part of a focal area for 20 Biodiversity field inventories in selected watersheds evolutionary diversification of plants, because at least 14 plant species are known only from the project area and nowhere else in the world. The herpetofaunal field survey documented five species not previously recorded in the study area, including one frog (amphibian) and four reptiles (Henríquez and Henríquez, this report). The mammal surveys found four new species for the study area (Girón et al., this report). The bird surveys reported 12 new species for southwestern El Salvador. In total, the taxa studied for this report number 2719 species (Table 3). Apparently 14 plant species and one vertebrate (a fish) found in the study area have never been detected elsewhere in the world. TABLE 3. SPECIES RECORDED IN THE IMCW PROJECT AREA, THROUGH 2007. 1 Nationally Globally Taxon All species threatened species threatened species Trees 584 17 23 Other plants (including 1287 78 1 orchids) Fish 220 51 4 Amphibians & reptiles 96 50 8 Mammals 101 18 1 Birds 431 188 0 All combined 2719 402 37 1 The species’ names are given in the following chapters. 3.1. IMPROVED FLORA AND FAUNA INVENTORIES The tree inventory improved dramatically in the study area, from 76% of the test species registered in the data base prior to field work, to 93% registered after completing the field phase. The fish inventory became available for the first time, with an estimate of 41% inventory completeness for the study area. The mammal inventory improved from 92% to 100%. The amphibians, reptiles, and birds inventories were already relatively complete (100% of test species recorded) before field work began, but those inventories improved greatly at the scale of watersheds (Table 4) and municipalities (Table 5), and several new species were recorded in the study area. Nonetheless, some watersheds and municipalities continue to be severely undersampled. The improvement in the overall coverage of inventory efforts and the relative completeness of the inventories is demonstrated graphically for the bird results (Fig. 3). For the first time, the state of the flora and fauna inventories have been evaluated at the level of ecosystems (Table 6) and in some cases, protected natural areas. For protected natural areas, only mammal and herpetofaunal inventories have been assessed for completeness (see Chapters 4 and 5). Individual areas have mammal inventories that are still very incomplete (generally 30–60% complete). The best inventoried protected natural area in southwestern El Salvador is El Imposible National Park, with 81% of the expected ―test‖ species documented. Biodiversity field inventories in selected watersheds 21 TABLE 4. LEVEL OF INVENTORY COMPLETENESS RECORDED BY WATERSHED. “Test” species recorded (%) 22 Watersheds Trees Fishes Amphibians & Reptiles Mammals Birds San Juan Barra Salada San Pedro Los Cóbanos San Julián Río Cauta Río Grande de Sonsonate Izalco Coatepeque Cara Sucia Barra de Santiago Complete Project Area 0 0 5 0 57 0 19 58 44 82 38 93 0 0 5 29 0 5 0 0 32 12 24 41 0 4 26 30 66 2 17 74 25 60 58 100 15 0 17 40 46 0 67 31 67 83 79 100 4 57 89 73 93 74 85 96 93 99 100 100 Biodiversity field inventories in selected watersheds COMBINED (proportion average) 4 12 28 33 52 16 38 52 52 67 60 86 TABLE 5. LEVEL OF INVENTORY COMPLETENESS IN THE MUNICIPALITIES. “Test” species recorded (%) Amphibians & Municipality Trees Fishes Mammals Birds Reptiles Acajutla 0 27 25 46 69 Apaneca 9 0 11 29 61 Armenia 0 0 0 19 0 Caluco 57 0 64 46 92 Chalchuapa 2 8 0 31 0 Concepción de Ataco 5 0 0 0 62 Cuisnahuat 0 0 0 0 0 Guaymango 0 0 0 0 0 Izalco 58 0 57 23 97 Juayúa 13 0 4 21 97 Jujutla 5 24 6 42 95 Nahuizalco 0 0 2 21 41 Nahulingo 0 0 0 0 27 Salcoatitán 0 0 0 0 0 San Antonio del Monte 0 0 0 0 0 San Francisco Menéndez 86 12 92 85 100 San Julián 1 0 30 0 8 San Pedro Puxtla 0 0 13 13 68 Santa Ana 45 0 13 60 77 Santa Catarina Masahuat 0 0 0 0 0 Santo Domingo de Guzmán 0 0 7 0 61 Sonsonate 3 8 43 10 82 Sonzacate 0 0 2 0 0 Tacuba 5 19 15 46 93 Tepecoyo 0 0 0 0 0 Biodiversity field inventories in selected watersheds COMBINED (proportion average) 32 22 4 52 8 13 0 0 47 27 34 13 5 0 0 75 8 19 39 0 14 29 0 35 0 23 TABLE 6. LEVEL OF INVENTORY COMPLETENESS RECORDED BY ECOSYSTEM. “Test” species recorded (%) 24 Ecosystem Trees Fishes Amphibians & Reptiles Terrestrial Mammals Birds Humid forest Dry forest Cloud forest Savanna (palm and morro) Freshwater Intertidal zone (rocks and beaches) Estuary Coral reef Marine 100 86 95 100 ------ ----63 -56 55 18 95 100 83 0 90 80 0 0 0 79 90 67 40 33 0 33 --- 98 97 98 35 84 81 83 64 64 Biodiversity field inventories in selected watersheds COMBINED (proportion average) 93 93 86 44 68 54 43 37 26 FIGURE 3. IMPROVEMENTS IN THE INVENTORY OF BIRDS DURING 2007, FOR WATERSHEDS (ABOVE) AND MUNICIPALITIES (BELOW). DARKER SHADING REPRESENTS MORE COMPLETE INVENTORIES. THE LEFTHAND COLUMN REPRESENTS THE STATE OF THE INVENTORY BEFORE FIELD WORK, AND THE RIGHTHAND COLUMN THE IMPROVED COVERAGE AFTER FIELD WORK. 3.2. PRIORITY AREAS FOR BIODIVERSITY CONSERVATION Several natural areas within the IMCW Project area are clearly priorities for biodiversity conservation, on a national and even global scale. As have several prior studies, the present inventory has demonstrated the presence of globally threatened species at most of the remaining natural areas. These areas have now been identified as globallyimportant Key Biodiversity Areas (Henríquez 2009) and Important Bird Areas (Komar & Ibarra Portillo 2009). In particular, the areas identified in those documents include Barra de Santiago, Plan de Amayo, San Marcelino, and Santa Rita reserves, as well as El Imposible and Los Volcanes National Parks. The latter park, along with surrounding areas that form the Apaneca-Ilamatepec Biosphere Reserve, were jointly identified as a Key Biodiversity Area. Much of the unprotected part of the Los Cóbanos peninsula was also identified as an Important Bird Area, because of the presence of a suite of dry forest bird species representative of the Pacific dry forest biome that stretches from northwestern México to Costa Rica. All of the aforementioned areas, including virtually all natural habitat remnants in the project area, are therefore of global importance for biodiversity conservation. Biodiversity field inventories in selected watersheds 25 Nonetheless, we considered the question: which areas within the study area stand out as having greater priority for biodiversity conservation? Our aim was to use nationally and globally threatened species as indicators of conservation importance. A condition for such an analysis is that the areas being compared are sufficiently well studied such that the value is not biased by the level of inventory effort. Unfortunately, for some of the desired comparisons, that condition was not met and the analysis could not be completed. For several taxonomic groups (trees, fish, amphibians, and reptiles), the biological inventories of watersheds were too variable or incomplete to permit comparisons. For mammals, only two watersheds could be analyzed, suggesting that the Cara Sucia watershed was more important than the Barra de Santiago watershed, probably due to the presence of the Santa Rita swamp forest in Cara Sucia. For birds, 10 watersheds were analyzed. Both Barra de Santiago and Cara Sucia had far greater importance than any other watershed, presumably because of El Imposible National Park (which straddles both watersheds). Barra de Santiago ranked slightly higher, probably because of the presence of more threatened waterbirds than in Cara Sucia. The next most important watersheds for birds were Coatepeque and Izalco, due to natural areas in Los Volcanes National Park and other areas of the Apaneca-Ilamatepec Biosphere Reserve. Although most groups could not be analyzed at the municipality level, the bird inventories were sufficiently complete in eight municipalities. Thus, it appears that San Francisco Menéndez (El Imposible National Park and Santa Rita Forest Protected Natural Area) and Jujutla (Barra de Santiago estuary) have considerably more conservation importance than Sonsonate (Los Cóbanos and Barra Salada Protected Natural Areas), which in turn is more important for birds than Tacuba, Izalco and Santa Ana (the latter two municipalities include Los Volcanes National Park and San Marcelino Complex of Protected Natural Areas). These results only partially mimic the results generated by analyzing birds at the watershed level. The inventories for the different taxonomic groups were more complete at the ecosystem level, permitting more useful analysis. Only fish data were excluded from this analysis (because of incomplete sampling), although current data strongly suggest a priority for coral reef habitat. The relative importance scores were not consistent among taxonomic groups, demonstrating the importance of including several taxonomic groups in the analysis. For example, the priorities for birds did not match the priorities for trees, mammals, amphibians, or reptiles (Table 7). The top-scoring ecosystems included humid forest (for trees, amphibians, and reptiles), dry forest (for mammals), and mangrove estuaries (for birds). When scores were combined, humid forest obtained the highest score. The humid forest ecosystem includes the permanently green forests growing at mid elevations, generally from 800 to 1800 meters above sea level; this is the area that has been most threatened by the expansion of coffee culture on montane hillsides. The cloud forest ecosystem scored consistently low across all groups. Although not part of the formal terms of reference for this study, some authors analyzed the relative conservation importance of the specific Protected Natural Areas studied. Sampling was sufficiently complete for such analysis only for amphibians and reptiles (birds were not considered, but may also have complete sampling). For the herpetofauna, El Imposible National Park scored highest, followed in declining order by Los Volcanes National Park, and then corridor sites Santa Rita, San Marcelino, Plan de Amayo, and Laguna de las Ninfas (See Chapter 4 for details). 26 Biodiversity field inventories in selected watersheds TABLE 7. IMPORTANCE RANKINGS BY ECOSYSTEM.* Amphibians & Combined Mammals Birds Reptiles (average rank) Humid forest 52 58 32 33 44 Dry forest 52 20 47 26 36 Cloud forest 16 12 -17 15 Savanna (palm and morro) 0 0 -5 2 Freshwater -26 -20 23 Intertidal zone ( beaches) -8 -13 11 Estuary ---39 39 *The rankings represent proportions of conservation-important indicator species (generally, threatened species) expected to be found in the study area. Ecosystem Trees Biodiversity field inventories in selected watersheds 27 4. RECOMMENDED CONSERVATION STRATEGIES Conservation strategy recommendations for estuaries focus mostly on education and awareness-building, while for upland habitats, forests need to be expanded and organized to connect fragments via altitudinal corridors, ideally along rivers. More flora and faunal inventory work is needed (especially flora), as well as long-term population monitoring, in order to track progress with climate change adaptability projects that should be implemented in the region. The recommendations from the thematic chapters are summarized by taxonomic group in Table 8. Some clear geographic conservation priorities are evident with respect to species of conservation importance, and in particular, globally threatened species. Threatened tree species, or plant species known only from southwestern El Salvador (unique world resources), are found mostly in the uplands of the Sierra de Bálsamo and Sierra de Apaneca. Remnant dry forests and limestone soil enclaves of the western end of the Sierra del Bálsamo at Plan de Amayo and nearby Protected Natural Areas stand out, as do the moist Sierra Madre de Chiapas forests of southfacing slopes at mid elevations in the Sierra de Apaneca, especially at El Imposible National Park. Also of importance, but less so, for plant conservation are the very humid montane forests (cloud forests) on the peaks of the Sierra de Apaneca, such as at Laguna de las Ranas, Cerro El Águila, Cerro Verde, and the Los Andes section of Los Volcanes National Park (collectively, the Apaneca-Ilamatepec Biosphere Reserve). The most important fish area in southwestern El Salvador appears to be the coral reef at Los Cóbanos, followed by freshwater habitats, in particular Lake Coatepeque. Globally threatened sea turtles occupy most of the area’s beaches for breeding, and also feed at Los Cóbanos reef. The globally vulnerable American Crocodile (Crocodylus acutus) is resident in the mangrove estuaries of the study area. Nationally threatened birds are concentrated in El Imposible National Park (Komar 2002). Relatively little natural area remains in the study area. Virtually every remnant natural area, as well as ecologically sustainable production areas such as shaded coffee plantations in the ecological corridors that connect the natural areas, present some justifications to protect local biodiversity. Site conservation priorities for fish and reptiles are very different than priorities for trees, amphibians, and birds, which makes prioritization particularly challenging. 28 Biodiversity field inventories in selected watersheds TABLE 8. SUMMARY OF RECOMMENDATIONS FOR CONSERVATION, ORGANIZED BY TAXONOMIC GROUP. Amphibians Actions Flora Fish Mammals Birds & Reptiles X X Fill geographic gaps X (coastal X X (atlas in the inventory (coastal zone) zone) mapping) X X Fill taxonomic gaps in X (Use (repeat the inventory, using (herbs, Anabat for during distinct detection bushes, high-flying spring methods epiphytes) bats) migration) Evaluate use of coffee plantations as X a biological corridor X Monitor populations X (frogs, sea of threatened species (esp. turtles, at permanent plots felines) crocodiles) Expand long-term monitoring of X common species Monitor habitat X quality using indicator (epiphytes) species Provide training to park rangers about the conservationX X important (threatened) species Provide X environmental X (especially education near (especially ref. coastal Protected Natural vipers) areas) Areas Undertake ecological research to better X X understand how species are using the environment Strengthen habitat and species protection at important X X conservation sites, and expand natural habitat cover Biodiversity field inventories in selected watersheds 29 Actions Flora Improve ecological function of rivers and riparian (and altitudinal) corridors Establish sustainable management programs at Lake Coatepeque and other lakes 5. Fish Amphibians & Reptiles Mammals X Birds X X RECOMMENDATIONS DEVELOPMENT FOR ECONOMIC The results of the taxonomic inventory provide abundant information about conservation priorities for biodiversity in the project area. Hopefully, the future economic development of the area can be sensitive to the needs of flora and fauna species, since this biodiversity is an integral part of the local natural heritage and a valuable natural resource that should be preserved and managed for the benefit of present and future generations. Biodiversity provides diverse benefits for all people (Martínez et al. 2009), and its conservation should be a priority not only in the Protected Natural Areas but also throughout the rural agricultural landscape (Harvey et al. 2008). Below are 10 suggestions for sustainable economic development suggested by the taxonomic inventory results. 1. Create financial incentives for the protection of natural habitats through the development of payments for environmental services, such as water conservation generated by forests, erosion control through abandonment of farming on steep slopes, and carbon sequestration by reforestation or by farming shaded permanent crops (coffee, cacao). 2. Develop the forestry industry. If extensive farmlands in the project area can be converted to sustainable forestry, biodiversity conservation can be benefited by increasing the functionality of ecological corridors. Forestry projects could be designed to benefit biodiversity by employing diversified plantings, including fruit trees or trees that otherwise provide foraging opportunities for wildlife. Forestry can also contribute to climate change adaptation and mitigation. 3. Keep urban development in the cities. Planned urban and rural development can help reduce the threat to biodiversity from new suburban residential areas sprouting up in former farmland, far from urban centers, and also prevent any further agricultural expansion. 4. Reduce agriculture’s impact on natural habitat, by adopting less intensive production techniques, such as organic production, and promoting certification programs that provide market incentives for adopting environmentally friendly practices. Such programs exist or are under development by SalvaNATURA for the major export crops in the project area, coffee and sugar cane. 30 Biodiversity field inventories in selected watersheds 5. Promote industries or businesses that provide jobs for the rural population, and otherwise reduce the threat of extractions of wildlife from their natural hábitats, such as wild animal farms (zoocriaderos). Wild species that could be farmed and exported for various uses and markets include: native palms, shrubs, ferns, orchids, and other flowers; freshwater fish such as tilapia and others; estuarine shell fish such as clams, mussels, conchas, crabs, and shrimp; crocodile and cayman, freshwater turtles, iguanas; deer, paca, armadillo, and rabbits. 6. Promote business practices that permit sustainable use of natural resources, such as ecotourism in the region’s national parks and Protected Natural Areas, and in estuaries or on beaches. 7. The national government could increase interest in sustainable tourism within southwestern El Salvador, by placing more signage on the highways for tourist attractions (such as the national parks), by improving some rural access roads to the parks, and by developing publicity campaigns for the natural areas, such as the biosphere reserves. 8. Biodiversity research, and scientific tourism in general, can have a multiplicative positive impact on local economies near the natural ecosystems of Protected Natural Areas. Not only does it bring some direct economic investment (construction of biological field stations, use of lodging facilities and local services by visiting scientists), but also can result in international publicity generated by publication of scientific investigation, which can greatly increase visitation from international ecotourists. 9. SalvaNATURA is currently developing a program that could help develop the market for sustainable tourism, and increase local pride in the native biodiversity. This program aims to reintroduce the locally extinct Scarlet Macaw (Ara macao) in the Barra de Santiago-El Imposible corridor (Bjork 2009), where the species can become a flagship for sustainable development in the corridor. Other high-profile reintroduction programs could be developed for species extirpated from the project area, such as Spider Monkeys (Ateles geoffroyi). These programs will require broad support and participation, not only from local communities but also local and national governments, donors, and businesses. 10. Business or agricultural activities that could harm the environment should be avoided. Examples include high-input crops (non-shade varieties of coffee, for example), or crops that may require further deforestation (such as biofuel crops), crops that require burning (certain varieties of sugar cane), factories that are not equipped to handle their waste properly or that require excessive quantities of water or fuel. Also not recommended are breeding programs of parrots for the pet industry, although at first glance, such an activity would seem to reduce pressure on the wild population. Nonetheless, it could also promote the practice of keeping parrots, which could create more demand for wild birds. Biodiversity field inventories in selected watersheds 31 CHAPTER 2: FLORA INVENTORY OF SOUTHWESTERN EL SALVADOR José Ledis Linares Conservation Science Department, SalvaNATURA. San Salvador, El Salvador linaresj_98@yahoo.com 1. INTRODUCTION The southwestern region of El Salvador, made up of the southern region of the Department of Ahuachapán, the southern edge of the Department of Santa Ana and the entire Department of Sonsonate, is an intricate network of many ecosystems and vegetation types immerse in a matrix of annual and permanent crops. It is almost impossible to know the distribution and extension of the types of original vegetation. However, there are still vestiges of what could have been the original vegetation. These small vegetation remnants and spots reveal that the region hosted and still hosts a substantial floral biodiversity. The flora inventory of El Salvador is far from being acceptable and even less complete. In spite of this, the aforementioned region is probably the best studied in the country, and even so, we continue to find new relatively large and noticeable tree species such as Dalbergia salvanaturae (Linares & Sousa 2008) endemic to El Imposible National Park. No accurate flora inventory of the area has yet been published and until the present study, there were only partial inventories of certain groups which are included in more general works such as the Flora appendix called ―Trees of the El Imposible Forest‖ by Reyna in Serrano et al. (1993), the list of trees prepared by Linares (2005) or the orchid inventory prepared by Hamer (1974, 1981). Most of the works conducted in the zone are undergraduate or graduate theses little known but valuable because of the information they provide such as the studies carried out at Cerro El Águila by Amaya et al. (2003) and Cerén et al. (2003). In other areas such as Las Lajas forest, we have the work of Komar & Herrera (1995) and the work of Villacorta (2000), and at El Imposible, the work of Ramírez (2001) also stands out in addition to the aforementioned work of Serrano (1993). To date, 1000 tree species are on record in El Salvador (Linares 2005) of which samples of 378 species were collected in the southwest region of the country with replicates deposited in local or foreign herbariums, until 2006. Of these collected tree samples, 185 were generalist species which are used as indicators of the level of completeness of an inventory. Prior to field work of the present study, the tree inventory was deemed 55% 32 Biodiversity field inventories in selected watersheds complete if we consider only those records supported by herbarium samples. The baseline study (Linares 2007) recognized that the inventory was incomplete since only 76% of the expected tree species in the area were recorded. This 76% also included studies or consultancies which did not have voucher specimens but were deemed reliable by the author. In total, the baseline study recognized 1716 flora species of which only 430 were trees and of these 255 were indicators of a complete inventory. Past studies on vegetation have been mainly focused on El Imposible National Park, which in view of its wealth of species, types of vegetation, easy access, and good state of conservation, has captured much attention since the 1980s. The primary objectives of this study were to determine the species, mainly trees, of interest for conservation in six watersheds of the southwestern region of El Salvador, as well as to try to fill information gaps regarding the presence and status of the populations of some tree species considered threatened, or otherwise important for the conservation of the biological diversity that still remains in the country. 2. METHODOLOGY 2.1. STUDY AREA The study was conducted in 6 of the 11 watersheds located in the southwestern region of El Salvador, which correspond to 7 of the 25 municipalities distributed among the Departments of Ahuachapán, Sonsonate, Santa Ana and La Libertad that make up the area of USAID’s Improved Management and Conservation of Critical Watersheds (―IMCW‖) Project. Table 9 and Fig. 4 detail the locations where the flora studies were carried out. 2.2. SAMPLING DESIGN Between June 2007 and January 2008, intensive searches were conducted based on 1000 m2 transects at each sampling site. In sites which comprised several types of forests, a transect was carried out in each forest type. Additionally, intensive searches were undertaken to locate species, either trees or other plants, that were not found in those transects. Samples that would allow proper identification were collected as needed. The samples or vouchers were deposited with the Herbarium of the Natural History Museum of El Salvador. Table 9 shows a detail of the sampling efforts and the number of species recorded in each natural area. Biodiversity field inventories in selected watersheds 33 TABLE 9. SAMPLING SITES FOR TREES AND TOTAL BOTANICAL SAMPLING EFFORT. Site Days Number of Transects (1000 m2 each) June 29 to July 3, July 10 to 14, from November 11 to 15, from December 26 to 30, 2007 and from January 8 to 12, 2008 25 2 Lava flows of varying successional stages and rermnants of tropical dry forest From July 27 to August 5, 2007 10 2 montane humid forest December 19 to 21, 27 to 28, 2007 5 1 Location Description Barra de Santiago Cerro Campana sector, Municipality of San Francisco Menéndez, high part of El Imposible (13.86° N; 89.91°W), Dept. of Ahuachapán Undisturbed montane humid forest. Cara Sucia San Fco. Menéndez sector, Municipality of San Francisco Menéndez, between cantones El Corozo and El Sacramento (13.84°N; 90.01°W), Dept. of Ahuachapán Evergreen broadleaf forest with anthropogenic disturbance due to the presence of a community within the area. El Imposible National Park Izalco Los Volcanes National Park Izalco 34 Dates Watershed María Auxiliadora sector, Municipality of Izalco, between the cantones of Chorro Abajo, Chorro Arriba, Shonshón, Teshcal, Las Marías, La Quebrada Española and Cuyagualo (13.89°N; 89.63°W), Dept. of Sonsonate Cerro Verde sector, Municipality of Santa Ana, Cantón Lomas de San Marcelino (13.82ºN; 89.62ºW ), Dept. of Santa Ana Biodiversity field inventories in selected watersheds TABLE 9. SAMPLING SITES FOR TREES AND TOTAL BOTANICAL SAMPLING EFFORT. Site Plan de Amayo Protected Natural Area San Marcelino Complex Protected Natural Area Cerro El Águila Protected Natural Area Watershed Location Description Dates Days Number of Transects (1000 m2 each) San Julián Municipality of Caluco, between the cantones of Plan de Amayo and El Castaño (13.41°N; 89.39°O), Dept. de Sonsonate Cliff vegetation, tropical dry forest (in the areas closest to the cliffs) and gallery forests (in the lower parts). August 22 to 31, 2007 and January 22 to 26, 2008 15 2 Vegetation succession on volcanic lava of different ages, deciduous forest, and evergreen broadleaf forest. December 12 to 16, 2007 5 1 Wooded summit consists of humid montane forest November 18 to 22, 2007 5 1 Municipalities of Izalco y Armenia, the cantones of Las Lajas, El Guayabo, Los Mangos, El Rosario, San Isidro and Las Marías, Dept. of Coatepeque Sonsonate. Municipalities of and Izalco Santa Ana and El Congo, between the cantones of Lomas de San Marcelino (13.49°N; 89.34°W), Montebello, El Rodeo and La Presa, Dept. of Santa Ana (13.69°N; 89.64°W) Río Grande Municipality of Juayúa (13.88°N; de 89.68°W), Dept. of Sonsonate Sonsonate Biodiversity field inventories in selected watersheds 35 2.3. LEVEL OF INVENTORY COMPLETENESS In order to make an analysis of the inventory progress or completeness in the municipalities and watersheds, I first prepared a list of indicator species for inventory completeness (Gómez de Silva & Medellín 2001). This list consists of generalist species, which are those that live in a variety of habitats and can be found in both disturbed and non-disturbed habitats. The relationship between the species that were recorded during the study and the list of expected generalist species represents an index of inventory completeness expressed as percent progress. The number of expected species recorded is divided by the total number of expected generalist species, which is 335 (listed in Linares 2007), and multiplied by one hundred. A single list of expected species was determined for watersheds and municipalities; however, separate lists of indicator species for inventory completeness were determined for each ecosystem or vegetation type (Table 10). The reason why each habitat has different indicator species for inventory completeness is that some generalist species of the low and medium altitude zones do not live in habitats of the higher altitude zones. 2.4. SELECTION OF SPECIES CONSIDERED IMPORTANT FOR CONSERVATION The main criterion to select species important for conservation is whether the species is included on the Red List of globally endangered species (IUCN 2008). We considered the tree Machaerium sp. to be an additional indicator of conservation importance, because we believe it meets the IUCN criteria for threatened status but the species has not yet been evaluated formally. All but one of the indicators are trees; the exception is a globally-vulnerable cycad Zamia herrerae, which is also included as an indicator in this analysis. 2.5. DATA SOURCES Most of the information on the species recorded in the project area comes from the databases of the Panamerican School of Agriculture (EAP, for its acronym in Spanish) and the Missouri Botanical Garden (MO) herbaria and from fieldwork of the present study. It was deemed that the majority of records in local herbaria are already included in the consulted databases because duplicates of the collected items were frequently sent to one of these foreign herbaria. Some of these data, specifically those from El Imposible and Cerro Verde, have been complemented with information that previously existed on these Protected Natural Areas (summarized in Linares 2007). The bibliographic information mainly comes from thesis works and consultancies carried out in the different Protected Natural Areas and cited in the Introduction. 36 Biodiversity field inventories in selected watersheds 7 1 2 4 3 6 5 FIGURE 4. STUDY AREA, WATERSHEDS, AND SAMPLING SITES FOR FLORA: (1) CERRO CAMPANA (EL IMPOSIBLE), (2) SAN FRANCISCO MENÉNDEZ SECTOR (EL IMPOSIBLE), (3) MARÍA AUXILIADORA SECTOR (LOS VOLCANES) (4) CERRO VERDE (LOS VOLCANES NATIONAL PARK), (5) PLAN DE AMAYO, (6) COMPLEJO SAN MARCELINO, (7) CERRO EL ÁGUILA. Biodiversity field inventories in selected watersheds 37 TABLE 10. INDICATOR SPECIES FOR INVENTORY COMPLETENESS OF TREES IN THE STUDY AREA. Species CF DF HF S Acacia angustissima (Mill.) Kuntze X Acacia cornigera (L.) Willd. X Acacia farnesiana (L.) Willd. X Acacia hindsii Benth. X Acacia polyphylla DC. X Aegiphila panamensis Moldenke X Albizia adinocephala (Donn. Sm.) Britton & Rose X X Albizia carbonaria Britton Albizia niopoides (Spruce ex Benth.) Burkart X X Alibertia edulis (Rich.) A. Rich. ex DC. X Allophylus occidentalis (Sw.) Radlk. Alstonia pittieri (Donn. Sm.) A.H. Gentry X Alvaradoa amorphoides Liebm. X Andira inermis (W. Wright) Kunth ex DC. X X Annona diversifolia Saff. X Annona glabra L. X Annona macroprophyllata Donn. Sm. X Annona reticulata L. Apeiba tibourbou Aubl. X X Aralia humilis Cav. X Ardisia compressa Kunth X X Ardisia paschalis Donn. Sm. X X Astronium graveolens Jacq. X X Banisteriopsis acapulcensis (Rose) Small Bauhinia cookii Rose X Bauhinia divaricata L. X Bauhinia pauletia Pers. X X Bauhinia ungulata L. X Bixa orellana L. X X Bocconia arborea S. Watson X Boehmeria caudata Sw. Boehmeria ulmifolia Wedd. X Bourreria huanita (Lex.) Hemsl. X Brachistus stramoniifolius (Kunth) Miers X Brosimum alicastrum Sw. X X Bunchosia nitida (Jacq.) DC. X Bursera simaruba (L.) Sarg. X X Byrsonima crassifolia (L.) Kunth X Caesalpinia eriostachys Benth. X Caesalpinia exostemma DC. X Calatola laevigata Standl. X X Calophyllum brasiliense var. rekoi (Standl.) Standl. X Calycophyllum candidissimun (Vahl) DC. X Capparis discolor Donn. Sm. X 38 Biodiversity field inventories in selected watersheds Species Capparis flexuosa (L.) L. Capparis frondosa Jacq. Capparis indica (L.) Druce Capparis mollicella Standl. Capparis odoratissima Jacq. Capparis pringlei Briq. Capparis quiriguensis Standl. Carica cauliflora Jacq. Carica papaya L. Cascabela ovata (Cav.) Lippold Cascabela thevetia (L.) Lippold Casearia aculeata Jacq. Casearia commersoniana Cambess. Casearia corymbosa Kunth Casearia sylvestris Sw. Casimiroa sapota Oerst. Cassia grandis L. f. Cassine xylocarpa Vent Castilla elastica Sessé ex Cerv. Cecropia obtusifolia Bertol. Cecropia peltata L. Cedrela odorata L. Cedrela salvadorensis Standl. Ceiba aesculifolia (Kunth) Britten & Baker f. Ceiba pentandra (L.) Gaertn. Celtis iguanaea (Jacq.) Sarg. Cestrum nocturnum L. Cestrum tomentosum L. f. Chiococca alba (L.) Hitchc. Chiococca pachyphylla Wernham Chrysobalanus icaco L. Chrysophyllum mexicanum Brandegee ex Standl. Citharexylum donnell-smithii Greenm. Clethra mexicana DC. Clibadium arboreum Donn. Sm. Clusia guatemalensis Hemsl. Coccoloba barbadensis Jacq. Coccoloba caracasana Meisn. Cochlospermum vitifolium (Willd.) Spreng. Colubrina arborescens (Mill.) Sarg. Colubrina heteroneura (Griseb.) Standl. Conocarpus erectus L. Conostegia xalapensis (Bonpl.) D. Don ex DC. Cordia alliodora (Ruiz & Pav.) Oken Cordia dentata Poir. Cordia panamensis L. Riley Cornutia pyramidata L. CF DF X HF S X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Biodiversity field inventories in selected watersheds X X X X X 39 Species Coutarea hexandra (Jacq.) K. Schum. Crateva tapia L. Critonia daleoides DC. Critonia hebebotrya DC. Critonia morifolia (Mill.) R.M. King & H. Rob. Cupania guatemalensis (Turcz.) Radlk. Curatella americana L. Dalbergia chontalensis Standl. & L.O. Williams Dendropanax arboreus (L.) Decne. & Planch. Dichapetalum donnell-smithii Engl. var donnell-smithii Diospyros salicifolia Humb. & Bonpl. ex Willd. Diphysa americana (Mill.) M. Sousa Drypetes lateriflora (Sw.) Krug & Urb. Dussia cuscatlanica (Standl.) Standl. & Steyerm. Ehretia latifolia DC. Enterolobium cyclocarpum (Jacq.) Griseb. Eremopsis triflosculosa (Kunth) Gleason subsp. triflosculosa Erythrina berteroana Urb. Eugenia jutiapensis Standl. & Steyerm. Eugenia lindeniana O. Berg Eugenia salamensis Donn. Sm. Eugenia sasoana Standl. & Steyerm. Exostema caribaeum (Jacq.) Roem. & Schult. Exostema mexicanum A. Gray Exothea paniculata (Juss.) Radlk. Ficus aurea Nutt. Ficus citrifolia Mill. Ficus costaricana (Liebm.) Miq. Ficus cotinifolia Kunth var. cotinifolia Ficus insipida Willd. Ficus maxima Mill. Ficus obtusifolia Kunth Ficus ovalis (Liebm.) Miq. Ficus pertusa L. f. Ficus trigonata L. Forchhammeria trifoliata Radlk. Garcia nutans Vahl Gaultheria erecta Vent. Genipa americana L. Gliricidia sepium (Jacq.) Kunth ex Walp. Godmania aesculifolia (Kunth) Standl. Guarea glabra Vahl Guazuma ulmifolia Lam. Guettarda deamii Standl. Guettarda macrosperma Donn. Sm. Guettarda subcapitata C.M. Taylor 40 CF DF X X X X X X X HF X S X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Biodiversity field inventories in selected watersheds X X X X X X X X Species Gyrocarpus americanus Jacq. Hamelia patens Jacq. Hauya elegans subsp. lucida (Donn. Sm. & Rose) P.H. Raven & Breedlove [= Hauya lucida Donn. Sm. & Rose] Heliocarpus mexicanus (Turcz.) Sprague Hemiangium excelsum (Kunth) A.C. Sm. Heteropterys laurifolia (L.) A. Juss. Hibiscus pernambucensis Arruda Hintonia latiflora (Sessé & Moc. ex DC.) Bullock Hintonia lumaeana (Baill.) Bullock Hymenaea courbaril L. Hyperbaena mexicana Miers Hyperbaena tonduzii Diels Inga flexuosa Schltdl. Inga pavoniana G. Don Inga punctata Willd. Inga vera Willd. Ipomoea wolcottiana Rose subsp. wolcottiana Jacaratia mexicana A. DC. Jacquinia longifolia Standl. Jacquinia nervosa C. Presl. Karwinskia calderonii Standl. Laguncularia racemosa (L.) C.F. Gaertn. Lasianthaea fruticosa (L.) K.M. Becker Ledenbergia macrantha Standl. Licania arborea Seem. Licania platypus (Hemsl.) Fritsch Licania retifolia S.F. Blake Lippia umbellata Cav. Lippia cardiostegia Benth. Lippia myriocephala Schltdl. & Cham. Lonchocarpus guatemalensis Benth. Lonchocarpus minimiflorus Donn. Sm. Lonchocarpus rugosus subsp. apricus (Lundell) M.Sousa Lonchocarpus salvadorensis Pittier Lonchocarpus santarosanus Donn. Sm. Luehea candida (Moc. & Sessé ex DC.) Mart. Lycianthes arrazolensis (J.M. Coult. & Donn. Sm.) Bitter Lycianthes heteroclita (Sendtn.) Bitter Lysiloma acapulcense (Kunth) Benth. Lysiloma auritum (Schltdl.) Benth. Lysiloma divaricatum (Jacq.) J.F. Macbr. Machaerium biovulatum Micheli Machaerium salvadorense (Donn. Sm.) Rudd Maclura tinctoria (L.) D. Don ex Steud. Malpighia glabra L. Malvaviscus arboreus Cav. var. arboreus CF DF X X HF S X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Biodiversity field inventories in selected watersheds X X X X X X X X X X X 41 Species Margaritaria nobilis L. f. Miconia argentea (Sw.) DC. Miconia laevigata (L.) D. Don Montanoa grandiflora Alaman es DC. Montanoa guatemalensis B.L. Rob. & Greenm. Montanoa hibiscifolia Benth. Montanoa tomentosa subsp.. xanthiifolia (Sch. Bip. ex C. Koch) V.A. Funk Muntingia calabura L. Myriocarpa longipes Liebm. Myrospermum frutescens Jacq. Myroxylon balsamum var. pereirae (Royle) Harms Nectandra martinicensis Mez Neea psychotrioides Donn. Sm. Ochroma pyramidale (Cav. ex Lam.) Urb. Ocotea veraguensis (Meisn.) Mez Omphalea oleifera Hemsl. Pachira aquatica Aubl. Pachythamnus crassirameus (B.L. Rob.) R.M. King & H. Rob. Parathesis vulgata Lundell Peltostigma pteleoides (Hook.) Walp. Perymenium grande Hemsl. var. grande Perymenium grande var. nelsonii (B.L. Rob. & Greenm.) J.J. Fay Phenax hirtus (Sw.) Wedd. Phenax mexicanus Wedd. Pilocarpus racemosus subsp. viridulus Kaastra Piper amalago L. Piper marginatum Jacq. Piper sanctum (Miq.) Schltdl. ex C. DC. Piper tuberculatum Jacq. Piptadenia obliqua (Pers.) J.F. Macbr. Pithecellobium dulce (Roxb.) Benth. Plumeria rubra forma acutifolia (Poir.) Woodson Poeppigia procera C. Presl Pogonopus speciosus (Jacq.) K. Schum. Posoqueria latifolia (Rudge) Roem. & Schult. Prockia crucis P. Browne ex L. Pseudobombax ellipticum (Kunth) Dugand [=Bombax ellipticum H.B.K.] Pseudobombax septenatum (Jacq.) Dugand Psidium guajava L. Psidium guianense Pers. Psychotria limonensis K. Krause Psychotria microdon (DC.) Urb. Psychotria pubescens Sw. 42 CF DF HF S X X X X X X X X X X X X X X X X X X X X X X Biodiversity field inventories in selected watersheds X X X X X X X X X X X X X X X X X X X X X X Species Quercus elliptica Née Quercus lancifolia Schltdl. & Cham. Quercus salicifolia Née Quercus sapotifolia Liebm. Quercus skinneri Benth. Randia armata (Sw.) DC. Randia grandifolia (Donn. Sm.) Standl. Robinsonella densiflora Fryxell Roldana petasioides (Greenm.) H. Rob. Rondeletia deamii (Donn. Sm.) Standl. Rondeletia laniflora Benth. Rondeletia thiemei Donn. Sm. Roupala glaberrima Pittier Ruprechtia costata Meisn. Samanea saman (Jacq.) Merr. Sapindus saponaria L. Sapium macrocarpum Müll. Arg. Sapranthus violaceus (Dunal) Saff. Schoepfia schreberi J.F. Gmel. Sciadodendron excelsum Griseb. Senna nicaraguensis (Benth.) H.S. Irwin & Barneby Senna pallida (Vahl) H.S. Irwin & Barneby Senna reticulata (Willd.) H.S. Irwin & Barneby Sideroxylon capiri subsp. tempisque (Pittier) T.D. Penn. Simarouba glauca DC. Sinclairia deamii (B.L. Rob. & Bartlett) Rydb. Sinclairia discolor Hook. & Arn. Sinclairia glabra (Hemsl) Rydb. Sinclairia sublobatum (B. L. Rob.) Rydb. Sloanea terniflora (Sessé & Moc. ex DC.) Standl. Solanum erianthum D. Don Solanum hazenii Britton Solanum lanceolatum Cav. Solanum umbellatum Mill. Spondias mombin L. Spondias purpurea L. Spondias radlkoferi Donn. Sm. Stemmadenia donnell-smithii (Rose) Woodson Stemmadenia eubracteata Woodson Stemmadenia obovata K. Schum. Stenocereus aragonii (F.A.C. Weber) Buxb. Styrax argenteus C. Presl Swartzia simplex (Sw.) Spreng. Swietenia humilis Zucc. Swietenia macrophylla King Synardisia venosa (Mast.) Lundell Tabebuia chrysantha (Jacq.) G. Nicholson CF X X X X X DF HF X X X X X S X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Biodiversity field inventories in selected watersheds X 43 Species CF Tabebuia donnell-smithii Rose Tabebuia impetiginosa (Mart. ex A. DC.) Standl. Tabebuia rosea (Bertol.) A. DC. Tabernaemontana amygdalifolia Jacq. Tecoma stans (L.) Juss. ex Kunth X Telanthophora arborescens (Steetz) H. Rob. & Brettell X Terminalia oblonga (Ruiz & Pav.) Steud. Thouinia velutina Radlk. Thouinia villosa DC. Thouinidium decandrum (Bonpl.) Radlk. Trema micrantha (L.) Blume var. micrantha Trema micrantha var strigillosa (Lundell) Standl. & X Steyerm. Trichilia americana (Sesse & Moc.) T.D. Penn. Trichilia havanensis Jacq. X Trichilia hirta L. Trichilia martiana C. DC. Trichospermum galeottii (Turcz.) Kosterm. Triplaris melaenodendron (Bertol.) Standl. & Steyerm. Trophis racemosa (L.) Urb. Turpinia occidentalis (Sw.) G. Don X Ulmus mexicana (Liebm.) Planch. X Urera baccifera (L.) Gaudich. ex Wedd. X Urera corallina (Liebm.) Wedd. X Urera eggersii Hieron. X Verbesina guatemalensis B.L. Rob. & Greenm. Verbesina turbacensis Kunth X Vernonia deppeana Less. Vernonia patens Kunth Viburnum hartwegii Benth. X Ximenia americana L. Xylosma chlorantha Donn. Sm. X Xylosma flexuosum (Kunth) Hemsl. X Zanthoxylum culantrillo Kunth Zanthoxylum microcarpum Griseb. Zapoteca formosa (Kunth) H.M. Hern. subsp. formosa Zinowiewia cuneifolia Lundell X * CF: Cloud Forest; DF: Dry Forest; HF: Humid forest; S: Savanna. 44 Biodiversity field inventories in selected watersheds DF X X X X X X X X X HF S X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 3. RESULTS AND DISCUSSION 3.1. SPECIES RECORDED During the study, we collected a total of 1164 herbarium samples coming from approximately 707 vascular plant species. With this new data, the floral inventory for the study area now presents 584 tree species (Table 11). This figure includes about 430 species that were already cited in previous reports or were collected previously in different locations of the project area. The remaining 154 were not recorded in the existing literature and/or were not included in the databases available at the beginning of the present study. During field work, botanical voucher specimens were collected for at least 415 tree species, giving priority to rare species or to those for which few specimens had been collected previously. The 584 tree species found in the project area represent approximately 58% of all trees known in the country. Many of the tree species found are completely restricted to the study area or their largest populations are known from the study area. At El Imposible National Park, the list of trees reached a total of 269 species, including the species reported in previous studies. A total of 190 species were recorded in Plan de Amayo, and 172 species in San Marcelino Complex Protected Natural Area. The list of trees from Cerro Verde reached a total of 94 species. A total of 84 species were recorded in the María Auxiliadora sector at Los Volcanes National Park, and 45 species on Cerro El Águila. 3.2. IMPORTANT SPECIES FOR CONSERVATION The study area presents 25 indicators of conservation importance (Table 12). The natural areas with the most threatened species are Plan de Amayo, El Imposible National Park, and Los Volcanes National Park with 12 species each, followed by San Marcelino with eight, and finally, El Águila with only two species. However, Plan de Amayo presents five species that are not shared with any of the other conservation areas, which also represent very rare species in El Salvador, such as Lonchocarpus molinae (Leguminosae). This is a species recorded in the country for the first time and also endemic to lowland karst (limestone) soil, which makes it very unlikely to find in another area of El Salvador. El Imposible National Park hosts 12 species of the IUCN red list and only has one of the 25 indicators, the Mahogany tree (Swietenia macrophylla), not shared with any other area of southwestern El Salvador. Biodiversity field inventories in selected watersheds 45 TABLE 11. TREE SPECIES RECORDED IN THE STUDY AREA. Family Species Indicator (I) Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Euphorbiaceae Euphorbiaceae Euphorbiaceae Cactaceae Achatocarpaceae Leguminosae Arecaceae Euphorbiaceae Verbenaceae Opiliaceae Opiliaceae Leguminosae Leguminosae Leguminosae Euphorbiaceae Rubiaceae Sapindaceae Betulaceae Apocynaceae Simaroubaceae Ulmaceae Rutaceae Rutaceae Anacardiaceae Leguminosae Annonaceae Annonaceae Annonaceae Annonaceae Annonaceae Annonaceae Annonaceae Annonaceae Tiliaceae Ulmaceae Acanthaceae Apocynaceae Acacia angustissima (Mill.) Kuntze Acacia cornigera (L.) Willd. Acacia farnesiana (L.) Willd. var. farnesiana Acacia hindsii Benth. Acacia polyphylla DC. Acacia riparia Kunth Acacia sp. Acalypha sp. Acalypha diversifolia Jacq. Acalypha villosa Jacq. Acanthocereus tetragonus (L.) Hummelinck Achatocarpus nigricans Triana Acosmium panamense (Benth.) Yakovlev Acrocomia aculeata (Jacq.) Lodd. ex Mart. Adelia barbinervis Schltdl. & Cham. Aegiphila panamensis Moldenke Agonandra loranthoides L.O. Williams Agonandra racemosa (DC.) Standl. Albizia adinocephala (Donn. Sm.) Britton & Rose Albizia carbonaria Britton Albizia niopoides (Spruce ex Benth.) Burkart Alchornea latifolia Sw. Alibertia edulis (Rich.) A. Rich. ex DC. Allophylus occidentalis (Sw.) Radlk. Alnus acuminata subsp. arguta (Schltdl.) Furlow Alstonia longifolia (A. DC.) Pichon Alvaradoa amorphoides Liebm. Ampelocera hottlei (Standl.) Standl. Amyris balsamifera L. Amyris guatemalensis Lundell Anacardium occidentale L. Andira inermis (W. Wright) DC. Annona cherimola Mill. Annona diversifolia Saff. Annona glabra L. Annona macroprophyllata Donn. Sm. Annona reticulata L. Annona sp. Annona squamosa L. Annonaceae indet. Apeiba tibourbou Aubl. Aphananthe monoica (Hemsl.) J.-F. Leroy Aphelandra schiedeana Schltdl. & Cham. Apocynaceae indet. I I 46 Biodiversity field inventories in selected watersheds I I I I I I I I I I I I I I I I Family Species Indicator (I) Araliaceae Myrsinaceae Myrsinaceae Myrsinaceae Bignoniaceae Apocynaceae Asteraceae Anacardiaceae Verbenaceae Verbenaceae Asteraceae Rubiaceae Malpighiaceae Leguminosae Leguminosae Leguminosae Aralia humilis Cav. Ardisia sp. Ardisia compressa Kunth Ardisia paschalis Donn. Sm. Arrabidaea costaricensis (Kraenzl.) A.H. Gentry Aspidosperma megalocarpon Müll. Arg. Asteraceae indet. Astronium graveolens Jacq. Avicennia bicolor Standl. Avicennia germinans (L.) Stearn Baccharis sp. Balmea stormae Martínez Banisteriopsis acapulcensis (Rose) Small Bauhinia cookii Rose Bauhinia divaricata L. Bauhinia pauletia Pers. Bauhinia seleriana Harms (= Bauhinia paradisii Standl. & L.O. Williams) Bauhinia ungulata L. Begonia calderonii Standl. Cydista sp. Berberis johnstonii Standl. & Steyerm. Bernardia sp. Bixa orellana L. Bixa urucurana Willd. Bocconia arborea S. Watson Boehmeria caudata Sw. Boehmeria ulmifolia Wedd. Bourreria huanita (Lex.) Hemsl. Brachistus stramoniifolius (Kunth) Miers Bravaisia integerrima (Spreng.) Standl. Brosimum alicastrum Sw. Buddleja americana L. Bunchosia nitida (Jacq.) DC. Bursera graveolens (Kunth) Triana & Planch. Bursera longicuspis Lundell Bursera simaruba (L.) Sarg. Byrsonima crassifolia (L.) Kunth Caesalpinia eriostachys Benth. Caesalpinia exostemma DC. Calatola laevigata Standl. Calliandra sp. nov. Callitropsis lusitanica (Mill.) D.P. Little Calophyllum brasiliense var. rekoi (Standl.) Standl. Calycophyllum candidissimun (Vahl) DC. Capparis discolor Donn. Sm. I Leguminosae Leguminosae Begoniaceae Bignoniaceae Berberidaceae Euphorbiaceae Bixaceae Bixaceae Papaveraceae Urticaceae Urticaceae Boraginaceae Solanaceae Acanthaceae Moraceae Loganiaceae Malpighiaceae Burseraceae Burseraceae Burseraceae Malpighiaceae Leguminosae Leguminosae Icacinaceae Leguminosae Cupressaceae Clusiaceae Rubiaceae Capparaceae Biodiversity field inventories in selected watersheds I I I I I I I I I I I I I I I I I I I I I I I I 47 Family Species Indicator (I) Capparaceae Capparaceae Capparaceae Capparaceae Capparaceae Capparaceae Capparaceae Capparaceae Solanaceae Caricaceae Caricaceae Apocynaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Rutaceae Leguminosae Celastraceae Moraceae Moraceae Cecropiaceae Meliaceae Meliaceae Meliaceae Meliaceae Bombacaceae Bombacaceae Ulmaceae Ulmaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Arecaceae Rubiaceae Rubiaceae Chrysobalanaceae Sapotaceae Verbenaceae Verbenaceae Capparis flexuosa (L.) L. Capparis frondosa Jacq. Capparis indica (L.) Druce Capparis mollicella Standl. Capparis odoratissima Jacq. Capparis pringlei Briq. Capparis quiriguensis Standl. Capparis sp. Capsicum ciliatum (Kunth) Kuntze Carica cauliflora Jacq. Carica papaya L. Cascabela ovata (Cav.) Lippold Casearia aculeata Jacq. Casearia arguta Kunth Casearia coronata Standl. & L.O. Williams Casearia tremula (Griseb.) Griseb. ex C. Wright Casearia commersoniana Cambess Casearia sp. Casearia corymbosa Kunth Casearia sylvestris Sw. var. sylvestris Casimiroa sapota Oerst. Cassia grandis L. f. Cassine xylocarpa Vent Castilla elastica Sessé ex Cerv. Cecropia obtusifolia Bertol. Cecropia peltata L. Cedrela fissilis Vell. Cedrela odorata L. Cedrela salvadorensis Standl. Cedrela tonduzii C. DC. Ceiba aesculifolia (Kunth) Britten & Baker f. Ceiba pentandra (L.) Gaertn. Celtis caudata Planch. Celtis iguanaea (Jacq.) Sarg. Cestrum dumetorum Schltdl. Cestrum nocturnum L. Cestrum pacayense Francey Cestrum racemosum Ruiz & Pav. Cestrum tomentosum L. f. Chamaedorea tepejilote Liebm. ex Mart. Chiococca alba (L.) Hitchc. Chiococca pachyphylla Wernham Chrysobalanus icaco L. Chrysophyllum mexicanum Brandegee ex Standl. Citharexylum costaricense Moldenke Citharexylum donnell-smithii Greenm. I I I I I I I 48 Biodiversity field inventories in selected watersheds I I I I I I I I I I I I I I I I I I I I I I I I I I Family Species Verbenaceae Clerodendrum pittieri Moldenke Clethraceae Clethra mexicana DC. Clethraceae Clethra sp. Clethraceae Clethra suaveolens Turcz. Asteraceae Clibadium arboreum Donn. Sm. Clusiaceae Clusia guatemalensis Hemsl. Cnidoscolus aconitifolius (Mill.) I.M. Johnst. subsp. Euphorbiaceae aconitifoliussp. Euphorbiaceae Cnidoscolus Polygonaceae Coccoloba acapulcensis Standl. Polygonaceae Coccoloba barbadensis Jacq. Polygonaceae Coccoloba caracasana Meisn. Polygonaceae Coccoloba montana Standl. Polygonaceae Coccoloba sp. nov. Polygonaceae Coccoloba venosa L. Cochlospermaceae Cochlospermum vitifolium (Willd.) Spreng. Leguminosae Cojoba arborea (L.) Britton & Rose Leguminosae Cojoba graciliflora (S.F. Blake) Britton & Rose Rhamnaceae Colubrina arborescens (Mill.) Sarg. Rhamnaceae Colubrina heteroneura (Griseb.) Standl. Combretaceae Combretum fruticosum (Loefl.) Stuntz Combretaceae Conocarpus erectus L. Melastomataceae Conostegia xalapensis (Bonpl.) D. Don ex DC. Boraginaceae Cordia alliodora (Ruiz et Pav.) Oken Boraginaceae Cordia collococca L. Boraginaceae Cordia dentata Poir. Boraginaceae Cordia panamensis L. Riley Boraginaceae Cordia salvadorensis Standl. Verbenaceae Cornutia pyramidata L. Chrysobalanaceae Couepia polyandra (Kunth) Rose Rubiaceae Coutarea hexandra (Jacq.) K. Schum. Capparaceae Crateva tapia L. Bignoniaceae Crescentia cujete L. Asteraceae Critonia sp. Asteraceae Critonia daleoides DC. Asteraceae Critonia hebebotrya DC. Asteraceae Critonia morifolia (Mill.) King et Rob Celastraceae Crossopetalum sp. nov. Euphorbiaceae Croton guatemalensis Lotsy Euphorbiaceae Croton reflexifolius Kunth Euphorbiaceae Croton sp. nov. Sapindaceae Cupania guatemalensis (Turcz.) Radlk. Sapindaceae Cupania mollis Standl. Dilleniaceae Curatella americana L. Annonaceae Cymbopetalum penduliflorum (Dunal) Baill. Leguminosae Dalbergia calycina Benth. Leguminosae Dalbergia chontalensis Standl. & L.O. Williams Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I I 49 Family Species Leguminosae Leguminosae Leguminosae Thymelaeaceae Leguminosae Araliaceae Dichapetalaceae Ebenaceae Ebenaceae Ebenaceae Leguminosae Leguminosae Euphorbiaceae Leguminosae Leguminosae Boraginaceae Leguminosae Dalbergia congestiflora Pittier Dalbergia melanocardium Pittier Dalbergia salvanaturae J. Linares et M. Sousa Daphnopsis witsbergeri Nevling, Matek. & Barringer Delonix regia (Bojer ex Hook.) Raf. Dendropanax arboreus (L.) DCne. et Planch. Dichapetalum donnell-smithii Engl. Diospyros morenoi A. Pool Diospyros nicaraguensis (Standl.) Standl. Diospyros salicifolia Humb. & Bonpl. ex Willd. Diphysa americana (Miller) M. Sousa Diphysa humilis Oerst. Drypetes lateriflora (Sw.) Krug & Urb. Dussia cuscatlanica (Standl.) Standl. & Steyerm. Dyphisa americana (Mill.) M. Sousa Ehretia latifolia DC. Enterolobium cyclocarpum (Jacq.) Griseb. Eremopsis triflosculosa (Kunth) Gleason subsp. triflosculosa Erythrina berteroana Urb. Erythroxylum areolatum L. Erythroxylum havanense Jacq. Erythroxylum rotundifolium Lunan Eugenia alfaroana Standl. Eugenia sp.1 Eugenia sp. 2 Eugenia jutiapensis Standl. & Steyerm. Eugenia lindeniana O. Berg Eugenia oerstediana O. Berg Eugenia quercetorum Standl. & L.O. Williams ex Barrie Eugenia salamensis Donn. Sm. var. salamensis Eugenia salamensis var. rensoniana (Standl.) McVaugh Eugenia sasoana Standl. & Steyerm. Euphorbia leucocephala Lotsy Euphorbia schlechtendalii Boiss. Euphorbia sp. Euphorbiaceae indet. (sp. nov.?) Exostema caribaeum (Jacq.) Roem. & Schult. Exostema mexicanum A. Gray Exothea paniculata (Juss.) Radlk. Eysenhardtia adenostylis Baill. Faramea occidentalis (L.) A. Rich. Ficus aurea Nutt. Ficus citrifolia Mill. Asteraceae Leguminosae Erythroxylaceae Erythroxylaceae Erythroxylaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Myrtaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Rubiaceae Rubiaceae Sapindaceae Leguminosae Rubiaceae Moraceae Moraceae 50 Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I Family Species Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Apocynaceae Onagraceae Euphorbiaceae Clusiaceae Ericaceae Rubiaceae Leguminosae Bignoniaceae Nyctaginaceae Nyctaginaceae Meliaceae Sterculiaceae Rubiaceae Rubiaceae Euphorbiaceae Hernandiaceae Hernandiaceae Rubiaceae Malvaceae Ficus costaricana (Liebm.) Miq. Ficus cotinifolia Kunth var. cotinifolia Ficus goldmanii Standl. Ficus insipida Willd. Ficus jimenezii Standl. Ficus maxima Mill. Ficus obtusifolia Kunth Ficus ovalis (Liebm.) Miq. Ficus trigonata L. Forchhammeria trifoliata Radlk. Fuchsia paniculata Lindl. Garcia nutans Vahl Garcinia intermedia (Pittier) Hammel Gaultheria erecta Vent. Genipa americana L. Gliricidia sepium (Jacq.) Kunth ex Walp. Godmania aesculifolia (Kunth) Standl. Grajalesia fasciculata (Standl.) Miranda Guapira witsbergeri Lundell Guarea glabra Vahl Guazuma ulmifolia Lam. Guettarda macrosperma Donn. Sm. Guettarda subcapitata C.M. Taylor Gymnanthes riparia (Schltdl.) Klotzsch Gyrocarpus americanus Jacq. Gyrocarpus jatrophifolius Domin Hamelia patens Jacq. Hampea stipitata S. Watson Hauya elegans subsp. lucida (Donn. Sm. & Rose) Onagraceae P.H. Raven & Breedlove Tiliaceae Heliocarpus mexicanus (Turcz.) Sprague Hippocrateaceae Hemiangium excelsum (Kunth) A.C. Sm. Malpighiaceae Heteropterys laurifolia (L.) A. Juss. Malvaceae Hibiscus pernambucensis Arruda Rubiaceae Hillia tetrandra Sw. Rubiaceae Hintonia latiflora (Sessé & Moc. ex DC.) Bullock Rubiaceae Hintonia lumaeana (Baill.) Bullock Malpighiaceae Hiraea sp. Malpighiaceae Hiraea velutina Nied. Chrysobalanaceae Hirtella racemosa var. hexandra (Willd. ex Roem. & Schult.) Prance Euphorbiaceae Hura polyandra Baill. Leguminosae Hymenaea courbaril L. Menispermaceae Hyperbaena mexicana Miers Menispermaceae Hyperbaena tonduzii Diels Aquifoliaceae Ilex discolor Hemsl. var discolor Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I I I I I I I I I 51 Family Aquifoliaceae Rutaceae Flacourtiaceae Celastraceae Meliaceae Melastomataceae Sapindaceae Sapotaceae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Convolvulaceae Amaranthaceae Amaranthaceae Amaranthaceae Rubiaceae Caricaceae Theophrastaceae Juglandaceae Rhamnaceae Rhamnaceae Rhamnaceae Lythraceae Combretaceae Asteraceae Phytolaccaceae Leguminosae Leguminosae Leguminosae Chrysobalanaceae Chrysobalanaceae Chrysobalanaceae Lauraceaeindet Lauraceae Verbenaceae Verbenaceae Verbenaceae Leguminosae Leguminosae Leguminosae Leguminosae 52 Species Ilex discolor var. tolucana (Hemsl.) Edwin ex T.R. Dudley Indeterminada Indeterminada Indeterminada Indeterminada Indeterminada Indeterminada Indeterminada Inga calderonii Standl. Inga flexuosa Schltdl. Inga oerstediana Benth. ex Seem. Inga pavoniana G. Don Inga punctacta Willd. Inga sapindoides Willd. Inga vera Willd. Ipomoea wolcottiana Rose subsp. wolcottiana Iresine diffusa Humb. & Bonpl. ex Willd. Iresine herrerae Conz. & S.F. Blake Iresine nigra Uline & W.L. Bray Ixora floribunda (A. Rich.) Griseb. Jacaratia mexicana A. DC. Jacquinia longifolia Standl. Juglans olanchana Standl. & L.O. Williams Karwinskia calderonii Standl. Krugiodendron ferreum (Vahl) Urb. Krugiodendron sp. nov. Lafoensia punicifolia DC. Laguncularia racemosa (L.) C.F. Gaertn. Lasianthaea fruticosa (L.) K.M. Becker Ledenbergia macrantha Standl. Lennea viridiflora Seem. Leucaena diversifolia (Schltdl.) Benth. Leucaena shannonii Donn. Sm. Licania arborea Seem. Licania platypus (Hemsl.) Fritsch Licania retifolia S.F. Blake Licaria capitata (Schltdl. & Cham.) Kosterm. Licaria sp. Lippia cardiostegia Benth. Lippia myriocephala Schltdl. & Cham. Lippia umbellata Cav. Lonchocarpus sp. Lonchocarpus acuminatus (Schltdl.) M. Sousa Lonchocarpus guatemalensis Benth. Lonchocarpus lanceolatus Benth. Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I Family Species Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Lonchocarpus michelianus Pittier Lonchocarpus minimiflorus Donn. Sm. Lonchocarpus molinae Standl. & L.O. Williams Lonchocarpus parviflorus Benth. Lonchocarpus peninsularis (Donn. Sm.) Pittier Lonchocarpus phaseolifolius Benth. Lonchocarpus purpusii Brandegee Lonchocarpus retiferus Standl. & L.O. Williams Lonchocarpus rugosus subsp. apricus (Lundell) M. Sousa Lonchocarpus salvadorensis Pittier Lonchocarpus sanctuarii Standl. & L.O. Williams Lonchocarpus santarosanus Donn. Sm. Lonchocarpus schiedeanus (Schltdl.) Harms Lonchocarpus stenophyllus M. Sousa Luehea candida (Moc. & Sessé ex DC.) Mart. Luffa operculata (L.) Cogn. Lycianthes arrazolensis (J.M. Coult. & Donn. Sm.) Bitter Lycianthes heteroclita (Sendtn.) Bitter Lycianthes orogenes Standl. & Steyerm. Lysiloma acapulcense (Kunth) Benth. Lysiloma auritum (Schltdl.) Benth. Lysiloma divaricatum (Jacq.) J.F. Macbr. Machaerium biovulatum Micheli Machaerium kegelii Meisn. Machaerium salvadorense (Donn. Sm.) Rudd Machaerium sp. Maclura tinctoria (L.) D. Don ex Steud. Malpighia glabra L. Malpighia sp. Malvaviscus arboreus Cav. var. arboreus Manilkara chicle (Pittier) Gilly Manilkara zapota (L.) P. Royen Margaritaria nobilis L. f. Matayba glaberrima Radlk. Maytenus chiapensis Lundell Maytenus sp. nov. Miconia sp. Miconia argentea (Sw.) DC. Miconia laevigata (L.) D. Don Montanoa grandiflora Alaman es DC. Montanoa guatemalensis B.L. Rob. & Greenm. Montanoa hibiscifolia Benth. Montanoa tomentosa subsp. xanthiifolia (Sch. Bip. ex C. Koch) V.A. Funk Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Tiliaceae Cucurbitaceae Solanaceae Solanaceae Solanaceae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Moraceae Malpighiaceae Malpighiaceae Malvaceae Sapotaceae Sapotaceae Euphorbiaceae Sapindaceae Celastraceae Celastraceae Melastomataceae Melastomataceae Melatomataceae Asteraceae Asteraceae Asteraceae Asteraceae Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I I 53 Family Species Myricaceae Morella cerifera (L.) Small Melastomataceae Mouriri myrtilloides subsp. parvifolia (Benth.) Morley Elaeocarpaceae Muntingia calabura L. Myrtaceae Myrcianthes fragrans (Sw.) McVaugh var. fragrans Myrtaceae Myrcianthes sp. Urticaceae Myriocarpa longipes Liebm. Leguminosae Myrospermum frutescens Jacq. Leguminosae Myroxylon balsamum var. pereirae (Royle) Harms Lauraceae Nectandra martinicensis Mez Lauraceae Nectandra mirafloris Lauraceae Nectandra salicifolia (Kunth) Nees Nyctaginaceae Neea psychotrioides Donn. Sm. Asteraceae Neurolaena lobata (L.) Cass. Bombacaceae Ochroma pyramidale (Cav. ex Lam.) Urb. Lauraceae Ocotea botrantha Rohwer Lauraceae Ocotea salvadorensis (Lundell) van der Werff Lauraceae Ocotea sinuata (Mez) Rohwer Lauraceae Ocotea veraguensis (Meisn.) Mez Euphorbiaceae Omphalea oleifera Hemsl. Euphorbiaceae Ophellantha spinosa Standl. Araliaceae Oreopanax xalapensis (Kunth) Decne. & Planch. Ochnaceae Ouratea lucens (Kunth) Engl. Bombacaceae Pachira aquatica Aubl. Pachythamnus crassirameus (B.L. Rob.) R.M. King Asteraceae & H. Rob. Palicourea padifolia (Willd. ex Roem. & Schult.) Rubiaceae C.M. Taylor & Lorence Myrsinaceae Parathesis columnaris Lundell Myrsinaceae Rutaceae Lauraceae Asteraceae Asteraceae Urticaceae Urticaceae Lauraceae Lauraceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Ulmaceae Simaroubaceae Rutaceae Pinaceae Piperaceae 54 Parathesis vulgata Lundell Peltostigma pteleoides (Hook.) Walp. Persea americana Mill. Perymenium grande Hemsl. var. grande Perymenium grande var. nelsonii (B.L. Rob. & Greenm.) J.J. Fay Phenax hirtus (Sw.) Wedd. Phenax mexicanus Wedd. Phoebe acuminatissima Lundell Phoebe sp. Phyllanthus elsiae Urb. Phyllanthus mocinianus Baill. Phyllanthus sp. Phyllostylon rhamnoides (J. Poiss.) Taub. Picramnia antidesma subsp. fessonia (DC.) W.W. Thomas Pilocarpus racemosus subsp viridulus Kaastra Pinus oocarpa Schiede ex Schltdl. Piper amalago L. Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I Family Species Indicator (I) Piperaceae Piperaceae Piperaceae Piperaceae Piperaceae Leguminosae Leguminosae Leguminosae Nyctaginaceae Nyctaginaceae Nyctaginaceae Nyctaginaceae Leguminosae Leguminosae Leguminosae Asteraceae Apocynaceae Asteraceae Polygonaceae Leguminosae Rubiaceae Rubiaceae Sapotaceae Sapotaceae Sapotaceae Flacourtiaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Bombacaceae Bombacaceae Myrsinaceae Myrtaceae Myrtaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Fagaceae Fagaceae Fagaceae Fagaceae Fagaceae Piper marginatum Jacq. Piper sanctum (Miq.) Schltdl. ex C. DC. Piper sp. Piper tuberculatum Jacq. Piper yzabalanum C. DC. Piptadenia obliqua (Pers.) J.F. Macbr. Piscidia carthagenensis Jacq. Piscidia grandifolia (Donn. Sm.) I.M. Johnst. Pisonia aculeata Jacq. Pisonia donnellsmithii Heimerl ex Standl. Pisonia macranthocarpa (Donn. Sm.) Donn. Sm. Pisonia sp. nov. Pithecellobium dulce (Roxb.) Benth. Pithecellobium microstachyum Standl. Platymiscium parviflorum Benth. Pluchea odorata (L.) Cass. Plumeria rubra forma acutifolia (Poir.) Woodson Podachaenium eminens (Lag.) Sch. Bip. Podopterus mexicanus Bonpl. Poeppigia procera C. Presl Pogonopus speciosus (Jacq.) K. Schum. Posoqueria latifolia (Rudge) Roem. & Schult. Pouteria compechiana (Kunth) Baehni Pouteria sapota (Jacq.) H.E. Moore & Stearn Pouteria viridis (Pittier) Cronquist Prockia crucis P. Browne ex L. Prunus sp. Prunus axitlana Standl. Prunus brachybotrya Zucc. Prunus salasii Standl. Prunus skutchii I.M. Johnst. Pseudobombax ellipticum (Kunth) Dugand Pseudobombax septenatum (Jacq.) Dugand Psidium guajava L. Psidium guineense Sw. Psidium sp, Psychotria sp. Psychotria costivenia Griseb. Psychotria limonensis K. Krause Psychotria microdon (DC.) Urb. Psychotria pubescens Sw. Quercus elliptica Née Quercus lancifolia Schltdl. & Cham. Quercus salicifolia Née Quercus sapotifolia Liebm. Quercus segoviensis Liebm. I I Biodiversity field inventories in selected watersheds I I I I I I I I I I I I I I I I I I 55 Family Species Indicator (I) Fagaceae Fagaceae Fagaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Myrsinaceae Asteraceae Quercus skinneri Benth. Quercus sp. Quercus tristis Liebm. Randia sp. 1 Randia sp. 2 Randia sp. 3 Randia aculeata L. Randia armata (Sw.) DC. Randia grandifolia (Donn. Sm.) Standl. Randia monantha Benth. Randia obcordata S. Watson Randia pleiomeris Standl. Randia thurberi S. Watson Rapanea sp. Rensonia salvadorica S.F. Blake Rhamnus sphaerosperma var. mesoamericana M.C. Johnst. & L.A. Johnst. Rhizophora mangle L. Robinsonella densiflora Fryxell Roldana petasioides (Greenm.) H. Rob. Rollinia membranacea Triana & Planch. Rondeletia cordata Benth. Rondeletia deamii (Donn. Sm.) Standl. Rondeletia laniflora Benth. Rondeletia thiemei Donn. Sm. Roupala glaberrima Pittier Ruprechtia chiapensis Lundell Ruprechtia costata Meisn. Samanea saman (Jacq.) Merr. Sapindus saponaria L. Sapium macrocarpum Müell. Arg. Sapranthus microcarpus (Donn. Sm.) R.E. Fr. Sapranthus palanga R.E. Fr. Sapranthus violaceus (Dunal) Saff. Saurauia kegeliana Schltdl. Schizolobium parahyba (Vell.) S.F. Blake Schoepfia schreberi J.F. Gmel. Schoepfia vacciniiflora Planch. ex Hemsl. Sciadodendron excelsum Griseb. Semialarium mexicanum (Miers) Mennega Senna nicaraguensis (Benth.) H.S. Irwin & Barneby Senna pallida (Vahl) H.S. Irwin & Barneby Senna reticulata (Willd.) H.S. Irwin & Barneby Sideroxylon sp. Sideroxylon capiri subsp. tempisque (Pittier) T.D. Penn. I Rhamnaceae Rhizophoraceae Malvaceae Asteraceae Annonaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Proteaceae Polygonaceae Polygonaceae Leguminosae Sapindaceae Euphorbiaceae Annonaceae Annonaceae Annonaceae Actinidiaceae Leguminosae Olacaceae Olacaceae Araliaceae Hippocrateaceae Leguminosae Leguminosae Leguminosae Sapotaceae Sapotaceae 56 Biodiversity field inventories in selected watersheds I I I I I I I I I I I I I I I I I I I I Family Sapotaceae Sapotaceae Simaroubaceae Rubiaceae Rubiaceae Asteraceae Asteraceae Asteraceae Asteraceae Elaeocarpaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Anacardiaceae Anacardiaceae Anacardiaceae Apocynaceae Apocynaceae Apocynaceae Apocynaceae Apocynaceae Cactaceae Sterculiaceae Loganiaceae Leguminosae Styrax Styracaceae Leguminosae Meliaceae Meliaceae Theaceae Symplocaceae Symplocaceae Myrsinaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Apocynaceae Sapindaceae Bombacaceae Species Sideroxylon persimile (Hemsl.) T.D. Penn. ssp. persimile Sideroxylon tepicense (Standl.) T.D. Penn. Simarouba glauca DC. Simira calderoniana (Standl.) Steyerm. Simira salvadorensis (Standl.) Steyerm. Sinclairia deamii (B.L. Rob. & Bartlett) Rydb. Sinclairia discolor Hook. & Arn. Sinclairia glabra (Hemsl) Rydb. Sinclairia sublobatum (B. L. Rob.) Rydb. Sloanea terniflora (Sessé & Moc. ex DC.) Standl. Solanum diphyllum L. Solanum erianthum D. Don Solanum hazenii Britton Solanum lanceolatum Cav. Solanum sp. Solanum umbellatum Mill. Spondias mombin L. Spondias purpurea L. Spondias radlkoferii Donn. Sm. Stemmadenia donnell-smithii (Rose) Woodson Stemmadenia eubracteata Woodson Stemmadenia obovata K. Schum. Stemmadenia pubescens Benth. Stemmadenia robinsonii Woodson Stenocereus aragonii (F.A.C. Weber) Buxb. Sterculia apetala (Jacq.) H. Karst. Strychnos brachistantha Standl. Styphnolobium sporadicum M. Sousa & Rudd Styrax argenteus C. Presl. Styrax warscewiczii Perkins Swartzia simplex var. ochnacea (DC.) R.S. Cowan Swietenia humilis Zucc. Swietenia macrophylla King. Symplocarpon sp. Symplococarpon purpusii (Brandegee) Kobuski Symplocos limoncillo Bonpl. Synardisia venosa (Mast.) Lundell Tabebuia chrysantha (Jacq.) G. Nicholson subsp. chrysantha Tabebuia donnell-smithii Rose Tabebuia impetiginosa (Mart. ex DC.) Standl. Tabebuia rosea (Bertol.) A. DC. Tabernaemontana amygdalifolia Jacq. Talisia olivaeformis (Kunth) Radlk. Tecoma stans (L.) Juss. ex Kunth Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I I I I I I I I I 57 Family Compositae Combretaceae Euphorbiaceae Sapindaceae Sapindaceae Sapindaceae Sapindaceae Sapindaceae Melastomaceae Boraginaceae Boraginaceae Anacardiaceae Ulmaceae Ulmaceae Meliaceae Meliaceae Meliaceae Meliaceae Meliaceae Meliaceae Tiliaceae Polygonaceae Moraceae Moraceae Staphyleaceae Ulmaceae Urticaceae Urticaceae Urticaceae Urticaceae Asteraceae Asteraceae Asteraceae Asteraceae Caprifoliaceae Hydrophyllaceae Celastraceae Olacaceae Verbenaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Rutaceae 58 Species Telanthophora arborescens (Steetz) H. Rob. & Brettell Terminalia oblonga (Ruiz & Pav.) Steud. Tetrorchidium sp. nov. Thouinia brachybotrya Donn. Sm. Thouinia velutina Radlk. Thouinia villosa DC. Thouinidium decandrum (Bonpl.) Radlk. Thouinidium sp. Tibouchina sp. Tournefortia acutiflora M. Martens & Galeotti Tournefortia glabra L. Toxicodendron striatum (Ruiz & Pav.) Kuntze Trema micrantha (L.) Blume var. micrantha Trema micrantha var. strigillosa (Lundell) Standl. & Steyerm. Trichilia americana (Sesse & Moc.) T.D. Penn. Trichilia glabra L. Trichilia havanensis Jacq. Trichilia hirta L. Trichilia martiana C. DC. Trichilia sp. Trichospermum galeotti (Turcz.) Kosterm. Triplaris melaenodendron (Bertol.) Standl. & Steyerm. Trophis mexicana (Liebm.) Bureau Trophis racemosa (L.) Urb. Turpinia occidentalis (Sw.) G. Don Ulmus mexicana (Liebm.) Planch. Urera baccifera (L.) Gaudich. ex Wedd. Urera corallina (Liebm.) Wedd. Urera eggersii Hieron. Urera sp. nov. Verbesina guatemalensis B.L. Rob. & Greenm. Verbesina turbacensis Kunth Vernonia deppeana Less. Vernonia patens Kunth Viburnum hartwegii Benth. Wigandia urens (Ruiz & Pav.) Kunth Wimmeria cyclocarpa Radlk. Ximenia americana L. Xolocotzia asperifolia Miranda Xylosma chlorantha Donn. Sm. Xylosma flexuosa (Kunth) Hemsl. Xylosma sp. Zanthoxylum aguilarii Standl. & Steyerm. Biodiversity field inventories in selected watersheds Indicator (I) I I I I I I I I I I I I I I I I I I I I I I I I I I I I Family Species Indicator (I) Rutaceae Rutaceae Rutaceae Rutaceae Zanthoxylum culantrillo Kunth Zanthoxylum kellermanii P. Wilson Zanthoxylum melanostictum Schltdl. & Cham. Zanthoxylum microcarpum Griseb. I Rutaceae Zanthoxylum nicaraguense Standl. & L.O. Williams Zanthoxylum riedelianum Engl. Zapoteca formosa (Kunth) H.M. Hern. subsp. Leguminosae formosa Celastraceae Zinowiewia cuneifolia Lundell *I=Indicator species for evaluating the completeness of an inventory. I Rutaceae Biodiversity field inventories in selected watersheds I I 59 60 Figure 5A. Diospyros morenoi (Zapote negro). Finca María Auxiliadora, Los Volcanes National Park, July 2007. Previously known only from Cinquera, Cabañas Department, El Salvador, and a locality in Nicaragua. Figure 5B. Pinus oocarpa (Pino). Los Volcanes National Park, Izalco Vocano, December 2007. Figure 5C. Guapira witsbergerii (Siete camisas rojo). Los Volcanes National Park, Cerro Verde, December 2007. First photographic record of this species endemic to the study area. Figure 5D. Triphora mexicana, a very rare orchid. Finca María Auxiliadora, Los Volcanes National Park. July 2007. Biodiversity field inventories in selected watersheds Figure 5E. Oxalis salvadorensis (Trébol de peña, or cliff clover), Plan de Amayo Protected Natural Area, August 2007. Endemic to the Project area. Figure 5F. Zamia herrerae (Palmita), Plan de Amayo Protected Natural Area, August 2007. Listed as Vulnerable in the IUCN Red List. Figure 5G. Peperomia linaresii. Plan de Amayo Protected Natural Area, August 2007. Very rare, known only from two localities, both within the Project area. Figure 5H. Balmea stormae, Cerro El Águila Protected Natural Area, November 2007. Listed in the IUCN Red List. Biodiversity field inventories in selected watersheds 61 62 Figure 5I. Phyllostylon rhamnoides, Plan de Amayo Protected Natural Area, August 2007. First record for El Salvador. Figure 5J. Tetrorchidium sp. Cerro El Águila Protected Natural Area, November 2007. Figure 5K. Phytolacca riviniodes. Cerro El Águila Protected Natural Area, November 2007. First record for El Salvador. Figure 5L. Flower of Cuscatlania vulcanicola (Cuscatlania), San Marcelino Complex Protected Natural Area, December 2007. Rediscovery, being the first record since 1922 of a species known only from El Salvador. Biodiversity field inventories in selected watersheds Figure 5M. Urera sp. El Imposible National Park, December 2007. Species new to science. Figure 5N. Prunus axitlana. El Imposible National Park, December 2007. Figure 5O. Sedum salvadorense. El Imposible National Park, December 2007. Very rare plant, known in the world only from the Project area, and never before photographed. Figure 5P. Bursera longicuspis (Jiote rojo). San Marcelino Complex Protected Natural Area, December 2007. First records for El Salvador. FIGURE 5. SELECTED PHOTOGRAPHS OF FLORA TAKEN DURING THE STUDY (by Frank Sullyvan Cardoza). Biodiversity field inventories in selected watersheds 63 TABLE 12. FLORA OF CONSERVATION IMPORTANCE IN THE PROJECT AREA. Species Risk* F** SM PA PNEI PNLV AG Aegiphila panamensis Moldenke VU A1c HF X Agonandra loranthoides L.O. VU C1 DF X Williams Bauhinia paradisi Standl. & L.O. CR C2a DF X Williams VU Cedrela odorata L. DF X X X X A1cd+2cd Cordia salvadorensis Standl. raro DF X Dalbergia retusa Hemsl. VU A1acd DF X Dichapetalum donnell-smithii VU D2 HF X X Engl. Eugenia salamensis Donn. Sm. EN C2a DF X X X X Juglans olanchana Standl. & EN C2a CF X L.O. Williams Lennea viridiflora Seem. VU A2c DF X Lonchocarpus minimiflorus EN C2b DF X X X X Donn. Sm. Lonchocarpus molinae Standl. & EN C2b HF X L.O. Williams Lonchocarpus phaseolifolius EN C2b DF X X Benth. Lonchocarpus retiferus Standl. EN C2a HF X & L.O. Williams Lonchocarpus sanctuarii Standl. CR C2b DF X X & L.O. Williams Lonchocarpus santarosanus VU B1+2c HF X X X Donn. Sm. Machaerium sp.*** raro HF X Parathesis vulgata Lundell EN C2a CF X X Persea schiedeana Nees VU A1c CF X Pisonia donnellsmithii Heimerl VU B1+2c HF X X X ex Standl. Quercus skinneri Benth. VU A1c CF X X X Swietenia humilis Zucc. VU A1cd DF X X VU Swietenia macrophylla King HF X A1cd+2cd Xolocotzia asperifolia Miranda EN C2a DF X Zamia herrerae S. Calderón & VU C1 HF X X Standl. * Risk codes come from IUCN (2007): CR=Critically Endangered, EN=Endangered, VU=Vulnerable, NT=Near-Threatened, DD=Data Deficient. The other codes presented are the criteria used to assign risk classifications and are explained in IUCN (2007). 64 Biodiversity field inventories in selected watersheds **F=Type of Forest; CF=Cloud Forest, DF=Dry Forest, HF=Humid Forest; SM=San Marcelino Complex Protected Natural Area, PA=Plan de Amayo Protected Natural Area, PNEI=El Imposible National Park, CV=Cerro Verde, AG=Cerro El Águila Protected Natural Area, PNLV=Los Volcanes National Park. ***Still not on the IUCN Red List, but should be evaluated. TABLE 13. INVENTORY COMPLETENESS FOR TREE SPECIES AND RELATIVE IMPORTANCE OF ECOSYSTEMS. Number of Level of Number of Expected Indicator Species Inventory Conservation Ecosystem Species Species for Recorded Completeness Important Recorded Inventory (%) Species Completeness Humid 249 135 135 100 13 Forest Dry Forest 281 166 194 86 13 Cloud 134 57 60 95 4 Forest Savanna 51 35 35 100 0 At the ecosystem level, the habitats presenting the largest amount of threatened and endangered species are humid forest and dry forest with 13 species each. Cloud forest only has four threatened species. The savanna did not present any globally threatened tree species (Table 13). Although our objective was to evaluate which watersheds and municipalities of the study area showed the highest indices of importance for tree conservation, it was impossible to make this analysis because the inventories for specific watersheds and municipalities are largely incomplete, with the exception of one watershed (Cara Sucia) and one municipality (San Francisco Menéndez). Comparisons based on incomplete inventories would produce misleading results. They reflect the level of research effort and not real values of conservation importance (Komar 2007). 3.3. SPECIES OF INTEREST Fourteen plant species are restricted globally to the project area, including 10 trees or shrubs and 4 herbaceous species (Table 14). Prior to this study, several of these endemic species were known only from El Imposible National Park. One of the important findings of this study is that nearly all of these species exist in other locations of southwestern El Salvador. New locations have been documented for Daphnopsis witsbergerii, the famous Guapira witsbergerii (―siete camisas rojo‖, Fig. 5C), and the recently named Dalbergia salvanaturae, three trees originally described for science from El Imposible and known Biodiversity field inventories in selected watersheds 65 only to exist in the study area. Furthermore, new localities were found for the herbaceous species Ageratum salvanaturae and Peperomia linaresii. Some plants found in the study area stand out either because they represent new records for El Salvador, for Central America, or because they are probably new species to science (Table 15). We believe that 11 of the species collected are indeed new for science, although it may take several years for their descriptions to be reviewed by the scientific community and published. Six of these have never been documented outside of the study area (see Table 14), but five are recognized by the author from his own collecting work in other parts of Central America. Although they were collected prior to the present study, they have yet to be described formally in the scientific literature (J. Linares, unpublished data). Others represent a second or third collection for El Salvador, or even more special, a second collection for science. For example, worth mentioning is the discovery of several specimens of Styphnolobium sporadicum (―almendro de montaña‖) previously known only from one collection in Colombia, one in Mexico and one in El Salvador. Today the species is known from two locations within the project area. TABLE 14. PLANTS ENDEMIC TO THE IMCW PROJECT AREA, WITH COLLECTION LOCATIONS FROM THE PRESENT STUDY. Collection Species Family Habit Locality No. El Imposible, Ageratum salvanaturae B. Asteraceae Herb Plan de Amayo, 12523 Smalla et N. Kilian & San Marcelino Besleria sp. nov. Gesneriaceae Herb Cerro El Aguila 12797 Calliandra sp. nov. Leguminosae Shrub El Imposible 13227 Dalbergia salvanaturae J. Leguminosae Tree El Imposible 12664 Linares et M. Sousa Daphnopsis witsbergeri Thymelaeaceae Tree El Imposible 12707 Nevling, Matek. & Barringer Eugenia alfaroana Standl. Myrtaceae Tree El Imposible 12162 Cerro Verde, El Guapira witsbergeri Lundell Nyctaginaceae Tree 12171 Imposible, Plan de Amayo Lonchocarpus stenophyllus Leguminosae Tree Plan de Amayo 12617 M. Sousa Maytenus sp. nov. Celastraceae Tree Los Volcanes 12259 Peperomia linaresii Véliz Piperaceae Herb Plan de Amayo 12533 Pisonia sp. nov. Nyctaginaceae Shrub San Marcelino 12944 Sedum salvadorense Standl. Crassulaceae Herb El Imposible 13040 Tetrorchidium sp. nov. Euphorbiaceae Tree Cerro El Aguila 12832 Urera sp. nov. Urticaceae Tree El Imposible 13228 66 Biodiversity field inventories in selected watersheds TABLE 15. BOTANICAL SPECIES OF INTEREST COLLECTED IN THE STUDY AREA. Criterion Species Family Locality New species for Besleria sp. Gesneriaceae El Águila science New species for Calliandra sp. Leguminosae El Imposible science New species for Coccoloba sp. Polygonaceae Plan de Amayo science New species for Crossopetalum sp. Celastraceae Plan de Amayo science New species for Croton sp. Euphorbiaceae Plan de Amayo science New species for Indet. Euphorbiaceae El Imposible science New species for El Imposible y Krugiodendron sp? Rhamnaceae science Plan de Amayo New species for Pisonia sp. Nyctaginaceae San Marcelino science New species for Tetrorchidium sp. Euphorbiaceae El Águila science New species for Urera sp. Urticaceae El Imposible science New for Central Iresine herrerae Conz. & Amaranthaceae El Imposible America S.F. Blake New for Central El Imposible y Manfreda pubescens Amaryllidaceae America Plan de Amayo First record in El Agonandra loranthoides Opiliaceae San Marcelino Salvador L.O. Williams First record in El Alysicarpus vaginalis (L.) Leguminosae Plan de Amayo Salvador DC. First record in El Amyris guatemalensis Rutaceae El Imposible Salvador Lundell First record in El Arrabidaea costaricensis Bignoniaceae San Marcelino Salvador (Kraenzl.) A.H. Gentry Eugenia quercetorum First record in El Standl. & L.O. Williams ex Myrtaceae Cerro Verde Salvador Barrie First record in El Lonchocarpus molinae Leguminosae Plan de Amayo Salvador Standl. & L.O. Williams First record in El Nectandra mirafloris van Lauraceae El Águila Salvador der Werff First record in El Philactis liebmannii (Klatt) Asteraceae El Imposible Salvador S.F. Blake First record in El Phyllostylon rhamnoides (J. Ulmaceae Plan de Amayo Salvador Poiss.) Taub. Biodiversity field inventories in selected watersheds 67 Criterion First record in El Salvador Rediscovery Rediscovery Species Phytolacca rivinoide Kunth & C.D. Bouché Cuscatlania vulcanicola Standl. Sedum salvadorense Standl. Family Locality Phytolaccaceae El Águila Nyctaginaceae San Marcelino Crassulaceae El Imposible Here, these species are presented as: Probable new species to science, new records for Central America, first records for El Salvador, and rediscoveries, according to the details provided below. PROBABLE NEW SPECIES TO SCIENCE Besleria sp. (Gesneriaceae, African violet family). During the field trip to Cerro El Águila Protected Natural Area, we observed several Besleria sp., an epyphite growing on large trees. The plants were losing their leaves and only some of them had fruit, therefore only a few botanical samples were collected. In 1993, the author collected a specimen at the same location, which was determined in 1996 by Dr. L. Skog of the U.S. National Herbarium, an expert on the family, to be a probable new species (L. Skog, pers. com.) but it has not been collected again in this area. The new specimens complement the previous collections in the same area because those only had flowers. Calliandra sp. (Leguminosae, Bean family). A shrub or small tree only observed in the path known as ―El Gallinero‖, on rocky and exposed ridges of El Imposible National Park. It is not more than 4 m high and is very similar to Calliandra mexicana with which it can be mistaken, but its leaves are glabrous with very marked nerves. If this is not a new species it should be classified at least as a subspecies given its geographic isolation and incipient morphological differentiation. According to Stevents et al. (2001), this species would be part of a group of species and subspecies distributed from the north of Mexico to Panama, and is poorly known taxonomically. The flowers are red and showy and the plant is known locally as ―barba‖ or ―barba de rey‖ (beard or king’s beard). Coccoloba sp. (Polygonaceae, ―Papaturro‖ family) Shrub or small tree known locally as ―papaturrito‖. It is particulary noticeable for its smallsized leaves, probably the smallest of the genus, at least in Central America. The Coccoloba nicaraguensis has leaves as small as our specimen but this species is only known from one single collection deposited with the herbarium of the Panamerican School of Agriculture and looks different. According to data from the park rangers of Plan de Amayo Protected Natural Area, this plant was observed by them for the first time in Caluco, Sonsonate, in May 2007. This probable new species could also be identified as Coccoloba acuminata, a species from the rain forests of the Atlantic or the Caribbean 68 Biodiversity field inventories in selected watersheds slope, from Mexico to Brazil. If this species is confirmed, our collection would be the first specimen on record for the Pacific slope area. Croton sp. (Euphorbiaceae, Poinsettia family) This tree is very closely related to Croton guatemalensis (Copalchí). The main difference between them is that it has hairs and scales instead of just scales as in C. guatemalensis. It was found in Plan de Amayo Protected Natural Area. Crossopetalum sp. (Rhamnaceae, ―Huilihuishte‖ family) This tree collected in Plan de Amayo Protected Natural Area does not match with any of the species known in the region. It could be a non-described species or a new country record. Euphorbiaceae indet. Samples of this tree species have not previously been collected in El Salvador. So far, we have not been able to establish the genus to which it belongs because female flowers or fruits are required. In any case, if it is not a new species for science, it would be a new species for Central America. It was found on the way down from Cerro La Cumbre, on a ridgetop, at El Imposible National Park. Krugiodendron sp. (Rhamnaceae, ―Huilihuishte‖ family) This tree was only known from a collection deposited with the herbarium of La Laguna Botanical Garden (LAGU) and was tagged as Karwinskia sp. To the author, it is obvious that it is not a Karwinskia; furthermore, it does not match with any of the Krugiodendron species from Mesoamerica described so far. During the present study, abundant material was collected from Plan de Amayo, Las Lajas forest, and El Imposible, but it is still necessary to collect material with flowers in good condition in order to be sure that this is a new species to science. It is a small tree found in rocky areas of the aforementioned locations. Maytenus sp. (Celastraceae) A small tree, collected in the higher part of Finca María Auxiliadora inside an active coffee plantation (buffer zone of Los Volcanes National Park). The flowers are greenish and inconspicuous. Only four trees of this apparently new species for science were observed, and botanical samples were collected. These trees were all at risk of being eliminated through normal coffee plantation maintenance, as they had grown up among the coffee shrubs. Pisonia sp. (Nyctaginaceae, Bougainvillea family). The plant is a spread-out shrub as opposed to other Pisonia species existing in the country, which are erect trees. E. Martínez from the National Herbarium of Mexico, a Mesoamerican tree expert, was consulted and has concluded that this tree could belong to an undescribed species, new to science. Only four individuals were observed at the top of Cerro Chino, in San Marcelino Complex Protected Natural Area. We did not determine a local name for this species, but other species of the genus are commonly called ―uña de gato‖ (cat´s claw) because of the curved thorns on the stem. This plant should be included Biodiversity field inventories in selected watersheds 69 on the IUCN Red List because its worldwide distribution may be limited to this location. Fertile botanical specimens were collected. Tetrorchidium sp. (Euphorbiaceae, Poinsettia family) This tree was found in Cerro El Águila Protected Natural Area, without any flower or fruit, but its vegetative features match with those of the Tetrorchidium genus, even though the size and shape of the leaves do not match with any of the five species of this genus currently on record for Central America. This genus was not recorded by Linares (2003 [2005]) on his list of trees. Only two sterile trees were observed and samples of each were collected (Fig. 5J). Urera sp. (Urticaceae, ―Ortiga‖ or Nettle family). It is a medium-sized tree from 6 to 10 m high that has only been found in the higher parts of the headwaters of the El Venado River at El Imposible National Park (Fig. 5M). Its inflorescence is 3–5 cm long and forms under the leaves, and for this reason it is difficult to observe in the field. As a result, there are no previous collections of this plant. It does not match any of the Urera tree species described for northern Central America. To classify this species it will be necessary to compare the material with collections housed in other countries. NEW RECORDS FOR CENTRAL AMERICA Two plants were found, one herbaceous and one arboreal, known until now from the Pacific coast of central and southern Mexico and reported here for the first time for Central America. These findings help us to confirm the links between the flora of southwestern El Salvador and that of the Pacific coast of central and southern Mexico. Iresine herrerae (Amaranthaceae, ―Amaranto‖ family). This species is a tree that grows more than 10 m high and has up to 20 cm diameter, which is significant for a genus mainly made up of shrubs and small trees. It was originally described as native to Puchutla on the Mexican Pacific slope, in the State of Oaxaca, the same as Manfreda pubescens (see previous paragraph). It was found at El Imposible National Park. Manfreda pubescens (Amarillydaceae, ―Búcaros‖ family). In El Salvador, the genus only has one additional species, Manfreda brachystachys that grows in the northern mountain range in pine-oak forests. Our species has wider leaves with pubescent and purple spots which give it a very colorful appearance. It was found in Plan de Amayo Protected Natural Area and El Imposible National Park. FIRST RECORDS FOR EL SALVADOR Agonandra loranthoides (Opiliaceae, ―Opilias‖ family). This tree is noted in the field for its small and fleshy leaves, reddish and slightly ribbed stem with bark exfoliation occurring in long, thin, and irregular plates. Several specimens were observed in San Marcelino Complex Protected Natural Area and even though they 70 Biodiversity field inventories in selected watersheds were sterile, their identification is accurate given the unmistakable characteristics described above. Samples had only been previously collected in Honduras, that is, it was considered endemic to that country. This is the first record outside of Honduras. The species was not known to the local guides and did not have a common name. This is the second location in the world for this species since it was only known to the Comayagua valley. This species is included on the IUCN Red List as Vulnerable. Botanical samples were collected. Alysicarpus vaginalis (Leguminosae). It is an invasive herbaceous plant native to the Old World and recently naturalized in Central America. It can be a very aggressive weed. It was found in Plan de Amayo Protected Natural Area. Amyris guatemalensis (Rutaceae, citrus family). This species is a tree that grows 5–10 m high. Until now, it was only known from semideciduous forests of Guatemala at altitudes below 400 m. It differs from Amyris balsamifera by having much larger folioles and opposing leaves. It was collected at El Imposible also at altitudes below 400 m. Arrabidaea costaricensis (Bignoniaceae, ―Maquilishuat‖ family). This plant was found as a small tree of a little more than 5 cm diameter. It was originally described as native to Costa Rica and then to Mexico, where it was described with the name of Arrabidaea erecta. It is easily recognized even in sterile state because it is the only tree or shrub species of a genus where all the rest are climbing vines. It was collected in Plan de Amayo (Municipality of Caluco, Department of Sonsonate) as well as in Las Lajas forest in San Marcelino Protected Natural Area. Eugenia quercetorum (Myrtaceae, Guava family). This species is a small tree collected previously only in Honduras. Only one specimen was observed at the foot of Cerro Verde, near the Izalco Volcano. Botanical samples were collected. This is the first record outside of Honduras. Lonchocarpus molinae (Leguminosae). This species, which is included on the IUCN red list, was described in Honduras but had not been collected in the region since 1950. It has been occasionally collected in Mexico. It is a conspicuous tree because its trunk has several thick ribs that resemble intertwined snakes, hence the common name ―culebro‖ (snake). It was found in Plan de Amayo Protected Natural Area. Nectandra mirafloris (Lauraceae, Avocado family). So far, this tree had only been collected in one specific location of Nicaragua. Several specimens were observed in this study in the cloud forest of Cerro El Águila Protected Natural Area. Samples were collected. Biodiversity field inventories in selected watersheds 71 Philactis leibmannii (Asteraceae, Sunflower family). This species is a small shrub known from Mexico and Guatemala. In the latter country it was reported in Santa Rosa, a department with which El Imposible forest shares many species. Therefore, it is not surprising that it was found there. It was collected at El Pedrerón lookout, on the way to La Fincona, in El Imposible National Park. Phyllostylon rhamnoides (Ulmaceae, Elm family). This tree is typical of dry or very dry forests and relatively common in other countries of Central America, but never reported in El Salvador. It was found in Plan de Amayo Protected Natural Area (Fig. 5I). Phytolacca rivinoides (Phytolaccaceae). This shrub with spread-out and fleshy stems is common to the Honduran and Guatemalan area of El Trifinio (Komar et al. 2006), but it had not been recorded before in El Salvador until it was collected on Cerro El Águila. It has distinct morphology because it is the only species of its genus which has the form of a shrub. Fertile botanical samples of this species were collected (Fig. 5K). REDISCOVERIES: Cuscatlania vulcanicola (Nyctaginaceae). This small herbaceous plant was described in 1923 from material collected in San Vicente, El Salvador, in 1922. Only two herbarium sheets were known with very little material (two small branches with some leaves and very few flowers). It had not been reported since from anywhere in the world. We found a population at San Marcelino Protected Natural Area (Las Lajas forest), with probably more than 30 plants. It is a rediscovery, a major botanical finding that emphasizes the importance of the locality as a reservoir of rare plants unique in the world. Botanical samples were collected and the finding was amply documented with photographs taken by Frank Sullyvan Cardoza (Fig. 5L). Sedum salvadorense (Crassulaceae). This potentially ornamental herbaceous plant was collected and described by Standley at Finca Las Colinas (wrongly cited as Colima) in January 1922. In 1999, Eliberto Sandoval collected some specimens at El Imposible National Park that were identified as Sedum sp. We have now documented the existence of this species in the area with several herbarium specimens and many photographs (Fig. 5O). We do not know of any records of the species in another country, and therefore, we consider it a rediscovery of an endemic species not reported since its discovery in 1922. 72 Biodiversity field inventories in selected watersheds 3.4. COMPLETENESS OF THE TREE INVENTORY The tree inventory for southwestern El Salvador made progress, from a completeness estimate of 76% prior to the study to a completeness estimate of 93% at the end of the present study. In other words, 312 of the 335 species that are widely distributed and considered to be indicators for inventory completeness have been recorded. The completeness of the tree inventory at the level of watersheds progressed significantly in the six watersheds where the flora studies were carried out. Currently, the watersheds with the most advanced tree inventories are Cara Sucia (82%), Izalco (58%), and San Julián (57%). In spite of the little time invested, the Coatepeque and Río Grande de Sonsonate watersheds also reported some progress although none of them exceeded 50% (Table 16). Through the present study, the tree inventory in the municipality of San Francisco Menéndez has almost been completed (Table 17). Another municipality that shows a high level of completeness is the municipality of Caluco, which prior to the research was at 0%. The municipality of Izalco shows a completeness level of 65%. The remaining municipalities show an inventory completeness level below 40%. Twelve municipalities do not have any records of trees in the data bases consulted (Table 17). At the ecosystem level, complete inventories have been attained for Humid Forest and Savanna (Table 13). The inventory of the Dry Forest is nearly complete, that is, 86% of the expected species are on record. The Cloud Forest is 95% complete. However, it is still possible to detect new ―unexpected‖ species in all of these ecosystems. TABLE 16. INVENTORY COMPLETENESS FOR TREE SPECIES AND RELATIVE IMPORTANCE OF WATERSHEDS. Inicial Level Final Level of Expected Registered of Inventory Inventory Watershed Registered Species Completeness Completeness Species (of 355) (%) (%) Barra de Santiago 128 128 38 38 Barra Salada 0 0 0 0 Cara Sucia 334 274 64 82 Izalco 225 193 15 58 Lago de 175 148 13 44 Coatepeque Los Cóbanos 1 1 0 0 Río Cauta 0 0 0 0 Río Grande de 44 44 13 19 Sonsonate San Juan 0 0 0 0 San Julian 295 190 1 57 San Pedro 79 62 5 5 Biodiversity field inventories in selected watersheds 73 TABLE 17. INVENTORY COMPLETENESS FOR TREE SPECIES AND RELATIVE IMPORTANCE OF MUNICIPALITIES. Expected Initial Level of Final Level of Species Species Inventory Inventory Municipality Recorded Recorded Completeness Completeness (of 355) (%) (%) Acajutla 4 4 1 1 Apaneca 50 29 9 9 Armenia 0 0 0 0 Caluco 295 190 0 57 Chalchuapa 12 12 6 16 Concepción de 16 16 5 5 Ataco Cuisnahuat 0 0 0 0 El Congo 200 143 10 43 Guaymango 0 0 0 0 Izalco 225 193 15 58 Juayua 79 45 7 13 Jujutla 18 18 5 5 Nahuizalco 0 0 0 0 Nahulingo 0 0 0 0 Salcoatitan 0 0 0 0 San Antonio del 0 0 0 0 Monte San Francisco 334 276 68 86 Menéndez San Julián 2 2 1 1 San Pedro Puxtla 0 0 0 0 Santa Ana 195 151 6 45 Santa Catarina 0 0 0 0 Masahuat Santo Domingo de 0 0 0 0 Guzmán Sonsonate 10 10 3 3 Sonzacate 0 0 0 0 Tacuba 16 16 5 5 Tepecoyo 0 0 0 0 74 Biodiversity field inventories in selected watersheds 4. RECOMMENDATIONS     Continue the flora inventory efforts in the southwest region of El Salvador with the goal of completing the inventories at the levels of ecosistema, watershed, municipality, and protected natural areas. Conduct new inventory efforts in the savannas of palm trees and ―morros‖ (Crescentia alata) and other ecosystems, with an emphasis on other plant groups such as herbaceous and shrublike species because samples of these were not taken in this study. Prepare sampling protocols for epyphitic species because they indicate to a large extent the degree of maturity and/or conservation of the ecosystems as well as the levels of degradation and/or contamination. Carry out training workshops for the park rangers of the Protected Natural Areas, focused on the threatened and endangered species recorded for those areas. Biodiversity field inventories in selected watersheds 75 CHAPTER 3: PRELIMINARY INVENTORY OF FISH SPECIES IN ELEVEN WATERSHEDS OF SOUTHWESTERN EL SALVADOR Francisco Chicas Batres School of Biology Faculty of Natural Sciences and Mathematics University of El Salvador francisco.chicas@ues.edu.sv José Alberto González Leiva Institute of Marine Sciences and Limnology of El Salvador University of El Salvador ICMARES/UES albergleiva@yahoo.es 1. INTRODUCTION 2008 Mesoamerica is considered one of the most important regions of the American continent for its great diversity of species (CEPF 2004, PNUMA 2004). From the moment it was formed, the region has functioned as the link between the two major continental masses, facilitating the migration processes of the terrestrial fauna (Jiménez 1994, Waid et al. 1999). Dynamic geological processes such as plate tectonics and volcanism have constantly modified the relief, allowing the formation of geographic barriers that along with the modification of niches have facilitated vicariant processes and dispersion events of many taxa, among them freshwater fish species (Concheiro Pérez et al. 2006, Hrbek 2007, Martin & Bermingham 1997, Morrone 2007, Perdices et al. 2002). It is estimated that the number of freshwater fish species of the Mesoamerican region reaches at least 446 (Myers 1966) with very specific distribution patterns in certain zones. Accordingly, Bussing (1976) proposed four icthyogeographic provinces (Fig. 6). One of them is the Chiapas–Nicaraguan province which comprises the entire Salvadoran territory and which is considered poor in species and with little endemism (Bussing 1998), a condition attributed to the ―recent‖ closing of the Isthmus of Panama that started approximately 3 million years ago (Bermingham et al. 1997). This may have been too recent to allow a higher degree of speciation of the representative genera of the group, as 76 Biodiversity field inventories in selected watersheds has occurred in North and South America. Perhaps for these reasons and because their size is smaller than the size of their congeners of other regions, the native populations of freshwater fish species of El Salvador have been more or less ignored. Their conservation has been similarly ignored, to the extent that the introduction of exotic species was recommended in order to diversify the diet and increase the income of the rural population (Hildebrand 1925, Lin 1957). FIGURE 6: ICHTHYOGEOGRAPHIC PROVINCES OF CENTRAL AMERICA Source: Bussing (1998). On the other hand, the marine ichthyofauna of El Salvador is included in the biogeographic region known as the Tropical Eastern Pacific (TEP), which extends from the southern part of Magdalena Bay (~ 24°N) in Baja California, passing through the Gulf of California and south along the continental coastline to approximately Cabo Blanco (4°S) in Perú (Zapata & Robertson 2006). This region also comprises five oceanic islands and groups of islands: Revillagigedos, Clipperton Island, Coco Island, Malpelo Island, and the Galapagos Islands (Fig. 7). The TEP has a coastal fish fauna that comprises approximately 1,285 known species, where the highest number (740–760 species) is located in Costa Rica and Panama. The TEP has the highest endemism rate, with 911 endemic species from coastal habitats (79%), when compared to any other tropical region of similar size (Robertson & Allen 2008). Biodiversity field inventories in selected watersheds 77 FIGURE 7. THE TROPICAL EASTERN PACIFIC BIOGEOGRAPHIC REGION Source: Zapata & Robertson 2006. In an analysis of the matter specifically made for El Salvador, we establish that several ichthyofauna studies which comprise two nationwide inventories have been carried out (Hildebrand 1925, Boeseman 1956), and more recently, Orellana (1992) published a work that compiles the research conducted in the country as of that year. With regard to lakes (lotic systems), the main studies are the research conducted by Monterrosa (1960), Johnson (1971a) and Ulloa & Castro (1985b) on Lake Gϋija; Lake Ilopango by Johnson & Argumedo (1971a), Godínez & Castro (1977), Ulloa & Melgar (1983), López (1995) and Huezo & Rosales (2001); Lake Coatepeque by Johnson & Argumedo (1971b); Laguna de Apastepeque by Johnson & Ramírez (1975); Laguna de Olomega by Johnson (1971b) and Castro & Godínez (1978); Laguna de Metapán by Ulloa & Castro (1985a); and the works of Castro (1983) and Ulloa et al. (1988) on the Cerrón Grande Reservoir. The studies of García (1974), García & Robinson (1974), Castro (1976) and Castro & Robinson (1976) on the Paz, Lempa and Grande de San Miguel rivers cover the lentic systems. One of the most recent studies on freshwater fish species was conducted by González (1995) and addressed endangered native fish species associated with the main continental water bodies of the country. Another was conducted by PREPAC (2005), which characterized and updated the state of use and conservation of the fishing resources of lakes and lagoons and some estuaries of the country. After an analysis of the IUCN criteria and the conservation status of the water bodies of the country, the study recommended to treat the native freshwater species as Vulnerable. With respect to marine fish, the most recent country-wide research was carried out by Fuentes and Hernández (2004), covering the entire Salvadoran coast and various bathimetric strata within the four fishing regions of the country. Other studies that include 78 Biodiversity field inventories in selected watersheds sampling on the continental shelf have been carried out to evaluate the effectiveness of circular fish hooks for catching migratory fish (Pacheco & Cokkom 2007) and in-shore species (Barahona & Henríquez 2007). The remaining research has been directed towards certain geographic areas of the country such as the Jiquilisco Bay (Calderón & Hernández 1974, Phillips & Ulloa 1981, Mejía & Chicas 2007, Méndez 2006); Estero de Jaltepeque (Candray & Ríos 2005, González & Ramírez 2007, Ulloa et al. 2003); Barra de Santiago (Orellana 1992, Vázquez 1995); and Gulf of Fonseca (Chicas et al. In preparation). The purpose of the present study was to evaluate the status of inventory completeness of the fish communities existing in the hydrographic watersheds of the southwestern region of El Salvador through a review of the studies carried out or published as of 2007 and a visual sampling report from the Los Cóbanos Reef System as of 2008, prepared by Enrique Barraza. 2. METHODOLOGY 2.1. STUDY AREA The study area comprises eleven hydrographic watersheds of southwestern El Salvador, located across 25 municipalities of the Departments of Ahuachapán, Sonsonate and part of Santa Ana (Table 18). The study area is characterized by a complex hydrological system formed by rivers with different flow intensity, lakes, and ponds. The area also includes estuarine ecosystems connected to the sea such as Barra de Santiago, Bocana San Juan, Cauta, and Barra Salada (Fig. 8). Los Còbanos ―watershed‖ is important because of it’s rocky reef ecosystem with coral formations that maintain a diversity of species. 2.2. INFORMATION COLLECTION The present report only consisted in a thorough bibliographic review of the fish studies carried out in the study area (Hildebrand 1925, Boeseman 1956, García & Robinson 1974, Orellana 1985, 1992, Jiménez 1986, Vásquez 1993, González 1995, MARN 2000b, 2005, Galdámez 2002, Reyes Bonilla & Barraza 2003, Fuentes & Hernández 2004, PREPAC 2005, ICMARES 2006, 2007, Segovia & Navarrete 2007, Schmitter-Soto 2007). The scope of these studies ranges from lists of species to deep-sea fishing research in the open sea where large fish species such as mahi-mahi or dolphinfish, shark and sword fish are caught. Additionally, the catalogued collections of the School of Biology of the University of El Salvador and the Natural History Museum of El Salvador as well as the references of other museums cited in scientific publications were verified. This information was used to prepare a species database, arranging them by ecosystem, watershed, and municipality. It is important to mention that georeferenced records are very few. Biodiversity field inventories in selected watersheds 79 TABLE 18. WATERSHEDS, MUNICIPALITIES, AND ECOSYSTEMS LOCATED IN THE DEPARTMENTS OF AHUACHAPAN, SANTA ANA, AND SONSONATE, CONSIDERED IN THE EVALUATION OF THE FISH INVENTORY. Departament Ahuachapán Santa Ana Watershed Municipality Ecosystem Cara Sucia, Barra de Santiago, *Bocana de San Juan, *Cauta, *San Pedro, *Grande de Sonsonate Coatepeque Concepción de Ataco, Guaymango, Jujutla, San Francisco Menéndez, San Pedro Puxtla, Tacuba Freshwater rivers and lakes, estuarine and marine environments El Congo Freshwater lake Acajutla, Armenia, Apaneca, Caluco, *Bocana de San Cuisnahuat, Santa Isabel Juan, Freshwater rivers, Ishuatán, Izalco, Juayúa, *Cauta, *San estuarine and Nahuizalco, Nahulingo, Pedro, mangrove forests Chalchuapa, San Antonio Sonsonate Los Cóbanos, environments, marine del Monte, San Julián, *Grande de environments, rocky Sonsonate, Izalco, Santa Catarina Masahuat, beach with coral Santo Domingo de Barra Salada y San outcrops, coral reef Guzmán, Sonsonate, Julián Salcoatitán y Sonzacate *Watersheds shared by the departments of Ahuachapán and Sonsonate. The classification criterion of Froese & Pauly (2007) was used in the ecological organization of the species on record. This classification establishes four types of habitats: Marine, Brackish, Reef-Associated, and Freshwater. Since a species may live in more than one habitat, we created combinations of those that can be inhabited by the same species, resulting in six categories: Reef, Marine, Marine-Reef, Marine-Brackish, Freshwater-Brackish, and Freshwater. The list of species was organized in alphabetical order by family and according to genus and species. 2.3. INVENTORY COMPLETENESS LEVEL The inventory completeness level was obtained from the relationship between the number of species recorded in the study area and the expected number of species to be found in the same area. The latter criterion was based on the records of generalist species found in other areas of the country as of 2007 and the reef fish studies of Los Cóbanos carried out by Segovia in 2007 (in preparation) and sampling undertaken by Barraza in the same area in 2008 (E. Barraza, unpublished data). 80 Biodiversity field inventories in selected watersheds FIGURE 8. STUDY AREA IN SOUTHWESTERN EL SALVADOR, WITH WATERSHEDS AND MUNICIPALITIES LABELED. Map by J. F. Gutiérrez. The works on regional fish distribution that refer to the presence of species in El Salvador (Villa 1971, 1982, Bussing 1998, Bussing & López 1993, Perdices et al. 2002, Smith & Bermingham 2005, Hernández 2006, Soto-Galera 2006, Ornelas-García et al. 2008, Robertson & Allen 2008, Froese & Pauly 2009) were consulted as well, comparing them with MARN records. As a result, the number of expected species in the study area is at least 541 (Table 19). 2.4. SELECTION OF IMPORTANT SPECIES FOR CONSERVATION Species considered important for conservation were selected according to their ecological and economic importance, vulnerability, and conservation status established in other studies and in the regional and worldwide databases (PREPAC 2005, IUCN 2007, 2009, Robertson & Allen 2008, Froese & Pauly 2009). For the present study, they were grouped in six categories: (A) included on the IUCN Red List, (B) freshwater species that are threatened or are indicative of environmental health, (C) endemic species, (D) living fossil, (E) scuba diving tourism, and (F) fishing importance. Biodiversity field inventories in selected watersheds 81 TABLE 19. LIST OF FISH SPECIES CONSIDERED TO BE INDICATORS OF COMPLETE SITE INVENTORIES IN SOUTHWESTERN EL SALVADOR, BUT NOT YET RECORDED. No. Family Scientific Name Habitat Trinectes fimbriatus Achiridae Marine 1 Trinectes fonsecensis Achiridae Marine-Brackish 2 Albula vulpes Albulidae Marine-Reef associated 3 Alopias pelagicus Alopiidae Reef associated 4 Ammodytoides gilli Ammodytidae Marine 5 Reef associated Antennarius avalonis Antennariidae 6 Reef associated Antennarius sanguineus Antennariidae 7 Reef associated Antennarius strigatus Antennariidae 8 Bagre pinnimaculatus Ariidae Freshwater-Brackish 9 Cathorops fuerthii Ariidae Marine-Brackish 10 Cathorops multiradiatus Ariidae Freshwater-Brackish 11 Cathorops steindachneri Ariidae Marine-Brackish 12 Marine-Brackish Notarius biffi Ariidae 13 Marine-Brackish Notarius planiceps Ariidae 14 Marine-Brackish Genus A platypogon Ariidae 15 Atherinella argentea Atherinopsidae Freshwater 16 Atherinella eriarcha Atherinopsidae Reef associated 17 Atherinella nepenthe Atherinopsidae Marine 18 Atherinella pachylepis Atherinopsidae Reef associated 19 Membras gilberti Atherinopsidae Marine 20 Reef associated Canthidermis maculata Balistidae 21 Reef associated Melichthys niger Balistidae 22 Ablennes hians Belonidae Marine-Reef associated 23 Tylosurus crocodilus fodiator Belonidae Reef associated 24 Marine Tylosurus pacificus Belonidae 25 Marine Strongylura scapularis Belonidae 26 Reef associated Entomacrodus chiostictus Blenniidae 27 Reef associated Hypsoblennius brevipinnis Blenniidae 28 Marine Bothus constellatus Bothidae 29 Marine Bothus leopardinus Bothidae 30 Marine Engyophrys sanctilaurentii Bothidae 31 Marine Monolene asaleus Bothidae 32 Marine Perissias taeniopterus Bothidae 33 Marine Brama dusumieri Bramidae 34 Marine Bregmaceros bathymaster Bregmacerotidae 35 Reef associated Grammonus diagramus Bythitidae 36 Reef associated Ogilbia ventralis Bythitidae 37 Marine Carangoides otrynter Carangidae 38 Marine Caranx lugubris Carangidae 39 Marine Caranx melanpigus Carangidae 40 82 Biodiversity field inventories in selected watersheds No. 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 Family Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carapidae Carapidae Carcharhinidae Carcharhinidae Carcharhinidae Carcharhinidae Centropomidae Centropomidae Centropomidae Chaenopsidae Chaenopsidae Characidae Cichlidae Cichlidae Cichlidae Cichlidae Cichlidae Cirrhitidae Clupeidae Clupeidae Clupeidae Clupeidae Clupeidae Clupeidae Congridae Congridae Congridae Congridae Congridae Congridae Coryphaenidae Scientific Name Chloroscombrus orqueta Decapterus maroadsi Decapterus macrosoma Hemicaranx leucurus Hemicaranx zelotes Naucrates ductor Selar crumenophthalmus Seriola lalandi Seriola peruana Seriola rivoliana Trachinotus kennedyi Trachinotus rhodopus Uraspis secunda Carapus dubius Carcharhinus albimarginatus Carcharhinus galapagensis Negaprion brevirostris Prionace glauca Rhizoprionodon longurio Centropomus unionensis Centropomus viridis Ekemblemaria myersi Emblemaria piratica Roeboides bouchellei Oreochromis mosambicus Oreochromis aureus Oreochromis urolepis hornorum Parachromis dovii Tilapia rendalli Oxycirrhites typus Etrumeus teres Harengula thrissina Odontognathus panamensis Opisthonema bulleri Opisthonema medirastre Opisthopterus dovii Ariosoma gilberti Chiloconger dentatus Gnathophis cinctus Gorgasia punctata Paraconger californiensis Rhynchoconger nitens Coryphaena equiselis Habitat Marine-Brackish Marine Reef associated Marine-Brackish Marine-Brackish Reef associated Reef associated Marine-Brackish Marine Reef associated Marine-Brackish Marine-Brackish Marine Marine Reef associated Reef associated Marine-Reef associated Marine Marine Marine-Brackish Marine Marine Marine Freshwater Freshwater Freshwater Freshwater Freshwater Freshwater Reef associated Marine Marine-Brackish Marine-Brackish Marine Marine Marine-Brackish Reef associated Marine Marine Marine Marine Marine Marine Biodiversity field inventories in selected watersheds 83 No. 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 84 Family Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Cynoglossidae Dactyloscopidae Dactyloscopidae Dactyloscopidae Dactyloscopidae Dactyloscopidae Dasyatidae Dasyatidae Diodontidae Echeneidae Echeneidae Echeneidae Engraulidae Engraulidae Engraulidae Engraulidae Engraulidae Engraulidae Engraulidae Engraulidae Engraulidae Exocoetidae Exocoetidae Exocoetidae Exocoetidae Exocoetidae Exocoetidae Exocoetidae Gerreidae Gerreidae Gerreidae Gerreidae Scientific Name Symphurus atramentatus Symphurus callopterus Symphurus chabanaudi Symphurus elongatus Symphurus fasciolaris Symphurus gorgonae Symphurus leei Symphurus melanurus Symphurus melasmatotheca Symphurus prolatinaris Symphurus undecimplerus Symphurus williamsi Dactyloscopus pectoralis Gillellus arenicola Gillellus searcheri Gillellus semicinctus Heteristius cinctus Dasyatis brevis Dasyatis dipterura Chilomycterus affinis Remora australis Remora osteochir Remorina albescens Anchoa mundeola Anchoa walkeri Anchoa argentivittata Anchoa curta Anchoa exigua Anchoa ischana Anchoa nasus Cetengraulis mysticetus Lycengraulis poeyi Cypselurus callopterus Exocoetus volitans Fodiator rostratus Hirundichthys marginatus Hirundichthys speculiger Oxyporrhamphus micropterus Prognichthys tringa Eucinostomus entomelas Eugerres axillaris Eugerres brevimanus Eugerres lineatus Habitat Reef associated Marine Marine-Brackish Marine-Brackish Marine-Brackish Marine Marine Marine-Brackish Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine Reef associated Marine Marine Marine Marine-Brackish Freshwater-Brackish Marine Freshwater-Brackish Marine Marine Marine-Brackish Marine-Brackish Marine-Brackish Marine Marine Marine Marine Marine Marine Marine Marine-Brackish Marine Marine Marine Biodiversity field inventories in selected watersheds No. 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 Family Ginglymostomatidae Gobiesocidae Gobiesocidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gymnuridae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Haemulidae Hemiramphidae Hemiramphidae Hemiramphidae Kyphosidae Labridae Labridae Labridae Labridae Labridae Labridae Labrisomidae Scientific Name Ginglymostoma cirratum Gobiesox papillifer Tomicodon petersii Akko rossi Chriolepis cuneata Elacatinus digueti Elacatinus puncticulatus Gobioides peruanus Gobiosoma etheostoma Gobiosoma nudum Gobiosoma paradoxum Gobulus hancocki Gymneleotris seminuda Microgobius brevispinis Microgobius cyclolepis Microgobius emblematicus Microgobius erectus Microgobius tabogensis Parrella maxillaris Gymnura marmorata Anisotremus dovii Anisotremus pacifici Conodon serrifer Haemulon maculicauda Haemulopsis axillaris Haemulopsis elongatus Haemulopsis nitidus Orthoptristis cantharinus Orthoptristis chalceus Pomadasys bayanus Pomadasys branickii Xenistius californiensis Hemiramphus saltator Hyporhamphus gilli Hyporhamphus rosae Kyphosus analogus Decodon melasma Halichoeres chierchiae Halichoeres melanotis Iniistius pavo Novaculichthys taeniourus Polylepion cruentum Dialommus macrocephalus Habitat Marine-Reef associated Marine Marine Marine Marine Marine-Reef associated Marine-Reef associated Freshwater-Brackish Marino Reef associated Marine-Reef associated Reef associated Reef associated Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine Reef associated Marine Marine Marine Marine Marine Freshwater-Brackish Marine-Brackish Marine Marine Marine Freshwater-Brackish Marine Reef associated Reef associated Reef associated Reef associated Reef associated Reef associated Marine Biodiversity field inventories in selected watersheds 85 No. 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 86 Family Labrisomidae Labrisomidae Labrisomidae Lophiidae Lutjanidae Lutjanidae Malacanthidae Malacanthidae Malacanthidae Microdesmidae Monacanthidae Moridae Mugilidae Mugilidae Mugilidae Mugilidae Mullidae Muraenesocidae Muraenidae Muraenidae Muraenidae Muraenidae Muraenidae Muraenidae Muraenidae Muraenidae Myliobatidae Myliobatidae Myliobatidae Myliobatidae Narcinidae Narcinidae Nettastomatidae Ogcocephalidae Ophichtidae Ophichtidae Ophichtidae Ophichtidae Ophichtidae Ophichtidae Ophichtidae Ophidiidae Ophidiidae Scientific Name Labrisomus multiporosus Malacoctenus ebisui Paraclinus mexicanus Lophiodes caulinaris Lutjanus jordani Lutjanus viridis Caulolatilus affinis Caulolatinus hubbsi Caulolatinus princeps Microdesmus dipus Aluterus monocerus Physiculus nematopus Chaenomugil proboscideus Joturus pichardi Mugil hospes Xenomugil thuborni Pseudupeneus grandiscuamis Cynoponticus coniceps Anarchias galapagensis Enchelycore octaviana Gymnomuraena zebra Gymnothorax panamensis Muraena argus Muraena lentiginosa Muraena clepsydra Uropterygius macrocephalus Manta birostris Mobula japanica Mobula thursoni Rhinoptera steindachneri Diplobatis ommata Narcine entemedor Hoplunnis sicarius Zalieutes elater Echiophis brunneus Myrichthys tigrinus Ophichthus frontalis Ophichthus longipenis Ophichthus mecopterus Ophichthus triserialis Ophichthus zophochir Brotula clarkae Lepophidium negropinna Habitat Reef associated Marine Reef associated Marine Reef associated Reef associated Marine-Brackish Marine Reef associated Marine Reef associated Marine Marine Freshwater-Brackish Marine-Brackish Marine Marine Marine Marine Reef associated Reef associated Marine Reef associated Reef associated Reef associated Reef associated Reef associated Reef associated Marine Marine Reef associated Marine Marine Marine Marine Reef associated Marine Marine Marine Marine Marine Marine Marine Biodiversity field inventories in selected watersheds No. 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 Family Ophidiidae Ophidiidae Opistognathidae Opistognathidae Ostraciidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Paralichthyidae Poeciliidae Poeciliidae Poeciliidae Poeciliidae Priacanthidae Pristidae Pristigasteridae Pristigasteridae Profundulidae Pseudocharchariidae Rhinobatidae Rhinobatidae Rhinobatidae Scaridae Scaridae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Scientific Name Lepophidium prorates Ophidion imitator Opisthognatus punctatus Opisthognatus scops Lactoria diaphana Ancylopsetta dendritica Citharichthys xanthostigma Cyclopsetta panamensis Cyclopsetta querna Etropus crossotus Etropus peruvianus Hippoglossina bollmani Hippoglossina tetrophthalma Paralichthys woolmani Syacium latifrons Syacium ovale Poecilia marcellinoi Poeciliosis pleurospilus Poecilia salvatoris Poeciliopsis turrubarensis Cookeolus japonicus Pristis pristis Neoopisthopterus tropicus Pliosteostoma lutipinnis Profundulus labialis Pseudocarcharias kamoharai Rhinobates leucorhynchus Rhinobatos glaucostigma Zapterix exasperata Nicholsina denticulata Scarus ghobban Bairdiella armata Corvula macrops Cynoscion albus Cynoscion phoxocephalus Elattarchus archidium Isopisthus remifer Larimus effulgens Larimus argenteus Larimus pacificus Menticirrhus panamensis Menticirrhus elongatus Menticirrhus nasus Habitat Marine Marine Marine Marine Reef associated Marine Marine Marine-Brackish Marine-Brackish Marine-Brackish Marine Marine Marine Marine-Brackish Marine Marine Freshwater Freshwater Freshwater Freshwater-Brackish Reef associated Freshwater-Brackish Marine-Brackish Marine Freshwater Marine Marine Marine Reef associated Reef associated Marine-Brackish Marine-Brackish Marine Marine-Brackish Marine Marine Marine-Brackish Marine Marine Marine Marine Marine Marine Biodiversity field inventories in selected watersheds 87 No. 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 88 Family Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Scombridae Scombridae Scombridae Scombridae Scombridae Scombridae Scombridae Scorpaenidae Scorpaenidae Scorpaenidae Scorpaenidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Scientific Name Menticirrhus paitensis Menticirrus undulatus Ophioscion imiceps Ophioscion scierus Ophioscion strabo Paralonchurus goodei Paralonchurus petersii Pareques viola Stellifer ericymba Stellifer fuerthii Stellifer illecebrosus Umbrina dorsalis Umbrina analis Umbrina bussingi Umbrina xanti Acanthocybium solandri Auxis rochei rochei Auxis thazard thazard Katsuwonus pelamis Sarda orientalis Scomber japonicus Thunnus albacares Pontinus sierra Scorpaena histrio Scorpaena russula Scorpaenodes xyris Alphestes multiguttatus Dermatolepis dermatolepis Diplectrum eumelum Diplectrum euryplectrum Diplectrum labarum Diplectrum macropoma Diplectrum pacificum Diplectrum rostrum Epinephelus acanthistius Epinephelus exsul Epinephelus niphobles Hemanthias peruanus Hemanthias signifer Mycteroperca xenarcha Pronotogrammus multifasciatus Pseudogramma thaumasium Serranus fasciatus Habitat Marine Marine Marine Marine Marine-Brackish Marine-Brackish Marine Reef associated Marine Marine Marine Marine Marine Marine Marine Marine Marine-Brackish Marine Marine Marine Marine Marine-Brackish Marine Marine Marine Reef associated Marine Reef associated Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine Marine-Brackish Marine Reef associated Marine-Reef associated Biodiversity field inventories in selected watersheds No. 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 Family Sparidae Sphyrnidae Sphyrnidae Sphyrnidae Stromateidae Syngnathidae Synodontidae Synodontidae Tetraodontidae Triakidae Trichiuridae Triglidae Triglidae Triglidae Triglidae Triglidae Triglidae Tripterygiidae Tripterygiidae Uranoscopidae Urolophidae Urolophidae Zanclidae Scientific Name Calamus brachysomus Sphyrna corona Sphyrna media Sphyrna mokarran Peprilus medius Syngnathus auliscus Synodus lacertinus Synodus sechurae Sphoeroides sechurae Mustelus dorsalis Trichiurus lepturus Bellator loxias Prionotus albirostris Prionotus birostratus Prionotus horrens Prionotus ruscarius Prionotus stephanophrys Axoclinus lucillae Enneanectes carminalis Kathetostoma averruncus Urobatis halleri Urotrygon rogersi Zanclus cornutus Habitat Reef associated Marine Marine Marine-Reef associated Marine Marine-Brackish Reef associated Marine Marine Marine Marine-Brackish Marine Reef associated Marine Marine Marine Marine-Brackish Reef associated Reef associated Marine Marine Marine Reef associated Biodiversity field inventories in selected watersheds 89 3. RESULTS 3.1. FISH SPECIES RECORDED IN THE STUDY AREA It was possible to establish that the study area presents 753 records of 220 fish species (Table 20) corresponding to 139 genera of 72 families. This value is equivalent to 40% of the nationwide list of fish species. Out of this total, 19 are primary freshwater species; 16 are from freshwater-brackish environments; 57 from marine-brackish or estuarine environments; 20 are strictly marine species; 13 are marine-reef species; and 89 are reef species. Most of the recorded species have nutritional and economic importance for the local communities engaged in the extraction and commercialization of this resource. These include commonly consumed species such as: snappers of the genus Lutjanus, jurels of the genera Caranx and Elagatis, bass of the genus Centropomus, ―mojarras‖ represented by the genera Eucinostomus currani and Diapterus peruvianus, ―roncadores‖ of the genera Pomadasys and Haemulopsis, mahi-mahi or dolphinfish (Coryphaena hippurus), and mackerel (Scomberomorus sierra). 3.2. SPECIES RECORDED BY ECOSYSTEM FRESHWATER ECOSYSTEM Lake Coatepeque and Laguna de Las Ninfas are among the most notable lotic ecosystems, complementing a very broad fluvial system formed by the Apancoyo, Grande de Sonsonate, Banderas, Las Marías and other minor rivers (Fig. 3-3). To date, the number of freshwater and freshwater-brackish species recorded in the area is 35. Most of them are native occurring in almost all the watersheds of the country, such as Astyanax fasciatus, Amphiolophus macracanthus, Rhamdia quelen, Poeciliopsis gracilis y P. sphenops. Eight families comprise the freshwater species: Atherinopsidae, Characidae, Cichlidae, Gobiidae, Eleotridae, Heptapteridae, Lepisosteidae, and Poecilidae. The freshwater-brackish species are frequently found in the upper parts of the estuaries or temporarily living there looking for food or the proper environmental conditions to spawn (Bussing 1998) when the salinity decreases as a result of the rains. They include Pacific four-eyed fish (Anableps dowei), freshwater mullet (Agonostomus monticola), Pacific sleeper (Gobiomorus maculatus), and marbled swamp eel (Synbranchus marmoratus). 90 Biodiversity field inventories in selected watersheds TABLE 20. FISH RECORDED IN SOUTHWESTERN EL SALVADOR, BY ECOSYSTEM, THROUGH 2008. Spanish Common Nº Family Scientific Name Name Coral Reef Acanthurus triostegus 1 Acanthuridae cirujano Acanthurus xanthopterus 2 Acanthuridae cirujano Prionurus laticlavius 3 Acanthuridae cirujano Prionurus punctatus 4 Acanthuridae cirujano Apogon dovii 5 Apogonidae cardenal Apogon pacificus 6 Apogonidae cardenal Apogon retrosella 7 Apogonidae cardenal Balistes polylepis 8 Balistidae tunco Pseudobalistes naufragium 9 Balistidae tunco Sufflamen verres 10 Balistidae tunco Ophioblennius steindachneri 11 Blenniidae señorita Plagiotremus azaleus 12 Blenniidae diente sable Alectis cilliaris 13 Carangidae jurel Caranx vinctus 14 Carangidae jurel Elagatis bipinnulata 15 Carangidae salmón Gnathonodon speciosus 16 Carangidae jurel Carcharhinus falciformis 17 Carcharhinidae tiburón Carcharhinus longimanus 18 Carcharhinidae tiburón Chaetodon humeralis 19 Chaetodontidae mariposa limón Johnrandallia nigrirostris 20 Chaetodontidae señorita Cirrhitus rivulatus 21 Cirrhitidae tigre Cirrihitichtys oxycephalus 22 Cirrhitidae tigre Diodon holocanthus 23 Diodontidae zorroespín Diodon hystrix 24 Diodontidae zorroespín Chaetodipterus zonatus 25 Ephippidae pañuelo Fistularia commersonii 26 Fistulariidae aguja Bathygobius andrei 27 Gobiidae mojarra Bathygobius ramosus 28 Gobiidae gobio Coryphopterus urospilus 29 Gobiidae gobio Microgobius miraflorensis 30 Gobiidae gobio Anisotremus caesius 31 Haemulidae ruco Anisotremus interruptus 32 Haemulidae melón Anisotremus taeniatus 33 Haemulidae melón Haemulon flaviguttatum 34 Haemulidae ruco Haemulon scudderi 35 Haemulidae ruco Haemulon sexfasciatum 36 Haemulidae ruco Haemulon steindachneri 37 Haemulidae naguilla Pomadasys macracanthus 38 Haemulidae ruco Pomadasys panamensis 39 Haemulidae ruco Biodiversity field inventories in selected watersheds 91 Nº Family Scientific Name 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 Holocentridae Holocentridae Kyphosidae Labridae Labridae Labridae Labridae Labridae Labrisomidae Labrisomidae Lutjanidae Lutjanidae Lutjanidae Lutjanidae Lutjanidae Lutjanidae Lutjanidae Lutjanidae Monacanthidae Mullidae Muraenidae Muraenidae Muraenidae Muraenidae Pomacanthidae Pomacanthidae Pomacanthidae Pomacentridae Pomacentridae Pomacentridae Pomacentridae Pomacentridae Pomacentridae Scaridae Scaridae Scaridae Scorpaenidae Serranidae Serranidae Serranidae Serranidae Serranidae Serranidae Signatidae Tetraodontidae Myripristis leiognathus Sargocentron suborbitalis Kyphosus elegans Bodianus diplotaenia Halichoeres dispilus Halichoeres nicholsi Halichoeres notospilus Thalassoma lucasanum Malacoctenus margaritae Malacoctenus tetranemus Hoplopagrus guentherii Lutjanus aratus Lutjanus argentiventris Lutjanus colorado Lutjanus guttatus Lutjanus novemfasciatus Lutjanus peru Lutjanus inermis Aluterus scriptus Mulloidichthys dentatus Echidna nebulosa Echidna nocturna Gymnothorax castaneus Gymnothorax dovii Chromis atrilobata Holacanthus passer Pomacanthus zonipectus Abudefduf concolor Abudefduf troschelii Microspathodon bairdii Microspathodon dorsalis Stegastes acapulcoensis Stegastes flavilatus Scarus compressus Scarus perrico Scarus rubroviolaceus Scorpaena mystes Alphestes immaculatus Cephalopholis panamensis Epinephelus analogus Paralabrax loro Paranthias colonus Serranus psittacinus Hippocampus ingens Arothron hispidus 92 Spanish Common Name salmonete salmonete chopa perico perico perico perico cortez sarado sarado sardo pargo pargueta pargo rojo lunarejo dentón guachinango pargo cuche salmonete culebra culebra morena morena castañuela muñeca rey ángel burra burra verde burra burra burrita burrita lora pez lora perico pez escorpión tigre señorita cabrilla cabrilla cabrilla loro caballo de mar globo Biodiversity field inventories in selected watersheds Nº Family Scientific Name 85 86 87 88 89 Tetraodontidae Tetraodontidae Tetraodontidae Tetraodontidae Tetraodontidae 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Carcharhinidae Clupeidae Coryphaenidae Dasyatidae Engraulidae Exocoetidae Fistulariidae Gerreidae Gobiesocidae Haemulidae Haemulidae Muraenidae Nematistiidae Ophichtidae Ophidiidae Rhincodontidae Sciaenidae Scombridae Scombridae Sphyraenidae Uranoscopidae 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Carangidae Carcharhinidae Carcharhinidae Carcharhinidae Echeneidae Ephippidae Gerreidae Gerreidae Hemiramphidae Kyphosidae Labridae Scaridae Serranidae Muraenidae Arothron meleagris Canthigaster punctatissima Sphoeroides angusticeps Sphoeroides annulatus Sphoeroides lobatus Marine Nasolamia velox Opisthonema libertate Coryphaena hippurus Dasyatis longa Anchoa mundeoloides Cheilopogon xenopterus Fistularia corneta Diapterus aureolus Arcos rhodospilus Microlepidotus brevipinnis Xenichthys xanti Gymnothorax equatorialis Nematistius pectoralis Myrophis vafer Lepophidium microlepis Rhincodon typus Larimus acclivis Euthynnus lineatus Scomberomorus sierra Sphyraena qenie Astroscopus zephyreus Marine–Reef associated Caranx sexfasciatus Carcharhinus leucas Carcharhinus limbatus Galeocerdo cuvier Echeneis naucrates Parapsettus panamensis Eucinostomus argenteus Gerres cinereus Hyporhamphus unifasciatus Sectator ocyurus Thalassoma grammaticum Scarus ghobban Rypticus nigripinnis Muraena lentiginosa Spanish Common Name globo globo tamboril tamboril tamboril Biodiversity field inventories in selected watersheds tiburón volador sardina boquita dorado raya sardina volador aguja iscanala calate ruco ruco morena gallo anguila congrio tiburon ballena pancha atún macarela picuda astrólogo jurel gambuso tiburón tiburón luna mojarra mojarra gorrión pichel cortez lora loro morena 93 Nº Family Scientific Name 15 16 17 18 Serranidae Serranidae Serranidae Sphyraenidae 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 Achiridae Achiridae Ariidae Batrachoididae Belonidae Belonidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carangidae Carcharhinidae Centropomidae Centropomidae Centropomidae Clupeidae Elopidae Engraulidae Engraulidae Engraulidae Engraulidae Gerreidae Gerreidae Gerreidae Gobiesocidae Gobiidae Haemulidae Hemiramphidae Labridae Labrisomidae Mugilidae Mugilidae Myliobatidae Narcinidae Paralichthyidae Polynemidae Alphestes multiguttatus Epinephelus itajara Epinephelus labriformis Sphyraena ensis Marine-Brackish (Estuarine) Achirus mazatlanus Achirus scutum Sciades seemanni Batrachoides waltersi Porichthys margaritatus Strongylura exilis Caranx caballus Caranx caninus Oligoplites altus Oligoplites saurus Oligoplites refulgens Selene brevoortii Selene orstedii Trachinotus paitensis Selene peruviana Carcharhinus porosus Centropomus armatus Centropomus medius Centropomus robalito Lile stolifera Elops affinis Anchoa lucida Anchoa panamensis Anchoa starksi Anchovia macrolepidota Diapterus peruvianus Eucinostomus currani Eucinostomus gracilis Gobiesox daedaleus Ctenogobius sagittula Haemulopsis leuciscus Hyporhamphus snyderi Halichoeres aestuaricola Malacoctenus zonifer Mugil cephalus Mugil curema Aetobatus narinari Narcine vermiculatus Citharichthys gilberti Polydactylus approximans 94 Spanish Common Name tigre mero mero picuda Biodiversity field inventories in selected watersheds lenguado lenguado bagre negro sapamiche sapamiche volador jurel jurel perro chucha chucha caballo papelillo palometa caballo frances punta robalo chichiguique aleta sardina macabilo sardina sardina sardina sardina bocona mojarra pampano mojarra calate guavina ruco fósforo perico trambollo lebriancha lisa gavilan raya eléctrica pez caite barbona Nº Family Scientific Name 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 Polynemidae Pristigasteridae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sciaenidae Sphyrnidae Stromateidae Synodontidae Synodontidae Urolophidae Urolophidae 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Anablepidae Ariidae Ariidae Ariidae Centropomidae Chanidae Eleotridae Eleotridae Engraulidae Gobiidae Gobiidae Gobiidae Lepisosteidae Mugilidae Poeciliidae Synbranchidae 1 2 3 4 5 6 7 8 9 10 Atherinopsidae Characidae Cichlidae Cichlidae Cichlidae Cichlidae Cichlidae Cichlidae Cichlidae Cichlidae Polydactylus opercularis Ilisha fuerthii Bairdiella ensifera Cynoscion reticulatus Cynoscion squamipinnis Cynoscion stolzmanni Micropogonias altipinnis Nebris occidentalis Odontoscion xanthops Paralonchurus dumerilii Stellifer crysoleuca Sphyrna lewini Peprilus snyderi Synodus evermanni Synodus scituliceps Urotrygon chilensis Urotrygon munda Freshwater–Brackish Anableps dowei Bagre panamensis Sciades troschelii Sciades guatemalensis Centropomus nigrescens Chanos chanos Dormitator latifrons Eleotris picta Anchoa spinifer Awaous banana Gobiomorus maculatus Gobionellus microdon Atractosteus tropicus Agonostomus monticola Poecilia sphenops Symbranchus marmoratus Freshwater Atherinella guatemalensis Astyanax fasciatus Amatitlania coatepeque Amatitlania nigrofasciata Amphilophus citrinellum Amphiolophus macracanthus Cichlasoma trimaculatum Parachromis managuense Parachromis motaguense Oreochromis niloticus niloticus Spanish Common Name barbona sardina pancha pancha rayada curvina curvina ratón guavina ratón pacún panchita tiburón martillo tilosa garrobo garrobo murciélago murciélago Biodiversity field inventories in selected watersheds cuatro ojos bagre tacazonte bagre robalo sábalo sambo ilama sardina talalais guavina luciernaga machorra tepemechín chimbolo anguila ejote plateada mojarra negra mojarra negra mojarra negra burra guapote guapote tigre guapote tigre tilapia 95 Nº Family Scientific Name 11 12 13 14 15 16 17 18 19 Cichlidae Gobiidae Heptapteridae Heptapteridae Poeciliidae Poeciliidae Profundulidae Profundulidae Sygnathidae Vieja guttalata Sicydium multipunctatum Rhamdia laticauda Rhamdia quelen Poecilia gillii Poeciliopsis gracilis Profundulus guatemalensis Profundulus punctatus Pseudophallus starksii Spanish Common Name burra culebra juilín juilín chimbolo bute chimbola chimbola anguila One species of special interest is the Tropical gar (Atractosteus tropicus) considered a living fossil, apparently confined to Zanjón del Chino, Department of Ahuachapán but occasionally it expands its distribution to the estuarine environments of Barra de Santiago (Solís 2006). Another is the native ―convict cichlid‖ Amatitlania coatepeque endemic to the Coatepeque Lake, considered until 2006 as Cichlasoma nigrofasciatum, but recent studies showed that this is a new species (Schmitter-Soto 2007). Other species have been introduced to diversify income and improve nutrition of the communities living near water bodies. We refer to the Nile tilapia (Oreochromis niloticus niloticus), guapote tigre (Parachromis managuense) and false yellowjacket cichlid (P. motaguense), recorded in the Cara Sucia and Coatepeque watersheds. Tilapia is from the African continent and the last two come from Nicaraguan lakes. ESTUARINE ECOSYSTEM The estuarine ecosystems, mainly those associated with mangrove ecosystems, are among the most productive in the world and are used by fish species for reproduction, feeding, and shelter (Day et al. 1989). Besides they function as biological corridors for movement and gene flow of freshwater and marine ichthyofauna (Yañez-Arancibia 1975). The present study reports 57 species recorded. Even though they are not strictly estuarine as a whole, the may occur frequently and they can even penetrate rivers with sufficient flow for short periods of time and during the highest tides. Schools of fish such as ―jurel‖ (Caranx caninus), ―mojarras‖ (Diapterus peruvianus and Eucinostomus currani), ―lisas‖ (Mugil curema), sardines and anchovies (Lile stolifera and Anchoa spp.), bass (Centropomus armatus, C. medius and C. robalito) can be frequently observed. Also representative of this ecosystem are the corvinas (Cynoscion reticulatus, C. phoxocephalus, C. squamipinnis, and C. stolzmani). MARINE ECOSYSTEM Given the continuity of the marine ecosystem habitats and the capacity of many species to move from one place to another, we recognize that the 39 species recorded only represents a small fraction of the species distributed in this habitat. These species are strictly marine, but can also be found in rocky environments and reefs. A wide variety of 96 Biodiversity field inventories in selected watersheds pelagic species of commercial importance occur in this habitat such as mackerel (Scomberomorus sierra), tuna (Euthynnus lineatus), and highly migratory species such as mahi-mahi or dolphinfish (Coryphaena hippurus). Consequently, fisheries productivity is very important in this zone, as shown in a study conducted to evaluate the by-catch ichthyofauna in industrial shrimping (Fuentes & Hernández 2004). This study showed that this coastal strip reported the highest volume of fish catch. Whale shark (Rhincodon typus), a species included on the IUCN Red List as Vulnerable, is also present. REEF ECOSYSTEM Coral reefs are considered among the most diverse and productive ecosystems in the world because their structure provides different habitats and microenvironments that facilitate the establishment of a complex food network. This complexity allows them to function as breeding, feeding, shelter, and reproduction zones for many marine organisms (Ruiz et al. 2003). Approximately 45 reef-forming coral species and approximately 25 km 2 of structural reefs are found in the TEP region. They are located in Baja California, Revillagigedos archipelago, Tres Marías Islands, Nayarit and the South Pacific Coast of Mexico, Los Cóbanos in El Salvador, Culebra Bay, Ballena Marine Park and Cocos Island in Costa Rica, as well as Coiba Island, Rey Island and San José in Panamá, and Galapagos Islands in Ecuador (Cortés 1997, Robertson & Allen 2008). The entire area is influenced by two zones where equatorial and temperate waters combine, bordering north with the California Current and south with the Peru Current (an extension of the Humboldt Current). The superficial current pattern of the zone varies bidirectionally, depending on the time of the year (Fisher et al. 1995). This condition turns the reef patches into strategic habitats that facilitate the dispersion of larvae of many coral, crustacean, echinoderm, mollusk, and fish species. It also intervenes in migratory processes, movement, and distribution of other fauna groups of regional and global importance, including marine turtles, whales, sharks, and tuna fish. The studies carried out in the reef zone of Los Cóbanos show records of 89 reef fish species and reef-associated fish species, even though the total number probably amounts to more than 170 species. We were able to identify 13 families directly depending on reef environments. Although some of them can sporadically penetrate the neighboring estuaries, they spend most of their life cycle in the reef. These families are: Acanthuridae, Balistidae, Cirrhitidae, Labridae, Labrisomidae, Pomacantidae, Pomacentridae, and Serranidae. Fishing this group is a significant means of support for the area’s coastal communities since th studies indicate that more than 800 artisanal fishermen benefit from the reef ecosystem (ICMARES 2006) and according to CENDEPESCA (2006), this group provided 23% of the total catch in the country for a total of U.S. $918,747. Biodiversity field inventories in selected watersheds 97 3.3. LEVEL OF INVENTORY COMPLETENESS The level of inventory completeness by watershed shows very low percentages with marked differences among them. Coatepeque reports the highest percentage, 32%; Los Cóbanos, 29%; Barra de Santiago, 24%; and Cara Sucia and Cauta report 12% and 5%, respectively, while the lowest values are reported by Grande de Sonsonate, 0.4%, and Barra Salada, 0.2%. The remaining watersheds, Bocana de San Juan, Izalco, and San Julián do not have any records (Table 21). TABLE 21. LEVEL OF FISH INVENTORY COMPLETENESS BY WATERSHED IN SOUTHWESTERN EL SALVADOR. Watershed Species Recorded Species Expected Level of Inventory Completeness (%) Los Cóbanos Barra de Santiago Barra Salada Cara Sucia Bocana de San Juan Cauta Grande de Sonsonate San Pedro Coatepeque Izalco San Julián 156 114 1 58 0 25 2 26 10 0 0 536 483 483 483 483 483 483 483 31 26 26 29 24 0 12 0 5 0 5 32 0 0 The inventory completeness by municipality is also very low since only 7 of the 25 municipalities in the study area presented records, illustrating the sparse coverage of prior research in the study area. The highest percentages were reported by El Congo, 32%; Acajutla, 27%; Jujutla, 24%; Tacuba and San Francisco Menéndez, 22% and 19%, respectively; while the lowest percentages were reported in Sonsonate and Chalchuapa reaching 8% each (Table 22). Given the spatial arrangement of the ecosystems, it is important to coordinate municipal efforts to improve land use planning. Knowing the completeness of the species inventory for a given ecosystem provides valuable information about its potential for maintaining biodiversity and presents an opportunity to evaluate the ecological functionality of the taxonomic groups and species in general with the purpose of identifying those with the highest relevance and providing guidance for conservation efforts. In this regard, the highest percentages for inventory completeness were reported for the Freshwater ecosystem, 62%; the Reef ecosystem reached 55%; and the Estuarine ecosystem, 56%. The lowest percentage was reported for the Marine ecosystem with only 17% (Table 23). 98 Biodiversity field inventories in selected watersheds TABLE 22. LEVEL OF FISH INVENTORY COMPLETENESS BY MUNICIPALITY IN SOUTHWESTERN EL SALVADOR. Species Species Level of Inventory Municipality Recorded Expected Completeness (%) Acajutla 142 529 27 Jujutla 118 485 24 San Francisco Menéndez 57 485 12 Sonsonate 41 529 8 El Congo 10 31 32 Tacuba 5 26 19 Chalchuapa 2 26 8 Apaneca 0 26 0 Armenia 0 26 0 Caluco 0 26 0 Concepción de Ataco 0 26 0 Cuisnahuat 0 26 0 Guaymango 0 26 0 Izalco 0 26 0 Juayúa 0 26 0 Nahuizalco 0 26 0 Nahulingo 0 26 0 Salcoatitán 0 26 0 San Antonio del Monte 0 26 0 San Julián 0 26 0 San Pedro Puxtla 0 26 0 Santa Catarina Masahuat 0 26 0 Santa Isabel Ishuatán 0 26 0 Santo Domingo de Guzmán 0 26 0 Sonzacate 0 26 0 Biodiversity field inventories in selected watersheds 99 TABLE 23. LEVEL OF FISH INVENTORY COMPLETENESS BY HABITAT IN SOUTHWESTERN EL SALVADOR. Ecosystem Species Recorded Total Indicator Species Level of Inventory Completeness (%) Freshwater Estuarine Marine Reef Total 35 57 39 89 220 56 102 222 161 541 63 56 18 55 41 3.4. IMPORTANT SPECIES FOR CONSERVATION The analysis of the information indicates that 51 species are important for conservation, of which the native freshwater species such as ―mojarra‖ Amatitlania coatepeque, a species endemic to Lake Coatepeque (Schmitter-Soto 2007) and the only vertebrate species restricted to El Salvador, have the highest ecological importance. Likewise, the tropical gar Atractosteus tropicus is unique because it represents a group of fish already extinct. The freshwater mullet Agonostomus monticola is also important because of it’s role in the diet of the local communities along the rivers where it dwells; it reaches up to 40 cm in length. This species lives in mountain rivers when it reaches maturity, but the larvae grow in coastal waters and estuaries. This species could be used as a bioindicator of environmental health. Other freshwater species such as ―plateada‖ (Astyanax fasciatus), ―juilín‖ (Rhamdia quelen), ―bute‖ (Poeciliopsis gracilis), ―mojarra negra‖ (Amphiolophus macracanthus), are also included owing to the deterioration experienced by their populations because they are still extracted without any fishing regulation. As a result, they were defined as vulnerable in the study carried out by PREPAC (2005). Other species important for conservation are strictly reef-dwelling, such as ―cirujanos‖ (Prionurus punctatus, Acanthurus xanthopterus), ―peces tigre‖ (Cirrithus rivulatus) and (Cirrhitichthys oxycephalus). The shark (Carcharhinus longimanus), hammerhead shark (Sphyrna lewini), whale shark (Rhincodon typus), and grouper (Epinephelus itajara), are included on the IUCN Red List as globally threatened. All snapper species recorded in the area have also been included as a priority for conservation since they have high value in the local market and hence generate income and provide food for more than 21 fishermen communities (Galdámez 2002, Rojas et al. 2004, CENDEPESCA 2006), but their extraction is not regulated in terms of catch volumes and commercialization sizes. The following species are included: Lutjanus aratus, L. argentiventris, L. colorado, L. guttatus, L. novemfasciatus, and L. peru. 100 Biodiversity field inventories in selected watersheds TABLE 24. CONSERVATION-IMPORTANT FISH IN SOUTHWESTERN EL SALVADOR. Spanish Nº Family Scientific Name Common Categories* Ecosystem Name Acanthurus 1 Acanthuridae cirujano E,1 Reef xanthopterus Prionurus punctatus 2 Acanthuridae cirujano E,1 Reef Carcharhinus 3 Carcharhinidae tiburón punta F Marine falciformis Marine 4 Carcharhinidae Carcharhinus leucas tiburón F,2,3 Carcharhinus Marine 5 Carcharhinidae tiburón F,2 limbatus Carcharhinus 6 Carcharhinidae tiburón A1; F,2 Marine Reef longimanus Carcharhinus 7 Carcharhinidae tiburón F Marine porosus Reef 8 Chaetodontidae Chaetodon humeralis mariposa E,1; F Johnrandallia Reef 9 Chaetodontidae mariposa E,1; F nigrirostris Astyanax fasciatus 10 Characidae plateada B; F,4 Freshwater Amatitlania Freshwater 11 Cichlidae mojarra C coatepeque Amatitlania Freshwater 12 Cichlidae chiva F,3 nigrofasciata Amphilophus Freshwater 13 Cichlidae mojarra F,3 macracanthus Freshwater Vieja guttulata 14 Cichlidae burrita F,1,3 Cirrhitichthys Reef 15 Cirrhitidae rayado E,1 oxycephalus Reef Cirrhitus rivulatus 16 Cirrhitidae tigre E,1 Dasyatis longa 17 Dasyatidae raya F,4 Marine Eleotris picta 18 Eleotridae ilama B Freshwater Freshwater19 Lepisosteidae Atractosteus tropicus machorra D Estuarine Lutjanus aratus 22 Lutjanidae pargo F,2 Reef Lutjanus 23 Lutjanidae pargueta F,1 Reef argentiventris Lutjanus colorado 24 Lutjanidae pargo rojo F,2 Reef Lutjanus guttatus 25 Lutjanidae lunarejo F,1,2 Reef Lutjanus 26 Lutjanidae pargo negro F,2 Reef novemfasciatus Lutjanus peru 27 Lutjanidae guachinango F,2 Reef Aetobatus narinari 28 Myliobatidae gavilán A,3; F,2 Marine Agonostomus 29 Mugilidae tepemechín B; F Estuarine monticola Poecilia sphenops 30 Poecilidae chimbolo B; F,3 Freshwater Poeciliopsis gracilis 31 Poecilidae chimbolo B; F,3 Freshwater Biodiversity field inventories in selected watersheds 101 Nº Family 32 Pomacanthidae 33 Pomacanthidae 34 Pomacentridae 35 Pomacentridae 36 Profundulidae 37 Profundulidae 38 Rhincodontidae Scientific Name Holocanthus passer Microspathodon bairdii Microspathodon dorsalis Pomacanthus zonipectus Profundulus guatemalensis Profundulus punctatus Rhincodon typus Spanish Common Name sarado Categories* Ecosystem E,1; F Reef Reef frentudo E frentudo E ángel E,1; F chimbola B; F,3 chimbola B, F3 tiburón ballena lora perico A,1; E,2 Reef Reef Freshwater Freshwater Marine Reef Scarus compressus E,3; F Reef Scarus perrico E,3 Scarus Reef 41 Scaridae lora E,3 rubroviolaceus Alphestes Reef 42 Serranidae tigre E,1 inmacularus Epinephelus Reef 43 Serranidae cabrilla E,2 analogus Reef Epinephelus itajara 44 Serranidae mero A,2; E,2 Epinephelus Reef 45 Serranidae cabrilla E,2 labriformis Epinephelus Reef 46 Serranidae mero E,2 niphobles Cephalopholis Reef 47 Serranidae mero A,2; E,2 panamensis Reef Paranthias colonus 48 Serranidae E,1 Reef Sphyrna lewini 49 Sphyrnidae charruda F Marine Urotrygon chilensis 50 Urotrygonidae raya F,4 Marine Urotrygon munda 51 Urotrygonidae raya F,4 *Criteria and justification for classifying species as important to conservation: A) Included in the IUCN Red List (2009): 1-Vulnerable at the global level, 2-Critically Endangered at the global level, 3-Near Threatened. B) Threatened freshwater species or environmental health indicator in El Salvador. C) Endemic to Lake Coatepeque, El Salvador. D) Living fossil. E) Scuba diving tourism: 1-and aquarium; 2-and fishery importance; 3-and substrateforming. F) Fishery or food importance: 1-and aquaculture; 2- and sport fishing; 3-and aquariums; 4-low importance 39 40 102 Scaridae Scaridae Biodiversity field inventories in selected watersheds 4. DISCUSSION The southwestern region of El Salvador is characterized by a complex network of aquatic systems that provide distinctive habitats, some of which function as migration corridors for the local and regional ichthyofauna. Recorded species richness is 220 species, which is relatively high when compared with other regions of the country. For example, Jiquilisco Bay, Jaltepeque Estuary, and the Gulf of Fonseca have reported totals of 56, 138, and 116 species, respectively (Calderón & Hernández 1974, Fuentes & Hernández 2004, Chicas et al. in prep.). Notwithstanding, the number of coastal fish species for this region is at least 541 (MARN 2003, Robertson & Allen 2008) and the inventory is southwestern El Salvador is only about 40% complete. The difference in the number of species is probably due to incomplete sampling. Not only is the study area highly diverse and important for maintaining biodiversity, but it also features a speciation site, because a ―mojarra‖ (Amatitlania coatepeque) is endemic to Lake Coatepeque. Mesoamerica is an important place for the evolution of certain groups of freshwater fish, particularly the family Cichlidae (Cocheiro Pérez et al. 2006). Special efforts should be made to characterize the ecology and population dynamics of this species in order to establish a sound and effective conservation program. Another conservation priority is the Tropical gar (Atractosteus tropicus), considered to be a living fossil (Mora et. al. 1997, Sangado-Maldonado et al. 2004), which has three separate populations in the Mesoamerican region, two of which are located in the Atlantic slope and one in the Pacific slope. The first of them can be found in the area from the Usumacinta River to the Coatzocoalcos River in southern Mexico and Guatemala; the other, further south in Lake Nicaragua and Río San Juan. On the Pacific slope, the population is distributed in a strip from the south of Chiapas to Río Negro in Nicaragua (Bussing 1987), and Froese & Pauly (2009) mention 5 reports for Costa Rica. The only place where the species can be found in El Salvador is Zanjón del Chino and adjoining Barra de Santiago (Solís 2006). Another important species is the freshwater mullet (Agonostomus monticola) in view of its important contribution to the local population’s nutrition (Phillips 1983, Cruz et. al. 1987), but lack of protection is decimating the fish population. Most records are from El Imposible National Park. On the other hand, freshwater ―mojarra‖ (Amphiolophus macracanthus, A. citrinellum and Vieja guttalata), ―guapotes‖ (Parachromis managuense, P. motaguense and Cichlasoma trimaculatum), among others, which occur in most of the lakes and ponds and are of nutritional importance, and indicators of environmental health such as ―bute‖ Poecilia sphenops, ―plateada‖ Astyanax fasciatus, ―burra‖ Amatitlania nigrofasciata, among others, are important because they are considered in need of critical conservation efforts (PREPAC 2005), due to population decreases. Los Cóbanos Reef System is a very particular case because coral reefs are considered among the most productive ecosystems of the world, providing habitat, feeding areas, Biodiversity field inventories in selected watersheds 103 shelter, and spawning areas for most coastal fish. As a consequence, they can sustain a high diversity of species such as ―burras‖ (Abudefduf concolor, A. troschelii, Stegastes acapulcoensis), ―cirujanos‖ (Prionurus punctatus, Acanthurus xanthopterus), ―pez tigre‖ (Cirritos rivulatus), groupers (Alphestes inmaculatus and Epinephelus itajara), ―señoritas‖ (Chaetodon humeralis and Johnrandallia nigrirostris), among others. The grouper recorded, Epinephelus itajara, and the sharks Carcharhinus longimanus, C. leucas, C. porosus, and C. limbatus, are species characterized by low fertility and late reproduction (Baum et al. 2003, Bishop et al. 2006), making them vulnerable to overexploitation. According to FAO (2009), the shark harvest in El Salvador decreased from 1,202 t in 2005 to 715 t in 2007. This problem hinders their recovery capacity, placing them in the threatened categories of the IUCN Red List (2008). 5. CONCLUSIONS The important sites for conservation are Los Cóbanos Reef System and Barra de Santiago estuary as well as Lake Coatepeque and the fluvial system because they function as bridges that facilitate the local and regional migration of the ichthyofauna of the ChiapasNicaraguan province. Los Cóbanos is the only major reef formation in the country and Barra de Santiago supports important mangrove forest coverage. For both cases, Art. 74 of the Environmental Law (MARN 2006) refers to mangroves and reefs as fragile ecosystems and therefore no disturbance whatsoever is allowed. This fragility is based on their functionality as feeding, shelter and reproduction zones for the fish species and most marine taxonomic groups. The high productivity of these ecosystems maintains a wide variety of species and contributes to the economic support and as a food source of the local population. The beautiful scenery can be used to promote environmentally sustainable tourism. The functional complexity of those ecosystems and anthropogenic damage also render them vulnerable, and for that reason, it is urgent to establish conservation measures to make them sustainable. The sparse coverage of research conducted in the zone explains why the level of fish inventory completeness in southwestern El Salvador is just 40%. The best inventoried watershed within the study area, Lake Coatepeque, probably has only 32% of its species identified. Three watersheds present no fish records at all. Only seven municipalities in the study area present fish records. The southwestern region of El Salvador is important for biodiversity maintenance and development of evolutionary speciation processes. There are records of 220 species, one of them is endemic to Lake Coatepeque (Amatitlania coatepeque) and another, the Tropical gar (Atractosteus tropics), represents a group of extinct fish species, and is restricted in El Salvador to Zanjón del Chino and Barra de Santiago, Ahuachapán. The location of El Salvador allows it to function as a native ichthyologic fauna reservoir, a condition that facilitates genetic material exchange among the populations distributed from the south of Mexico to the north of Costa Rica. Additionally, the native and exotic ichthyofauna is very important as a source of food and income for the local population. 104 Biodiversity field inventories in selected watersheds 6. RECOMMENDATIONS Recommendations for improved ichthyofauna management in the watersheds of southwestern El Salvador: 1. Conduct field research on the distribution and abundance of the ichthyofauna of the area with emphasis on watersheds with few records in order to increase the level of inventory completeness. The studies should include molecular markers to establish phylogenetic relationships and distributional ranges, and the nation-wide update of species taxonomy. 2. Conduct ichthyoplancton studies, reproductive and feeding studies of the fish populations associated with the aquatic ecosystems such as estuaries, reefs, and adjacent marine waters in order to identify the areas used for spawning, reproduction, and feeding. This will elucidate species interactions and contribute to applying management and conservation strategies with an ecosystem focus. 3. The conservation programs of important sites for biodiversity management, mainly Los Cóbanos Complex and Barra de Santiago, should be strengthened, since these sites present a diversity of aquatic environments that maintain a wide variety of species and facilitate migration processes. Therefore, specific research aimed at key groups of the system, such as fish, is critical to the implementation of management alternatives with a biological foundation. 4. The ecosystemic functionality of the fluvial system made up of the river watersheds of San Julián, San Pedro, Cara Sucia, Bocana de San Juan, Cauta, Grande de Sonsonate, and Barra Salada, which as a whole form a biological corridor that facilitates local migration and provides a habitat for the species, should be conserved. 5. Establish a comprehensive watershed management program for Lake Coatepeque and other lakes in the study area since they provide habitat to native freshwater fish species, which are important food sources for local communities. Biodiversity field inventories in selected watersheds 105 CHAPTER 4: INVENTORY OF AMPHIBIANS AND REPTILES IN SOUTHWESTERN EL SALVADOR Vladlen Henríquez & Xiomara Henríquez Conservation Science Department, SalvaNATURA San Salvador, El Salvador vhenriquez@salvanatura.org 1. INTRODUCTION The southwestern region of El Salvador is a mosaic of heterogeneous vegetation communities surrounded by agricultural landscapes. The area presents a wide variety of rivers and a few lakes that together with the different types of vegetation form ideal habitats for many amphibians and reptiles (herpetofauna). However, there is not enough information about many of these sites with regard to which and how many herpetofauna species are present, which makes it difficult to make decisions and take actions concerning biodiversity protection and conservation of these zones. In El Salvador, 132 herpetofauna species have been detected so far (Köhler et al. 2006, Herrera et al. 2007, Henríquez & Vaquerano 2008). As of 2006, 91 species were on record in the area of the Improved Management and Conservation of Critical Watersheds (IMCW) Project. The zone is formed by eleven watersheds in addition to several natural areas that function as refuges for threatened or endangered herpetofauna species. Herpetofauna research in this area has been conducted mainly at El Imposible and Los Volcanes National Parks, where national and foreign researchers have made several collecting expeditions, as recently as 2006 (Henríquez 2004, Henríquez & Komar 2006, Köhler et al. 2006). The main objectives of this study were: to determine species of interest for conservation in 11 watersheds of the southwestern region of El Salvador; to fill information gaps regarding the presence and status of the herpetofauna populations in the municipalities and watersheds that form the IMCW Project area; and to use the results as a decision-making tool for biodiversity protection and conservation. 106 Biodiversity field inventories in selected watersheds 2. METHODOLOGY 2.1. STUDY AREA The study was conducted in seven of the 11 watersheds in the southwestern region of El Salvador, which correspond to 11 of the 25 municipalities distributed among the Departments of Ahuachapán, Sonsonate, Santa Ana and La Libertad that make up the area of the MMCC Project. Fig. 9 and Table 25 show a detail of the sites where the herpetofauna studies were carried out. 3 1 2 8 4 5 6 7 FIGURE 9. MAP OF THE SAMPLING SITES OF THE STUDY OF AMPHIBIANS AND REPTILES DURING 2007: (1) EL IMPOSIBLE NATIONAL PARK, (2) LAGUNA DE LAS NINFAS, (3) LOS ANDES SECTOR, LOS VOLCANES NATIONAL PARK, (4) LAVAS DE IZALCO/FINCA MARÍA AUXILIADORA (LOS VOLCANES NATIONAL PARK), (5) SAN MARCELINO COMPLEX, (6) PLAN DE AMAYO, (7) LOS CÓBANOS, (8) SANTA RITA FOREST. Biodiversity field inventories in selected watersheds 107 TABLE 25. DESCRIPTION OF THE SAMPLING SITES AND EFFORT FOR THE HERPETOFAUNA STUDY. Study Site El Imposible National Park El Imposible National Park 108 Watershed Barra de Santiago Barra de Santiago Los Volcanes National Park Izalco Los Volcanes National Park Santa Ana Geographic Location Cerro Campana Sector, Municipality of Tacuba, between the cantones of El Sincuyo and La Montaña (13.86° N; 89.91°O), Dept. of Ahuachapán San Francisco Menéndez Sector, Municipality of San Francisco Menéndez, between the cantones of El Corozo and El Sacramento (13.84°N; 90.01°O), Dept. of Ahuachapán María Auxiliadora Sector, Municipality of Izalco, between the cantones of Chorro Abajo, Chorro Arriba, Shonshón, Teshcal, Las Marías, La Quebrada Española and Cuyagualo (13.89°N; 89.63°O), Dept. of Sonsonate Los Andes Sector, Municipalities of Santa Description Date Days Hours of Transects June 25 to July 4 and July 10 to 19, 2007 20 50 July 27 to August 05 and August 01 to October 10, 2007 20 69 Undisturbed very humid mountain forest. Evergreen riparian broadleaf forest with anthropogenic disturbance as there is a community within the area Lava flows that have different succesional stages and remnants of Tropical Dry Forest. Very humid montane forest with Biodiversity field inventories in selected watersheds Study Site Santa Rita y Zanjón El Chino Protected Natural Area Plan de Amayo Protected Natural Area San Marcelino Complex Protected Natural Area Watershed Geographic Location Description Ana and Chalchuapa, between the cantones of Palo de Campana, Potrero Grande Arriba and Buenos Aires (13.86°N; 89.63°O), Dept. of Santa Ana some sites affected by the Santa Ana volcano eruption in 2005 and other sites with Cypress (Cupressus lusitanicus) plantations. Cara Sucia Municipality of San Francisco Menéndez, between the cantones of Cara Sucia, La Hachadura and Garita Palmera (13.80°N; 90.06°O), Dept. of Ahuachapán San Julián Municipality of Caluco, between the cantones of Plan de Amayo and El Castaño, Dept. of Sonsonate Cliff vegetation, tropical dry forest (in the parts closest to the cliff) and moist tropical forest (in the lower parts) Lago de Coatepeque and Izalco Municipalities of Izalco and Armenia, between the cantones of Las Lajas, El Guayabo, Los Mangos, El Rosario, San Isidro and Successional vegetation on volcanic lava of different ages, Dry Tropical Dry Forest Tropical evergreen floodplain forest, surrounded by intensive agricultural activity. Biodiversity field inventories in selected watersheds Date Days Hours of Transects August 08 to 17 and October 13 to 22, 2007 20 63 August 22 to 31 and October 26 to November 04, 2007 20 64 September 18 to 27 and December 03 to 12, 2007 20 77 109 Study Site Watershed Geographic Location Description Las Marías, Departament of Sonsonate. Municipalities of Santa Ana and El Congo, between the cantones of Lomas de San Marcelino, Montebello, El Rodeo and La Presa, Dept. of Santa Ana (13.69°N; 89.64°O) (deciduous and subdeciduous). Laguna de Las Ninfas Protected Natural Area Río Grande de Sonsonate Municipality of Apaneca (13.87°N; 89.80°O), Dept. of Ahuachapán Los Cóbanos Marine Park Protected Natural Area* Los Cóbanos Municipality of Acajutla, Dept. of Sonsonate (13°12´N; 89°30´0) Forested zone around the lake consists of montane humid forest with an area of 80 mz. Rocky beach with volcanic origens as well as mangroves, rivers, estuaries, and agriculture fields. *Data collected by Karla Lara Sandoval. 110 Biodiversity field inventories in selected watersheds Date Days Hours of Transects September 08 to 17 and November 20 to 29, 2007 20 72 August 20 to 29, 2007 10 15 2.2. SAMPLING DESIGN Extensive searches were conducted based on transects with a duration of 1 hour, separated by 15 minutes between each one, without repeating transects. The field work was carried out during 120 field days (June to December 2007) with sampling schedules from 8:00 to 13:00 hours and from 20:00 to 22:00 hours. The study included the rainy season, the transition from the rainy to the dry season, and the beginning of the dry season. The sampling sites were selected based on conditions for higher probability of finding amphibians and reptiles. For reptiles, we selected exposed sites with trunks and shrubs useful for hiding; for amphibians, very humid places, swampy areas in pastures or inside forests as well as small pools in ravines and streams. This report includes observations of amphibians and reptiles by other field teams working at the protected natural areas of Los Cóbanos Marine Park, Laguna de las Ranas, and Barra Salada. It also includes species identified through photographs taken by other researchers at sites where herpetofauna studies were not conducted. Table 25 shows a detail of the sampling effort and the number of species recorded in each natural area. 2.3. LEVEL OF INVENTORY COMPLETENESS In order to analyze progress or completeness of the inventories in the municipalities and watersheds, we first prepared a list of species indicative of a complete inventory (a method adapted from Gómez de Silva & Medellín 2001). This list usually consists of generalist species which are those that live in a variety of habitats and eat a variety of food types, and are found in both disturbed and undisturbed habitats (for example, the black iguana Ctenosaura similis). We prepared lists of expected generalist species for each municipality, watershed, natural ecosystem, and protected natural area. We compared the species recorded during the study to the list of expected generalist species in order to obtain an inventory completeness index score for each site (Table 26). The index score is the percentage of expected species that have been recorded; the number of expected species recorded is divided by the total number of expected generalist species, and multiplied by one hundred. A common list of 46 expected species was determined for watersheds and municipalities; however, specific lists of indicator species for inventory completeness were determined for each ecosystem and natural area and they differ from one ecosystem or site to another. The reason why each habitat has different indicator species for inventory completeness is that some generalist species of the low and medium elevational zones do not live in the higher zones. Also, we used species accumulation curves to determine the degree of inventory completeness. Data from the curves were used to generate species richness estimates based on ACE (Abundance-based Coverage Estimator), ICE (Incidence-based Coverage Estimator), Chao 1, Chao 2, Jack-Knife 1 and Jack-Knife 2 indices, calculated with the program EstimateS® version 7.51 (Colwell 2006). Biodiversity field inventories in selected watersheds 111 TABLE 26. INDICATOR SPECIES FOR INVENTORY COMPLETENESS. Species HF DF FE CF S RIZ Dermophis x x x x mexicanus Craugastor x x rhodopis Craugastor x x x rupinius Rhinella marina x x x x x Incilius coccifer x x x x Incilius luetkenii x x x x Agalychnis x x moreletii Scinax staufferi x x x x Smilisca baudinii x x x x Trachycephalus x x x x venulosus Leptodactylus x x x x fragilis Leptodactylus x x x x melanonotus Engystomops x x x x pustulosus Lithobates forreri x x x x Lithobates x x x maculatus Crocodylus acutus x Caiman crocodilus x Chelonia mydas x Eretmochelys x imbricata Lepidochelys x olivacea Dermochelys x coriacea Gonatodes x x x albogularis Phyllodactylus x x x tuberculosus Basiliscus vittatus x x x Ctenosaura similis x x x x Iguana iguana x x x Norops crassulus x Norops x x macrophallus Norops sericeus x x x x x Norops serranoi x x x Sceloporus x x x malachiticus Sceloporus x x x x x squamosus 112 Biodiversity field inventories in selected watersheds E CR x x x x x x x x x x x x Species HF DF FE CF S RIZ E CR Mabuya x x x unimarginata Ameiva undulata x x x x x Aspidoscelis x x x x x deppii Boa constrictor x x x x x Conophis concolor x x x x x x Drymobius x x margaritiferus Drymarchon x x x x melanurus Leptodeira x x x septentrionalis Masticophis x x x x x mentovarius Mastigodryas x dorsalis Ninia sebae x x Oxybelis aeneus x x x x x Oxybelis fulgidus x Scolecophis x x atrocinctus Senticolis triaspis x x x Spilotes pullatus x x x x Stenorrhina x x x x freminvilli Trimorphodon x x x x biscutatus Micrurus x x x x x nigrocinctus Cerrophidion x godmani Crotalus simus x x x HF: Humid forest; DF: Dry forest; FE: Freshwater Ecosystems; CF: Cloud Forest; S: Savanna; RIZ: Rocky Intertidal Zone; E: Estuaries; CR: Coral Reefs Biodiversity field inventories in selected watersheds 113 2.4. SELECTION OF SPECIES CONSIDERED IMPORTANT FOR CONSERVATION The main criterion to select species important for conservation is whether the species is included on the Red List of Endangered Species (UICN 2008) or if it is considered a threatened species in the country according to MARN (2004) or Greenbaum & Komar (2005) (Table 27). 2.5. DATA SOURCES Most of the information on the species recorded in the project zone comes from the field stage of the study; however, these data have been complemented with information already existing from each Protected Natural Area and other parts of the study area. The bibliographic information, summarized in Komar 2007, comes from books (Leenders 2003, Köhler et al. 2006) and reports from consultancies carried out in the different Protected Natural Areas (Rivera Muñoz 2000, Paz Quevedo & Ventura Centeno 2003, Herrera & Valle Andrade 2003, Henríquez & Komar 2006) as well as from other publications (Herrera et al. 2005, Herrera et al. 2007). 3. RESULTS AND DISCUSSION 3.1. SPECIES RECORDED The present study has recorded 74 amphibian and reptile species, bringing the total recorded for the IMCW project area to 96 species, including 91 species recorded in previous studies. The 22 amphibian and 74 reptile species represent 73% of the 132 species recorded in El Salvador (Table 28). None of the recorded species has a distribution that is fully restricted to El Salvador, however, in the country two species have only been recorded in the project area. They are the lizard Celestus atitlanensis and the toad Incilius valliceps. Some of the species recorded are restricted in El Salvador to the western region of the country (including some regions outside of the study area), such as the Keeled Helmeted Basilisk (Corytophanes percarinatus), the anole lizard Norops crassulus, and the Godman’s Montane Pitviper (Cerrophidion godmani). Other species found present few records in the country, such as the burrowing toad (Rhinophrynus dorsalis), freshwater turtles Trachemys venusta and Staurotypus salvinii, the American Crocodile (Crocodylus acutus), the Spectacled Cayman (Caiman crocodilus), a skink (Mesoscincus managuae), and several snakes (Mastigodryas melanolomus, Imantodes gemmistratus, Leptodrymus pulcherrimus, Rhadinaea pilonaorum, Scaphiodontophis annulatus, Scolecophis atrocinctus, Sibon nebulatus, Tantilla taeniata, Tropidodipsas fischeri, Loxocemus bicolor and Atropoides nummifer). Four species of sea turtles are known in the study area: the Green Sea Turtle (Chelonia mydas), Hawksbill Sea Turtle (Eretmochelys imbricate), Olive Ridley Sea Turtle (Lepidochelys olivacea), and Leatherback Sea Turtle (Dermochelys coriacea); all are endangered species worldwide. 114 Biodiversity field inventories in selected watersheds TABLE 27. LIST OF INDICATOR SPECIES FOR CONSERVATION IMPORTANT SITES. National Status (according National Status Family Species to Greenbaum & Komar (according to MARN 2004) 2005) Dermophis mexicanus Caeciliidae Not Threatened Least Concern Oedipina taylori Plethodontidae Threatened Vulnerable Incilius canaliferus Not Threatened Endangered Bufonidae Incilius valliceps Endangered Critically Endangered Agalychnis moreletii Not Threatened Endangered Dendropsophus Hylidae Threatened Vulnerable robertmertensi Trachycephalus venulosus Not Threatened Vulnerable Gastrophryne usta Threatened Vulnerable Microhylidae Hypopachus barberii Threatened Endangered Rhinophrynus dorsalis Rhinophrynidae Threatened Endangered Chelonia mydas Endangered Vulnerable Eretmochelys imbricata Cheloniidae Endangered Vulnerable Lepidochelys olivacea Endangered Endangered Dermochelys coriacea Dermochelydae Endangered Critically Endangered Trachemys venusta Emydidae Endangered Endangered Staurotypus salvinii Kinosternidae Threatened Endangered Caiman crocodilus Alligatoridae Endangered Critically Endangered Crocodylus acutus Crocodylidae Endangered Vulnerable Celestus atitlanensis Anguidae Threatened Endangered Coleonyx mitratus Eublepharidae Not Threatened Endangered Corytophanes percarinatus Threatened Vulnerable Iguanidae Iguana iguana Threatened Least Concern Scincidae Colubridae Mesoscincus managuae Coniophanes fissidens Not Threatened Not Threatened Endangered Vulnerable Biodiversity field inventories in selected watersheds Global Status (according to IUCN 2008) Vulnerable Data Deficient Least Concern Least Concern Critically Endangered Least Concern Least Concern Least Concern Endangered Least Concern Endangered Critically Endangered Endangered Critically Endangered Near-threatened Near-threatened Near-threatened Vulnerable Least Concern Least Concern Least Concern Least Concern Least Concern Not Evaluated 115 Family Colubridae Elapidae Viperidae 116 Not Threatened Not Threatened Not Threatened Threatened Not Threatened Endangered Threatened Not Threatened Threatened Threatened Threatened National Status (according to Greenbaum & Komar 2005) Endangered Vulnerable Vulnerable Vulnerable Endangered Critically Endangered Endangered Vulnerable Vulnerable Endangered Endangered Threatened Endangered Not Evaluated Not Threatened Not Threatened Threatened Threatened Not Threatened Threatened Not Threatened Threatened Threatened Not Threatened Threatened Threatened Threatened Threatened Endangered Vulnerable Vulnerable Endangered Vulnerable Endangered Vulnerable Endangered Vulnerable Vulnerable Endangered Endangered Endangered Least Concern Not Evaluated Not Evaluated Not Evaluated Not Evaluated Not Evaluated Not Evaluated Not Evaluated Least Concern Not Evaluated Not Evaluated Not Evaluated Near-threatened Not Evaluated Not Evaluated National Status (according to MARN 2004) Species Coniophanes piceivittis Drymarchon melanurus Imantodes gemmistratus Lampropeltis triangulum Leptodeira nigrofasciata Leptodrymus pulcherrimus Mastigodryas melanolomus Oxybelis fulgidus Pliocercus elapoides Rhadinaea godmani Rhadinaea pilonaorum Scaphiodontophis annulatus Scolecophis atrocinctus Senticolis triaspis Sibon anthracops Sibon nebulatus Spilotes pullatus Tantilla taeniata Trimorphodon biscutatus Tropidodipsas fischeri Micrurus nigrocinctus Pelamis platurus Atropoides nummifer Agkistrodon bilineatus Cerrophidion godmani Crotalus simus Biodiversity field inventories in selected watersheds Global Status (according to IUCN 2008) Least Concern Least Concern Not Evaluated Not Evaluated Least Concern Not Evaluated Least Concern Not Evaluated Not Evaluated Not Evaluated Not Evaluated TABLE 28. HERPETOFAUNA SPECIES RECORDED IN THE STUDY AREA. Order Family Species Dermophis mexicanus Gymnophiona Caeciliidae Oedipina taylori Plethodontidae Craugastor rhodopis Craugastoridae Craugastor rupinius Rhinella marina Incilius canaliferus Incilius coccifer Bufonidae Incilius luetkenii Incilius valliceps Agalychnis moreletii Dendropsophus robertmertensi Scinax staufferi Anura Hylidae Smilisca baudinii Trachycephalus venulosus Engystomops pustulosus Leiuperidae Leptodactylus fragilis Leptodactylidae Leptodactylus melanonotus Gastrophryne usta Microhylidae Hypopachus barberi* Lithobates forreri Ranidae Lithobates maculatus Rhinophrynus dorsalis Rhinophrynidae Caiman crocodilus Alligatoridae Crocodylia Crocodylus acutus* Crocodylidae Chelonia mydas* Eretmochelys imbricata Cheloniidae Lepidochelys olivacea* Dermochelys coriacea* Dermochelydae Testudines Trachemys venusta Emydidae Rhinoclemmys pulcherrima Geoemydidae Kinosternon scorpioides Kinosternidae Staurotypus salvinii Celestus atitlanensis* Anguidae Coleonyx mitratus Eublepharidae Gonatodes albogularis Hemidactylus frenatus Gekkonidae Phyllodactylus tuberculosus Gymnophthalmus speciosus* Gymnophthalmidae Basiliscus vittatus Corytophanes percarinatus Ctenosaura similis Squamata Iguana iguana* Norops crassulus Iguanidae Norops macrophallus Norops sericeus Norops serranoi Sceloporus malachiticus Sceloporus squamosus Mabuya unimarginata Scincidae Biodiversity field inventories in selected watersheds 117 Order Species Mesoscincus managuae* Sphenomorphus assatus Ameiva undulata Teiidae Aspidoscelis deppii Lepidophyma smithii Xantusiidae Boa constrictor Boidae Coniophanes fissidens Coniophanes piceivittis Conophis concolor Drymarchon melanurus Drymobius margaritiferus Imantodes gemmistratus Lampropeltis triangulum Leptodeira annulata Leptodeira nigrofasciata Leptodeira septentrionalis Leptodrymus pulcherrimus Matigodryas dorsalis* Mastigodryas melanolomus* Masticophis mentovarius Ninia sebae Oxybelis aeneus Colubridae Oxybelis fulgidus Pliocercus elapoides* Rhadinaea godmani* Rhadinaea pilonaorum* Scaphiodontophis annulatus Scolecophis atrocinctus Senticolis triaspis Sibon anthracops Sibon nebulatus* Spilotes pullatus Stenorrhina freminvillii Tantilla brevicauda Tantilla taeniata* Trimorphodon biscutatus Tropidodipsas fischeri* Tropidodipsas sartorii Micrurus nigrocinctus Elapidae Pelamis platurus* Leptotyphlops goudotii* Leptotyphlopidae Loxocemus bicolor* Loxocemidae Atropoides nummifer Agkistrodon bilineatus Cerrophidion godmani Viperidae Crotalus simus* Porthidium ophryomegas * Species registered in prior studies but not during the present study. Note: Scientific names according to Köhler et al. (2006). Recent changes in scientific name and families according to IUCN (2009). 118 Family Biodiversity field inventories in selected watersheds Figure 10A. Taylor’s Worm Salamander (Oedipina taylori), El Imposible National Park, Cerro Campana sector, July 2007. Figure 10B. Red Toad (Incilius canaliferus), San Marcelino Complex Protected Natural Area, September 2007. Figure 10C. Black-eyed Tree Frog Figure 10D. Yellow Cricket Tree Frog (Agalychnis moreletii), El Imposible (Dendropsophus robertmertensi), Santa National Park, December 2006. Rita and Zanjón El Chino Protected Natural Area, October 2007. Figure 10E. Milk Frog (Trachycephalus Figure 10F. Narrow-mouthed Toad venulosus), Plan de Amayo Protected (Gastrophryne usta), Plan de Amayo Natural Area, August 2007. Protected Natural Area, August 2007. Biodiversity field inventories in selected watersheds 119 Figure 10G. Burrowing Toad (Rhinophrynus dorsalis), Santa Rita and Zanjón El Chino Protected Natural Area, October 2007. Figure 10H. Salvin’s Snapping Turtle (Staurotypus salvinii), Santa Rita and Zanjón El Chino Protected Natural Area, August 2007. Figure 10I. Cayman (Caiman crocodilus), Santa Rita and Zanjón El Chino Protected Natural Area, August 2007. Figure 10J. Leopard Gecko (Coleonyx mitratus), Santa Rita and Zanjón El Chino Protected Natural Area, October 2007. Figure 10K. Keeled Helmeted Basilisk Figure 10L. Painted Whiptail Lizard (Corytophanes percarinatus), El (Ameiva undulata), Plan de Amayo Imposible National Park, Cerro Protected Natural Area, August 2007. Campana sector, July 2007. 120 Biodiversity field inventories in selected watersheds Figure 10M. Boa constrictor (Boa Figure 10N. Yellowbelly Snake constrictor), Plan de Amayo Protected (Coniophanes fissidens), Santa Rita and Natural Area, October 2007. Zanjón El Chino Protected Natural Area, August 2007. Figure 10O. Central American Tree Figure 10P. Milk Snake (Lampropeltis Snake (Imantodes gemmistratus), Plan triangulum), Plan de Amayo Protected de Amayo Protected Natural Area, Natural Area, November 2004. October 2007. Figure 10Q. Black-banded Cat-eyed Snake (Leptodeira nigrofasciata), Plan de Amayo Protected Natural Area, November 2007. Figure 10R. Black-banded Snake (Scolecophis atrocinctus), El Imposible National Park, Cerro Campana sector, July 2007. Biodiversity field inventories in selected watersheds 121 Figure 10S. Green Rat Snake (Senticolis Figure 10T Cope´s Snail Sucker Snake triaspis), Plan de Amayo Protected (Sibon anthracops), Plan de Amayo Natural Area, October 2007. Protected Natural Area, July 2007. By Luis Girón Figure 10U. Tiger Rat Snake (Spilotes pullatus), Santa Rita and Zanjón El Chino Protected Natural Area, August 2007. By Xiomara Henríquez Figure 10V. Merten´s Centipede Snake (Tantilla brevicauda). Los Volcanes National Park, Los Andes sector, October 2007. Figure 10W. Mexican Cantil (Agkistrodon bilineatus), Santa Rita and Zanjón El Chino Protected Natural Area, August 2007. Figure 10X. Godman´s Montane Pit Viper (Cerrophidion godmani), Los Volcanes National Park, Los Andes sector, October 2007. FIGURE 10. SELECTED PHOTOGRAPHS OF AMPHIBIANS AND REPTILES TAKEN DURING THE STUDY. (Photos by Vladlen Henríquez except where indicated.) 122 Biodiversity field inventories in selected watersheds 3.2 IMPORTANT SPECIES FOR CONSERVATION The study area presents 50 species which are on the lists of threatened species nationwide and worldwide. In the case of worldwide threatened species, eight species were recorded in the project area including two amphibians and six reptiles. There are 42 other species which are included on the list of nationally endangered species (MARN 2004, Greenbaum & Komar 2005). See Table 27. 3.3 NEW RECORDS FOR THE STUDY AREA Of the 73 species recorded during the present study, five are new records for the project area and for the departments where they were recorded. These include Narrow Mouthed Toad (Gastrophryne usta), an exotic (non-native) gecko Hemidactylus frenatus, the Blackbanded Cat-eyed Snake (Leptodeira nigrofasciata), and False Coral Snake (Sibon anthracops). The Terrestrial Snail Sucker (Tropidodipsas sartorii), a snake encountered on the Santa Ana Volcano, is a new record for the project area but was previously recorded in the northwest zone of the Santa Ana department. Including these five species, a total of 20 new reports were recorded at the level of department. First records for the Department of Ahuachapán include cricket tree frog (Dendropsophus robertmertensi), swamp tree frog (Scinax staufferi), milk frog (Trachycephalus venulosus), white lipped frog (Leptodactylus fragilis), narrow mouthed toad (Gastrophryne usta), burrowing toad (Rhinophrynus dorsalis), Mesoamerican slider (Trachemys venusta), Pacific coast giant musk turtle (Staurotypus salvinii), black-striped snake (Coniophanes piceivittis), annulated cat-eyed snake (Leptodeira annulata), neckbanded snake (Scaphiodontophis annulatus), common cantil (Agkistrodon bilineatus), and Godman’s montane pitviper (Cerrophidion godmani). First records for the Department of Sonsonate include: Narrow-mouthed Toad (Gastrophryne usta), Hawksbill Sea Turtle (Eretmochelys imbricata), Gecko (Hemidactylus frenatus), False Coral Snake (Lampropeltis triangulum), Black-banded Cateyed Snake (Leptodeira nigrofasciata), Brown Vinesnake (Oxybelis aeneus), and False Coral Snake (Sibon anthracops). 3.4 LEVEL OF INVENTORY COMPLETENESS In general, the level of herpetofauna inventory completeness increased in nine watersheds in 2007. Currently, the watersheds with the most advanced inventories are Izalco, San Julián, Cara Sucia, and Barra de Santiago. Notwithstanding the sampling efforts made in the Río Grande de Sonsonate and Coatepeque watersheds, their levels of inventory completeness is below 50% (Table 29). The inventory of the municipality of San Francisco Menéndez was almost completed (92%) with this study. The municipality of Caluco shows a high level of inventory completeness (65%) whereas before this study it was at 0%. The municipality of Izalco also shows a relatively high completeness level of 57%. The remaining municipalities show inventory completeness levels below 40%. Ten municipalities do not have any herpetofauna records (Table 30). Biodiversity field inventories in selected watersheds 123 At the level of ecosystems, six present almost complete inventories (80–90%) as is the case of humid forest, freshwater ecosystems, and the rocky intertidal zone. The dry forest presents a relatively complete inventory, that is, 100% of the expected species are recorded. However, it is still possible to detect new ―unexpected‖ species. The intertidal zone presents an almost complete inventory but no sampling efforts were made during this study. Research has yet to be conducted in savannas and estuaries (Table 31). TABLE 29. INVENTORY COMPLETENESS FOR AMPHIBIANS AND REPTILES AND RELATIVE IMPORTANCE OF WATERSHEDS. Watershed Cara Sucia Barra de Santiago San Juan Río Cauta San Pedro Río Grande Sonsonate Izalco San Julian Lago de Coatepeque Barra Salada Los Cóbanos 124 Species Recorded Expected Species Recorded (of 53) Initial level of Final Level of Inventory Inventory Completeness Completeness (%) (%) 52 32 51 60 74 42 31 49 58 50 0 3 15 0 1 14 0 2 23 0 2 26 0 2 0 10 9 10 17 2 57 43 39 35 47 32 74 66 32 22 26 13 23 25 8 2 2 0 4 0 27 16 26 30 4 Biodiversity field inventories in selected watersheds Proportion of Conservation Important Species (%) TABLE 30. INVENTORY COMPLETENESS FOR AMPHIBIANS AND REPTILES AND RELATIVE IMPORTANCE OF MUNICIPALITIES. Expected Initial Level of Final Level of Proportion of Species Species Inventory Inventory Conservation Municipality Recorded Recorded Completeness Completeness Important (of 53) (%) (%) Species (%) San Francisco 67 49 74 92 68 Menéndez Concepción 0 0 0 0 de Ataco Apaneca 9 6 6 11 2 San Pedro 13 7 7 13 2 Puxtla Guaymango 0 0 0 0 0 Tacuba 13 8 1 15 12 Jujutla 10 3 3 6 10 Acajutla 24 13 13 25 6 San Antonio 0 0 0 0 0 del Monte Salcoatitán 0 0 0 0 0 Santa 0 0 0 0 0 Catarina Masahuat Izalco 40 30 29 57 24 Juayúa 2 2 1 4 2 Nahuizalco 1 1 1 2 0 San Julián 24 16 16 30 10 Caluco 41 34 32 64 20 Armenia 0 0 0 0 0 Tepecoyo 0 0 0 0 0 Santo 3 3 1 7 2 Domingo de Guzmán Cuisnahuat 0 0 0 0 0 Nahulingo 0 0 0 0 0 Sonsonate 30 23 19 43 8 Sonzacate 1 1 2 2 0 Santa Ana 7 7 7 13 8 Chalchuapa 0 0 0 0 0 Biodiversity field inventories in selected watersheds 125 TABLE 31. INVENTORY COMPLETENESS FOR AMPHIBIANS AND REPTILES AND RELATIVE IMPORTANCE OF ECOSYSTEMS. No. of Proportion of Level of Expected Indicator Conservation Species Inventory Ecosystem Species Species for Important Recorded Completeness Recorded Inventory Species (%) Completeness (%) Humid 58 42 44 95 58 Forest 39 36 36 100 20 Dry Forest Freshwater Ecosystems Cloud Forest Savanna Rocky Intertidal Zone Estuaries Coral Reefs 45 36 40 90 26 13 10 12 83 12 0 0 20 0 0 8 8 10 80 8 9 9 15 60 1 0 0 0 0 2 With regards to the Protected Natural Areas, the analysis of inventory completeness based on the expected generalist species indicates that El Imposible National Park reports the largest number of recorded species (57 species) and the highest percentage of inventory completeness (85%). It also reports the largest number of threatened and endangered species (23 species). The rest of the natural areas have similar quantities of important species for conservation and level of inventory completeness (see Table 32). Therefore, more field efforts should be made to complete the inventories of the study areas. We obtain the same result from an analysis based on species accumulation curves and wealth estimators. The species accumulation curves of the Protected Natural Areas indicate that most of them do not show a trend towards stabilization and therefore it is necessary to make more sampling efforts (Fig. 11). Only the curve of Laguna de las Ninfas Protected Natural Area shows a trend towards stabilization. The wealth estimators indicate that the inventory of some of the Protected Natural Areas could be almost complete. The wealth estimators of El Imposible National Park indicate that up to 58 species could be recorded of which 57 are already on record. The estimators of Los Volcanes National Park indicate that up to 49 species could be recorded of which 46 are already on record, and consequently, their inventory is almost complete. Some species are yet to be recorded in San Marcelino Complex, Santa Rita and Plan de Amayo Protected Natural Areas (Table 4-9). The wealth estimators of Laguna de Las Ninfas Protected Natural Area indicate that up to 9 species could be recorded but 10 species are already on record and it is possible that additional field work could discover more. 126 Biodiversity field inventories in selected watersheds TABLE 32. INVENTORY COMPLETENESS FOR AMPHIBIANS AND REPTILES AND RELATIVE IMPORTANCE OF PROTECTED NATURAL AREAS. Protected Natural Area El Imposible National Park Los Volcanes National Park Laguna de Las Ninfas San Marcelino Complex Santa Rita y Zanjón El Chino Plan de Amayo No. of Level of Proportion of Indicator Inventory Conservation Species for Completeness Important Inventory (%) Species (%) Completeness Species Recorded Expected Species Recorded 57 39 46 85 46 46 35 45 78 34 10 8 12 67 6 40 32 41 78 22 45 33 42 79 28 41 29 41 71 16 Biodiversity field inventories in selected watersheds 127 45 Number of Species 40 35 SMC 30 PA 25 LC 20 SR LVNP 15 EINP 10 5 0 1 3 5 7 9 11 13 15 17 19 Sampling Days FIGURE 11. ACCUMULATION CURVE OF HERPETOFAUNA SPECIES FROM EACH PROTECTED NATURAL AREA STUDIED. TABLE 33. ESTIMATORS OF HERPETOFAUNA SPECIES RICHNESS IN EACH PROTECTED NATURAL AREA STUDIED. Estimated Richness Richness Estimator EINP LVNP SMC SR LLN PA ACE 51 39 35 49 9 44 ICE 51 43 46 52 8 50 Chao 1 50 37 33 43 8 41 Chao 2 52 38 35 57 7 56 Jack-Knife 1 49 42 37 55 9 47 Jack-Knife 2 58 49 42 54 8 57 EINP = El Imposible National Park LVNP = Los Volcanes National Park SMC = San Marcelino Complex Protected Natural Area SR = Santa Rita Protected Natural Area LLN = Laguna de Las Ninfas Protected Natural Area PA = Plan de Amayo Protected Natural Area LC = Los Cóbanos Marine Park Protected Natural Area 128 Biodiversity field inventories in selected watersheds LC 22 20 22 19 19 23 80 Number of Species 70 60 50 40 30 20 10 0 1 9 17 25 33 41 49 57 65 73 81 89 97 105 113 121 Sampling days FIGURE 12. ACCUMULATION CURVE FOR HERPETOFAUNA SPECIES FROM SOUTHWESTERN EL SALVADOR. As we have seen in the case of the Protected Natural Areas, the use of species accumulation curves is another way to determine inventory completeness. Only the data obtained during the field stage for 74 species were used to analyze the entire project area. The curve demonstrates that stabilization has not been achieved after 120 days of sampling (Fig. 12). Richness estimators based on the curve suggest that southwestern El Salvador presently has between 84 and 95 herpetofauna species (Table 34). Considering previous studies, 96 species are already recorded. This suggests that all amphibians and reptiles in the area have already been recorded. However, at a finer scale such as at the level of watershed, municipality, ecosystems, or individual natural areas, the curves and estimators show that additional field work is required. For ecosystems, the species accumulation curve did not approach stabilization (Fig. 13). Richness estimators for the Dry Forest indicate that up to 39 species should be found; for Humid Forest up to 74, for Cloud Forest up to 17, and for Freshwater Ecosystems, up to 64 (Table 4-9). In the case of Dry Forest, considering previous studies, the species list already reached a total of 39, and therefore it is likely that the inventory for this habitat is already complete. The estimators for the Humid Forest, Cloud Forest, and Freshwater Ecosystems indicate that other species are yet to be recorded. Biodiversity field inventories in selected watersheds 129 50 45 Number of Species 40 Humid Forest 35 Freshwater Ecosystems Dry Forest 30 25 20 Cloud Forest 15 10 5 0 1 4 7 10 13 16 19 22 25 28 Sampling Days FIGURE 13. ACCUMULATION CURVE FOR HERPETOFAUNA SPECIES FROM EACH ECOSYSTEM STUDIED. TABLE 34. ESTIMATORS OF HERPETOFAUNA SPECIES RICHNESS IN EACH ECOSYSTEM STUDIED. Richness Estimator ACE ICE Chao 1 Chao 2 Jack-Knife 1 Jack-Knife 2 130 Estimated Richness Project Area 87 86 86 84 89 95 Dry Forest Humid Forest Cloud Forest 34 36 31 33 37 39 66 74 67 70 64 71 15 17 13 13 17 15 Biodiversity field inventories in selected watersheds Freshwater Ecosystems 49 62 43 56 55 64 3.5 IMPORTANCE OF THE AREAS FOR CONSERVATION The level of inventory completeness in watersheds and municipalities is not adequate to evaluate the relative importance of these areas for herpetofauna conservation. However, at the level of Protected Natural Area and ecosystem, the inventories are relatively complete (67%–85% for Protected Natural Areas and five ecosystems have reached a completeness level of 80%), suggesting that the proportion of indicator species for conservation recorded in these areas will be useful as an importance index. Among the Protected Natural Areas, El Imposible reports the highest importance index followed by Los Volcanes, Santa Rita, San Marcelino Complex, Plan de Amayo, and Laguna de Las Ninfas (Table 32). Among the ecosystems, the Humid Forest presents the largest number of threatened and endangered species with a total of 28 species (Table 31). It also presents the highest species richness (58 species). The Freshwater ecosystem presents 12 species important for conservation and the second richest ecosystem with 45 species. The Dry Forest presents 10 threatened species and 39 species in total. The Cloud Forest presents six threatened species (out of 13 total). The habitat of the Intertidal and Rocky Zone reports only two endangered species; however, it should be considered that this type of habitat is only used by sea turtles to lay their eggs and by some lizard species which are not threatened. The Coral Reef and the Savanna do not yet have formal herpetofauna records. Considering that the inventories of the five ecosystems are well advanced, it is unlikely that the order of importance will change if more studies are conducted. However, our understanding of the conservation status of each species could change, which could modify the relative importance of each ecosystem. 4. RECOMMENDATIONS Below are some recommendations for educational and scientific purposes:     Further efforts should be made to carry out inventories of the coastal zone habitats such as reefs, estuaries, intertidal and rocky zone, and savanna since these habitats were not included in the sampling. It is particularly urgent to conduct studies in the savannas since it is a distinct and reduced ecosystem of the study area that could have species important for biodiversity maintenance in the study area. Monitoring protocols should be prepared for the eight globally threatened herpetofauna species that have been recorded in the project area. Training workshops for park rangers in the Protected Natural Areas should be provided, focusing on the threatened and endangered species recorded in those areas. Park rangers and members of the communities near the Protected Natural Areas should receive environmental education about venomous snakes and first-aid training. Biodiversity field inventories in selected watersheds 131 CHAPTER 5: INVENTORY OF BIRDS IN SOUTHWESTERN EL SALVADOR Oliver Komar SalvaNATURA, Conservation Science Department San Salvador, El Salvador okomar@salvanatura.org 1. INTRODUCTION Of all of the classes of flora and fauna, birds (Aves) are the best known, not only by scientists but also by the general public. They are more easily seen and heard than mammals, amphibians, reptiles, insects and fish. As intelligent vertebrates, they capture people’s attention and imagination more so than other life forms, such as plants and insects. At the level of sites, such as particular Protected Natural Areas or forests, birds are frequently more completely documented then other kinds of animals or plants. Most species have been studied in some depth, and virtually all species were long ago described for science. Numerous books have been written to help people know birds; all of El Salvador’s birds are illustrated in several field guides written for the public (such as Howell and Webb 1995). That cannot be said for some other groups of animals and especially for plants. In fact, annually, several new plants, insects, fish, and amphibians are still being described for science in Central America. The rate of discovery for other vertebrates (reptiles and mammals) is lower, but still higher than for birds. This high level of knowledge available for birds allows this taxonomic group to serve as good indicators for environmental change or for relative importance of sites as biodiversity refuges. There are three characteristic criteria for identifying a good indicator species: it must be easily identifiable, detectable, and sensitive to change. Birds can typically be identified in the field by experienced observers, and there is abundant literature available to assist biologists in learning to identify them. Most birds are reasonably easy to detect, being generally showy creatures, and often advertising their own presence by singing. Finally, many birds are sensitive to changes in the environment: birds are diverse creatures and in virtually any bird community, one can find at least some species that are sensitive to ecological change. The mere presence of certain generalist species can signal perturbations in the environment; population declines can reflect disturbances such as contamination or habitat fragmentation. The presence of certain habitat specialists can signal sites of conservation value. Thus, birds can be indicators of sites with high quality habitat, or sites useful as biological corridors. In order to be able to use birds as indicators for site quality, they must be inventoried and monitored. 132 Biodiversity field inventories in selected watersheds The birds of southwestern El Salvador, in the area where USAID has sponsored the Improved Management and Conservation of Critical Watersheds (IMCW) project, have been studied frequently since 1925. Through 2006, there were at least 25 field studies, generating 21,724 records of birds, based on approximately 1,388 days of field observations (detailed in Komar 2007b). All of this work documented 413 bird species, including 100% of the 74 widespread, generalist bird species that serve as indicators of a complete inventory. A baseline study has documented the relative completeness of the bird inventory in each of 11 watersheds and in each of 25 municipalities within southwestern El Salvador (Komar 2007b). The mean completeness of the inventories, based on the presence of expected generalist species was 68% for watersheds, and 36% for municipalities. While the bird inventory for the entire region of southwestern El Salvador appeared to be complete, the inventories for specific sites within that region were considerably less complete. Some entire municipalities and some specific Protected Natural Areas within southwestern El Salvador had never had their birds documented at all. The IMCW project was interested in determining if specific watersheds or municipalities within southwestern El Salvador were more important than others for conserving biodiversity. Comparison of sites, using indicator species, requires the existence of complete inventories at the site level (Remsen 1994, Komar 2003). Otherwise, the sites with more complete inventories will simply appear to be of greater value, regardless of their true relative value. The purpose of the present study was to advance the state of the bird inventories for a series of sites and ecosystems in southwestern El Salvador, such that relative value for biodiversity conservation could be assessed. 2. STUDY AREA AND METHODS The study area included seven of the 11 watersheds within the IMCW project area (Fig.14). The Río Banderas watershed was divided into three watersheds for the purpose of biological analysis: the lower Río Banderas (coastal plain), the Izalco watershed which drains most of Los Volcanes National Park, and the San Julian watershed which drains the northwestern sector of the Sierra del Bálsamo. The Los Cóbanos watershed actually consists of a series of approximately seven minor watersheds, each emptying into the Pacific Ocean, and almost indistinguishable on the ground. The Barra de Santiago watershed was divided into two: ―Cara Sucia,‖ which drains the western two-thirds of El Imposible National Park and ―Barra de Santiago‖ which drains the eastern third of the national park and several valleys to the east. Most of the rivers in the Cara Sucia watershed are drained or go underground before they reach the estuary. The IMCW project area encompasses 28 municipalities, although three only marginally. Field work was carried out at eight sites located within eight municipalities (Table 35). Not reported here are the results of monitoring of bird populations carried out during 2007 and 2008, at numerous additional sites. These results also contribute to the ongoing inventory but will be reported separately. Below I describe the process for creating lists of three types of indicator bird species, used for the evaluation of the completeness of the inventory work or the evaluation of site Biodiversity field inventories in selected watersheds 133 importance for the conservation of biodiversity. First is the list of widespread, common species that are guaranteed to occur in any reasonably large and heterogeneous area, such as a municipality or a watershed. These species serve as indicators of inventory completeness at the site level (Gómez de Silva & Medellín 2001, Komar 2007). Second is the list of common ecosystem indicators, which can be generalist or specialist species, and that are expected to be found in any existing patch of appropriate habitat. These species serve as indicators of inventory completeness at the ecosystem level. Third is the list of threatened species, which should serve as indicators of conservation importance for the sites where they occur (Komar 2007). For example, conservation priority should be assigned to a site with higher richness of threatened species. This is an important distinction, since some sites may have higher overall species richness but have low richness for conservation-importance indicator species; such sites should not be mistakenly considered to be important for conservation simply for having high numbers of species and high levels of biodiversity (Komar 2003). 3 7 8 4 1 2 6 5 FIGURE 14. MAP OF BIRD INVENTORY FIELD SITES DURING 2007: (1) SANTO DOMINGO DE GUZMÁN, (2) PLAN DE AMAYO, (3) LAGUNA DE LAS NINFAS, (4) LAVAS DE IZALCO/FINCA MARÍA AUXILIADORA, (5) BARRA SALADA, (6) LOS CÓBANOS, (7) LAGUNA DE LAS RANAS, (8) BOSQUE SANTA RITA. 134 Biodiversity field inventories in selected watersheds TABLE 35. LOCALITIES WHERE BIRD INVENTORY FIELD WORK WAS CARRIED OUT BY THE IMCW PROJECT. Site Santo Domingo de Guzmán Watershed Municipality San Pedro Santo Domingo de Guzmán Ecosystems Freshwater wetlands (Riparian forest and rivers) Plan de Amayo San Julian Caluco Dry forest Laguna de las Ninfas Río Grande de Sonsonate Apaneca Humid montane forest Lavas de Izalco/Finca María Auxiliadora Izalco Izalco Dry forest Barra Salada Barra Salada Sonsonate Estuary (including mangrove forest) Los Cóbanos Los Cóbanos Acajutla y Sonsonate Savanna, dry forest, and beach Laguna de las Ranas Río Grande de Sonsonate Juayúa Cara Sucia San Francisco Menéndez Bosque Santa Rita Humid montane forest Freshwater wetlands (swamp forest) Dates (in 2007) Field team1 4 days (16–19 July) JF, CF, JJ, ST, IV 12 days (20–24 June, 15–21 October) 11 days (29 June to 8 July and then 20 July) 20 days (3–12 August and 1–10 November) 19 days (3–12 September and 15–23 December) 20 days (20–29 August, and 3–12 December) CF, JF, JJ, ST, IV, CZ JF, CF, KL, ST, IV, CZ CF, LA, JF, JJ. LS, IV, CZ CF, JJ, OK, LS, ST, IV, CZ CF, JJ, OK, KL, LS, ST, IV, CZ 10 days (1–10 October) CF, JF, JJ, ST, IV, CZ 20 days (17–26 September, 14– 23 November) JF, CF, JJ, KL, LS, IV, CZ 1 LA=Lety Andino, CF=Carlos Funes, JF=Jesse Fagan, JJ=Jorge Jiménez, OK=Oliver Komar, KL=Karla Lara, LS=Lya Samayoa, ST=Sofía Trujillo, IV=Iselda Vega, CZ=Carlos Zaldaña Biodiversity field inventories in selected watersheds 135 2.1. INDICATORS OF INVENTORY COMPLETENESS Following the methodology of Gómez de Silva and Medellín (2001), I first considered which resident bird species are widespread generalists and expected in virtually any watershed or municipality of the study area. Such species can be considered ―test‖ species, for testing how complete inventory efforts have been at any given site. To determine the test species appropriate for sites within the IMCW project area, I selected species from the revised bird list of El Salvador (Komar et al. 2009), which includes a total of 548 species each classified for status as habitat generalists or specialists, residents or migrants, and for threat at the national level. To build the list of test species, I first selected species classified as permanent residents or partially migratory residents that are also forest generalists. From the resulting list of 94 permanent residents and seven partially migratory residents, I eliminated 57 species classified as nationally threatened (including critically endangered, endangered, vulnerable, or near-threatened), because these are rare species not likely to be widespread in the study area. Of the remaining 44 species, I eliminated 4 species restricted to highlands above 1000 m elevation. I considered if some of the species on the remaining list may be restricted only to the lowest elevations, but although several are more common in the lowlands, none were restricted to them. As would be predicted, the remaining 40 species are widespread in El Salvador and should be found in every municipality or watershed within Ahuachapán and Sonsonate. Although these species are all forest species, they can be found in disturbed forests including tree plantations and urban parks, and are by no means restricted to natural areas or Protected Natural Areas. I then added to the list of test species resident birds that live in open areas rather than forests. I culled the overall bird list for resident species (including species that are partially migratory) classified as open area generalists. From the resulting lists of 49 permanent residents and 5 partially migratory resident species, I eliminated 12 threatened and nearthreatened species, and 3 species restricted to high altitudes, generating a list of 39 species that should be registered at any given non-forest site within Ahuachapán or Sonsonate. Several species were eliminated from this list for various reasons. The Tropical Mockingbird (Mimus gilvus) was considered to be restricted to very low elevations close to the coast. The Rock Pigeon (Columba livia) and House Sparrow (Passer domesticus), both introduced species, were considered to be restricted to urban and other non- natural habitats. The Nutting’s Flycatcher (Myiarchus nuttingi) is probably restricted to low elevations (<500 m) and is of uncertain occurrence in the study area. The Zone-tailed Hawk (Buteo albonotatus) has a breeding range in El Salvador restricted to areas outside the study area. The resulting list of open area generalist species, useful as indicators for inventory completeness (i.e., test species) was 34 species. The combination of forest and open-area site inventory indicators was 74 species (Table 36). The species in Table 36 should be registered in any complete inventory of a heterogeneous area with both forest and non-forest habitats. All of these species should occupy every municipality or watershed in the study area. These are all common, widespread species, and are useful for identifying gaps in the inventory. Nonetheless, an area with 100% of these species recorded may still have an incomplete inventory, as rare species may still not be included in a site’s inventory despite being present and despite all of the common species having been detected. 136 Biodiversity field inventories in selected watersheds TABLE 36. THE 74 RESIDENT BIRD SPECIES, EXPECTED IN ANY MUNICIPALITY OR WATERSHED OF THE STUDY AREA, THAT SERVE AS INDICATORS OF INVENTORY COMPLETENESS (“TEST” SPECIES) AT THE SITE LEVEL. Family Species Spanish name English name Tinamidae Crypturellus cinnamomeus Tinamú Canelo Thicket Tinamou Cracidae Ortalis leucogastra Chachalaca Vientreblanco White-bellied Chachalaca Odontophoridae Colinus cristatus Codorniz-cotui Centroamericana Cathartidae Coragyps atratus Zopilote Negro Black Vulture Cathartidae Cathartes aura Aura Cabecirroja Turkey Vulture Accipitridae Buteogallus anthracinus Aguililla Negra Menor Common Black-Hawk Accipitridae Buteo magnirostris Aguililla Caminera Roadside Hawk Accipitridae Buteo nitidus Aguililla Gris Gray Hawk Falconidae Herpetotheres cachinnans Halcón Guaco (Guas) Laughing Falcon Columbidae Patagioenas flavirostris Paloma Morada Red-billed Pigeon Columbidae Zenaida asiatica Paloma Aliblanca White-winged Dove Columbidae Columbina inca Tórtola Colilarga Inca Dove Columbidae Columbina talpacoti Tortolita Rojiza Ruddy Ground-Dove Columbidae Leptotila verreauxi Paloma Arroyera (Rodadora) White-tipped Dove Psittacidae Aratinga canicularis Perico Frentinaranja Orange-fronted Parakeet Psittacidae Brotogeris jugularis Periquito Barbinaranja Orange-chinned Parakeet Cuculidae Piaya cayana Chocolatero, Piscoy Squirrel Cuckoo Cuculidae Morococcyx erythropygus Cuco-terrestre Menor Lesser Ground-Cuckoo Cuculidae Crotophaga sulcirostris Pijuyo Groove-billed Ani Strigidae Glaucidium brasilianum Tecolotito Común (Aurorita) Ferruginous Pygmy-Owl Strigidae Ciccaba virgata Búho Café Mottled Owl Caprimulgidae Nyctidromus albicollis Tapacaminos Pucuyo (Caballero) Common Pauraque Apodidae Streptoprocne rutila Vencejo Cuellicastaño Chestnut-collared Swift Apodidae Streptoprocne zonaris Vencejo Cuelliblanco White-collared Swift Apodidae Chaetura vauxi Vencejo de Vaux Vaux’s Swift Crested Bobwhite Biodiversity field inventories in selected watersheds 137 Family Species Spanish name English name Trochilidae Campylopterus hemileucurus Fandanguero Morado Violet Sabrewing Trochilidae Chlorostilbon canivetii Esmeralda de Canivet Canivet's Emerald Trochilidae Amazilia beryllina Colibrí de Berilo Berylline Hummingbird Trochilidae Amazilia rutila Colibrí Canelo Cinnamon Hummingbird Trogonidae Trogon violaceus Trogón Violaceo (Coa Pechiamarilla) Violaceous Trogon Trogonidae Trogon elegans Trogón Elegante (Coa Pechiroja) Elegant Trogon Momotidae Momotus momota Momoto Coroniazul (Talapo) Blue-crowned Motmot Momotidae Eumomota superciliosa Momoto Cejiturqueza (Torogoz) Turquoise-browed Motmot Picidae Melanerpes aurifrons Carpintero Frentidorado Golden-fronted Woodpecker Picidae Piculus rubiginosus Carpintero Olivaceo Golden-olive Woodpecker Furnariidae Xiphorhynchus flavigaster Trepatroncos Piquiclaro Ivory-billed Woodcreeper Thamnophilidae Thamnophilus doliatus Batará Barrada Barred Antshrike Tyrannidae Tolmomyias sulphurescens Picoplano Ojiblanco Yellow-olive Flycatcher Tyrannidae Contopus cinereus Pibí Tropical Tropical Pewee Tyrannidae Myiarchus tuberculifer Copetón Triste Dusky-capped Flycatcher Tyrannidae Pitangus sulphuratus Luis Grande (Cristo Fue) Great Kiskadee Tyrannidae Megarynchus pitangua Luis Piquigrueso Boat-billed Flycatcher Tyrannidae Myiozetetes similis Luis Gregario (Chio) Social Flycatcher Tyrannidae Tyrannus melancholicus Tirano Tropical Tropical Kingbird Incertae sedis Pachyramphus aglaiae Cabezón Degollado Rose-throated Becard Incertae sedis Tityra semifasciata Titira Enmascarada (Torreja) Masked Tityra Vireonidae Cyclarhis gujanensis Vireón Cejirrufo Rufous-browed Peppershrike Corvidae Calocitta formosa Urraca-hermosa Cariblanca (Urraca) Hirundinidae Progne chalybea Martín Pechigris 138 White-throated MagpieJay Biodiversity field inventories in selected watersheds Gray-breasted Martin Family Species Spanish name English name Hirundinidae Stelgidopteryx serripennis Golondrina-aliserrada Norteña Northern Rough-winged Swallow Troglodytidae Campylorhynchus rufinucha Guacalchía (Matraquita Nuquirrufa) Rufous-naped Wren Troglodytidae Thryothorus modestus Saltapared Sencillo Plain Wren Troglodytidae Troglodytes aedon Saltapared Continental Norteño House Wren Sylviidae Polioptila albiloris Perlita Cejiblanca Turdidae Turdus grayi Zorzal Pardo Clay-colored Robin Parulidae Euthlypis lachrymosa Chipe Roquero Fan-tailed Warbler Parulidae Basileuterus rufifrons Chipe Gorrirrufo Rufous-capped Warbler Thraupidae Thraupis episcopus Tángara Azuligris Blue-gray Tanager Thraupidae Thraupis abbas Tángara Aliamarilla Yellow-winged Tanager Emberizidae Volatinia jacarina Semillero Brincador Blue-black Grassquit Emberizidae Sporophila torqueola Semillero Collarejo White-collared Seedeater Emberizidae Aimophila ruficauda Cardinalidae Saltator coerulescens Saltator Grisáceo (Dichoso-fuí) Grayish Saltator Cardinalidae Saltator atriceps Saltator Cabecinegro (Chepito) Black-headed Saltator Cardinalidae Passerina caerulea Picogrueso Azul Blue Grosbeak Icteridae Dives dives Tordo Cantor Melodious Blackbird Icteridae Quiscalus mexicanus Zanate Mayor (Zanate♀, Clarinero♂) Great-tailed Grackle Icteridae Molothrus aeneus Vaquero Ojirrojo Bronzed Cowbird Icteridae Icterus pustulatus Bolsero Dorsirrayado Streak-backed Oriole Icteridae Icterus pectoralis Icteridae Icterus gularis Bolsero de Altamira Altamira Oriole Icteridae Amblycercus holosericeus Cacique Piquiclaro Yellow-billed Cacique Fringillidae Euphonia affinis Eufonia Gorjinegro Scrub Euphonia Fringillidae Carduelis psaltria Dominico Dorsioscuro Lesser Goldfinch White-lored Gnatcatcher Zacatonero Cabecirrayado Bolsero Pechimanchado Biodiversity field inventories in selected watersheds Stripe-headed Sparrow Spot-breasted Oriole 139 To evaluate the level of completeness for the bird inventory at the level of ecosystems (or habitats), first I adopted as the ecosystems of interest the eight ecosystems identified as priority conservation objects for the study area by the IMCW project (Komar & Girón 2007). These included cloud forest, humid montane forest, dry forest, estuarine habitat (including mangrove forest and mudflats), beaches, savannas, and coral reefs. Also included were freshwater wetlands, a broad category that includes lakes, rivers, swamp forest (such as at Bosque Santa Rita, which is seasonally flooded), and riparian habitat at the edges of waterways. I assembled lists of habitat specialists and generalists that are sufficiently common to be expected in every existing patch of natural habitat within each of these ecosystem categories. For forest ecosystems, these lists tended to include all 40 of the forest generalists included in Table 36, but in the case of forest habitats restricted to narrow elevational ranges, such as cloud forest and mangrove, not all 40 were included. Added to the lists of indicators were specialist species found only in the given habitat, if the species is sufficiently common to be expected in every patch. For non-forest ecosystems, the 34 open-area generalists were likely to all be included, as well as a few additional open-area specialist species if these are quite common or widespread across patches. The resulting lists of inventory indicator species are presented in Table 37. 2.2. INDICATORS OF CONSERVATION IMPORTANCE Globally-threatened species are often considered the best indicators for identifying sites of conservation importance; such species are listed annually by the World Conservation Union (IUCN 2008). Globally-threatened species are used worldwide as principal indicators for identifying Important Bird Areas, Key Biodiversity Areas, and Alliance for Zero Extinction sites. Relatively few globally-threatened species, however, occur in El Salvador. In fact, no resident bird species recorded in the study area are globallythreatened, and just one, the Great Curassow (Crax rubra), is classified as globally nearthreatened (several migratory visitors—non-residents—are also globally near-threatened). The curassow is believed to be restricted in El Salvador to the forests of El Imposible National Park. Another group of species sometimes used as indicators of conservation importance are migratory birds. U.S.-based bird conservation donors tend to support projects that aim to protect migratory birds that breed in northern latitudes and winter in Central America. Such species are often thought to be vulnerable because their migratory habits expose them to diverse threats in different places at different times. The accumulation of threats on the breeding grounds, migratory stopover sites, and wintering grounds suggests that these species should be protected wherever they go, to minimize the accumulation of threats. However, most migratory species are actually well adapted to diverse conditions, including severe habitat disturbance. Most of these species are widely distributed, with abundant populations, and are not considered threatened in any of the areas where they breed or winter. The Swainson’s Thrush (Catharus ustulatus), for example, has an estimated North American population of over 100 million individuals (Rich et al. 2004). Nearly 200 migratory species visit El Salvador each year. Just a few of these species have been classified in continent-wide conservation plans and analyses as of conservation concern (Kushlan et al. 2002, Rich et al. 2004). 140 Biodiversity field inventories in selected watersheds TABLE 37. BIRD SPECIES USED AS “TEST SPECIES” TO INDICATE RELATIVE LEVELS OF INVENTORY COMPLETENESS AT THE ECOSYSTEM LEVEL, IN SOUTHWESTERN EL SALVADOR. Species English name Crypturellus cinnamomeus Ortalis leucogastra Dendrortyx leucophrys Colinus cristatus Dactylortyx thoracicus Sula granti Sula leucogaster Pelecanus occidentalis Phalacrocorax brasilianus Fregata magnificens Ardea herodias Ardea alba Egretta thula Egretta tricolor Bubulcus ibis Butorides virescens Nyctanassa violacea Eudocimus albus Coragyps atratus Cathartes aura Pandion haliaetus Elanus leucurus Accipiter striatus Buteogallus anthracinus Buteo magnirostris Thicket Tinamou White-bellied Chachalaca Buffy-crowned Wood-Partridge Crested Bobwhite Singing Quail Nazca Booby Brown Booby Brown Pelican Neotropic Cormorant Magnificent Frigatebird Great Blue Heron Great Egret Snowy Egret Tricolored Heron Cattle Egret Green Heron Yellow-crowned Night-Heron White Ibis Black Vulture Turkey Vulture Osprey White-tailed Kite Sharp-shinned Hawk Common Black-Hawk Roadside Hawk Humid forest Dry forest x x Cloud forest Estuaries & mangroves Beaches Freshwater Savannas wetlands Coral reef x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x x x 141 Species English name Buteo platypterus Buteo nitidus Buteo brachyurus Buteo swainsoni Buteo albonotatus Buteo jamaicensis Micrastur semitorquatus Caracara cheriway Herpetotheres cachinnans Falco sparverius Falco columbarius Falco peregrinus Burhinus bistriatus Pluvialis squatarola Charadrius wilsonia Charadrius semipalmatus Charadrius vociferus Himantopus mexicanus Jacana spinosa Actitis macularius Tringa melanoleuca Tringa semipalmata Tringa flavipes Numenius phaeopus Arenaria interpres Calidris alba Calidris pusilla Broad-winged Hawk Gray Hawk Short-tailed Hawk Swainson’s Hawk Zone-tailed Hawk Red-tailed Hawk Collared Forest-Falcon Crested Caracara Laughing Falcon American Kestrel Merlin Peregrine Falcon Double-striped Thick-knee Black-bellied Plover Wilson’s Plover Semipalmated Plover Killdeer Black-necked Stilt Northern Jacana Spotted Sandpiper Greater Yellowlegs Willet Lesser Yellowlegs Whimbrel Ruddy Turnstone Sanderling Semipalmated Sandpiper 142 Humid forest x x x x x Dry forest x x x Cloud forest Estuaries & mangroves Beaches x x x x x Freshwater Savannas wetlands x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x x x x x x x x x x x x x x x x Coral reef Species English name Calidris mauri Calidris minutilla Limnodromus griseus Larus atricilla Larus pipixcan Chlidonias niger Sterna hirundo Thalasseus maximus Thalasseus sandvicensis Thalasseus elegans Rynchops niger Stercorarius pomarinus Stercorarius parasiticus Patagioenas flavirostris Zenaida asiatica Zenaida macroura Columbina inca Columbina passerina Columbina minuta Columbina talpacoti Leptotila verreauxi Geotrygon albifacies Aratinga canicularis Brotogeris jugularis Piaya cayana Morococcyx erythropygus Crotophaga sulcirostris Western Sandpiper Least Sandpiper Short-billed Dowitcher Laughing Gull Franklin’s Gull Black Tern Common Tern Royal Tern Sandwich Tern Elegant Tern Black Skimmer Pomarine Jaeger Parasitic Jaeger Red-billed Pigeon White-winged Dove Mourning Dove Inca Dove Common Ground-Dove Plain-breasted Ground-Dove Ruddy Ground-Dove White-tipped Dove White-faced Quail-Dove Orange-fronted Parakeet Orange-chinned Parakeet Squirrel Cuckoo Lesser Ground-Cuckoo Groove-billed Ani Humid forest Dry forest Cloud forest Estuaries & mangroves x x x x x x x x x x x Beaches Freshwater Savannas wetlands Coral reef x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x x x x x x 143 Species English name Megascops cooperi Glaucidium brasilianum Ciccaba virgata Chordeiles acutipennis Nyctidromus albicollis Chaetura vauxi Campylopterus rufus Campylopterus hemileucurus Colibri thalassinus Anthracothorax prevostii Chlorostilbon canivetii Amazilia beryllina Amazilia rutila Lampornis viridipallens Eugenes fulgens Archilochus colubris Trogon melanocephalus Trogon violaceus Trogon elegans Momotus momota Eumomota superciliosa Ceryle alcyon Chloroceryle americana Aulacorhynchus prasinus Melanerpes aurifrons Piculus rubiginosus Pacific Screech-Owl Ferruginous Pygmy-Owl Mottled Owl Lesser Nighthawk Common Pauraque Vaux’s Swift Rufous Sabrewing 144 Violet Sabrewing Green Violet-ear Green-breasted Mango Canivet's Emerald Berylline Hummingbird Cinnamon Hummingbird Green-throated Mountain-gem Magnificent Hummingbird Ruby-throated Hummingbird Black-headed Trogon Violaceous Trogon Elegant Trogon Blue-crowned Motmot Turquoise-browed Motmot Belted Kingfisher Green Kingfisher Emerald Toucanet Golden-fronted Woodpecker Golden-olive Woodpecker Humid forest Cloud forest x Dry forest x x x x x x x x x x x x Estuaries & mangroves x x x x x Beaches Freshwater Savannas wetlands x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x Coral reef Species English name Dryocopus lineatus Xiphorhynchus flavigaster Lepidocolaptes affinis Thamnophilus doliatus Grallaria guatimalensis Camptostoma imberbe Elaenia flavogaster Elaenia frantzii Zimmerius vilissimus Todirostrum cinereum Rhynchocyclus brevirostris Tolmomyias sulphurescens Contopus cinereus Empidonax flaviventris Empidonax traillii Empidonax minimus Empidonax flavescens Myiarchus tuberculifer Myiarchus tyrannulus Pitangus sulphuratus Megarynchus pitangua Myiozetetes similis Myiodynastes luteiventris Tyrannus melancholicus Tyrannus verticalis Tyrannus tyrannus Tyrannus forficatus Lineated Woodpecker Ivory-billed Woodcreeper Spot-crowned Woodcreeper Barred Antshrike Scaled Antpitta Northern Beardless-Tyrannulet Yellow-bellied Elaenia Mountain Elaenia Paltry Tyrannulet Common Tody-Flycatcher Eye-ringed Flatbill Yellow-olive Flycatcher Tropical Pewee Yellow-bellied Flycatcher Willow Flycatcher Least Flycatcher Yellowish Flycatcher Dusky-capped Flycatcher Brown-crested Flycatcher Great Kiskadee Boat-billed Flycatcher Social Flycatcher Sulphur-bellied Flycatcher Tropical Kingbird Western Kingbird Eastern Kingbird Scissor-tailed Flycatcher Humid forest Dry forest x x Cloud forest Estuaries & mangroves x Beaches Freshwater Savannas wetlands Coral reef x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x x x x x 145 Species English name Pachyramphus aglaiae Rose-throated Becard Tityra semifasciata Masked Tityra Vireo pallens Mangrove Vireo Vireo flavifrons Yellow-throated Vireo Vireo solitarius Blue-headed Vireo Vireo gilvus Warbling Vireo Vireo leucophrys Brown-capped Vireo Vireo flavoviridis Yellow-green Vireo Hylophilus decurtatus Lesser Greenlet Cyclarhis gujanensis Rufous-browed Peppershrike Cyanocitta stelleri Steller’s Jay Calocitta formosa White-throated Magpie-Jay Cyanocorax melanocyaneus Bushy-crested Jay Progne chalybea Gray-breasted Martin Tachycineta albilinea Mangrove Swallow Tachycineta thalassina Violet-green Swallow Stelgidopteryx serripennis Northern Rough-winged Swallow Petrochelidon pyrrhonota Cliff Swallow Hirundo rustica Barn Swallow Campylorhynchus rufinucha Rufous-naped Wren Thryothorus maculipectus Spot-breasted Wren Thryothorus rufalbus Rufous-and-white Wren Thryothorus pleurostictus Banded Wren Thryothorus modestus Plain Wren Troglodytes aedon House Wren Troglodytes rufociliatus Rufous-browed Wren Ramphocaenus melanurus Long-billed Gnatwren 146 Humid forest x Dry forest x x Cloud forest Estuaries & mangroves x x x Beaches Freshwater Savannas wetlands x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x x x Coral reef Species English name Polioptila caerulea Polioptila albiloris Myadestes occidentalis Blue-gray Gnatcatcher White-lored Gnatcatcher Brown-backed Solitaire Orange-billed NightingaleThrush Ruddy-capped NightingaleThrush Swainson’s Thrush Black Robin Clay-colored Robin White-throated Robin Tropical Mockingbird Blue-and-white Mockingbird Cedar Waxwing Tennessee Warbler Crescent-chested Warbler Yellow Warbler Catharus aurantiirostris Catharus frantzii Catharus ustulatus Turdus infuscatus Turdus grayi Turdus assimilis Mimus gilvus Melanotis hypoleucus Bombycilla cedrorum Vermivora peregrina Parula superciliosa Dendroica petechia Dendroica petechia xantholora Dendroica magnolia Dendroica virens Dendroica townsendi Mniotilta varia Setophaga ruticilla Helmitheros vermivorum Seiurus aurocapillus Seiurus noveboracensis Humid forest x Dry forest x x Cloud forest Beaches Freshwater Savannas wetlands x x x x x x x x x x x x x x Coral reef x x x x x x x x x x x x x x x x x x x x x x x Mangrove Warbler Magnolia Warbler Black-throated Green Warbler Townsend’s Warbler Black-and-white Warbler American Redstart Worm-eating Warbler Ovenbird Northern Waterthrush Estuaries & mangroves x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds x 147 Species English name Geothlypis trichas Wilsonia pusilla Wilsonia canadensis Myioborus miniatus Euthlypis lachrymosa Basileuterus rufifrons Icteria virens Habia rubica Habia fuscicauda Piranga rubra Piranga ludoviciana Piranga bidentata Piranga leucoptera Thraupis episcopus Thraupis abbas Cyanerpes cyaneus Volatinia jacarina Sporophila torqueola Sporophila minuta Diglossa baritula Melozone leucotis Aimophila ruficauda Ammodramus savannarum Saltator coerulescens Saltator atriceps Pheucticus ludovicianus Cyanocompsa parellina Common Yellowthroat Wilson’s Warbler Canada Warbler Slate-throated Redstart Fan-tailed Warbler Rufous-capped Warbler Yellow-breasted Chat Red-crowned Ant-Tanager Red-throated Ant-Tanager Summer Tanager Western Tanager Flame-colored Tanager White-winged Tanager Blue-gray Tanager Yellow-winged Tanager Red-legged Honeycreeper Blue-black Grassquit White-collared Seedeater Ruddy-breasted Seedeater Cinnamon-bellied Flowerpiercer White-eared Ground-Sparrow Stripe-headed Sparrow Grasshopper Sparrow Grayish Saltator Black-headed Saltator Rose-breasted Grosbeak Blue Bunting 148 Humid forest x x x x x Dry forest Cloud forest Estuaries & mangroves Beaches Freshwater Savannas wetlands x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Biodiversity field inventories in selected watersheds Coral reef Species English name Passerina caerulea Passerina cyanea Passerina ciris Spiza americana Agelaius phoeniceus Sturnella magna Dives dives Quiscalus mexicanus Molothrus aeneus Icterus maculialatus Icterus spurius Icterus pustulatus Icterus pectoralis Icterus gularis Icterus galbula Amblycercus holosericeus Euphonia affinis Euphonia hirundinacea Chlorophonia occipitalis COUNT Blue Grosbeak Indigo Bunting Painted Bunting Dickcissel Red-winged Blackbird Eastern Meadowlark Melodious Blackbird Great-tailed Grackle Bronzed Cowbird Bar-winged Oriole Orchard Oriole Streak-backed Oriole Spot-breasted Oriole Altamira Oriole Baltimore Oriole Yellow-billed Cacique Scrub Euphonia Yellow-throated Euphonia Blue-crowned Chlorophonia Humid forest Dry forest x x x Cloud forest Estuaries & mangroves Beaches Freshwater Savannas wetlands x x x x x Coral reef x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 110 55 71 66 x x x x x 83 106 x 46 Biodiversity field inventories in selected watersheds 11 149 A third option, and one which should be followed in El Salvador, is to use nationally threatened bird species as the indicators for conservation importance of sites (Komar 2002). Komar et al. (2009) recently reevaluated all El Salvador birds to determine the national threat status. They produced a list of 246 nationally threatened species, based on international standards promoted by the World Conservation Union (IUCN 2003), and including migratory species recognized as of conservation importance by any of the several continental wide conservation plans developed for North American migratory birds. This large list of rare and threatened species permits more options for prioritizing areas for conservation than would a very small list of globally threatened or near-threatened bird species (Komar 2002). Not all of the country’s nationally threatened bird species are found in the study area, as some are restricted to other areas such as the pine-oak forests and cloud forests along the Honduran border. Approximately 188 nationally threatened species are useful indicators for prioritizing sites of national conservation importance within the study area (Table 38). These species can be stratified by level of threat. For example, 31 are critically endangered, 69 endangered, and 88 vulnerable. 2.3. SOURCE OF LOCALITY DATA At the start of field work, SalvaNATURA’s flora y fauna database included 21,724 avian locality records from the Project area, including mostly unpublished observations from field notes, but also mist net captures and some museum specimens (Table 39). The 14,679 observation records represented over 76,000 individual birds. During field work, thousands of new locality records were collected, and the database for the project area was increased to 25,792 avian records. Of these, 4,068 were collected during the field study by project staff. The 1,767 new observation records represent 6,083 individual birds. 2.4. FIELD METHODS New information on bird distribution and abundance was collected in the field via observations and net captures. Observations were made during 296 hours by walking through the study areas during most mornings, covering new territory each day as much as possible, noting all birds identified in a field notebook. Observers varied, but generally two people per field trip were assigned to this task, and they worked together, such that each observation hour represents a team of two observers working for one hour. Observations effort was distributed among all of the field sites (Table 40). When possible, species observed were documented by photographs. Birds were captured in standard, black nylon, 36 mm mesh mist nets, measuring 12 m long by 2.6 m high purchased from the Association of Field Ornithologists (USA). The nets were placed at ground-level, in various configurations adapted to the terrain, considered by the field team to be useful for maximizing captures. No attempt was made to standardize mist net effort among sites, since the purpose was to contribute to the advancement of the existing inventory, rather than compare capture rates among sites. Nonetheless, mist-netting effort (net-hours) were recorded so as to better interpret the results from each site (Table 40). Once captured, birds were identified in 150 Biodiversity field inventories in selected watersheds TABLE 38. BIRDS OF NATIONAL CONSERVATION CONCERN (188 SPECIES), USEFUL AS INDICATORS FOR SITE PRIORITIZATION IN THE STUDY AREA. English Family Species Spanish name Status1 name Anatidae Cairina moschata Pato Real Muscovy Duck VU Cracidae Penelope purpurascens Pavo Cojolito Crested Guan CR Cracidae Penelopina nigra Pajuil (Pava Negra) Highland Guan EN Cracidae Crax rubra Hocofaisán (Pajuil) Great Curassow CR Podicipedidae Tachybaptus dominicus Zambullidor Menor Least Grebe EN Podicipedidae Podilymbus podiceps Zambullidor Piquipinto Pied-billed Grebe EN Pelicanidae Pelecanus erythrorhynchos Pelicano Blanco Americano American White Pelican VU Pelicanidae Pelecanus occidentalis Pelicano Café Brown Pelican VU Anhingidae Anhinga anhinga Pato Aguja Anhinga CR Fregatidae Fregata magnificens Fragata Magnificent Frigatebird EN Ardeidae Botaurus pinnatus Avetoro Neotropical Pinnated Bittern CR Ardeidae Ixobrychus exilis Avetorito Americano Least Bittern CR Ardeidae Tigrisoma mexicanum Garza Tigre Gorjinuda Bare-throated Tiger-Heron EN Ardeidae Egretta thula Garza Nivea Snowy Egret EN Ardeidae Egretta caerulea Garza Azul Little Blue Heron EN Ardeidae Egretta tricolor Garza Tricolor Tricolored Heron CR Ardeidae Egretta rufescens Garza Rojiza Reddish Egret VU Ardeidae Nycticorax nycticorax Garza-nocturna Coroninegra Blackcrowned Night-Heron EN Biodiversity field inventories in selected watersheds 151 Family Species Spanish name English name Status1 Ardeidae Nyctanassa violacea Garza-nocturna Coroniclara Yellowcrowned Night-Heron VU Ardeidae Cochlearius cochlearius Garza Cucharón Boat-billed Heron EN Threskiornithidae Eudocimus albus Ibis Blanco White Ibis VU Threskiornithidae Platalea ajaja Espátula Rosada Roseate Spoonbill EN Ciconiidae Mycteria americana Cigueña Americana (Pulido) Wood Stork EN Cathartidae Sarcoramphus papa Zopilote Rey (Rey Zope) King Vulture CR Accipitridae Leptodon cayanensis Milano Cabecigris Gray-headed Kite EN Accipitridae Elanoides forficatus Milano Tijereta Swallow-tailed Kite EN Accipitridae Elanus leucurus Milano Coliblanco White-tailed Kite VU Accipitridae Rostrhamus sociabilis Milano Caracolero Snail Kite EN Accipitridae Ictinia plumbea Milano Plomizo Plumbeous Kite EN Accipitridae Busarellus nigricollis Aguililla Canela Black-collared Hawk CR Accipitridae Geranospiza caerulescens Gavilán Zancudo Crane Hawk CR Accipitridae Leucopternis albicollis Aguililla Blanca White Hawk CR Accipitridae Buteogallus urubitinga Aguililla Negra Mayor Great BlackHawk EN Accipitridae Parabuteo unicinctus Aguililla de Harris Harris’s Hawk EN Accipitridae Buteo swainsoni Aguililla de Swainson Swainson’s Hawk VU Accipitridae Buteo albicaudatus Aguililla Coliblanca White-tailed Hawk CR 152 Biodiversity field inventories in selected watersheds Family Species Spanish name English name Status1 Accipitridae Spizaetus tyrannus Águila Tirana (Águila Crestada Negra) Black HawkEagle CR Accipitridae Spizaetus ornatus Águila Elegante Ornate HawkEagle CR Falconidae Micrastur ruficollis Halcón-selvático Barrado Barred Forest-Falcon EN Falconidae Caracara cheriway Caracara Común (Querque) Crested Caracara VU Falconidae Falco rufigularis Halcón Murcielaguero Bat Falcon CR Rallidae Laterallus ruber Polluela Rojiza Ruddy Crake VU Rallidae Aramides axillaris Rascón Cuellirrufo Rufousnecked Wood-Rail VU Rallidae Aramides cajanea Rascón Cuelligris Gray-necked Wood-Rail CR Rallidae Porzana flaviventer Polluela Pechiamarilla Yellowbreasted Crake EN Rallidae Porphyrio martinica Gallineta Morada Purple Gallinule VU Rallidae Gallinula chloropus Gallineta Común Common Moorhen EN Rallidae Fulica americana Gallareta Americana American Coot CR Aramidae Aramus guarauna Carao (Garza Caracolera) Limpkin VU Burhinidae Burhinus bistriatus Alcaraván Americana Doublestriped Thickknee VU Charadriidae Pluvialis squatarola Chorlo Gris Black-bellied Plover VU Charadriidae Pluvialis dominica Chorlo Dorado Americano American Golden-Plover EN Charadriidae Charadrius alexandrinus Chorlito Niveo Snowy Plover CR Charadriidae Charadrius wilsonia Chorlito Piquigrueso Wilson’s Plover EN Biodiversity field inventories in selected watersheds 153 Family Species Spanish name English name Status1 Charadriidae Charadrius vociferus Chorlito Tildío Killdeer VU Haematopodidae Haematopus palliatus Ostrero Americano American Oystercatcher CR Recurvirostridae Himantopus mexicanus Cadelero Americano (Policía) Black-necked Stilt EN Recurvirostridae Recurvirostra americana Avoceta Americana American Avocet VU Scolopacidae Tringa solitaria Playero Solitario Solitary Sandpiper EN Scolopacidae Tringa incana Playero Vagabundo Wandering Tattler VU Scolopacidae Tringa melanoleuca Patamarilla Mayor Greater Yellowlegs VU Scolopacidae Tringa semipalmata Playero Pihuihui Willet VU Scolopacidae Tringa flavipes Patamarilla Menor Lesser Yellowlegs VU Scolopacidae Bartramia longicauda Zarapito Ganga Upland Sandpiper EN Scolopacidae Numenius phaeopus Zarapito Trinador Whimbrel EN Scolopacidae Numenius americanus Zarapito Piquilargo Long-billed Curlew CR Scolopacidae Limosa fedoa Picopando Canelo Marbled Godwit EN Scolopacidae Arenaria interpres Vuelvepiedras Rojizo Ruddy Turnstone EN Scolopacidae Aphriza virgata Playero de Marejada Surfbird EN Scolopacidae Calidris canutus Playero Gordo Red Knot EN Scolopacidae Calidris alba Playero Blanco Sanderling EN Scolopacidae Calidris pusilla Playerito Semipalmado Semipalmated Sandpiper VU Scolopacidae Calidris mauri Playerito Occidental Western Sandpiper EN 154 Biodiversity field inventories in selected watersheds Family Species Spanish name English name Status1 Scolopacidae Calidris minutilla Playerito Mínimo Least Sandpiper VU Scolopacidae Calidris himantopus Playero Zancudo Stilt Sandpiper VU Scolopacidae Tryngites subruficollis Playerito Pradero Buff-breasted Sandpiper CR Scolopacidae Limnodromus griseus Costurero Piquicorto Short-billed Dowitcher EN Scolopacidae Gallinago delicata Agachona de Wilson Wilson’s Snipe VU Scolopacidae Phalaropus tricolor Falarapo de Wilson Wilson’s Phalarope EN Scolopacidae Phalaropus lobatus Falarapo Cuellirojo Red-necked Phalarope VU Scolopacidae Phalaropus fulicarius Falarapo Piquigrueso Red Phalarope VU Laridae Larus pipixcan Gaviota de Franklin Franklin’s Gull VU Laridae Sternula antillarum Golondrinamarina Mínima Least Tern CR Laridae Gelochelidon nilotica Golondrinamarina Piquigruesa Gull-billed Tern EN Laridae Chlidonias niger Golondrinamarina Negra Black Tern VU Laridae Sterna dougallii Golondrinamarina Rosada Roseate Tern EN Laridae Sterna forsteri Golondrinamarina de Forster Forster’s Tern VU Laridae Thalasseus maximus Golondrinamarina Real Royal Tern VU Laridae Thalasseus elegans Golondrinamarina Elegante Elegant Tern VU Laridae Rynchops niger Rayador Americano Black Skimmer CR Columbidae Columbina minuta Tórtola Pechilisa Plainbreasted Ground-Dove EN Biodiversity field inventories in selected watersheds 155 Family Species Spanish name English name Status1 Columbidae Claravis pretiosa Tórtola Azul Blue GroundDove EN Columbidae Geotrygon albifacies Paloma-perdiz Cariblanca White-faced Quail-Dove VU Columbidae Geotrygon montana Paloma-perdiz Rojiza Ruddy QuailDove VU Psittacidae Aratinga holochlora Perico Gorjirrojo Green Parakeet EN Psittacidae Aratinga strenua Perico Verde Centroamericano Pacific Parakeet VU Psittacidae Amazona albifrons Loro Frentiblanco White-fronted Parrot VU Psittacidae Amazona auropalliata Loro Nuquiamarillo Yellow-naped Parrot EN Cuculidae Coccyzus minor Cuco Manglero Mangrove Cuckoo VU Tytonidae Tyto alba Lechuza de Campanario Barn Owl VU Strigidae Megascops trichopsis Tecolote Bigotudo Whiskered Screech-Owl EN Strigidae Pulsatrix perspicillata Búho de Anteojos Spectacled Owl VU Strigidae Bubo virginianus Búho Cornudo Great Horned Owl EN Strigidae Ciccaba nigrolineata Búho Blanquinegro Black-andwhite Owl CR Caprimulgidae Chordeiles acutipennis Chotacabras Menor Lesser Nighthawk EN Nyctibiidae Nyctibius jamaicensis Bienparado Norteño Northern Potoo VU Apodidae Cypseloides niger Vencejo Negro Black Swift VU Apodidae Aeronautes saxatalis Vencejo Gorjiblanco Whitethroated Swift VU Apodidae Panyptila cayennensis Vencejo-tijereta Menor Lesser Swallow-tailed Swift EN 156 Biodiversity field inventories in selected watersheds Family Species Spanish name English name Status1 Trochilidae Anthracothorax prevostii Mango Pechiverde Greenbreasted Mango VU Trochilidae Abeillia abeillei Colibrí Barbiesmeralda Emeraldchinned Hummingbird VU Trochilidae Hylocharis eliciae Zafiro Gorjiazul Blue-throated Goldentail VU Trochilidae Hylocharis leucotis Colibrí Orejiblanco White-eared Hummingbird VU Trochilidae Lampornis viridipallens Colibrí-serrano Gorjiverde Greenthroated Mountain-gem VU Trochilidae Lampornis amethystinus Colibrí-serrano Gorjiamatisto Amethystthroated Hummingbird VU Trochilidae Eugenes fulgens Colibrí Magnífico Magnificent Hummingbird VU Trochilidae Doricha enicura Tijereta Centroamericana Slender Sheartail EN Trochilidae Tilmatura dupontii Colibrí Colipinto Sparklingtailed Hummingbird EN Trochilidae Atthis ellioti Zumbador Centroamericano Wine-throated Hummingbird EN Trogonidae Trogon melanocephalus Trogón Cabecinegro Black-headed Trogon VU Momotidae Hylomanes momotula Momoto Enano Tody Motmot CR Alcedinidae Ceryle torquatus Martín-pescador Collarejo Ringed Kingfisher CR Alcedinidae Chloroceryle aenea Martín-pescador Enano American Pygmy Kingfisher VU Ramphastidae Aulacorhynchus prasinus Tucaneta Verde (Tucán Verde) Emerald Toucanet VU Ramphastidae Pteroglossus torquatus Tucancillo Collarejo (Pico de Navaja) Collared Aracari VU Biodiversity field inventories in selected watersheds 157 Family Species Spanish name English name Status1 Picidae Veniliornis fumigatus Carpintero Café Smoky-brown Woodpecker EN Picidae Dryocopus lineatus Carpintero Lineado Lineated Woodpecker VU Picidae Campephilus guatemalensis Carpintero Piquiclaro Pale-billed Woodpecker EN Furnariidae Synallaxis erythrothorax Guitío Pechirrufo Rufousbreasted Spinetail EN Furnariidae Dendrocincla homochroa Trepatroncos Rojizo Ruddy Woodcreeper CR Furnariidae Sittasomus griseicapillus Trepatroncos Olivaceo Olivaceous Woodcreeper EN Furnariidae Xiphocolaptes promeropirhynchus Trepatroncos Gigante Strong-billed Woodcreeper EN Furnariidae Dendrocolaptes sanctithomae Trepador Barrado Northern BarredWoodcreeper CR Furnariidae Lepidocolaptes souleyetii Trepatroncos Corona-rayada Streakheaded Woodcreeper VU Formicariidae Grallaria guatimalensis Hormiguerocholino Escamoso Scaled Antpitta EN Tyrannidae Mionectes oleagineus Mosquero Vientre-ocre Ochre-bellied Flycatcher VU Tyrannidae Zimmerius vilissimus Mosquerito Cejiblanco Paltry Tyrannulet VU Tyrannidae Oncostoma cinereigulare Picocurvo Norteño Northern Bentbill VU Tyrannidae Rhynchocyclus brevirostris Picoplano de Anteojos Eye-ringed Flatbill EN Tyrannidae Platyrinchus cancrominus Picochato Rabón Stub-tailed Spadebill CR Tyrannidae Contopus cooperi Pibí Boreal Olive-sided Flycatcher VU Tyrannidae Empidonax traillii Mosquero Saucero Willow Flycatcher VU Tyrannidae Sayornis nigricans Mosquero Negro Black Phoebe EN 158 Biodiversity field inventories in selected watersheds Family Species Spanish name English name Status1 Tyrannidae Attila spadiceus Atila Rabadillabrillante Bright-rumped Attila EN Tyrannidae Myiarchus tyrannulus Copetón Tirano Browncrested Flycatcher EN Incertae sedis Pachyramphus major Cabezón Cuelligris Gray-collared Becard VU Pipridae Chiroxiphia linearis Saltarín Colilargo (Toledo) Long-tailed Manakin VU Vireonidae Vireo pallens Vireo Manglero Mangrove Vireo VU Vireonidae Vireo bellii Vireo de Bell Bell’s Vireo EN Vireonidae Vireo leucophrys Vireo Gorripardo Browncapped Vireo EN Vireonidae Vireolanius pulchellus Vireón Esmeraldo Green ShrikeVireo CR Troglodytidae Salpinctes obsoletus Saltapared Roquero Rock Wren EN Troglodytidae Troglodytes rufociliatus Saltapared Cejirrufo Rufousbrowed Wren VU Sylviidae Ramphocaenus melanurus Soterillo Picudo Long-billed Gnatwren VU Turdidae Sialia sialis Azulejo Gorjicanelo Eastern Bluebird VU EN Turdidae Catharus frantzii Zorzalito de Frantzius Ruddycapped NightingaleThrush Turdidae Hylocichla mustelina Zorzalito Maculado Wood Thrush VU Turdidae Turdus infuscatus Zorzal Negro Black Robin VU Turdidae Turdus assimilis Zorzal Gorjiblanco Whitethroated Robin VU Turdidae Turdus rufitorques Zorzal Cuellirufo Rufouscollared Robin EN Mimidae Melanotis hypoleucus Mulato Pechiblanco Blue-andwhite Mockingbird VU Biodiversity field inventories in selected watersheds 159 160 Family Species Spanish name English name Status1 Parulidae Vermivora pinus Chipe Aliazul Blue-winged Warbler VU Parulidae Vermivora chrysoptera Chipe Alidorado Goldenwinged Warbler EN Parulidae Parula superciliosa Chipe Cejiblanco Crescentchested Warbler VU Parulidae Dendroica petechia xantholora Chipe de Manglar Mangrove Warbler VU Parulidae Dendroica chrysoparia Chipe Caridorado Goldencheeked Warbler CR Parulidae Dendroica occidentalis Chipe Cabeciamarillo Hermit Warbler VU Parulidae Dendroica graciae Chipe de Grace Grace’s Warbler EN Parulidae Protonotaria citrea Chipe Protonotario Prothonotary Warbler VU Parulidae Helmitheros vermivorum Chipe Gusanero Worm-eating Warbler VU Parulidae Oporornis formosus Chipe de Kentucky Kentucky Warbler VU Parulidae Geothlypis poliocephala Mascarita Piquigruesa Gray-crowned Yellowthroat VU Parulidae Wilsonia canadensis Chipe Collarejo Canada Warbler VU Parulidae Cardellina rubrifrons Chipe Carirrojo Red-faced Warbler VU Parulidae Myioborus pictus Pavito Aliblanco Painted Redstart VU Parulidae Basileuterus culicivorus Chipe Coronadorada Goldencrowned Warbler EN Thraupidae Habia rubica Tángarahormiguera Coronirroja Red-crowned Ant-Tanager EN Biodiversity field inventories in selected watersheds Family Species Spanish name English name Status1 Thraupidae Piranga bidentata Tángara Dorsirrayada Flame-colored Tanager VU Emberizidae Amaurospiza concolor Semillero Azul Blue Seedeater EN Diglossa baritula Picaflor Vientrecanelo Cinnamonbellied Flowerpiercer VU Emberizidae Melozone leucotis Rascador Orejiblanco White-eared GroundSparrow VU Cardinalidae Passerina ciris Colorín Sietecolores Painted Bunting EN Cardinalidae Spiza americana Arrocero Americano Dickcissel VU Icteridae Icterus wagleri Bolsero de Wagler Black-vented Oriole EN Icteridae Cacicus melanicterus Cacique Mexicano Yellowwinged Cacique CR Fringillidae Euphonia hirundinacea Eufonia Gorjiamarillo Yellowthroated Euphonia VU Fringillidae Euphonia elegantissima Eufonia Capuchaazul Elegant Euphonia VU Fringillidae Chlorophonia occipitalis Clorofonia Coroniazul Blue-crowned Chlorophonia VU Emberizidae 1 CR=Critically Endangered, EN=Endangered, VU=Vulnerable. Source: Komar et al. (2009). the hand and recorded in the field notes. Each species was documented by digital photographs. Most individuals were released unharmed after capture, although some individuals, generally one or two per species at each site, were collected as voucher specimens and deposited in the scientific collections of the El Salvador Natural History Museum and the Univeristy of Kansas Natural History Museum and Biodiversity Research Center (Lawrence, Kansas, USA). All handling and collecting of birds was carried out with authorization from the Ministry of Environment and Natural Resources. Biodiversity field inventories in selected watersheds 161 TABLE 39. TYPES OF AVIAN LOCALITY RECORDS AVAILABLE FOR THE PRESENT ANALYSIS. IMCW Record type Ahuachapán Sonsonate Santa Ana Project Area Prior to Field Study Museum specimens 467 876 327 1,670 Photographic records 136 8 271 415 Mist net captures 1,708 1,484 1768 4,960 Observational records 10,470 2,122 2087 14,679 Museum specimens 126 266 14 406 Mist net captures 667 547 681 1895 Observational records 491 1197 79 1767 Total record count 14,065 6,500 5,227 25,792 During Field Study Source: SalvaNATURA Flora & Fauna Database. TABLE 40. SAMPLING EFFORT FOR BIRDS AT EACH STUDY SITE DURING 2007 FIELD WORK. Observation Mist-netting Site hours hours Santo Domingo de Guzmán 21 0 Plan de Amayo 50 837 Laguna de las Ninfas 15 1,545 Lavas de Izalco/Finca María Auxiliadora 10 1,940 Barra Salada 35 1,536 Los Cóbanos 63 1,933 Laguna de las Ranas 30 567 Bosque Santa Rita 72 2,366 Totals 296 10,724 162 Biodiversity field inventories in selected watersheds 3. RESULTS AND DISCUSSION 3.1. SPECIES RECORDED At the start of the study (mid 2007), 412 bird species (75% of the 548 species recorded in the country) were recorded within the IMCW project area, including all of the 74 inventory indicator (test) bird species, as well as 167 conservation important species. In general, the Project area was already well inventoried for birds, with recording localities scattered across all 11 watersheds and 16 of the 25 municipalities (Komar 2007). Contemporaneous with this study, one new species was added to the national bird list, and shortly after the field study, two additional species were discovered for the national list. All three were wandering migratory species that were recorded inside the project area: one was a Magellanic Penguin (Spheniscus magellanicus) washed ashore at Barra de Santiago in June 2007 and found by park guards (reported in Jones and Komar 2008); the other was a White-eyed Vireo (Vireo griseus) captured at a bird monitoring station in El Imposible National Park in March 2008, by IMCW project workers (Lety Andino, pers. comm.). The third was a group of Black-vented Shearwaters (Puffinus opisthomelas) found in off-shore waters of Los Cóbanos Protected Natural Area by Ministry of Environment and Natural Resources staff in October 2008 (Ricardo Ibarra, pers. comm.). Studies of marine birds off Los Cóbanos have since discovered at least four additional species for the study area (R. Ibarra, pers. comm.). Although the ornithology team in this study did not detect any new species for El Salvador, they did detect many new species for the project area, and for the Ahuachapán and Sonsonate departments, despite the inventory having already been relatively complete (100% of test species had already been recorded). The proportion of test species recorded serves as a relative indicator, or index, of inventory completeness, but the possibility of finding new, rare species once all of the test species are encountered always exists. The team detected 12 new bird species for the Project Area, bringing the total bird list for the area to 431 species (Table 41), representing 79% of the national list. Biodiversity field inventories in selected watersheds 163 164 Figure 15A.Wine-throated Hummingbird (Atthis ellioti), Laguna de Las Ranas. October 2007. By: Carlos Funes. Figure 15B. Emerald-chinned Hummingbird (Abeillia abeillei), Laguna de Las Ranas. October 2007. By: Carlos Funes. Figure 15C. Great Horned Owl (Bubo virginianus), Plan de Amayo. October 2007. By: Carlos Funes. Figure 15D. Emerald Toucanet (Aulacorhynchus prasinus), Laguna de Las Ranas, October 2007. By: Carlos Funes. Biodiversity field inventories in selected watersheds Figure 15E. Eggs of the White-collared Swift (Streptoprocne zonaris), found in Santo Domingo de Guzmán. July 2007. By: Carlos Funes. Figure 15F. Scaled Antpitta (Grallaria guatimalensis), Laguna de las Ninfas. July 2007. By: Karla Lara. Figure 15G. Rock Wren (Salpinctes obsoletus), an individual captured in the lava field of Volcán de Izalco. August 2007. By: Carlos Funes. Figure 15H. Tricolored Munia (Lonchura malacca), Santa Rita Protected Natural Area. September 2007. By: Carlos Funes. Biodiversity field inventories in selected watersheds 165 166 Figure 15I. Boat-billed Heron (Cochlearius cochlearius), Santa Rita Protected Natural Area, September 2007. By: Carlos Funes. Figure 15J. American Pygmy Kingfisher (Chloroceryle aenea) female, Santa Rita Protected Natural Area, September 2007. By: Carlos Funes. Figure 15K. Rose-throated Becard (Pachyramphus aglaiae) female, Santa Rita Protected Natural Area, November 2007. By: Jorge Jiménez. Figure 15L. Yellow-winged Cacique (Cacicus melanicterus), Los Cóbanos, August 2007. By: Carlos Funes. Biodiversity field inventories in selected watersheds Figure 15M. Wandering Tattler (Tringa incana), Los Cóbanos Protected Natural Area. December 2007. By: Karla Lara. Figure 15N. Pomarine Jaeger (Stercorarius pomarinus), Los Cóbanos Protected Natural Area. December 2007. By: Karla Lara. Figure 15O. Green-breasted Mango (Anthracothorax prevostii), Los Cóbanos. August 2007. By: Carlos Funes. Figure 15P. Purple Gallinule (Porphyrio martinica), Playa El Zope, Los Cóbanos Protected Natural Area. August 2007. By: Carlos Funes. FIGURE 15. SELECTED PHOTOGRAPHS OF BIRDS TAKEN DURING THE FIELD STUDY. Biodiversity field inventories in selected watersheds 167 TABLE 41. LIST OF 431 BIRD SPECIES RECORDED IN THE PROJECT AREA (order follows AOU Checklist, www.aou.org). Species Spanish name English name Seasonality Habitat Elevation Crypturellus cinnamomeus Tinamú Canelo Thicket Tinamou Resident Forest Generalist Lowlands Dendrocygna autumnalis Pijiji Aliblanco (Pishishe) Black-bellied Whistling-Duck Resident Cairina moschata Pato Real Muscovy Duck Resident Anas americana Pato Chalcuan American Wigeon Visitor Anas platyrhynchos Cerceta Cuello Negro Mallard Migratory Vagrant Anas discors Cerceta Aliazul, Pajarilla Blue-winged Teal Visitor Anas clypeata Pato Cucharón Norteño Northern Shoveler Visitor Oxyura jamaicensis Pato Tepalcate Ruddy Duck Status Uncertain Ortalis leucogastra Chachalaca Vientre-blanco White-bellied Chachalaca Resident Forest Generalist Lowlands Penelope purpurascens Pavo Cojolito Crested Guan Resident Forest Specialist Lowlands Penelopina nigra Pajuil (Pava Negra) Highland Guan Resident Forest Specialist Highlands Crax rubra Hocofaisán (Pajuil) Great Curassow Resident Forest Specialist Lowlands Dendrortyx leucophrys Gallina-de-monte Centroamericana Buffy-crowned Wood-Partridge Resident Forest Generalist Highlands 168 Biodiversity field inventories in selected watersheds Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Species Spanish name English name Seasonality Habitat Elevation Colinus cristatus Codorniz-cotui Centroamericana Crested Bobwhite Resident Open Area Generalist Lowlands Dactylortyx thoracicus Codorniz Silbadora Singing Quail Resident Forest Generalist Highlands Tachybaptus dominicus Zambullidor Menor Least Grebe Resident Podilymbus podiceps Zambullidor Piquipinto Pied-billed Grebe Resident and migratory Wetlands Specialist Wetlands Specialist Spheniscus magellanicus Pingüino de Magellanes Magellanic Penguin Migratory Vagrant Marine Waters Lowlands Puffinus pacificus Pardela Colicuña Wedge-tailed Shearwater Visitor Marine Waters Lowlands Puffinus opisthomelas Pardela Mexicana Black-vented Shearwater Visitor Marine Waters Lowlands Puffinus lherminieri Pardela de Audubón Audubon’s Shearwater Visitor Marine Waters Lowlands Phaethon aethereus Rabijunco Piquirrojo Red-billed Tropicbird Non-migratory Vagrant Marine Waters Lowlands Sula granti Bobo Enmascarado de Grant Nazca Booby Visitor Marine Waters Lowlands Sula nebouxii Bobo Patiazul Blue-footed Booby Visitor Marine Waters Lowlands Sula leucogaster Bobo Vientre-blanco Brown Booby Visitor Marine Waters Lowlands Pelecanus erythrorhynchos Pelicano Blanco Americano American White Pelican Visitor Pelecanus occidentalis Pelicano Café Brown Pelican Visitor Biodiversity field inventories in selected watersheds Wetlands Specialist Wetlands Specialist Lowlands Lowlands Lowlands Lowlands 169 Species Spanish name English name Seasonality Phalacrocorax brasilianus Cormorán Neotropical Neotropic Cormorant Resident Anhinga anhinga Pato Aguja Anhinga Resident Fregata magnificens Fragata Magnificent Frigatebird Visitor Ixobrychus exilis Avetorito Americano Least Bittern Resident Tigrisoma mexicanum Garza Tigre Gorjinuda Bare-throated Tiger-Heron Resident Ardea herodias Garzon Cenizo Great Blue Heron Visitor Ardea alba Garza Grande Great Egret Egretta thula Garza Nivea Snowy Egret Egretta caerulea Garza Azul Little Blue Heron Visitor Egretta tricolor Garza Tricolor Tricolored Heron Resident and migratory Egretta rufescens Garza Rojiza Reddish Egret Visitor Bubulcus ibis Garza Ganadera (Garrapatera) Cattle Egret Resident Butorides virescens Garza Verde Green Heron Resident and migratory 170 Resident and migratory Resident and migratory Biodiversity field inventories in selected watersheds Habitat Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Elevation Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Species Spanish name English name Seasonality Nycticorax nycticorax Garza-nocturna Coroninegra Black-crowned Night-Heron Resident Nyctanassa violacea Garza-nocturna Coroniclara Yellow-crowned Night-Heron Visitor Cochlearius cochlearius Garza Cucharón Boat-billed Heron Resident Eudocimus albus Ibis Blanco White Ibis Resident Platalea ajaja Espátula Rosada Roseate Spoonbill Resident Mycteria americana Cigueña Americana (Pulido) Wood Stork Coragyps atratus Zopilote Negro Black Vulture Cathartes aura Aura Cabecirroja Turkey Vulture Cathartes burrovianus Aura Sabanera Lesser Yellow-headed Vulture Sarcoramphus papa Zopilote Rey (Rey Zope) Pandion haliaetus Habitat Elevation Resident and migratory Resident and migratory Resident and migratory Non-migratory Vagrant Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Open Area Generalist Open Area Generalist Open Area Specialist King Vulture Resident Forest Generalist Altitude Generalist Gavilán Pescador Osprey Visitor Wetlands Specialist Lowlands Leptodon cayanensis Milano Cabecigris Gray-headed Kite Resident Forest Specialist Lowlands Chondrohierax uncinatus Milano Piquiganchudo Hook-billed Kite Visitor Forest Generalist Altitude Generalist Biodiversity field inventories in selected watersheds Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Altitude Generalist Altitude Generalist Lowlands 171 Species Spanish name English name Seasonality Habitat Elevation Elanoides forficatus Milano Tijereta Swallow-tailed Kite Transient Forest Generalist Altitude Generalist Elanus leucurus Milano Coliblanco White-tailed Kite Resident Rostrhamus sociabilis Milano Caracolero Snail Kite Status Uncertain Harpagus bidentatus Milano Bidentado Double-toothed Kite Non-migratory Vagrant Forest Generalist Altitude Generalist Ictinia mississippiensis Milano de Misisipi Mississippi Kite Transient Forest Generalist Altitude Generalist Ictinia plumbea Milano Plomizo Plumbeous Kite Breeding Visitor Forest Specialist Lowlands Busarellus nigricollis Aguililla Canela Black-collared Hawk Extinct (Locally) Forest Specialist Lowlands Circus cyaneus Gavilán Rastrero Northern Harrier Visitor Open Area Generalist Lowlands Accipiter striatus Gavilán Pajarero Sharp-shinned Hawk Visitor Forest Generalist Accipiter cooperi Gavilán de Cooper Cooper’s Hawk Visitor Forest Generalist Geranospiza caerulescens Gavilán Zancudo Crane Hawk Resident Forest Generalist Lowlands Leucopternis albicollis Aguililla Blanca White Hawk Resident Forest Specialist Lowlands Buteogallus anthracinus Aguililla Negra Menor Common Black-Hawk Resident Forest Generalist Lowlands 172 Biodiversity field inventories in selected watersheds Open Area Generalist Wetlands Specialist Lowlands Lowlands Altitude Generalist Altitude Generalist Species Spanish name English name Seasonality Habitat Elevation Buteogallus urubitinga Aguililla Negra Mayor Great Black-Hawk Resident Forest Generalist Lowlands Parabuteo unicinctus Aguililla de Harris Harris’s Hawk Resident Buteo magnirostris Aguililla Caminera Roadside Hawk Resident Buteo platypterus Aguililla Aluda Broad-winged Hawk Visitor Forest Generalist Highlands Buteo nitidus Aguililla Gris Gray Hawk Resident Forest Generalist Lowlands Buteo brachyurus Aguililla Colicorta Short-tailed Hawk Resident and migratory Forest Generalist Altitude Generalist Buteo swainsoni Aguililla de Swainson Swainson’s Hawk Transient Buteo albicaudatus Aguililla Coliblanca White-tailed Hawk Resident Buteo albonotatus Aguililla Aura Zone-tailed Hawk Resident and migratory Open Area Generalist Lowlands Buteo jamaicensis Aguililla Colirroja Red-tailed Hawk Resident and migratory Forest Generalist Highlands Spizaetus tyrannus Águila Tirana (Águila Crestada Negra) Black Hawk-Eagle Resident Forest Generalist Altitude Generalist Spizaetus ornatus Águila Elegante Ornate Hawk-Eagle Extinct (Locally) Forest Specialist Lowlands Micrastur ruficollis Halcón-selvático Barrado Barred Forest-Falcon Resident Forest Generalist Altitude Generalist Micrastur semitorquatus Halcón-selvático Barrado Collared Forest-Falcon Resident Forest Generalist Lowlands Biodiversity field inventories in selected watersheds Open Area Specialist Open Area Generalist Open Area Generalist Open Area Generalist Lowlands Lowlands Lowlands Lowlands 173 Species Spanish name English name Seasonality Habitat Elevation Caracara cheriway Caracara Común (Querque) Crested Caracara Resident Open Area Generalist Lowlands Herpetotheres cachinnans Halcón Guaco (Guas) Laughing Falcon Resident Forest Generalist Lowlands Falco sparverius Cernícalo Américano American Kestrel Resident and migratory Open Area Specialist Altitude Generalist Falco columbarius Esmerejón Merlin Visitor Open Area Generalist Lowlands Falco rufigularis Halcón Murcielaguero Bat Falcon Resident Open Area Generalist Lowlands Falco peregrinus Halcón Peregrino Peregrine Falcon Visitor Open Area Generalist Lowlands Aramides axillaris Rascón Cuellirrufo Rufous-necked Wood-Rail Resident Forest Specialist Aramides cajanea Rascón Cuelligris Gray-necked Wood-Rail Resident Wetlands Specialist Porzana carolina Polluela Sora Sora Visitor Wetlands Specialist Lowlands Porphyrio martinica Gallineta Morada Purple Gallinule Resident Wetlands Specialist Lowlands Gallinula chloropus Gallineta Común Common Moorhen Resident and migratory Wetlands Specialist Lowlands Fulica americana Gallareta Americana American Coot Resident and migratory Wetlands Specialist Altitude Generalist 174 Biodiversity field inventories in selected watersheds Altitude Generalist Altitude Generalist Species Spanish name English name Seasonality Habitat Elevation Heliornis fulica Pájaro-cantil Sungrebe Non-migratory Vagrant Wetlands Specialist Lowlands Aramus guarauna Carao (Garza Caracolera) Limpkin Resident Wetlands Specialist Lowlands Burhinus bistriatus Alcaraván Americana Double-striped Thick-knee Resident Pluvialis squatarola Chorlo Gris Black-bellied Plover Visitor Pluvialis dominica Chorlo Dorado Americano American Golden-Plover Transient Charadrius collaris Chorlito Collarejo Collared Plover Visitor Charadrius alexandrinus Chorlito Niveo Snowy Plover Visitor Charadrius wilsonia Chorlito Piquigrueso Wilson’s Plover Resident and migratory Charadrius semipalmatus Chorlo Semipalmado Semipalmated Plover Visitor Charadrius vociferus Chorlito Tildío Killdeer Visitor Haematopus palliatus Ostrero Americano American Oystercatcher Resident Himantopus mexicanus Cadelero Americano (Policía) Black-necked Stilt Resident Jacana spinosa Jacana Mesoamericana Northern Jacana Resident Biodiversity field inventories in selected watersheds Open Area Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands 175 176 Species Spanish name English name Seasonality Actitis macularius Playero Alzacolita (Alzaculito) Spotted Sandpiper Visitor Tringa solitaria Playero Solitario Solitary Sandpiper Visitor Tringa incana Playero Vagabundo Wandering Tattler Visitor Tringa melanoleuca Patamarilla Mayor Greater Yellowlegs Visitor Tringa semipalmata Playero Pihuihui Willet Visitor Tringa flavipes Patamarilla Menor Lesser Yellowlegs Visitor Bartramia longicauda Zarapito Ganga Upland Sandpiper Transient Numenius phaeopus Zarapito Trinador Whimbrel Visitor Numenius americanus Zarapito Piquilargo Long-billed Curlew Visitor Limosa fedoa Picopando Canelo Marbled Godwit Visitor Arenaria interpres Vuelvepiedras Rojizo Ruddy Turnstone Visitor Aphriza virgata Playero de Marejada Surfbird Transient Calidris canutus Playero Gordo Red Knot Visitor Biodiversity field inventories in selected watersheds Habitat Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Open Area Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Elevation Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Species Spanish name English name Seasonality Habitat Elevation Calidris alba Playero Blanco Sanderling Visitor Wetlands Specialist Lowlands Calidris pusilla Playerito Semipalmado Semipalmated Sandpiper Visitor Wetlands Specialist Lowlands Calidris mauri Playerito Occidental Western Sandpiper Visitor Calidris minutilla Playerito Mínimo Least Sandpiper Visitor Calidris melanotos Playero Pectoral Pectoral Sandpiper Transient Calidris himantopus Playero Zancudo Stilt Sandpiper Transient Limnodromus griseus Costurero Piquicorto Short-billed Dowitcher Visitor Gallinago delicata Agachona de Wilson Wilson’s Snipe Visitor Phalaropus lobatus Falarapo Cuellirojo Red-necked Phalarope Visitor Marine Waters Lowlands Phalaropus fulicarius Falarapo Piquigrueso Red Phalarope Migratory Vagrant Marine Waters Lowlands Leucophaeus atricilla Gaviota Reidora Laughing Gull Visitor Leucophaeus pipixcan Gaviota de Franklin Franklin’s Gull Transient Larus delawarensis Gaviota Piquianillada Ring-billed Gull Migratory Vagrant Biodiversity field inventories in selected watersheds Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Wetlands Specialist Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands 177 Species Spanish name English name Seasonality Chlidonias niger Golondrina-marina Negra Black Tern Transient Sterna dougallii Golondrina-marina Rosada Roseate Tern Migratory Vagrant Sterna hirundo Golondrina-marina Común Common Tern Visitor Thalasseus maximus Golondrina-marina Real Royal Tern Visitor Thalasseus sandvicensis Golondrina-marina de Sandwich Sandwich Tern Visitor Marine Waters Lowlands Thalasseus elegans Golondrina-marina Elegante Elegant Tern Rayador Americano Black Skimmer Stercorarius pomarinus Salteador Pomarino Paloma Doméstica (Paloma de Castillo) Pomarine Jaeger Rock Pigeon Resident Marine Waters Wetlands Specialist Marine Waters Open Area Generalist Lowlands Rynchops niger Transient Resident and migratory Transient Patagioenas flavirostris Paloma Morada Red-billed Pigeon Resident Forest Generalist Patagioenas fasciata Paloma Encinera Band-tailed Pigeon Zenaida asiatica Paloma Aliblanca White-winged Dove Resident Resident and migratory Zenaida macroura Paloma Huilota Mourning Dove Visitor Columbina inca Tórtola Colilarga Inca Dove Resident Columbina passerina Tórtola Común Common Ground-Dove Resident Columbina minuta Tórtola Pechilisa Plain-breasted Ground-Dove Resident Forest Specialist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Specialist Open Area Specialist Columba livia 178 Biodiversity field inventories in selected watersheds Habitat Wetlands Specialist Marine Waters Wetlands Specialist Wetlands Specialist Elevation Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Altitude Generalist Altitude Generalist Highlands Altitude Generalist Lowlands Lowlands Lowlands Lowlands Species Spanish name English name Seasonality Columbina talpacoti Tortolita Rojiza Ruddy Ground-Dove Resident Claravis pretiosa Leptotila verreauxi Geotrygon albifacies Geotrygon montana Tórtola Azul Paloma Arroyera (Rodadora) Paloma-perdiz Cariblanca Paloma-perdiz Rojiza Blue Ground-Dove White-tipped Dove White-faced Quail-Dove Ruddy Quail-Dove Resident Resident Resident Resident Open Area Generalist Forest Specialist Forest Generalist Forest Generalist Forest Generalist Aratinga holochlora Perico Gorjirrojo Green Parakeet Status Uncertain Forest Specialist Aratinga strenua Perico Verde Centroamericano Pacific Parakeet Resident Forest Generalist Aratinga canicularis Perico Frentinaranja Orange-fronted Parakeet Resident Forest Generalist Brotogeris jugularis Amazona albifrons Amazona auropalliata Piaya cayana Coccyzus americanus Periquito Barbinaranja Loro Frentiblanco Loro Nuquiamarillo Chocolatero, Piscoy Cuco Piquiamarillo Orange-chinned Parakeet White-fronted Parrot Yellow-naped Parrot Squirrel Cuckoo Yellow-billed Cuckoo Forest Generalist Forest Generalist Forest Generalist Forest Generalist Forest Generalist Coccyzus minor Cuco Manglero Mangrove Cuckoo Tapera naevia Cuco Rayado Striped Cuckoo Resident Resident Resident Resident Transient Resident and migratory Resident Forest Generalist Lowlands Lowlands Highlands Lowlands Altitude Generalist Altitude Generalist Altitude Generalist Lowlands Lowlands Lowlands Lowlands Lowlands Altitude Generalist Lowlands Morococcyx erythropygus Cuco-terrestre Menor Lesser Ground-Cuckoo Resident Forest Generalist Lowlands Geococcyx velox Correcaminos Menor Lesser Roadrunner Resident Crotophaga sulcirostris Pijuyo Groove-billed Ani Resident Tyto alba Lechuza de Campanario Barn Owl Resident Biodiversity field inventories in selected watersheds Habitat Forest Generalist Open Area Generalist Open Area Generalist Open Area Generalist Elevation Lowlands Highlands Altitude Generalist Lowlands 179 Species Spanish name English name Seasonality Habitat Elevation Megascops cooperi Megascops trichopsis Pulsatrix perspicillata Tecolote de Cooper Tecolote Bigotudo Búho de Anteojos Pacific Screech-Owl Whiskered Screech-Owl Spectacled Owl Resident Resident Resident Forest Generalist Forest Specialist Forest Specialist Bubo virginianus Búho Cornudo Great Horned Owl Resident Forest Generalist Glaucidium brasilianum Tecolotito Común (Aurorita) Ferruginous Pygmy-Owl Resident Forest Generalist Ciccaba virgata Búho Café Mottled Owl Resident Forest Generalist Ciccaba nigrolineata Búho Blanquinegro Black-and-white Owl Resident Pseudoscops clamator Búho-cornudo Cariblanco Striped Owl Resident Chordeiles acutipennis Chotacabras Menor Lesser Nighthawk Resident and migratory Common Pauraque Resident Chuck-will’s-widow Forest Specialist Open Area Specialist Open Area Specialist Open Area Generalist Forest Generalist Lowlands Highlands Lowlands Altitude Generalist Lowlands Altitude Generalist Lowlands Caprimulgus carolinensis Tapacaminos Pucuyo (Caballero) Tapacaminos Carolinense Caprimulgus vociferus Tapacaminos Cuerprihuio Whip-poor-will Nyctibius jamaicensis Bienparado Norteño Northern Potoo Transient Resident and migratory Resident Cypseloides niger Vencejo Negro Black Swift Status Uncertain Streptoprocne rutila Vencejo Cuellicastaño Chestnut-collared Swift Resident Streptoprocne zonaris Vencejo Cuelliblanco White-collared Swift Resident Chaetura vauxi Vencejo de Vaux Vaux’s Swift Resident Nyctidromus albicollis 180 Biodiversity field inventories in selected watersheds Forest Generalist Forest Generalist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Lowlands Lowlands Lowlands Lowlands Altitude Generalist Lowlands Highlands Altitude Generalist Lowlands Altitude Generalist Species Spanish name English name Seasonality Aeronautes saxatalis Vencejo Gorjiblanco White-throated Swift Visitor Panyptila cayennensis Vencejo-tijereta Menor Lesser Swallow-tailed Swift Resident Panyptila sanctihieronymi Vencejo-tijereta Mayor Great Swallow-tailed Swift Resident Campylopterus rufus Fandanguero Rufo Rufous Sabrewing Resident Open Area Generalist Open Area Generalist Open Area Generalist Forest Specialist Campylopterus hemileucurus Fandanguero Morado Violet Sabrewing Resident Forest Generalist Florisuga mellivora Jacobino Nuquiblanco White-necked Jacobin Non-migratory Vagrant Forest Generalist Lowlands Colibri thalassinus Orejavioleta Verde Green Violetear Resident Open Area Specialist Highlands Anthracothorax prevostii Mango Pechiverde Green-breasted Mango Resident and migratory Forest Generalist Lowlands Abeillia abeillei Colibrí Barbiesmeralda Resident Forest Specialist Highlands Chlorostilbon canivetii Hylocharis eliciae Amazilia beryllina Esmeralda de Canivet Zafiro Gorjiazul Colibrí de Berilo Emerald-chinned Hummingbird Canivet's Emerald Blue-throated Goldentail Berylline Hummingbird Resident Resident Resident Lowlands Lowlands Lowlands Amazilia rutila Colibrí Canelo Cinnamon Hummingbird Resident Lampornis viridipallens Colibrí-serrano Gorjiverde Resident Highlands Lampornis amethystinus Colibrí-serrano Gorjiamatisto Green-throated Mountain-gem Amethyst-throated Hummingbird Forest Generalist Forest Specialist Forest Generalist Open Area Generalist Forest Generalist Resident Forest Generalist Highlands Eugenes fulgens Colibrí Magnífico Magnificent Hummingbird Resident Open Area Specialist Highlands Biodiversity field inventories in selected watersheds Habitat Elevation Highlands Lowlands Highlands Highlands Altitude Generalist Lowlands 181 Species Spanish name English name Seasonality Habitat Elevation Heliomaster longirostris Picolargo Coroniazul Long-billed Starthroat Resident Lowlands Heliomaster constantii Picolargo Coronioscuro Plain-capped Starthroat Resident Doricha enicura Tijereta Centroamericana Slender Sheartail Status Uncertain Tilmatura dupontii Colibrí Colipinto Sparkling-tailed Hummingbird Status Uncertain Forest Generalist Open Area Generalist Open Area Specialist Open Area Specialist Archilochus colubris Colibrí Gorjirrubi Ruby-throated Hummingbird Visitor Atthis ellioti Zumbador Centroamericano Wine-throated Hummingbird Resident Selasphorus platycercus Zumbador Coliancho Broad-tailed Hummingbird Trogon melanocephalus Hylomanes momotula Trogón Cabecinegro Trogón Violaceo (Coa Pechiamarilla) Trogón Elegante (Coa Pechiroja) Momoto Enano Momotus momota Highlands Highlands Altitude Generalist Black-headed Trogon Non-migratory Vagrant Resident Open Area Specialist Open Area Generalist Forest Generalist Violaceous Trogon Resident Forest Generalist Lowlands Elegant Trogon Resident Forest Generalist Lowlands Tody Motmot Resident Forest Specialist Momoto Coroniazul (Talapo) Blue-crowned Motmot Resident Forest Generalist Lowlands Altitude Generalist Eumomota superciliosa Momoto Cejiturqueza (Torogoz) Turquoise-browed Motmot Resident Megaceryle torquata Martín-pescador Collarejo Ringed Kingfisher Resident Megaceryle alcyon Martín-pescador Norteño Belted Kingfisher Visitor Trogon violaceus Trogon elegans 182 Forest Generalist Lowlands Biodiversity field inventories in selected watersheds Open Area Generalist Wetlands Specialist Wetlands Specialist Highlands Highlands Lowlands Lowlands Lowlands Lowlands Species Spanish name English name Seasonality Habitat Elevation Chloroceryle amazona Martín-pescador Amazona Amazon Kingfisher Resident Chloroceryle americana Martín-pescador Verde Green Kingfisher Resident Chloroceryle aenea Martín-pescador Enano American Pygmy Kingfisher Resident Wetlands Specialist Wetlands Specialist Forest Generalist Notharchus macrorhynchos Buco Collarejo White-necked Puffbird Status Uncertain Forest Specialist Lowlands Aulacorhynchus prasinus Tucaneta Verde (Tucán Verde) Emerald Toucanet Resident Forest Generalist Highlands Lowlands Lowlands Lowlands Collared Aracari Resident Forest Generalist Lowlands Melanerpes formicivorus Tucancillo Collarejo (Pico de Navaja) Carpintero Arlequín Acorn Woodpecker Resident Forest Specialist Melanerpes aurifrons Carpintero Frentidorado Golden-fronted Woodpecker Resident Forest Generalist Sphyrapicus varius Chupasavia Vientre-amarillo Yellow-bellied Sapsucker Visitor Forest Generalist Veniliornis fumigatus Carpintero Café Smoky-brown Woodpecker Resident Forest Specialist Colaptes rubiginosus Carpintero Olivaceo Golden-olive Woodpecker Resident Forest Generalist Dryocopus lineatus Carpintero Lineado Lineated Woodpecker Resident Forest Generalist Highlands Altitude Generalist Altitude Generalist Lowlands Altitude Generalist Altitude Generalist Campephilus guatemalensis Carpintero Piquiclaro Pale-billed Woodpecker Resident Forest Specialist Lowlands Synallaxis erythrothorax Guitío Pechirrufo Rufous-breasted Spinetail Resident Open Area Specialist Lowlands Dendrocincla homochroa Trepatroncos Rojizo Ruddy Woodcreeper Resident Forest Specialist Lowlands Sittasomus griseicapillus Trepatroncos Olivaceo Olivaceous Woodcreeper Forest Generalist Lowlands Glyphorynchus spirurus Trepatroncos Piquicuna Wedge-billed Woodcreeper Resident Non-migratory Vagrant Forest Specialist Lowlands Pteroglossus torquatus Biodiversity field inventories in selected watersheds 183 Species Spanish name English name Seasonality Habitat Elevation Xiphocolaptes promeropirhynchus Trepatroncos Gigante Strong-billed Woodcreeper Resident Forest Specialist Highlands Dendrocolaptes sanctithomae Trepatroncos Barrado Northern Barred-Woodcreeper Resident Forest Specialist Lowlands Xiphorhynchus flavigaster Trepatroncos Piquiclaro Ivory-billed Woodcreeper Resident Forest Generalist Lowlands Lepidocolaptes souleyetii Trepatroncos Corona-rayada Streak-headed Woodcreeper Resident Forest Generalist Lowlands Lepidocolaptes affinis Trepatroncos Corona-punteada Spot-crowned Woodcreeper Resident Forest Generalist Highlands Thamnophilus doliatus Batará Barrada Barred Antshrike Resident Forest Generalist Altitude Generalist Grallaria guatimalensis Hormiguero-cholino Escamoso Scaled Antpitta Resident Forest Generalist Highlands Camptostoma imberbe Mosquerito Lampiño Norteño Northern Beardless-Tyrannulet Resident Forest Specialist Lowlands Myiopagis viridicata Elenia Verdosa Greenish Elaenia Resident Lowlands Elaenia flavogaster Mosquero Elenia Ventriamarillo Yellow-bellied Elaenia Resident Elaenia frantzii Mionectes oleagineus Elenia Serrana Mosquero Vientre-ocre Mountain Elaenia Ochre-bellied Flycatcher Resident Resident Forest Generalist Open Area Generalist Forest Generalist Forest Generalist Zimmerius vilissimus Mosquerito Cejiblanco Paltry Tyrannulet Resident Forest Generalist Oncostoma cinereigulare Todirostrum cinereum Picocurvo Norteño Espatulilla Común Northern Bentbill Common Tody-Flycatcher Resident Resident Forest Specialist Forest Specialist Rhynchocyclus brevirostris Picoplano de Anteojos Eye-ringed Flatbill Resident Forest Specialist Tolmomyias sulphurescens Picoplano Ojiblanco Yellow-olive Flycatcher Resident Forest Generalist Lowlands Platyrinchus cancrominus Picochato Rabón Stub-tailed Spadebill Resident Forest Specialist Lowlands Onychorhynchus coronatus Mosquero Real Royal Flycatcher Status Uncertain Forest Specialist Lowlands 184 Biodiversity field inventories in selected watersheds Lowlands Highlands Lowlands Altitude Generalist Lowlands Lowlands Altitude Generalist Species Spanish name English name Seasonality Habitat Contopus cooperi Pibí Boreal Olive-sided Flycatcher Transient Forest Generalist Contopus sordidulus Pibí Occidental Western Wood-Pewee Transient Forest Generalist Contopus virens Pibí Oriental Eastern Wood-Pewee Forest Generalist Contopus cinereus Pibí Tropical Tropical Pewee Forest Generalist Lowlands Empidonax flaviventris Mosquero Vientre-amarillo Yellow-bellied Flycatcher Transient Resident and migratory Visitor Altitude Generalist Altitude Generalist Lowlands Forest Generalist Empidonax alnorum Mosquero Ailero Alder Flycatcher Transient Forest Generalist Lowlands Altitude Generalist Empidonax traillii Mosquero Saucero Willow Flycatcher Visitor Empidonax albigularis Mosquero Gorjiblanco White-throated Flycatcher Status Uncertain Empidonax minimus Empidonax hammondii Empidonax flavescens Empidonax fulvifrons Sayornis nigricans Attila spadiceus Mosquero Mínimo Mosquero de Hammond Mosquero Amarillento Mosquero Pechicanelo Mosquero Negro Atila Rabadilla-brillante Least Flycatcher Hammond's Flycatcher Yellowish Flycatcher Buff-breasted Flycatcher Black Phoebe Bright-rumped Attila Visitor Visitor Resident Resident Resident Resident Open Area Generalist Open Area Specialist Forest Generalist Forest Specialist Forest Generalist Forest Specialist Especialista ríos Forest Generalist Myiarchus tuberculifer Copetón Triste Dusky-capped Flycatcher Resident Forest Generalist Myiarchus nuttingi Copetón de Nutting Nutting's Flycatcher Resident Myiarchus crinitus Copetón Viajero Great Crested Flycatcher Myiarchus tyrannulus Copetón Tirano Brown-crested Flycatcher Visitor Resident and migratory Biodiversity field inventories in selected watersheds Open Area Generalist Forest Generalist Forest Specialist Elevation Lowlands Highlands Lowlands Highlands Highlands Highlands Lowlands Lowlands Altitude Generalist Lowlands Lowlands Lowlands 185 Species Spanish name English name Seasonality Habitat Elevation Pitangus sulphuratus Luis Grande (Cristo Fue) Great Kiskadee Resident Open Area Generalist Lowlands Megarynchus pitangua Luis Piquigrueso Boat-billed Flycatcher Resident Forest Generalist Myiozetetes similis Luis Gregario (Chio) Social Flycatcher Resident Forest Generalist Altitude Generalist Lowlands Myiodynastes luteiventris Papamoscas Vientre-amarillo Sulphur-bellied Flycatcher Breeding Visitor Forest Generalist Lowlands Tyrannus melancholicus Tirano Tropical Tropical Kingbird Resident Tyrannus verticalis Tirano Occidental Western Kingbird Visitor Tyrannus tyrannus Tirano Viajero Eastern Kingbird Transient Tyrannus forficatus Tirano-tijereta Rosado Scissor-tailed Flycatcher Visitor Pachyramphus major Pachyramphus aglaiae Cabezón Cuelligris Cabezón Degollado Gray-collared Becard Rose-throated Becard Resident Resident Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Forest Specialist Forest Generalist Tityra semifasciata Titira Enmascarada (Torreja) Masked Tityra Resident Forest Generalist Chiroxiphia linearis Saltarín Colilargo (Toledo) Long-tailed Manakin Forest Specialist Pipra mentalis Saltarín Cabecirrojo Red-capped Manakin Forest Generalist Lowlands Vireo griseus Vireo pallens Vireo Ojoblanco Vireo Manglero White-eyed Vireo Mangrove Vireo Resident Non-migratory Vagrant Migratory Vagrant Resident Highlands Lowlands Altitude Generalist Lowlands Lowlands Lowlands Vireo bellii Vireo de Bell Bell’s Vireo Visitor Vireo flavifrons Vireo plumbeus Vireo Gorgiamarillo Vireo Plomizo Yellow-throated Vireo Plumbeous Vireo Visitor Status Uncertain Forest Generalist Forest Specialist Open Area Generalist Forest Generalist Forest Specialist 186 Biodiversity field inventories in selected watersheds Lowlands Lowlands Altitude Generalist Lowlands Lowlands Lowlands Highlands Species Spanish name English name Seasonality Habitat Vireo solitarius Vireo Solitario Blue-headed Vireo Visitor Forest Generalist Vireo gilvus Vireo Gorjeador Warbling Vireo Visitor Forest Generalist Vireo leucophrys Vireo philadelphicus Vireo Gorripardo Vireo Filadelfia Brown-capped Vireo Philadelphia Vireo Resident Visitor Forest Specialist Forest Generalist Vireo olivaceus Vireo Ojirrojo Red-eyed Vireo Transient Forest Generalist Vireo flavoviridis Hylophilus decurtatus Vireolanius pulchellus Vireo Amarillo-verdoso Verdillo Menor Vireón Esmeraldo Yellow-green Vireo Lesser Greenlet Green Shrike-Vireo Breeding Visitor Resident Resident Forest Generalist Forest Generalist Forest Specialist Cyclarhis gujanensis Vireón Cejirrufo Rufous-browed Peppershrike Resident Forest Generalist Calocitta formosa Urraca-hermosa Cariblanca (Urraca) White-throated Magpie-Jay Resident Open Area Generalist Altitude Generalist Altitude Generalist Highlands Highlands Altitude Generalist Lowlands Lowlands Lowlands Altitude Generalist Altitude Generalist Cyanocorax melanocyaneus Chara Centroamericana Bushy-crested Jay Resident Forest Generalist Highlands Cyanolyca pumilo Chara de Niebla Black-throated Jay Resident Progne chalybea Martín Pechigris Gray-breasted Martin Resident Highlands Altitude Generalist Tachycineta bicolor Golondrina Arbolera Tree Swallow Visitor Tachycineta albilinea Golondrina Manglera Mangrove Swallow Resident Tachycineta thalassina Golondrina Cariblanca Violet-green Swallow Visitor Stelgidopteryx serripennis Golondrina-aliserrada Norteña Northern Rough-winged Swallow Resident Forest Specialist Open Area Generalist Open Area Generalist Wetlands Specialist Open Area Generalist Open Area Generalist Biodiversity field inventories in selected watersheds Elevation Lowlands Lowlands Altitude Generalist Altitude Generalist 187 Species Spanish name English name Seasonality Riparia riparia Golondrina Riverena Bank Swallow Transient Petrochelidon pyrrhonota Golondrina Risquera Cliff Swallow Transient Petrochelidon fulva Golondrina Pueblera Cave Swallow Visitor Hirundo rustica Golondrina Ranchera Barn Swallow Visitor Campylorhynchus rufinucha Guacalchía (Matraquita Nuquirrufa) Rufous-naped Wren Resident Salpinctes obsoletus Saltapared Roquero Rock Wren Resident Thryothorus maculipectus Saltapared Pechimanchado Spot-breasted Wren Resident Forest Specialist Lowlands Thryothorus rufalbus Saltapared Rufiblanco Rufous-and-white Wren Resident Forest Generalist Altitude Generalist Thryothorus pleurostictus Saltapared Vientre-barrado Banded Wren Resident Forest Specialist Lowlands Thryothorus modestus Saltapared Sencillo Plain Wren Resident Open Area Generalist Troglodytes aedon Saltapared Continental Norteño House Wren Resident Forest Generalist Troglodytes rufociliatus Saltapared Cejirrufo Rufous-browed Wren Resident Forest Specialist Altitude Generalist Altitude Generalist Highlands Henicorhina leucophrys Saltapared-selvático Pechigris Gray-breasted Wood-Wren Resident Forest Specialist Highlands Ramphocaenus melanurus Soterillo Picudo Long-billed Gnatwren Resident Forest Specialist Lowlands Polioptila caerulea Polioptila albiloris Sialia sialis Perlita Grisilla Perlita Cejiblanca Azulejo Gorjicanelo Blue-gray Gnatcatcher White-lored Gnatcatcher Eastern Bluebird Visitor Resident Resident Forest Generalist Forest Generalist Forest Specialist Lowlands Lowlands Highlands 188 Biodiversity field inventories in selected watersheds Habitat Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Specialist Elevation Lowlands Lowlands Lowlands Altitude Generalist Lowlands Altitude Generalist Species Spanish name English name Seasonality Habitat Elevation Myadestes occidentalis Clarín Jilguero (Guardabarranco) Brown-backed Solitaire Resident Forest Generalist Highlands Catharus aurantiirostris Zorzalito Piquinaranja Resident Forest Generalist Highlands Catharus frantzii Zorzalito de Frantzius Resident Forest Specialist Highlands Catharus ustulatus Zorzalito de Swainson Swainson’s Thrush Visitor Forest Generalist Catharus guttatus Zorzalito Colirrufo Hermit Thrush Migratory Vagrant Forest Generalist Hylocichla mustelina Zorzalito Maculado Wood Thrush Visitor Forest Generalist Turdus infuscatus Zorzal Negro Black Thrush Resident Forest Specialist Turdus grayi Zorzal Pardo Clay-colored Thrush Resident Forest Generalist Turdus assimilis Zorzal Gorjiblanco White-throated Thrush Resident Turdus rufitorques Zorzal Cuellirufo Rufous-collared Robin Resident Forest Specialist Open Area Specialist Dumetella carolinensis Pájaro-gato Gris Gray Catbird Migratory Vagrant Mimus gilvus Cenzontle Sureño Tropical Mockingbird Resident Melanotis hypoleucus Mulato Pechiblanco Blue-and-white Mockingbird Resident Anthus rubescens Bisbita Americana American Pipit Migratory Vagrant Bombycilla cedrorum Ampelis Americano Cedar Waxwing Visitor Forest Generalist Vermivora pinus Chipe Aliazul Blue-winged Warbler Visitor Forest Generalist Orange-billed NightingaleThrush Ruddy-capped NightingaleThrush Biodiversity field inventories in selected watersheds Forest Generalist Open Area Generalist Forest Specialist Open Area Generalist Altitude Generalist Highlands Altitude Generalist Highlands Altitude Generalist Highlands Highlands Altitude Generalist Lowlands Highlands Altitude Generalist Altitude Generalist Highlands 189 Species Spanish name English name Seasonality Habitat Elevation Vermivora chrysoptera Chipe Alidorado Golden-winged Warbler Visitor Forest Generalist Vermivora peregrina Chipe Peregrino Tennessee Warbler Visitor Forest Generalist Highlands Altitude Generalist Vermivora celata Chipe Corona-naranja Orange-crowned Warbler Migratory Vagrant Vermivora ruficapilla Chipe de Nashville Nashville Warbler Visitor Parula superciliosa Parula americana Chipe Cejiblanco Parula Norteña Crescent-chested Warbler Northern Parula Parula pitiayumi Parula Tropical Tropical Parula Dendroica petechia Chipe Amarillo Yellow Warbler Resident Migratory Vagrant Non-migratory Vagrant Visitor Dendroica petechia xantholora Chipe de Manglar Mangrove Warbler Dendroica pensylvanica Chipe Flanquicastaño Dendroica magnolia 190 Open Area Generalist Open Area Generalist Forest Specialist Forest Generalist Highlands Highlands Forest Generalist Highlands Lowlands Altitude Generalist Lowlands Resident Forest Specialist Lowlands Chestnut-sided Warbler Migratory Vagrant Forest Generalist Chipe de Magnolia Magnolia Warbler Visitor Forest Generalist Dendroica tigrina Chipe Atigrado Cape May Warbler Migratory Vagrant Forest Generalist Dendroica caerulescens Chipe Azuloso Black-throated Blue Warbler Migratory Vagrant Dendroica chrysoparia Chipe Caridorado Golden-cheeked Warbler Visitor Open Area Generalist Forest Specialist Dendroica virens Chipe Dorsiverde Black-throated Green Warbler Visitor Forest Generalist Dendroica townsendi Dendroica occidentalis Chipe de Townsend Chipe Cabeciamarillo Townsend’s Warbler Hermit Warbler Visitor Visitor Forest Generalist Forest Specialist Dendroica fusca Chipe Gorjinaranja Blackburnian Warbler Transient Forest Generalist Biodiversity field inventories in selected watersheds Forest Generalist Altitude Generalist Lowlands Altitude Generalist Altitude Generalist Highlands Altitude Generalist Highlands Highlands Altitude Generalist Species Spanish name English name Seasonality Habitat Elevation Dendroica graciae Dendroica discolor Chipe de Grace Chipe Pradeño Grace’s Warbler Prairie Warbler Resident Migratory Vagrant Forest Specialist Forest Generalist Mniotilta varia Chipe Trepador Black-and-white Warbler Visitor Forest Generalist Setophaga ruticilla Pavito Migratorio American Redstart Visitor Forest Specialist Helmitheros vermivorum Chipe Gusanero Worm-eating Warbler Visitor Forest Generalist Limnothlypis swainsonii Chipe de Swainson Swainson’s Warbler Migratory Vagrant Forest Generalist Seiurus aurocapilla Chipe-suelero Coronado Ovenbird Visitor Forest Generalist Seiurus noveboracensis Chipe-suelero Chaquero Northern Waterthrush Visitor Forest Specialist Seiurus motacilla Chipe-suelero Arroyero Louisiana Waterthrush Visitor Forest Specialist Oporornis formosus Chipe de Kentucky Kentucky Warbler Visitor Forest Generalist Oporornis philadelphia Chipe Llorón Mourning Warbler Transient Forest Generalist Oporornis tolmiei Chipe de Tolmie MacGillivray’s Warbler Visitor Geothlypis trichas Mascarita Común Common Yellowthroat Visitor Geothlypis poliocephala Mascarita Piquigruesa Gray-crowned Yellowthroat Resident Wilsonia citrina Chipe Encapuchado Hooded Warbler Visitor Highlands Lowlands Altitude Generalist Lowlands Altitude Generalist Altitude Generalist Altitude Generalist Lowlands Altitude Generalist Altitude Generalist Altitude Generalist Altitude Generalist Altitude Generalist Altitude Generalist Altitude Generalist Biodiversity field inventories in selected watersheds Open Area Generalist Open Area Generalist Open Area Specialist Forest Generalist 191 192 Species Spanish name English name Seasonality Habitat Wilsonia pusilla Chipe de Wilson Wilson’s Warbler Visitor Forest Generalist Wilsonia canadensis Chipe Collarejo Canada Warbler Transient Forest Generalist Cardellina rubrifrons Myioborus miniatus Euthlypis lachrymosa Basileuterus culicivorus Chipe Carirrojo Pavito Gorjigris Chipe Roquero Chipe Corona-dorada Red-faced Warbler Slate-throated Redstart Fan-tailed Warbler Golden-crowned Warbler Visitor Resident Resident Resident Forest Specialist Forest Generalist Forest Generalist Forest Specialist Basileuterus rufifrons Chipe Gorrirrufo Rufous-capped Warbler Resident Forest Generalist Icteria virens Gritón Pechiamarillo Yellow-breasted Chat Visitor Open Area Generalist Lowlands Habia rubica Tángara-hormiguera Coronirroja Red-crowned Ant-Tanager Resident Forest Specialist Altitude Generalist Habia fuscicauda Tángara-hormiguera Gorjirroja Red-throated Ant-Tanager Resident Forest Specialist Lowlands Piranga rubra Piranga olivacea Tángara Roja Tángara Escarlata Summer Tanager Scarlet Tanager Visitor Transient Forest Generalist Forest Generalist Piranga ludoviciana Tángara Occidental Western Tanager Visitor Forest Generalist Piranga bidentata Piranga leucoptera Tángara Dorsirrayada Tángara Aliblanca Flame-colored Tanager White-winged Tanager Resident Resident Thraupis episcopus Tángara Azuligris Blue-gray Tanager Resident Thraupis abbas Tángara Aliamarilla Yellow-winged Tanager Resident Forest Specialist Forest Generalist Open Area Generalist Open Area Generalist Lowlands Lowlands Altitude Generalist Highlands Highlands Cyanerpes cyaneus Mielero Patirrojo Red-legged Honeycreeper Resident and migratory Biodiversity field inventories in selected watersheds Forest Generalist Elevation Altitude Generalist Altitude Generalist Highlands Highlands Lowlands Highlands Altitude Generalist Lowlands Altitude Generalist Lowlands Species Spanish name English name Seasonality Volatinia jacarina Semillero Brincador Blue-black Grassquit Resident Sporophila torqueola Semillero Collarejo White-collared Seedeater Resident Sporophila minuta Semillero Pechicanelo Ruddy-breasted Seedeater Resident Amaurospiza concolor Semillero Azul Blue Seedeater Resident Tiaris olivaceus Semillero Olivaceo Yellow-faced Grassquit Resident Diglossa baritula Picaflor Vientre-canelo Atlapetes albinucha Melozone biarcuata Melozone leucotis Saltón Nuquiblanco Rascador Patilludo Rascador Orejiblanco Cinnamon-bellied Flowerpiercer White-naped Brush-Finch Prevost’s Ground-Sparrow White-eared Ground-Sparrow Aimophila ruficauda Zacatonero Cabecirrayado Stripe-headed Sparrow Resident Aimophila rufescens Zacatonero Rojizo Rusty Sparrow Resident Ammodramus savannarum Gorrión Chapulín Grasshopper Sparrow Visitor Melospiza lincolnii Gorrión de Lincoln Lincoln’s Sparrow Visitor Zonotrichia capensis Gorrión Chingolo Rufous-collared Sparrow Resident Saltator coerulescens Saltator Grisáceo (Dichoso-fuí) Grayish Saltator Resident Biodiversity field inventories in selected watersheds Resident Resident Resident Resident Habitat Elevation Open Area Generalist Open Area Generalist Open Area Specialist Altitude Generalist Forest Specialist Open Area Generalist Open Area Specialist Forest Generalist Forest Generalist Forest Specialist Open Area Generalist Open Area Generalist Open Area Specialist Open Area Generalist Open Area Generalist Open Area Generalist Lowlands Lowlands Altitude Generalist Highlands Highlands Highlands Highlands Highlands Lowlands Highlands Altitude Generalist Highlands Highlands Altitude Generalist 193 Species Spanish name English name Seasonality Habitat Saltator atriceps Saltator Cabecinegro (Chepito) Black-headed Saltator Resident Forest Generalist Rose-breasted Grosbeak Visitor Forest Generalist Blue Bunting Resident Resident and migratory Forest Specialist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Open Area Specialist Open Area Specialist Cyanocompsa parellina Picogrueso Pechirrosado (Puñalada) Colorín Azulinegro Passerina caerulea Picogrueso Azul Blue Grosbeak Passerina cyanea Colorín Azul Indigo Bunting Visitor Passerina ciris Colorín Sietecolores (Sietecolores) Painted Bunting Visitor Spiza americana Arrocero Americano Dickcissel Visitor Agelaius phoeniceus Tordo Sargento Red-winged Blackbird Resident Sturnella magna Pradero Común Eastern Meadowlark Resident Dives dives Tordo Cantor Melodious Blackbird Resident Quiscalus mexicanus Zanate Mayor (Zanate♀, Clarinero♂) Great-tailed Grackle Resident Molothrus aeneus Vaquero Ojirrojo Bronzed Cowbird Resident Icterus wagleri Icterus maculialatus Bolsero de Wagler Bolsero Guatemalteco Black-vented Oriole Bar-winged Oriole Resident Resident Icterus spurius Bolsero Castaño Orchard Oriole Visitor Pheucticus ludovicianus 194 Biodiversity field inventories in selected watersheds Forest Generalist Open Area Generalist Open Area Generalist Forest Generalist Forest Generalist Open Area Generalist Elevation Altitude Generalist Altitude Generalist Lowlands Lowlands Lowlands Lowlands Lowlands Lowlands Altitude Generalist Altitude Generalist Altitude Generalist Lowlands Highlands Highlands Lowlands Species Spanish name English name Seasonality Icterus pustulatus Bolsero Dorsirrayado Streak-backed Oriole Resident Icterus pectoralis Bolsero Pechimanchado Spot-breasted Oriole Resident Icterus gularis Bolsero de Altamira Altamira Oriole Resident Icterus galbula Bolsero de Baltimore Baltimore Oriole Visitor Amblycercus holosericeus Cacique Piquiclaro Yellow-billed Cacique Resident Cacicus melanicterus Cacique Mexicano Yellow-winged Cacique Resident Euphonia affinis Eufonia Gorjinegro Scrub Euphonia Resident Open Area Generalist Forest Generalist Euphonia hirundinacea Eufonia Gorjiamarillo Yellow-throated Euphonia Resident Forest Specialist Euphonia elegantissima Chlorophonia occipitalis Carduelis notata Eufonia Capucha-azul Clorofonia Coroniazul Dominico Cabecinegro Elegant Euphonia Blue-crowned Chlorophonia Black-headed Siskin Resident Resident Resident Carduelis psaltria Dominico Dorsioscuro Lesser Goldfinch Resident Passer domesticus Gorrión Doméstico House Sparrow Resident Lonchura malacca Capuchino de Cabeza Negra Tricolored Munia Non-migratory Vagrant Forest Generalist Forest Generalist Forest Specialist Open Area Generalist Open Area Generalist Open Area Generalist Biodiversity field inventories in selected watersheds Habitat Elevation Open Area Generalist Open Area Generalist Open Area Generalist Open Area Generalist Altitude Generalist Altitude Generalist Altitude Generalist Altitude Generalist Forest Generalist Lowlands Lowlands Lowlands Altitude Generalist Highlands Highlands Highlands Altitude Generalist Lowlands Lowlands 195 3.2. INVENTORY COMPLETENESS It is important to evaluate completeness of the inventories before comparing the sites, because species richness is influenced by inventory effort (Komar 2002, 2007). Before the field study began, the baseline bird inventory of the watersheds within the IMCW Project area was substantially incomplete, with only 5 of the 11 watersheds above 80% completeness (based on the presence of 74 test species). After field work, 10 of the 11 watersheds now have reasonably complete inventories, averaging 86% complete, and can be compared for conservation importance (Fig. 16, Table 42). The only watershed that will be not be analyzed, for lack of data, is the Bocana San Juan Watershed. In seven watersheds, more than 85% of the test species have been recorded. In the three watersheds with 57% to 74% of test species recorded, the watersheds lack middle and higher elevations, such that some of the test species may actually be absent, thus these areas may have more complete inventories than indicated here. Prior to field work, the level of inventory completeness (or effort) was a reasonably good predictor of the number of threatened bird species recorded from each watershed (linear regression R-squared=0.466, P=0.021, statistically signficant), indicating that lists of threatened species were not comparable among watersheds. But after completing the field work and eliminating the one watershed with a very poor inventory, the level of inventory completeness is no longer a good predictor of the number of threatened bird species present (R2=0.281, P=0.115, not statistically significant). Now we can assume that the number of threatened birds present is relatively accurate and can be used as an index of relative conservation importance to prioritize the 10 watersheds for conservation action. FIGURE 16. INCREMENT IN INVENTORY COMPLETENESS INDEX FOR BIRDS IN WATERSHEDS OF SOUTHWESTERN EL SALVADOR, RESULTING FROM FIELD STUDY. 196 Biodiversity field inventories in selected watersheds TABLE 42. INVENTORY COMPLETENESS FOR BIRDS AND RELATIVE IMPORTANCE OF WATERSHEDS. Inventory Inventory Conservation Conservation test test important important species species species species Watershed recorded recorded recorded recorded through through through 2006 through 2007 2006 (%) 2007 (%) (%) (%) Bocana San Juan 4 4 5 5 Barra Salada 22 57 15 26 Los Cóbanos 66 73 25 30 San Julián 74 93 15 17 Cauta 74 74 21 21 San Pedro 76 89 12 22 Grande de Sonsonate 82 85 24 27 Izalco 86 96 26 31 Coatepeque 93 93 39 39 Cara Sucia 99 99 72 72 Barra de Santiago 100 100 71 74 Full Project Area 100 100 The 25 municipalities have widely varying levels of inventory effort, and in fact nine municipalities still present no data on bird presence. After completing the inventory field work, eight municipalities now have inventories with more than 75% of test species recorded (i.e., inventory completeness), including six with more than 90%. Prior to field work, only five municipalities had inventory completeness above 75% (Table 43). The inventory completeness level for all municipalities combined is a very good predictor for the proportion of threatened bird species present (R2=0.632, F=39.44, P<0.001), which demonstrates that the varying levels of effort significantly biases the apparent relative importance of the municipalities for threatened bird species. If we eliminate municipalities with <60% inventory completeness, inventory effort is still a good predictor (R2=0.369, F=6.44, P=0.028). But if we consider only municipalities with >75% inventory completeness, the relative completeness is a very poor predictor of threatened bird species present (R2=0.046, F=0.29, P=0.609), such that this group of municipalities can be compared. The mean completeness score for this group is 92%. Biodiversity field inventories in selected watersheds 197 TABLE 43. INVENTORY COMPLETENESS FOR BIRDS AND RELATIVE IMPORTANCE OF MUNICIPALITIES. Inventory Inventory Conservation Conservation test test important important species species species species Municipality recorded recorded recorded recorded through through through 2006 through 2007 2006 (%) 2007 (%) (%) (%) Acajutla 66 69 30 33 Apaneca 45 61 13 18 Armenia 0 0 0 0 Caluco 74 92 14 15 Chalchuapa 0 0 0 0 Concepción de Ataco 62 62 9 9 Cuisnahuat 0 0 0 0 Guaymango 0 0 0 0 Izalco 93 97 26 28 Juayúa 61 97 15 18 Jujutla 95 95 57 57 Nahulingo 27 27 0 0 Nahuizalco 41 41 1 1 Salcoatitán 0 0 0 0 San Antonio del Monte 0 0 0 0 San Francisco Menéndez 100 100 66 68 San Julián 8 8 3 3 San Pedro Puxtla 68 68 13 13 Santa Ana 77 77 32 32 Sta. Catarina Masahuat 0 0 0 0 Santo Domingo de Guzmán 0 61 0 2 Sonsonate 66 82 18 39 Sonzacate 0 0 0 0 Tacuba 93 93 30 30 Tepecoyo 0 0 0 0 Full Project Area 100 198 100 Biodiversity field inventories in selected watersheds Inventory effort for the eight ecosystems varies widely, according to the proportion of test species that have been recorded in each ecosystem (Table 44). The differing levels of effort appear to bias the relative importance scores for the ecosystems, although the linear regression (threatened bird species richness regressed against proportion of test species recorded) is not quite statistically significant (R 2=0.442, F=4.751, P=0.072). Removing the two poorly-studied ecosystems (savanna and coral reef) removes most of the bias. The remaining six ecosystems all have inventory completeness scores above 80%. For this group of reasonably completely studied ecosystems, the conservation importance score (threatened species richness) is not at all predicted by the level of inventory effort (regression results: R2=0.013, F=0.0054, P=0.828), suggesting that the numbers of threatened bird species recorded can be compared among sites and used as an indicator of conservation importance. TABLE 44. INVENTORY COMPLETENESS FOR BIRDS AND RELATIVE IMPORTANCE OF ECOSYSTEMS. Conservation Inventory test important species Ecosystem species recorded recorded through through 2007 (%) 2007 (%) Humid Forest 98 33 Dry Forest 97 26 Cloud Forest 98 17 Estuaries (and Mangroves) 83 39 Beaches 81 13 Freshwater Wetlands (and swamp forests) 84 20 Savanna 35 5 Coral Reef 64 2 3.3. EVALUATION OF CONSERVATION IMPORTANCE AMONG AREAS AND ECOSYSTEMS. The Cara Sucia and the Barra de Santiago watersheds (both include parts of El Imposible National Park and parts of Barra de Santiago estuary) have the highest conservation importance, with more than 70% of the area’s nationally threatened bird species present. They are distantly followed by the Coatepeque and Izalco watersheds (which combined include Los Volcanes National Park and the San Marcelino Protected Natural Area complex, each a part of the Apaneca-Ilamatepec Biosphere Reserve), which have 39% and 31% of the threatened bird species present, respectively. The greater importance for birds of the El Imposible area with respect to the San Marcelino area was previously noted by Komar & Herrera (1995). Other watersheds have fewer threatened birds present (Table 42). Biodiversity field inventories in selected watersheds 199 Among municipalities, San Francisco Menéndez (El Imposible National Park and Santa Rita Protected Natural Area) and Jujutla (Barra de Santiago estuary and Protected Natural Area) have considerably more conservation importance than Sonsonate (Los Cóbanos and Barra Salada Protected Natural Areas), which in turn is more important for birds than Tacuba, Izalco and Santa Ana (the latter two municipalities include Los Volcanes National Park and San Marcelino Complex of Protected Natural Areas). The lowest importance scores were for Juayúa (Laguna de las Ranas) and Caluco (Plan de Amayo Protected Natural Area) (Table 43). Estuaries (Mangroves), such as at Barra de Santiago, are the most important ecosystem for bird conservation, with 39% of the area’s threatened bird species, more than humid forest (33%) and dry forest (26%; Table 44). These ecosystems are followed in importance by freshwater wetlands, cloud forest, and beaches. 4. RECOMMENDATIONS I present below two types of recommendations: those that relate to bird conservation strategies, resulting from the results reported herein, and those that relate to filling gaps in the existing bird inventory. Although field surveys carried out during 2007 were intensive, and the study area has been extensively studied for birds previously (Komar 2007), there are still noteworthy gaps in the ornithological inventory. The field surveys reconfimed what was generally already known about the Study Area. The natural habitats are highly fragmented and of very reduced size. Most birds found in these habitats are generalist species adapted to rapidly changing landscapes and disturbance. Conserving the few habitat specialist species that persist in southwestern El Salvador will require great efforts to increase natural vegetation cover, especially forest cover but also natural savanna vegetation. Ideally, reforestation or habitat recuperation can be organized so as to create natural habitat corridors connecting patches of habitat, such as the mangrove estuaries to the humid forests that still remain in the upper watersheds. The small fragments of natural habitat are subject to numerous threats (Komar & Girón 2007), one of which is global climate change which is probably already causing ecosystem adjustments that will stress the remaining wildlife populations. Such large scale landscape processes stress the need for a major effort to invest in re-greening the landscape. The best way to allow the ecosystems to adapt to climate change is to provide space across an altitudinal gradient for natural ecosystems to migrate as temperature regimes change. Where are the priority sites for implementing habitat conservation? While the mangrove ecosystems are home to the most threatened bird species, many of those species are waterbirds and migratory shorebirds, neither of which will benefit greatly from reforestation or expanding habitat corridors. Waterbirds and shorebirds are indeed highly threatened, because of their habitat specializations. Many shorebirds migrate to El Salvador from Arctic breeding grounds which are themselves threatened by climate change and by anthropogenic activities such as oil drilling. The recommended conservation strategy for these birds in El Salvador is to increase education for local 200 Biodiversity field inventories in selected watersheds fishermen and tourists. As users of the estuaries learn about the special challenges these birds face, they may increase their appreciation for the birds and take more care to protect them. The next highest priorities are in the humid forest and dry forests ecosystems. These are the areas that most require efforts to expand natural areas and connect them via corridors. Especially useful would be to reforest the riparian areas, connecting the remnant transition forests behind the mangroves to small patches of dry forest and further upstream, to humid forest. Also important for the long-term conservation of birds in southwestern El Salvador is to expand bird monitoring efforts. Long-term monitoring of bird populations is needed to determine if birds are able to withstand the various stresses, and can indicate to conservation managers if efforts are not enough. Current monitoring efforts carried out by SalvaNATURA at the two national parks in the Study Area are not sufficient, and additional monitoring stations are needed both as study replicates, and also to monitor birds in additional habitats, such as in mangrove forests and on mudflats (where shorebirds feed). The 2007 field inventories failed to ―complete‖ the local bird inventories, in part, because of the seasonality of the field work. Sites were visited twice (two different seasons) during the second semester of the year, but bird communities vary across the entire year, and a third visit (at least) would be required to cover all seasons important for documenting bird communities. For example, none of the sites were visited during spring (April and early May), when spring migration is in full swing and when most local resident birds are beginning their breeding season. The results indicate that only one of the watersheds and only one municipality have complete inventories (100% of test species registered), therefore considerable work across the study area is still needed to complete the inventories. More complete inventories could reveal different or additional conservation priorities within the study area. They could also provide more information useful for valuing the area’s natural resources, such as by identifying localities where species attractive for ecotourism can be found reliably by tourists. Given that southwestern El Salvador is arguably the best inventoried area in the country (for birds), I recommend using this area to initiate a pilot project to take the bird inventory to a new level. In many countries, ornithologists have used an inventory method known as atlas mapping, in particular for documenting the local breeding ranges of resident bird species. This method consists of dividing the study area into a series of grid squares, and then mobilizing many observers (ornithologists and bird enthusiasts) to visit each grid to determine the bird species that breed within the grid. A reasonable grid size would be 5 x 5 km, or 25 km 2. Biologists documenting breeding birds could also document other types of flora and fauna during their visits to each grid. The atlas would determine much more accurately the actual distribution and range size for each species in the study area. It would also be useful for collecting data on elevational ranges for species, which are needed to determine long-term adaptability to projected climate change for each species. Biodiversity field inventories in selected watersheds 201 CHAPTER 6: INVENTORY OF TERRESTRIAL MAMMALS IN SOUTHWESTERN EL SALVADOR Luis Girón, Melissa Rodríguez, and James G. Owen Conservation Science Department SalvaNATURA, San Salvador, El Salvador legiron@salvanatura.org 1. INTRODUCTION El Salvador’s natural areas are highly fragmented and anthropogenically disturbed, which complicates the study of wild mammals. Mammalian inventories have not been completed in any of the country’s natural areas. The first terrestrial mammal collection efforts in southwestern El Salvador were carried out in 1927 and 1942 (Burt and Stirton 1961). The second phase of collections was carried out by Heinz Felten, between 1952 and 1954 (Felten 1956). Then, during a third collection phase between 1988 and 2005, James G. Owen invested the largest effort so far to document mammals of El Salvador. The most recent efforts documenting mammals in the southwestern region of the country, mainly bats, began in 2003 with the Neobats–El Salvador program carried out by Luis Girón of SalvaNATURA. Neobats–El Salvador provided data up to 2006 (unpublished data). The information from mammal studies carried out previously in southwestern El Salvador was systematized prior to field work, generating a list of 97 mammal species recorded in the study area (Girón & Owen 2007). The purpose of this study was to complement these inventories with field work and make an evaluation of mammals in selected watersheds, municipalities, ecosystems, and Protected Natural Areas. 2. METHODOLOGY 2.1. STUDY AREA The study was conducted in eight Protected Natural Areas, in seven of the 11 watersheds of southwestern El Salvador (Fig. 17). These sites correspond to eight of the 25 municipalities located within the IMCW project area, within the Departments of 202 Biodiversity field inventories in selected watersheds Ahuachapán, Sonsonate, and Santa Ana (Tabla 5-1). The ecosystems included Dry Forest, Humid Forest, Cloud Forest, and a Savanna dominated by Crescentia alata. 2.2. FIELD METHODOLOGY During June–December 2007, traps were placed at each site, usually two harp traps to capture bats, 50 Sherman live traps for small rodents, five Tomahawk traps for mediumsized mammals, and 12 camera traps. We also established observation transects at each site (except for Laguna de las Ninfas). Sampling efforts per site are given in Table 45. In order to calculate the relative frequency and relative abundance, each day was used as a sampling unit. The relative frequency was obtained by multiplying the number of samples (days) by 100, in which each one of the species appeared, divided by the total number of samples. The relative abundance was obtained by dividing the number of each individual species observed between the total numbers of individuals observed of all of the species multiplying the result by 100. 1 2 3 4 8 5 6 7 FIGURE 17. MAP OF SITES SAMPLED FOR MAMMALS IN 2007: (1) EL IMPOSIBLE NATIONAL PARK, (2) SIERRA DE APANECA WEST (LAGUNA DE LAS NINFAS), (3) SIERRA DE APANECA EAST (LAGUNA DE LAS RANAS), (4) LOS VOLCANES NATIONAL PARK, (5) SAN MARCELINO COMPLEX, (6) PLAN DE AMAYO, (7) LOS CÓBANOS, (8) BOSQUE SANTA RITA. Biodiversity field inventories in selected watersheds 203 TABLE 45. SITES AND SAMPLING EFFORTS FOR MAMMALS. Specific Site Municipality Departament Latitude and Longitude Laguna de las Ninfas Apaneca Ahuachapán 13.8733º -89.80557º 1500 5 0 320 20 12 40 Laguna de las Ranas Chalchuapa Santa Ana 13.90391º -89.72281º 1750 10 10 320 20 8 88 Caluco Sonsonate 335 5 10 220 15 5 64 Acajutla Sonsonate 3 10 10 480 40 16 80 Santa Rita San Francisco Menéndez Ahuachapán 33 10 10 162 20 6 81 San Marcelino Izalco Sonsonate 917 5 10 240 16 8 64 Los Andes Santa Ana Santa Ana 20 20 550 55 22 184 Cerro Verde Santa Ana Santa Ana 40 36 850 80 36 352 Site Sierra de Apeneca West Sierra de Apaneca East Plan de Amayo Los Cóbanos Los Volcanes La Fincona El Imposible San Benito San Francisco Menéndez San Francisco Menéndez Ahuachapán Ahuachapán Cerro Campana Tacuba Ahuachapán Guayapa San Francisco Menéndez Ahuachapán 13.69088º -89.64741º 13.52764º -89.80955º 13.80º -90.06º 13.81040º -89.57049º 13.86848º -89.62009º 13.82611º -89.62389º 13.84320º -89.97761º 13.82712º -89.94712º 13.85675º -89.91398º 13.82778º -89.93663º Elevation Days TH RTN TTN HTN CTN 1735 2000 738 730 1350 478 TOTAL 105 106 3,142 266 113 953 TH: Transect hours; RTN: Rodent trap nights; TTN: Tomahawk trap nights; HTN: Harp trap nights; CTN: Camera trap days & nights. 204 Biodiversity field inventories in selected watersheds 2.3. LEVEL OF INVENTORY COMPLETENESS A list of 48 generalist species expected in the area was created, to determine the degree of inventory completeness in the study area’s municipalities and watersheds (Table 46), modifying a method proposed by Gómez de Silva and Medellín (2001). In ecological terms, the generalist species are those that exist in a variety of habitats and eat varied foods that are found in both disturbed and pristine habitats. These species are expected to be present in every site in the departments of Ahuachapán, Santa Ana, and Sonsonate. We compared the generalist species recorded with the list of expected generalist species for each municipality, watershed, ecosystem, and protected natural area, to calculate the percentage level of inventory completeness. As a complement, we used species accumulation curves and to estimate the inventory completeness. From the curves, we generated various estimates of species richness, using ACE (Abundance-based Coverage Estimator), ICE (Incidence-based Coverage Estimator), Chao 1, Chao 2, Jack-Knife 1 and Jack-Knife 2 methods, all calculated with the EstimateS version 7.51 program (Colwell 2006). The accumulation curves were obtained taking each day as a sampling unit, using data from all of the combined techniques, to achieve a similar sampling effort for each sampling day. 2.4. EVALUATION OF THE RELATIVE IMPORTANCE OF SITES We prepared a list of 34 species of conservation importance that can be found in southwestern El Salvador (Table 47). For each municipality, watershed, ecosystem, and Protected Natural Area, we determined the percentage of these species that have been recorded. This percentage represents the relative importance of a site for mammal conservation. Since the level of inventory completeness can influence the proportion of conservation-important species reported (Komar 2007), we only evalutated the importance of the sites when these showed an inventory completeness of more than 70%. 3. RESULTS AND DISCUSSION 3.1. RECORDED SPECIES We recorded 1296 individuals representing 72 species, of which 49 were captured, 17 were photographed by camera traps, 21 were observed in transects, and five were recorded only outside of standardized sampling. We recorded 16 species at Sierra de Apaneca West, 27 at Plan de Amayo, 16 at Los Cóbanos, 13 at Santa Rita, 21 at Sierra de Apaneca East, 22 at Los Volcanes, 21 at San Marcelino, and 54 at El Imposible. Regarding ecosystems, 44 species were recorded in Dry Forest, 44 in Humid Forest, 30 in Cloud Forest, and 17 in Savanna. The species that showed the greatest relative frequency were the Central American Agouti (Dasyprocta punctata) which appeared in 46% of the samples followed by the Nine-banded Armadillo (Dasypus novemcinctus) and Common Opposum (Didelphis marsupialis), which appeared in 40% and 31% of the samples, respectively. The species with greatest relative abundance, both bats, were the Jamaican Fruit-eating Bat (Artibeus jamaicensis), with 17% of all records, followed by the Hairy-legged Myotis (Myotis keaysi) with 13% of all records (Table 48). Biodiversity field inventories in selected watersheds 205 TABLE 46. INDICATOR SPECIES (MAMMALS) OF INVENTORY COMPLETENESS. Family Species Spanish Name Didelphidae Didelphis marsupialis Tacuazín Negro Didelphidae Didelphis virginianus Tacuazín Blanco Didelphidae Marmosa mexicana Tacuazín Ratón Didelphidae Phylander opossum Hurón o Cuatro Ojos Dasypodidae Dasypus novemcinctus Cuzuco Emballonuridae Saccopteryx bilineata Murciélago Grande de Líneas Blancas Mormoopidae Pteronotus davyi Murciélago de Espalda Desnuda Phyllostomidae Phyllostomus discolor Murciélago Hoja de Lanza Menor Phyllostomidae Glossophaga soricina Murciélago Longi Rostro Común Phyllostomidae Glossophaga leachii Murciélago Longirostro Gris Phyllostomidae Glossophaga commissarisi Murciélago Longorostro de Commissaris Phyllostomidae Carollia subrufa Murciélago de Cola Corta Gris Phyllostomidae Carollia perspicillata Murciélago Frutero Común de Cola Corta Phyllostomidae Sturnira lilium Murciélago Frutero Común de Hombros Amarillos Phyllostomidae Artibeus intermedius Murciélago Frutero Intermedio Phyllostomidae Artibeus jamaicensis Murciélago Frutero de Jamaica Phyllostomidae Artibeus toltecus Murciélago Frutero Tolteca Phyllostomidae Artibeus phaeotis Murciélago Peludo de Ojos Grandes Phyllostomidae Uroderma bilobatum Murciélago Constructor de Tiendas Común Phyllostomidae Artibeus helleri Murciélago de Heller Phyllostomidae Chiroderma villosum Murciélago Frutero de Velvety Phyllostomidae Desmodus rotundus Vampiro Común 206 Biodiversity field inventories in selected watersheds Family Species Spanish Name Vespertilionidae Myotis nigricans Murciélago Negruzco Común Vespertilionidae Myotis keaysi Murciélago de Patas Peludas Vespertilionidae Rhogeessa tumida Murciélago Enano de Alas Negras Molossidae Molossus ater Murciélago Mastín Negro Molossidae Molossus molossus Murciélago Mastín Común Sciuridae Sciurus variegatoides Ardilla Gris Geomyidae Orthogeomys grandis Taltuza Heteromyidae Liomys salvini Ratón Bolsero Cricetidae Sigmodon hispidus Rata Algodonera Cricetidae Reithrodontomys gracilis Ratón Cosechador Cricetidae Peromyscus mexicanus Ratón Venado Mexicano Cricetidae Baiomys musculus Ratón Bolsero del sur Cricetidae Ototylomys phyllotis Rata Escaladora de Orejas Grandes Cricetidae Nyctomys sumichrasti Ratón Cricetidae Rattus rattus Rata Negra Cricetidae Mus musculus Ratón Casero Erethizontidae Sphiggurus mexicanus Puerco Espín o Zorro Espín Dasyproctidae Dasyprocta punctata Cotuza Leporidae Sylvilagus floridanus Conejo Silvestre Canidae Urocyon cinereoargenteus Zorra Gris Procyonidae Procyon lotor Mapache Mustelidae Mustela frenata Comadreja Mephitidae Mephitis macroura Zorrillo Listado Mephitidae Spilogale angustifrons Zorrillo Manchado Felidae Puma yagouaroundi Gato Zonto Cervidae Odocoileus virginianus Venado Cola Blanca Biodiversity field inventories in selected watersheds 207 TABLE 47. MAMMALS THAT ARE INDICATORS OF IMPORTANT SITES FOR CONSERVATION.1 Species Spanish Name National Status (MARN 2004) Chironectes minimus Tacuazín de Agua Endangered Tamandua mexicana Oso Hormiguero Threatened Rhynchonycteris naso Murciélago Threatened Saccopteryx leptura Murciélago Pequeño de Líneas Blancas Threatened Peropteryx macrotis Murciélago Threatened Balantiopteryx plicata Murciélago de Alas de Saco Gris Threatened Noctilio albiventris Murciélago Pescador Pequeño Threatened Mormoops megalophylla Murciélago Rostro de Fantasma Threatened Pteronotus personatus Murciélago Bigotudo Pequeño Threatened Pteronotus gymnonotus Murciélago de Espalda Desnuda Grande Threatened Micronycteris microtis Murciélago Orejón Común Threatened Lonchorhina aurita Murciélago Orejón Nariz de Espada Threatened Macrophyllum macrophyllum Murciélago de Patas Largas Threatened Trachops cirrhosus Murciélago con Verrugas en el Labio Threatened Chrotopterus auritus Murciélago Falso Vampiro Threatened Leptonycteris yerbabuenae Murciélago Magueyero Endangered 208 Biodiversity field inventories in selected watersheds Red List (IUCN 2008) Vulnerable Species Spanish Name National Status (MARN 2004) Artibeus hartii Murciélago Frutero Pigmeo Threatened Uroderma magnirostrum Murciélago Constructor de Tiendas Café Diaemus youngi Vampiro de Ala Blanca Bauerus dubiaquercus2 Murciélago de Van Gelder Lasiurus blossevilli Murciélago Rojo Eumops auripendulus Murciélago Threatened Eumops underwoodi Murciélago de Underwood Endangered Neotoma mexicana Rata de Bosque Mexicana Threatened Cuniculus paca Tepezcuintle Threatened Eira barbara Tayra o Perico Ligero Endangered Lontra longicaudis Nutria Endangered Puma concolor Puma Endangered Leopardus pardalis Ocelote Endangered Leopardus wiedii Tigrillo Endangered Tapirus bairdi Tapir o Danta Tayassu tajacu Cuche de Monte Red List (IUCN 2008) Endangered Endangered Endangered 1 Threatened or endangered at the national level (MARN 2004, modified by the authors). Species recently reported for the country that has not been evaluated for the national level threatened species list. 2 Biodiversity field inventories in selected watersheds 209 TABLE 48. INDIVIDUAL MAMMALS COUNTED DURING THE STUDY. Ecosystem RA (%) (%) DF HF CF S WA PA LV LC SR EA SM EI Didelphis marsupialis 31 3 11 24 6 3 1 5 2 3 1 3 5 24 44 Didelphis virginiana 16 2 1 9 9 6 1 8 6 1 2 7 25 Philander opossum 6 1 9 3 9 Scientific Name Marmosa mexicana1 3 1 Dasypus novemcinctus (Fig. 18A) 40 4 Tamandua mexicana (Fig. 18B) 1 0 Saccopteryx bilineata 1 0 Balantiopteryx plicata 4 1 Pteronotus parnellii (Fig. 18C) 1 0 Pteronotus davyi 4 1 Pteronotus personatus 1 0 Trachops cirrhosus (Fig. 18D) 1 0 1 Micronycteris microtis (Fig. 18E) 2 1 1 Phyllostomus discolor 1 0 1 210 Sampled Sites RF 12 3 18 2 32 57 1 1 1 1 1 1 15 18 15 2 1 1 14 2 3 5 2 1 3 2 2 12 1 25 Total 2 1 1 1 1 5 2 1 1 1 1 6 1 1 Biodiversity field inventories in selected watersheds 8 1 6 7 1 Ecosystem RF RA (%) (%) Anoura geoffroyi 1 0 Glossophaga leachii 13 2 10 6 4 Glossophaga commissarisi 12 2 12 10 2 Glossophaga soricina 5 1 2 Choeroniscus godmani 1 0 Carollia subrufa 6 1 Scientific Name DF HF CF Sampled Sites S WA PA LV LC SR EA SM 1 3 1 2 2 2 2 2 3 1 Artibeus intermedius2 6 1 5 10 20 5 14 24 2 7 1 1 2 15 1 1 14 52 222 2 5 18 2 19 25 2 1 7 5 1 43 32 Artibeus jamaicensis 23 17 147 Artibeus lituratus 9 1 11 Artibeus toltecus 10 2 14 Artibeus aztecus 2 0 Artibeus phaeotis 1 0 1 1 1 Artibeus hartii 1 0 1 1 1 Artibeus helleri 1 0 1 1 Uroderma bilobatum 1 0 5 7 2 4 117 5 32 7 Total 1 1 9 EI 2 3 1 3 3 1 1 Biodiversity field inventories in selected watersheds 1 3 1 211 Ecosystem RF RA (%) (%) Chiroderma salvini 2 0 Centurio senex (Fig. 18F) 3 1 8 Sturnira lilium 11 1 3 Sturnira ludovici 8 2 Diphylla ecaudata 1 0 1 Desmodus rotundus 6 2 15 2 4 Natalus stramineus 4 0 1 2 2 Eptesicus furinalis 1 0 1 Eptesicus fuscus 1 0 Bauerus dubiaquercus (Fig. 51G) 1 0 Myotis elegans 2 0 5 Myotis keaysi 26 13 8 Myotis nigricans 2 1 Rhogeessa tumida 4 0 Scientific Name 212 DF HF CF 1 2 8 4 9 16 Sampled Sites S WA PA LV LC SR EA SM 2 2 1 2 2 1 2 2 14 3 8 4 2 2 1 31 34 7 2 1 3 8 8 9 17 9 25 1 1 4 21 3 5 1 1 1 2 100 2 1 1 Total 1 1 3 62 EI 1 Biodiversity field inventories in selected watersheds 35 1 3 3 3 5 67 170 6 1 7 2 4 RF RA (%) (%) Sciurus deppei 12 1 Sciurus variegatoides 9 1 Orthogeomys grandis 4 Liomys salvini Scientific Name Ecosystem DF Sampled Sites HF CF 1 16 1 6 2 3 1 0 1 1 3 11 1 9 6 Oryzomys couesi 1 0 Ototylomys phyllotis (Fig. 18H) 4 0 Tylomys nudicaudatus 2 0 Neotoma mexicana (Fig. 18I) 1 0 Nyctomys sumichrasti 7 1 Peromyscus mexicanus (Fig. 18J) 18 8 Reithrodontomys mexicanus 1 0 Sigmodon hispidus 5 0 WA 1 PA 2 LV 2 LC SR EA 12 3 1 1 2 1 4 1 1 SM 4 2 4 11 1 1 5 7 4 16 1 1 1 5 4 14 84 8 1 1 2 8 15 63 3 3 4 1 1 7 62 14 14 98 1 3 20 6 1 1 3 1 1 46 1 1 1 4 3 Biodiversity field inventories in selected watersheds Total 17 1 1 1 EI 1 3 Coendou mexicanus1 Dasyprocta punctata (Fig. 18K) S 9 7 1 2 6 1 1 74 98 213 Ecosystem Sampled Sites RF RA (%) (%) DF HF Cuniculus paca (Fig. 18L) 21 4 5 50 Urocyon cinereoargenteus (Fig. 18M) 18 2 3 Puma yaguarondi (Fig. 18N) 2 0 1 Leopardus wiedii (Fig. 18Ñ) 1 0 1 1 1 Mephitis macroura1 1 1 1 Spilogale angustifrons1 1 1 1 Mustela frenata1 1 Scientific Name S WA PA LV LC SR EA SM EI 55 23 3 1 1 2 21 3 1 1 1 1 0 1 1 Potos flavus (Fig. 18O) 13 1 6 5 Bassariscus sumichrasti 8 1 2 7 Nasua narica (Fig. 18P) 13 3 Procyon lotor (Fig. 18Q) 17 2 18 Sylvilagus floridanus 6 1 1 Odocoileus virginianus (Fig. 18R) 19 2 1 8 5 3 1 45 2 4 2 13 4 7 1 18 Biodiversity field inventories in selected watersheds 1 16 4 9 4 1 Total 55 29 1 Eira barbara 214 CF 2 1 1 1 11 19 8 9 45 45 4 24 7 2 14 32 Ecosystem Sampled Sites RF RA (%) (%) DF HF Tayassu tajacu (Fig. 18S) 4 1 1 6 Total records - - 370 480 376 71 70 168 214 71 54 91 Species recorded during the study - - 44 44 31 17 16 27 22 16 13 Total species3 - - 78 70 56 25 18 31 47 25 40 Scientific Name CF S WA PA LV LC SR EA SM EI Total 7 7 68 575 1296 21 21 53 72 21 43 72 101 RF (%) = Relative Frequency, RA (%) = Relative Abundance, DF = Dry Forest, HF = Humid Forest, CF= Cloud Forest, S= Savanna, WA=West Apaneca, PA = Plan de Amayo, LV = Los Volcanes (only in cloud forest), LC = Los Cóbanos, SR = Santa Rita, EA=East Apaneca, SM = San Marcelino, EI = El Imposible. 1 Reported outside of sampling. 2 A. intermedius may be a subspecies of A. lituratus (Reid 1997). 3 Species totals take into account prior investigations. Biodiversity field inventories in selected watersheds 215 Figure 18A. Nine-banded Armadillo (Dasypus novemcinctus), El Imposible National Park, taken with camera traps, November 2007. Figure 18B. Northern Tamandua (Tamandua mexicana), El Imposible National Park, taken with camera traps, December 2007. Figure 18C. Common Moustached Bat (Pteronotus parnellii), San Marcelino Complex Protected Natural Area, October 2007, by Melissa Rodríguez. Figure 18D. Fringe-lipped Bat (Trachops cirrhosus), Santa Rita Protected Natural Area, Sepember 2007, by Melissa Rodríguez. Figure 18E. Common Big-eared Bat Figure 18F. Wrinkle-faced bat (Centurio (Micronycteris microtis), El Imposible senex), El Imposible National Park, National Park, October 2007, by Stefany December 2007, by Melissa Rodríguez. Henríquez. 216 Biodiversity field inventories in selected watersheds Figure 18G. Van Gelder´s Bat (Bauerus dubiaquercus), Imposible National Park, Cerro Campana sector, December 2007, by Luis Girón. Figure 18H. Big-eared Climbing Rat (Ototylomys phyllotis), San Marcelino Complex Protected Natural Area, October 2007, by Luis Girón. Figure 18I. Mexican Wood Rat (Neotoma mexicana), Cerro Verde, Los Volcanes National Park, September 2007, by Luis Giròn. Figure 18J. Mexican Deer Mouse (Peromyscus mexicanus), Laguna Las Ranas Protected Natural Area, October 2007, by Melissa Rodríguez. Figure 18K. Central American Agouti (Dasyprocta punctata), Laguna Las Ranas Protected Natural Area, taken with camera traps, October 2007. Figure 18L. Paca (Cuniculus paca), El Imposible National Park, taken with camera traps, December 2007. Biodiversity field inventories in selected watersheds 217 Figure 18M. Gray Fox (Urocyon cinereoargenteus), Los Volcanes National Park, Los Andes sector, taken with camera traps, August 2007. Figure 18Ñ. Margay (Leopardus wiedii), Plan de Amayo Protected Natural Area, taken with camera traps, July 2007. Figure 18P. Coati (Nasua narica), El Imposible National Park, taken with camera traps, December 2007. 218 Figure 18N. Jaguarundi (Puma yagouaroundi), Plan de Amayo Protected Natural Area, taken with camera traps, July 2007. Figure 18O. Kinkajou (Potos flavus), El Imposible National Park, November 2007, by Luis Girón. Figure 18Q. Northern Raccoon (Procyon lotor), El Imposible National Park, taken with camera traps, October 2007. Biodiversity field inventories in selected watersheds Figure 18R. White-tailed Deer (Odocoileus virginianus), Los Volcanes National Park, Cerro Verde, taken with camera traps, September 2007. Figure 18S. Collared Peccary (Tayassu tajacu), El Imposible National Park, taken with camera traps, November 2007. FIGURE 18. SELECTED MAMMAL PHOTOGRAPHS TAKEN DURING THE STUDY. 3.2. STATUS OF INVENTORY COMPLETION Four new species, all bats, were recorded in the study area for the first time (Micronycteris microtis, Pteronotus parnelli, Pteronotus personatus, and Bauerus dubiaquercus), increasing the list of mammals for the study area to 101 species (Table 49). Bauerus dubiaquercus was new for El Salvador (Girón, Owen & Rodríguez in press). Considering the list of 48 indicator species used to evaluate inventory completion, we estimated that the level of inventory completeness for the study area increased from 92% to 100%. The Protected Natural Areas with the most complete inventories are: El Imposible (81%), Santa Rita (63%), and Los Volcanes (52%) (Table 50). Inventory completeness indices have been calculated for nine watersheds. The watersheds with the highest level of inventory completeness are: Cara Sucia (83%), Barra de Santiago (79%), Grande de Sonsonate (67%), and Coatepeque (67%) (Tabla 51). Indices of completeness were calculated for 16 municipalities. Those with the highest indices are San Francisco Menéndez (85%), Santa Ana (60%), Tacuba (46%), Acajutla (46%), and Caluco (46%) (Table 52). Among ecosystems, the habitats with the highest inventory completeness indices are: Dry Forest (90%), Humid Forest (79%), and Cloud Forest (67%) (Table 53). Biodiversity field inventories in selected watersheds 219 TABLE 49. LIST OF TERRESTRIAL MAMMAL SPECIES RECORDED IN SOUTHWESTERN EL SALVADOR THROUGH 2007. Order Family Scientific Name Spanish Common Name English Common Name Source DIDELPHIMORPHA DIDELPHIDAE Didelphis marsupialis Tacuazín Negro Common Opossum Present Study Didelphis virginiana Tacuazín Blanco Virginia Opossum Present Study Philander opossum Tacuazín Cuatro Ojos o Hurón Gray Four-eyed Opossum Present Study Marmosa mexicana Tacuazín Ratón Mexican Mouse Opossum Present Study SalvaNATURA Chironectes minimus Tacuazín de Agua Water Opossum Database XENARTHRA DASYPODIDAE Dasypus Cusuco Nine-banded Armadillo Present Study novemcinctus MYRMECOPHAGIDAE Tamandua mexicana Oso Hormiguero Northern Tamandua Present Study INSECTIVORA SORICIDAE SalvaNATURA Cryptotis parva Musaraña Listada Least Shrew Database Néstor Herrera, Cryptotis merriami Musaraña de Orejas Cortas Merriam´s Short-eared Shrew unpublished data CHIROPTERA EMBALLONURIDAE Murciélago Grande de Líneas Saccopteryx bilineata Greater White-lined Bat Present Study Blancas Balantiopteryx plicata Murciélago Gris de Alas de Saco Gray Sac-winged Bat Present Study SalvaNATURA Peropteryx macrotis Murciélago Lesser Doglike Bat Database MORMOOPIDAE Pteronotus parnellii Murciélago Bigotudo Común Common Mustached Bat Present Study 220 Baseline inventory of flora and fauna Order Family Scientific Name Pteronotus davyi Pteronotus personatus NOCTILIONIDAE Spanish Common Name Murciélaguito de Espalda Desnuda English Common Name Davy's Naked-backed Bat Source Present Study Murciélago Bigotudo Lesser Mustached Bat Present Study Noctilio leporinus Murciélago Pescador Fishing Bat Noctilio albiventris Murciélago Pescador Pequeño Lesser Fishing Bat SalvaNATURA Database SalvaNATURA Database PHYLLOSTOMIDAE Fringe-lipped Bat Present Study Micronycteris microtis Murciélago con Verrugas en el Labio Murciélago Orejudo Común Common Big-eared Bat Chrotopterus auritus Murciélago Falso Vampiro Woolly False Vampire Bat Murciélago Hoja de Lanza Menor Murciélago Longirostro Común de Piernas Peludas Murciélago Longirostro gris Murciélago Longirostro de Commissari Murciélago Longirostro Común Pale Spear-nosed Bat Present Study SalvaNATURA Database Present Study Geoffroy's Tailless Bat Present Study Gray Long-tounged Bat Commissaris's Long-tounged Bat Pallas's Long-tounged Bat Present Study Murciélago de Godman Godman’s Whiskered Bat Present Study Trachops cirrhosus Phyllostomus discolor Anoura geoffroyi Glossophaga leachii Glossophaga commissarisi Glossophaga soricina Choeroniscus godmani Carollia perspicillata Carollia subrufa Artibeus intermedius Artibeus jamaicensis Artibeus lituratus Artibeus toltecus Artibeus aztecus Murciélago Frutero Común de Cola Corta Murciélago de Cola Corta Común Murciélago Frutero Intermedio Murciélago Frutero de Jamaica Murciélago Frutero Mayor Murciélago Frutero Tolteca Murciélago Frutero Azteca Seba's Short-tailed Bat Gray Short-tailed Bat Intermediate Fruit-eating Bat Jamaican Fruit-eating Bat Great Fruit-eating Bat Toltec Fruit-eating Bat Aztec Fruit-eating Bat Baseline inventory of flora and fauna Present Study Present Study SalvaNATURA Database Present Study Present Study Present Study Present Study Present Study Present Study 221 Order Family Scientific Name Artibeus phaeotis Artibeus hartii Artibeus helleri Uroderma bilobatum Chiroderma salvini Spanish Common Name Murciélago Peludo de Ojos Grandes Murciélago Frutero Pigmeo Murciélago de Heller Murciélago Constructor de Tiendas Común Murciélago de Ojos Grandes de Salvin English Common Name Source Hairy Big-eyed Bat Present Study Pygmy Fruit-eating Bat Heller’s Broad-nosed Bat Present Study Present Study Common Tent-making Bat Present Study Salvin's Big-eyed Bat Present Study Chiroderma villosum Murciélago Frutero de Velvety Velvety Fruit-eating Bat Centurio senex Murciélago de Cara Arrugada Murciélago Frutero Común de Hombros Amarillos Wrinkle-faced Bat SalvaNATURA Database Present Study Little Yellow Shouldered Bat Present Study Sturnira lilium Sturnira ludovici Diphylla ecaudata Desmodus rotundus Vampiro de Patas Peludas Vampiro Común Highland Yellow-shouldered Bat Hairy-legged Vampire Bat Common Vampire Bat Murciélago de Orejas de Embudo Mexican Funnel-eared Bat Present Study Murciélago Vespertino Menor Murciélago Café Gande Murciélago de Van Gelder Murciélago Amarillo del sur Murciélago Amarillo Centroamericano Murciélago Elegante Murciélago de patas peludas Murciélago Negruzco Común Murciélago Enano de Alas Negras Argentine Brown Bat Big Brown Bat Van Gelder´s Bat Southern Yellow Bat Elegant Myotis Hairy-legged Myotis Black Myotis Black-winged Little Yellow Bat Present Study Present Study Present Study Girón 2005 SalvaNATURA Database Present Study Present Study Present Study Present Study Murciélago Cara de Perro de Greenhall's Dog-faced Bat Girón 2005 Murciélago Frutero de Occidente Present Study Present Study Present Study NATALIDAE Natalus stramineus VESPERTILIONIDAE Eptesicus furinalis Eptesicus fuscus Bauerus dubiaquercus Lasiurus ega Lasiurus intermedius Myotis elegans Myotis keaysi Myotis nigricans Rhogeessa tumida Central American Yellow Bat MOLOSSIDAE Molossops greenhalli* 222 Baseline inventory of flora and fauna Order Family Scientific Name Spanish Common Name Greenhall English Common Name Molossus molossus Murciélago Mastín Común Pallas's Mastiff Bat Molossus ater Murciélago Mastín Negro Black Mastiff Bat Eumops underwoodi Murciélago de Underwood Underwood’s Bonneted Bat SalvaNATURA Database SalvaNATURA Database SalvaNATURA Database Moloso Cola de Ratón Mediano Broad-eared Bat Girón 2005 Ardilla Cuzca o Ardilla Café Ardilla Gris Deppe's Squirrel Variegated Squirrel Present Study Present Study Taltuza Giant Pocket Gopher Present Study Ratón Bolsero Salvin's Spiny Pocket Mouse Present Study Ratón Casero House Mouse Rata Negra House Rat, Black Rat Ratón Bolsero del Sur Southern Pygmy Mouse Ratón Arrocero de Coues Coues’ Rice Rat Oryzomys rostratus Ratón Arrocero Rusty Rice Rat Ototylomys phyllotis Rata Escaladora de Orejas Grandes Big-eared Climbing Rat Present Study Ratón Arrocera Pigmea del Norte Northern Pygmy Rice Rat Ibarra 2005 Nyctinomops laticaudatus* Source RODENTIA SCIURIDAE Sciurus deppei Sciurus variegatoides GEOMYIDAE Orthogeomys grandis HETEROMYIDAE Liomys salvini CRICETIDAE Mus musculus Rattus rattus Baiomys musculus Oryzomys couesi Olygoryzomys fulvescens Baseline inventory of flora and fauna SalvaNATURA Database SalvaNATURA Database SalvaNATURA Database Present Study SalvaNATURA Database 223 Order Family Scientific Name Tylomys nudicaudus Neotoma mexicana Nyctomys sumichrasti Peromyscus mexicanus Reithrodontomys gracilis Reithrodontomys mexicanus Rheomys thomasi Sigmodon hispidus ERETHIZONTIDAE Sphiggurus mexicanus DASYPROCTIDAE Dasyprocta punctata CUNICULIDAE Cuniculus paca CARNIVORA CANIDAE Canis latrans Urocyon cinereoargenteus Spanish Common Name Rata Escaladora del norte Rata Mexicana de Bosque Ratón Jocotero English Common Name Northern Climbing Rat Mexican Wood Rat Vesper Rat Source Present Study Present Study Present Study Ratón Venado Mexicano Mexican Deer Mouse Present Study Ratón Cosechador Slender Harvest Mouse SalvaNATURA Database Ratón Cosechador Mexicano Mexican Harvest Mouse Present Study Ratón de Agua Thomas's Water Mouse Rata Algodonera Hispid Cotton Rat SalvaNATURA Database Present Study Puerco Espín o Zorro Espín Mexican Porcupine Present Study Cotuza Central American Agouti Present Study Tepezcuintle Paca Present Study Coyote Coyote Nestor Herrera, unpublished data Zorra Gris o gato de monte Gray Fox Present Study Yaguarundi o Gato Zonto Jaguarundi Puma Puma Ocelote Ocelot Present Study Néstor Herrera, unpublished data SalvaNATURA Database FELIDAE Puma yagouaroundi Puma concolor Leopardus pardalis 224 Baseline inventory of flora and fauna Order Family Scientific Name Leopardus wiedii Spanish Common Name Tigrillo English Common Name Margay, Tree Ocelot Source Present Study Mephitis macroura Zorrillo Listado Common Hog-nosed Skunk Conepatus leuconotus Zorrillo Lomo Blanco Hooded Skunk Spilogale angustifrons Zorrilo Manchado Spotted Skunk Present Study Néstor Herrera, Data not published Present Study Comadreja Andina (Comadreja) Tayra Long-tailed Weasel Tayra Nutria Neotropical River Otter Micoleón Kinkajou Present Study Cacomistle, Guayanoche Central American Cacomistle Present Study Pezote Mapache Coatimundi Northern Raccoon Present Study Present Study Conejo Silvestre Eastern Cottontail Present Study Venado Cola Blanca White-tailed Deer Present Study Cuche de monte Collared Peccary Present Study Tapir o Danta Baird´s Tapir SalvaNATURA Database MEPHITIDAE MUSTELIDAE Mustela frenata Eira barbara Lontra longicaudis Rivera Muñoz 2000 Present Study SalvaNATURA Database PROCYONIDAE Potos flavus Bassariscus sumichrasti Nasua narica Procyon lotor LAGOMORPHA LEPORIDAE Sylvilagus floridanus ARTIODACTYLA CERVIDAE Odocoileus virginianus TAYASSUIDAE Tayassu tajacu PERISODACTYLA TAPIRIDAE Tapirus bairdi 9 27 101 Baseline inventory of flora and fauna 225 The records deposited in the SalvaNATURA database for this study come from specimens fround in the following museums: American Museum of Natural History (AMNH), Carnegie Museum of Natural History (CM), Museo de Historia Natural de El Salvador (MUHNES), Harrison Museum (HM), Royal Ontario Museum of Natural History, Texas Cooperative Wildlife Collection (TCWC), Museum of Vertebrate Zoology (MVZ) y The Museum of Natural History of the University of Kansas (KU). Furthermore, data has been taken from the SalvaNATURA/WCS Neobats-El Salvador project. * = Ultrasound record. Never collected in the country. 226 Baseline inventory of flora and fauna TABLE 50. LEVEL OF MAMMAL PROTECTED NATURAL AREAS. Species Recorded in the Study Species Recorded in the Site El Imposible Los Cóbanos Los Volcanes Plan de Amayo San Marcelino 53 Santa Rita Protected Natural Area INVENTORY COMPLETENESS Inventory Completeness IN THE Conservation Important Species % Previous % Current % Previous % Current 72 63 81 32 41 16 25 2 40 3 6 23 47 33 52 15 21 27 31 23 52 0 12 21 43 42 63 12 12 13 40 52 63 3 9 21 2 35 0 0 18 0 29 0 3 103 92 100 47 65 Sierra de Apaneca 21 East Sierra de Apaneca 16 West TOTAL7 73 7 The entire study area. TABLE 51. INVENTORY COMPLETENESS FOR MAMMALS AND RELATIVE IMPORTANCE OF WATERSHEDS. Watershed Barra de Santiago Barra Salada Bocana San Juan Cara Sucia Cauta Coatepeque Grande de Sonsonate Izalco Los Cóbanos San Julián San Pedro Inventory Completeness Conservation Important Species % Previous % Current % Previous % Current 54 0 15 58 0 46 42 21 2 19 13 79 0 15 83 0 67 67 31 40 46 17 18 0 0 35 0 18 3 0 3 0 3 24 0 0 41 0 21 6 0 6 18 3 Baseline inventory of flora and fauna 227 TABLE 52. INVENTORY COMPLETENESS FOR MAMMALS AND RELATIVE IMPORTANCE OF MUNICIPALITIES. Municipality Acajutla Apaneca Armenia Atiquizaya Caluco Chalchuapa Concepción de Ataco Cuisnahuat El Congo Guaymango Izalco Juayua Jujutla Nahuizalco Nahulingo Salcoatitán San Antonio del Monte San Francisco Menéndez San Julián San Pedro Puxtla Santa Ana Santa Catarina Masahuat Sto. Domingo de Guzmán Sonsonate Sonzacate Tacuba Tepecoyo Inventory Completeness Conservation Important Species % Previous % Current % Previous % Current 17 0 0 0 19 4 0 0 0 0 2 17 23 21 0 0 0 56 0 13 52 0 0 2 0 21 0 46 29 19 27 46 31 0 0 19 0 23 21 42 21 0 0 0 85 0 13 60 0 0 10 0 46 0 6 0 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 18 0 0 12 0 0 3 0 0 0 9 3 3 0 12 0 0 0 6 0 0 0 9 0 0 0 0 44 0 0 12 0 0 3 0 26 0 TABLE 53. INVENTORY COMPLETENESS FOR MAMMALS AND RELATIVE IMPORTANCE OF ECOSYSTEMS. Inventory Completeness Ecosystem Humid forest Cloud forest Dry forest Freshwater Intertidal Mangrove Savanna 228 Conservation Important Species % Previous % Current % Previous % Current 48 50 69 33 0 33 2 79 67 90 33 0 42 40 15 9 24 6 0 6 3 32 21 47 6 0 9 6 Baseline inventory of flora and fauna Another way of calculating inventory completeness is by using species accumulation curves. For this study the accumulation curve is still not stabilized after 98 days of standardized sampling efforts (Fig.19). Based on this curve, different estimators project from 91 up to 138 species of land mammals in the southwestern region of El Salvador (Table 54). On the other hand, taking into account that we have already recorded 101 species and we are still missing 12 species of conservation importance that are potentially present in the area, we can estimate the existence of at least 113 terrestrial mammal species in the study area. In any case, we expect additional species, particularly bats and rodents, even though 100% of the expected generalists have already been detected. FIGURE 19. SPECIES ACCUMULATION CURVE OF TERRESTRIAL MAMMALS IN SOUTHWESTERN EL SALVADOR. For ecosystems, the accumulation curves also do not become stabilized (Fig. 20). For Dry Forest, the richness calculators estimate there could be up to 69 species, for the Humid Forest up to 75, for the Cloud Forest up to 43 and for Savanna up to 26 (Table 54). In the case of Dry Forest, 78 species have been recorded including previous studies (greater than the estimated richness) and very few expected species are still missing to complete the inventory; it is a similar case with the Humid Forest and the Cloud Forest, reporting 70 and 57 species, respectively. Savanna reports 25 species whereas a maximum of 26 are estimated. Baseline inventory of flora and fauna 229 TABLE 54. ESTIMATORS OF TERRESTRIAL MAMMAL SPECIES RICHNESS IN THE STUDY AREA AND IN EACH ECOSYSTEM. Estimated Richness (Colwell 2006) Richness Estimator ACE ICE Chao 1 Chao 2 Jack-Knife 1 Jack-Knife 2 Average Total Area Dry Forest Humid Forest Cloud Forest Savanna 92 95 138 99 91 107 104 67 63 67 59 60 69 64 54 67 53 68 62 75 63 34 40 33 36 40 43 38 20 23 19 21 23 26 22 The fact that both the Dry Forest and Cloud Forest present lists with more species than were estimated from the species accumulation curve, suggests that some of the species reported in the past are already locally extinct. Some examples of possibly extinct species are: Chrotopterus auritus collected only once in 1927 at the Barra de Santiago, Reithrodontomys gracilis collected for the last time in 1958 at Lake Coatepeque and Tapirus bairdi for which there is a report of a skull found in 1987 near El Imposible National Park although it is unknown when the specimen was alive. Although there are no formal records, elderly persons living in the area still remember when the jaguar (Pantera onca) existed (Karla Lara, pers. comm.) and it is also possible that the Spider Monkey Ateles geoffroyi lived in the area many years ago. FIGURE 20. ACCUMULATION CURVES OF TERRESTRIAL MAMMAL SPECIES BY ECOSYSTEM. 230 Baseline inventory of flora and fauna 3.3. SPECIES OF CONSERVATION IMPORTANCE Twenty-two threatened species, indicators of conservation importance, have been recorded in the study area. Examples include the Margay Cat (Leopardus wiedii), Ocelot (Leopardus pardalis), Central American River Otter (Lontra longicaudis), and the Baird’s Tapir (Tapirus bairdi). There are 11 threatened species and seven endangered species at the national level (MARN 2004), and also one globally endangered species (IUCN 2008) (See Table 47). Below is a description of some important species: Trachops cirrhosus (Fig. 18D). Up until 2004 this species was only known in El Salvador from three sites in the departments of La Libertad and San Miguel. On 19 September 2007, we found the second record for Santa Rita and the second record since 1953 in the country. The first record from Santa Rita was in June 2005 during the Neobats-El Salvador project. The species is considered nationally threatened (MARN 2004). Micronycteris microtis (Fig. 18E). This species had not been captured in El Salvador since 1978 and was only known in El Salvador from four sites in the departments of San Miguel and La Libertad. We recorded the species at Plan de Amayo on 21 July 2007 and at El Imposible National Park on 31 October 2007. These are the first records for the project area and for the departments of Sonsonate and Ahuachapán, respectively. This species is considered nationally threatened (MARN 2004). Bauerus dubiaquercus (Fig. 18G). On 15 December 2007, this species was recorded for the first time in El Salvador at El Imposible National Park (Girón et al. in press). Previously, its known distribution range was highly fragmented, with populations known from Jalisco, Guerrero and the Island of Tres Marías in Mexico, and on the Caribbean slopes of Honduras and Costa Rica. It has a globally Near-Threatened status (IUCN 2008). Neotoma mexicana (Fig. 18I). This species had only been collected in Montecristo and at the Santa Ana Volcano. On 9 September 2007, we found an individual at Cerro Verde, the first collection of the species since 1979. This rodent is considered nationally endangered (MARN 2004). Leopardus wiedii (Fig. 18Ñ). This species was only documented from five sities around the country: Cerro Cacahuatique, Montecristo, Jucuarán, La Montañona, and El Imposible. We provide the first record for Plan de Amayo, from a camera trap on 22 July 2007. 3.4. EVALUATION OF CONSERVATION IMPORTANCE FOR THE SITES AND ECOSYSTEMS UNDER STUDY. Of the list of 34 threatened species identified as indicators of site importance, the proportion actually recorded in the project area increased from 47% to 65%. At the watershed level, only two have reasonably complete inventories, permitting comparisons of site importance. The Cara Sucia watershed has 41% indicators for site importance, Baseline inventory of flora and fauna 231 while the Barra de Santiago has only 24% (Table 51). Only one municipality has a reasonably complete inventory, therefore the relative importance of the municipalities cannot be analyzed. Regarding ecosystems, only two relatively complete inventories can be compared. The Dry Forest (47%) is more important than the Humid Forest (32%) (Table 53). Only one Protected Natural Area has a nearly complete inventory, therefore the Protected Natural Areas cannot be analyzed for relative importance. 4. RECOMMENDATIONS To complement the efforts made and complete the inventories, we recommend working at the following sites: San Marcelino, Plan de Amayo and in the coffee plantations of the Apaneca mountain range to see if mammal species may be using the scattered remnants of natural forest and shaded plantations as a biological corridor. For example, the Large Brown Bat (Eptesicus fuscus) has been recorded at the eastern extreme of the mountain range (Santa Ana Volcano) and at west side of the range, at Laguna de Las Ninfas. San Marcelino and Plan de Amayo are important sites for mammal conservation but have not been studied in depth and their inventories are incomplete. Permanent monitoring stations with camera traps should be established in the natural areas where felines have been detected; little is known about their populations and all of the members of the cat family are nationally threatened, which makes it important to study these species. The Anabat acoustic sampling technique should be carried out to detect high flying bats as these are captured only on very rare occasions, such as the Greenhall Dog-faced Bat (Molossops greenhalli), which has only been recorded in El Salvador by that technique. It is very important to monitor and carry out ecological studies of some mammal species that enter into conflict with humans due to their interference with agricultural crops. This situation is mentioned constantly in communities adjoining the natural areas and we recommend studying this problem. If it is documented that some species may cause more harm to the crops and therefore generate economic losses, then a strategy could be established to minimize the problem without harming the mammal populations involved. Some mammal population controls may be warranted, because of the demise of natural predators, such as the jaguar (Panthera onca) and the puma (Puma concolor). 232 Baseline inventory of flora and fauna REFERENCES Amaya, J.A., S.M. García & T.A. Valladares. 2003. Estudio de la composición florística del estrato arbustivo en el Cerro ―El Águila‖ Municipio de Chalchuapa, Santa Ana.Tésis de Licenciatura. El Salvador: Universidad de El Salvador. 49 p. Barahona, D. E. & A. V. Henríquez. 2007. Captura incidental de tortugas marinas en la pesca de palangre de fondo con anzuelos ―j‖ y anzuelos circulares 13 y 14, y su efectividad en la pesca de peces demersales de la costa salvadoreña. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 98 pp. Baum, J.K., Myers, R.A., Kehler, D.G., Worm, B., Harley, S.J., & Doherty, P.A. 2003. Collapse and conservation of shark populations in the Northwest Atlantic. Science 299:389-392. Bishop S.D.H, Francis M.P, Duffy C. &, Montgomery J.C. 2006. Age, growth, maturity, longevity and natural mortality of the shortfin mako (Isurus oxyrinchus) in New Zealand waters. Marine and Freshwater Research 57: 143–154. Bjork, R. D. 2009. Reintroduction of the Scarlet Macaw to El Salvador. Parrots International Magazine 1, on-line (3 pp.). Boeseman, M. 1956. Sobre una colección de peces de la República de El Salvador. Comunicaciones, Instituto Tropical de Investigaciones Científicas, Universidad de El Salvador, El Salvador. 4: (2/3). 75–88 pp. Burt, W. H. & R. A. Stirton. 1961. The mammals of El Salvador. Miscellaneous Publications Museum of Zoology, University of Michigan 117: 1–69. Bussing, W. A. 1976. Geographic distribution of the San Juan Ichthyofauna of Central America with remarks on its origin and ecology, pp. 157–175 In T.B. Thorson (eds). Investigations of Nicaraguan Lakes. School of Life Sciences, Univ. Nebraska. Bussing, W. A. 1987. Los peces de las aguas continentales de Costa Rica. Editorial Universidad de Costa Rica. 287 pp. Bussing, W. A. 1998. Peces de las aguas continentales de Costa Rica/Freshwater fishes of Costa Rica. Editorial Universidad de Costa Rica. 448 pp. Bussing, W. A. & M. I. López. 1993. Peces demersales y pelágicos costeros del Pacífico de Centroamérica Meridional. Rev. Biol. Trop. Publicación Especial: 1–164 pp. Cabrera Martínez, G. I. & D. I. Mejía Marroquín. 1998. Vegetación del bosque las Lajas, Refugio de Vida Silvestre del Complejo San Marcelino, Santa Ana - Sonsonate, El Salvador. Tesis de Licenciatura. El Salvador: Universidad de El Salvador. 86 p. Baseline inventory of flora and fauna 233 Calderón, M. G. & R. A. Hernández. 1974. Estudio biológico pesquero de la Bahía de Jiquilisco. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 207–218 pp. Candray, M. A. & C. W. Ríos. 2005. Composición y abundancia de la ictiofauna con importancia comercial en el Estero de Jaltepeque, San Luís La Herradura, Departamento de La Paz, El Salvador. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 55 pp. Castro, R. A. 1976. Ictiofauna y algunas determinaciones físico-químicas en el Río Lempa. Tesis para optar al grado de Licenciatura en Biología, Departamento de Biología, Facultad de Ciencias y Humanidades, Universidad de El Salvador. 79 pp. Castro, R. A. 1983. Los métodos y aperos de pesca más importantes utilizados en los lagos, lagunas y embalses de El Salvador, su eficiencia en lo económico y en la captura por especie. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 20 pp. Castro R. A. & J. P. Robinson. 1976. Estudio Preliminar Biológico Pesquero del Río Lempa. Dirección General de Recursos Naturales Renovables, MAG, Volumen III, El Salvador. 188 pp. Castro R. A. & F. Godínez. 1978. Evaluación pesquera en el Lago de Ilopango y Laguna de Olomega. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 36 pp. Centro de Desarrollo de la Pesca y la Acuicultura. 2006. Estadísticas Pesqueras y Acuícola, Vol. 33. MAG, El Salvador. 70 pp. Cerén, J.G.; G.N. López & D. Delcid. 2003. Estudio de la composición florística del estrato herbáceo en el área natural del Cerro ―El Águila‖.Tesis de Licenciatura. El Salvador: Universidad de El Salvador. 86 p. Chicas, F. A., J. A. González, & W. E. Ramírez. Ecología y manejo de las comunidades de peces del Golfo de Fonseca, Departamento de La Unión, El Salvador. Centro de Investigaciones Científicas, Universidad de El Salvador, (en Prensa). Colwell, R. K. 2006. Statistical estimation of species richness and shared species from samples. University of Connecticut, United States. www.viceroy.eeb.uconn.edu/estimate Concheiro Pérez, G. A., L. Oldlich, G. Ortí, G. E. Bermingham, I. Doadrio, & R. Zardoya. 2005. Phylogeny and biogeography of 91 species of heroine cichlids (Teleostei: Cichlidae) based on sequences of the cytochrome b gene. Molecular Phylogenetics and Evolution 43 (2007) 91–110. Cortes, J. 1997. Biology and ecology of eastern Pacific coral reefs. Coral Reefs 16: S39– S46. 234 Baseline inventory of flora and fauna Cruz, G. A. 1987. Reproductive biology and feeding habits of Cuyamel, Joturus pichardi and Tepemechín, Agonostomus monticola (Pisces: Mugilidae) from Rio Plátano, Mosquitia, Honduras. Bull. Mar. Sci. 40: 63–72. Day, J. W., C. A. Hall, W. M. Kemp & A. Yáñez-Arancibia. 1989. Estuarine Ecology. John Willey & Sons, Inc. 557 pp. Felten, H. 1956. Quirópteros en El Salvador. Revista Comunicaciones. Instituto Tropical de Investigaciones Científicas. pp. 153–170. Fischer, W., F. Krupp, W. Shneider, C. Sommer, K.E. Carpenter, & V.H. Niem. 1995. Guía FAO para la identificación de especies para los fines de la pesca. Pacifico Centro-Oriental. Volumen II. Vertebrados. Parte I. Roma FAO, 647–1200 pp. Francis, M. P., & Duffy, C. 2005. Length at maturity in three pelagic sharks (Lamna nasus, Isurus oxyrinchus, and Prionace glauca) from New Zealand. Fishery Bulletin 103: 489– 500. Froese, R. & D. Pauly (Eds). 2007. Fishbase. World Wide Web electronic publication. www.fishbase.org, versión (04/2008). Fuentes, R., C. I. & N. R. Hernández R. 2004. Distribución y abundancia de la ictiofauna con importancia comercial asociada a la pesca de arrastre de camarones Peneidos (Penaeus stylirrostris, P. vannamei, P. occidentalis, P. californiensis y P. brevirostris) en la Costa Salvadoreña. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 85 pp. Galdámez, A. 2002. Composición de los peces capturados y la madurez sexual de las especies de ―pargo‖ (Lutjanidae) en Los Cóbanos, Sonsonate. Trabajo de graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 97 pp. García, R. C. 1974. Inventario y esfuerzo de pesca en el Río Grande de San Miguel. Servicio Piscícola, Ministerio de Agricultura y Ganadería. 28 pp. García, R. C. & J. P. Robinson. 1974. Estudio Biológico Pesquero del Río Paz. Dirección General de Recursos Naturales Renovables. MAG, El Salvador. 42 pp. García Rivas, J. R. 1997. Vegetación Arbórea de Las Lajas, Refugio de Vida Silvestre, Complejo San Marcelino, Sonsonate, El Salvador. Tesis de Licenciatura. El Salvador: Universidad de El Salvador. 117 p. Garrison, G. 2005. Peces de la Isla del Coco/ Isla del Coco Fishes. 2da edición, Santo Domingo de Heredia, Costa Rica: Instituto Nacional de Biodiversidad, INBIO, 430 pp. Gierloff-Emden, H. G. 1958. Los ríos de El Salvador y sus subcuencas tributarias. Comunicaciones, Instituto Tropical de Investigaciones Científicas. 7, 3/4: 23–35 pp. Baseline inventory of flora and fauna 235 Girón, L. 2005. Identificación y distribución de los murciélagos del sector Los Andes del volcán de Santa Ana, Complejo Los Volcanes, Santa Ana, El Salvador. Tesis de licenciatura. Universidad de El Salvador. 76 pp. Girón, L. E. & J.G. Owen. 2007. Status of the inventory of terrestrial mammalian fauna. Pages 74–84 in Komar, O., editor, Preliminary inventory in eleven hydrographic watersheds of southwest El Salvador. USAID IMCW Project /Education for Conservation Colection No. 3. Girón, L. E., Owen, J. G., & Rodríguez, M. E. In press. Van Gelder´s Bat: first record of Bauerus dubiaquercus (Chiroptera: Vespertilionidae) from El Salvador, Central America. Southwest Naturalist. Godínez, J. F. & R. Castro. 1977. Estudio sobre selección de redes agalleras, su introducción en el Lago de Ilopango y Laguna de Metapán. 71 pp. Gómez de Silva, H. & R. A. Medellín. 2001. Evaluating completeness of species lists for conservation and macroecology: a case study of Mexican land birds. Conservation Biology 15: 1384–1395. González, J. C. 1977. La vegetación arbórea del pedregal de San Isidro: un análisis florístico y cuantitativo. Tesis de Licenciatura, Universidad de El Salvador, San Salvador. 39 p. González, R. G. 1995. Los peces nativos en vías de extinción en las aguas continentales de El Salvador, Proyecto Fortalecimiento a la Acuicultura, Programa Regional de Apoyo al Desarrollo de la Pesca en el Istmo Centroamericano (PRADEPESCA), El Salvador. 69 pp. González, J. A. & W. E. Ramírez. 2007. Ictiofauna del Estero Jaltepeque, con Énfasis en Doce Especies de Importancia Comercial, El Salvador. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 78 pp. Greenbaum E. & O. Komar. 2005. Threat assessment and conservation prioritization of the herpetofauna of El Salvador. Biodiversity and Conservation 14: 2377–2395. Harvey, C. A., Komar, O., Chazdon, R., Ferguson, B. G., Finegan, B., Griffith, D. M., Martínez-Ramos, M., Morales, H., Nigh, R., Soto-Pinto, L., van Breugel, M., & Wishnie, M. H. 2008. Integrating agricultural landscapes with biodiversity conservation in the Mesoamerican hotspot. Conservation Biology 22: 8–15. Henríquez, V. 2004. Sistema de monitoreo de indicadores biológicos del Área Natural Protegida Complejo Los Volcanes. Estudio de anfibios y reptiles del Complejo Los Volcanes. Informe de consultoría para CATIE. 72 pp. Henríquez, V. 2009. Las KBAs de El Salvador: Establecimiento de una Línea Base de Áreas Claves para la Biodiversidad. SalvaNATURA, San Salvador, El Salvador 29 pp. Available at www.salvanatura.org. 236 Baseline inventory of flora and fauna Henríquez, V. & O. Komar. 2006. Los anfibios y reptiles del Parque Nacional Los Volcanes, El Salvador: Sus valores para la conservación y el ecoturismo. Forma parte de: Komar, O., J. P. Arce, J. R. Smith & I. Varley, 2006. Evaluación de la biodiversidad del Parque Nacional Los Volcanes, El Salvador. Informe de consultoría para el Banco Interamericano de Desarrollo. SalvaNATURA Programa de Ciencias para la Conservación, San Salvador, El Salvador y NatureServe, Washington, D.C., USA. 18 pp. Henríquez, V. & R. Vaquerano. 2008. Sphaerodactylus glaucus (Cope, 1865) (Sauria, Gekkonidae) nuevo registro para la herpetofauna de El Salvador. Acta Zoológica Mexicana 24 (1): 237– 238. Hernández, R. R. 2006. Diversidad biológica en Chiapas. Capitulo 5, Diversidad de peces continentales en Chiapas. Colegio de la Frontera Sur (ECOSUR).195–221 pp. Herrera, N., V. Henríquez, and E. Greenbaum. 2007. New department and country records for amphibians and reptiles from El Salvador. Herpetological Review 38:222– 226. Herrera, N. & N. Valle Andrade. 2003. Estudio de Flora y Fauna Vertebrada del Área Natural Protegida Plan de Amayo, Caluco, departamento de Sonsonate, El Salvador. Para Asociación AGAPE de El Salvador. Hildebrand, S. F. 1925. Fishes of the Republic of El Salvador, Central America. Bull. of Bureau of Fisheries, Dept. of Commerce. U.S.A., Document N. 985. 41: 237–287 pp. Howell, S. N. G. & S. Webb. 1995. A guide to the birds of Mexico and Central America. Oxford Univ. Press, New York. Hrbek T.; J. Seckinger & A. Meyer 2007. A phylogenetic and biogeographic perspective on the evolution of poeciliid fishes. Molecular Phylogenetics and Evolution 43 (2007) 986–998. Huezo Chinchilla, H. L. & T. A. Rosales Menéndez. 2001. Evaluación de la selectividad de redes agalleras y determinación de la madurez sexual de los cíclicos comerciales del Lago de Ilopango, El Salvador. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 65 p. Ibarra, R. 2005. Flora y Fauna Vertebrada del Área Natural Protegida ―Santa Águeda-El Zope‖ (Complejo Los Cóbanos), Sonsonate, El Salvador. Ministerio de Medio Ambiente y Recursos Naturales. 44 pp. ICMARES/UES. 2006. Diagnostico de la línea base de las condiciones Biofísicas, Socioeconómicas e Institucionales del Sistema Arrecifal de Los Cóbanos. Instituto de Ciencias del Mar y limnología, Universidad de El Salvador, El Salvador, C.A. 2/3: 77– 83 pp. Baseline inventory of flora and fauna 237 ICMARES/UES. 2007. Evaluación del área de dragado en la playa Las Veraneras, Sistema Arrecifal Los Cóbanos, Departamento de Sonsonate, El Salvador. Universidad de El Salvador. 54 pp. IUCN. 2003. Guidelines for Application of IUCN Red List Criteria at Regional Levels: Version 3.0. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. ii + 26 pp. IUCN. 2007. 2007 IUCN Red List of Threatened Species. <www.redlist.org> IUCN, Gland, Switzerland and Cambridge, UK. Accessed 23 June 2008. IUCN. 2009. 2009 IUCN Red List of Threatened Species. <www.redlist.org> IUCN, Gland, Switzerland and Cambridge, UK. Accessed 10 July 2009. Jiménez, D. J. 1986. Efecto de la luz en la pesca de dos especies de sardinas en el estero de la Barra de Santiago. Tesis para optar al grado de Licenciatura en Biología, Facultad de Ciencias y Humanidades, Universidad de El Salvador 78 pp. Jiménez-Ramón, J. A. 1994. Los Manglares del Pacífico Centroamericano. Editorial Heredia, Editorial Fundación UNA, Organización para Los Estudios Tropicales (OTS) Costa Rica, 352 pp. Johnson, K. 1971a. Reporte preliminar sobre la pesca del Lago de Gϋija, El Salvador. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 45 pp. Johnson, K. 1971b. A preliminary report on the fishery of Olomega Lagoon, El Salvador. Report presented to the Ministerio de Agricultura of El Salvador. 20 pp. Johnson, K. & A. Argumedo. 1971a. Lago de Ilopango, 1971, un estudio preliminar de la pesca del Lago. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 22 pp. Johnson, K. & A. Argumedo. 1971b. La pesca en el Lago de Coatepeque. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 15 pp. Johnson, K. & M. Ramírez. 1975. Estudio de la pesca en la Laguna de Apastepeque. Dirección General de Recursos Naturales Renovables, MAG. El Salvador. 40 pp. Jones, H. L. & O. Komar. 2008. Central America (in The Regional Reports: The Nesting Season, June and July 2007). North American Birds 61: 648–651. Köhler, G., M. Veselý, & E. Greenbaum. 2006. The amphibians and reptiles of El Salvador. Krieger Press, Melbourne, Florida. 238 pp. Komar, O. 2002. Priority conservation areas for birds in El Salvador. Animal Conservation 5: 173–183. Komar, O. 2003. En las listas faunísticas, no todas las especies son iguales: las aves de El Salvador como un ejemplo. Pp. 57–61 En H Gómez de Silva & A. Oliveras de Ita (eds.). Conservación de aves: experiencias en México. Cipamex. México, D.F. 238 Baseline inventory of flora and fauna Komar, O., editor. 2007. Preliminary inventory in eleven hydrographic watersheds in southwest El Salvador. Status of the flora and fauna inventory through 2006. USAID El Salvador IMCW Project Conservation Education Collection No. 3. 130 pp. Komar, O. 2007b. Status of the inventory of birds. Pp. 85–130 en Komar, O. (ed.) Preliminary inventory in eleven hydrographic watersheds in southwest El Salvador: Status of the flora and fauna inventory through 2006. USAID El Salvador IMCW Project Conservation Education Collection No. 3. Komar, O., Borjas, G., Cruz, G. A., Eisermann, K., Herrera, N., Linares, J. L., Escobar, C. E., & Girón, L. 2006. Evaluación ecológica rápida en la propuesta área protegida trinacional Montecristo en territorio guatemalteco y hondureño. SalvaNATURA Informe de Consultoría, para el Banco Interamericano de Desarrollo y la Comisión Trinacional Plan Trifinio. 170 pp. Komar, O. & L. E. Girón. 2007. Prioridades para la conservación de la biodiversidad en el suroccidente de El Salvador. USAID Proyecto MMCC/Colección Educación para la Conservación 2. 109 pp. Komar, O. & N. Herrera. 1995. Diversidad de Avifauna en el Parque Nacional El Imposible y el Refugio de Vida Silvestre del Complejo San Marcelino. El Salvador. Working Series (Wildlife Conservation Society, Bronx, NY) No. 4. 76 pp. Komar, O., N. Herrera, R. Ibarra Portillo & L. Girón. In prep. Lista Roja de aves de El Salvador. SalvaNATURA Biodiversity Series No. 3, San Salvador. Komar, O., & Ibarra-Portillo, R. 2009. El Salvador. Pp. 197–204 in C. Devenish, D. F. Díaz Fernández, R. P. Clay, I. Davidson & I. Yépez Zabala, Eds. Important Bird Areas Americas - Priority sites for biodiversity conservation. Quito, Ecuador: BirdLife International (BirdLife Conservation Series No. 16). Kushlan, J. A., M. J. Steinkamp, K. C. Parsons, J. Capp, et al. 2002. Waterbird Conservation for the Americas: The North American Waterbird Conservation Plan, Version 1. Washington, D.C., U.S.A. Leenders, T. 2003. Los Anfibios y Reptiles. Pp. 158–161. En: Alvarez, J. M. y Komar, O. (Eds): El Parque Nacional El Imposible y su Vida Silvestre. Biodiversity Series No. 2. San Salvador. Lin, S. Y. 1957. Estudio general de los recursos de la pesquería regional de El Salvador, MAG, El Salvador. 57 pp. Linares, J. L. 2003 [2005]. Listado comentado de los árboles nativos y cultivados en la república de El Salvador. Ceiba 44(2):105–268. Linares, J. L. 2007. Status of the inventory of flora (tres and orchids). Pp. 24–55 in Komar, O. (editor.). Preliminary inventory in eleven hydrographic watersheds of southweset El Salvador: Status of the flora and fauna inventory through 2006. USAID El Salvador IMCW Project Conservation Education Collection No. 3. Baseline inventory of flora and fauna 239 Linares, J. L. & M. Sousa. 2008. Nuevas especies de Dalbergia (Leguminosae: Papilionoideae: Dalbergieae) en México y Centroamérica. Ceiba 48(1-2):61–82 López, T. N. 1995. Estudio del desarrollo gonadal y análisis del contenido estomacal de la ―mojara‖ (Cichlasoma guija) en el Lago de Ilopango. Tesis para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 50 pp. Martínez, C. E., Henríquez Cisneros, V. E., & Komar, O. 2009. Conozcamos la Biodiversidad. Manuales del Programa de Capacitaciones Comunidades y Biodiversidad (SalvaNATURA, El Salvador) Componente 2, 41 pp. Mejía, L. A. & F. A. Chicas. 2007. La pesca de sardinas y otros peces juveniles en la Bahía de Jiquilisco, El Salvador. Rev. Quehacer Científico. Vol. 6, Año 4. Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 19–26 pp, Méndez, M. F. 2006. Inventario de Ictiofauna de Salamar, Cuenca Sur de la Sierra de Jucuarán, Departamento de Usulután, El Salvador. Programa de Ciencias para la Conservación. SalvaNATURA, San Salvador, El Salvador. Informe de consultoría Anexo 3. 12 pp. MARN (Ministerio de Medio Ambiente y Recursos Naturales). 2000a. Estrategia Nacional de Diversidad Biológica. Ministerio de Medio Ambiente y Recursos Naturales, San Salvador, El Salvador. MARN (Ministerio de Medio Ambiente y Recursos Naturales). 2000b. Colección de CD’s Medio Ambiente: CD 2 Sistema de Información Ambiental del Ministerio y Atlas de Índices de Sensibilidad de Toda la Zona Costera El Salvador C.A. MARN (Ministerio de Medio Ambiente y Recursos Naturales). 2004. Listado Oficial de Especies de Fauna Silvestre Amenazada o en Peligro de Extinción en El Salvador. Diario Oficial (San Salvador, El Salvador) Tomo No. 363, Número 78. Acuerdo No. 10. pp. 5–27. MARN (Ministerio de Medio Ambiente y Recursos Naturales). 2005. Estado del conocimiento de la Biodiversidad en El Salvador, Documento Final. Informe técnico preparado por Melibea Gallo, 154 pp. MARN (Ministerio de Medio Ambiente y Recursos Naturales). 2006. Ley del Medio Ambiente y sus Reglamentos. Ministerio de Medio Ambiente y Recursos Naturales, 282 pp. Monterrosa, E. 1960. Observaciones sobre la pesca con redes agalleras en el Lago de Gϋija. Dirección General de Recursos Naturales Renovables, MAG, El Salvador. 3 pp. Mora, M., J. Cabrera & G. Galeano. 1997. Reproducción y alimentación del gaspar Atractosteus tropicus (Pisces: Lepisosteidae) en el Refugio Nacional de Vida Silvestre Caño Negro, Costa Rica. Rev. Biol. Trop. 45: 861–866 pp. 240 Baseline inventory of flora and fauna Morrone, J. J. 2007. Hacia una biogeografía Evolutiva. Rev. Chil. Hist. Nat. Santiago de Chile. Vol. 80. Nº 4. Orellana, J. J. 1985. Peces Marinos de Los Cóbanos, Peces de El Salvador. FUNDACION SIGMA. New Cork. 126 pp. Orellana, J. J. 1992. Inventario preliminar de los peces de agua dulce y marinos de El Salvador, América Central. Secretaria Ejecutiva del Medio Ambiente (SEMA), El Salvador. 150 pp. Pacheco, S. R. & S. N. Cokkom. 2007. Efectividad de anzuelos circulares números 15 y 16 con respecto a anzuelos ―j‖ en la pesca con palangre de especies altamente migratorias y su incidencia en la captura de especies no objetivo, dentro de la zona económica exclusiva de El Salvador. Trabajo de Graduación para optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador, 53 pp. Paz Quevedo, O. W. & N. E. Ventura Centeno. 2004. Protección y Conservación del Área Natural Protegida Santa Rita y Laguna Gamboa. Inventario de Flora y Fauna del Área Natural Protegida Santa Rita. Ahuachapán. Para ADESCONE. 51 pp. Perdices, A. B., E. A. Birmingham, A. B. Montilla, A. B. & I. B. Doatrio. 2002. Evolutionary history of the genus Rhamdia (Teleostei: Pimelodidae) in Central América. Molecular Phylogenetics and Evolution 25 (2002) 172–189 pp. Phillip, D.A. 1983. Reproduction and feeding of the mountain mullet, Agonostomus monticola, in Trinidad, West Indies. Environmental Biology of Fishes 37: 47–55 Phillips, P. & J. B. Ulloa. 1981. Estudio de las Poblaciones de Peces de la Bahía de Jiquilisco. El Salvador. 45 pp. PNUMA (Programa de las Naciones Unidas para el Medio Ambiente). 2004. Geo Centroamérica: Perspectivas del Medio Ambiente 2004. 68 pp. PREPAC (Plan Regional de Pesca y Acuicultura Continental, El Salvador). 2005. Inventario de cuerpos de agua continentales de El Salvador, con énfasis en la pesca y la acuicultura. 477 pp. Ramírez, C. R. 2001. Vegetation of a Subtropical Pre-Montane Moist Forest in Central America. Tesis de grado Doctor of Philosophy. Facultad de Biología, Universidad de New York. Consultada el 22 de marzo de 2002. Disponible en: http://it.stlawu.edu/~cram/El Imposible/Diss.html Reid, F. A. 1997. Mammals of Central America and Southeast Mexico. Oxford University Press, New York. Remsen, J. V. Jr. 1994. Use and misuse of bird lists in community ecology and conservation. Auk 111: 225–227. Baseline inventory of flora and fauna 241 Reyes Bonilla, H. & J. E. Barraza. 2003. Corals and associated marine communities from El Salvador. En: J. Cortés (ed.). Coral reefs of Latin America. Elsevier (Amsterdam, Holanda), 331–340 pp. Rich, T. D., Beardmore, C. J., Berlanga, H., Blancher, P. J., Bradstreet, M. S. W., Butcher, G. S., Demarest, D. W., Dunn, E. H., Hunter, W. C., Iñigo-Elias, E. E., Kennedy, J. A., Martell, A. M., Panjabi, A. O., Pashley, D. N., Rosenberg, K. V., Rustay, C. M., Wendt, J. S., & Will, T. C. 2004. Partners in Flight North American landbird conservation plan. Cornell Lab of Ornithology, Ithaca, New York. Rivera Muñoz, J. R. 2000. Conservación, protección y recuperación de los recursos naturales del Refugio de Vida Silvestre Complejo San Marcelino. Estudio de fauna de las clases Amphibia, Reptilia, Aves y Mammalia en el Complejo San Marcelino. Asociación Salvadoreña de Conservación del Medio Ambiente. Fondo Iniciativas para las América El Salvador – FIAES. 36 pp. Robertson, D. R. & G. R. Allen, 2002. Peces costeros del Pacífico Oriental Tropical: Un Sistema de Información. Instituto Smithsonian de Investigaciones Tropicales, Balboa, República de Panamá. Robertson, D. R. & G. R. Allen. 2006. Peces Costeros del Pacífico Oriental Tropical: Un Sistema de Información. Instituto Smithsonian de Investigaciones tropicales, Balboa, República de Panamá. Robertson, D. R & G. R. Allen. 2008. Shorefishes of the Tropical Eastern Pacific online information system. Version 1.0. Smithsonian Tropical Research Institute, Balboa, Panamá. www.neotropicalfishes.org/sftep, www.stri.org/sftep. Rojas, M. J., E. Maravilla & F. A. Chicas. 2004. Hábitos alimentarios del ―pargo mancha‖ Lutjanus guttatus (Pisces: Lutjanidae) En los Cóbanos y Puerto de La Libertad, El Salvador Rev. Biol. Trop., Vol. 52, Nº 1, 163–170. Ruiz, L. J. Torres de J. A. Mendez, E. & A. S. Prieto. 2003. Aspectos biométricos de Sparisoma aurofrenatum (Pisces: Scaridae) de un parche arrecifal del Parque Nacional Mochima, Venezuela. Biol., dic. 2003, vol, Nº 3, 161–170. Salgado-Maldonado, G., F. Moravec, G. Cabañas-Carranza, R. Aguilar-Aguilar, P. SánchezNava, R. Báez-Valé, & T. Scholz. 2004. Helminth parasites of the tropical gar, Atractosteus tropicus Gill from Tabasco, Mexico. Journal of Parasitology 90: 260–265. Schmitter-Soto, J. J. 2007. A systematic revision of the genus Archocentrus (Perciformes: Cichlidae), with the description of two new genera and six new species. Zootaxa 1603: 1–78. Segovia Prado, J. V. Fauna ictiológica del sistema arrecifal Los Cóbanos, El Salvador. (En preparación). Segovia, J. V. & M. Navarrete. 2007. Biodiversidad en parches de coral hermatípicos (Porites lobata y Pocilopora sp.) a nivel de Ecosistema en el Sistema Arrecifal Los Cóbanos, Departamento de Sonsonate, El Salvador. Trabajo de Graduación para 242 Baseline inventory of flora and fauna optar al grado de Licenciatura en Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 70 pp. Solís, I. B., 2006. Distribución y abundancia del pez machorra, Atractosteus tropicus, en el Zanjon el Chino, Municipio de San Francisco Menéndez, Ahuachapán, El Salvador. Escuela de Biología, Facultad de Ciencias Naturales y Matemática, Universidad de El Salvador. 46 pp. Soto-Galera, E. 2006. Ictiofauna de los corredores biológicos Sierra Madre Sur y Selva Maya Zoque. Instituto politécnico Nacional. Informe final SNIB-CONABIO proyecto Nº Y026. México D.F., 14 pp. Stevens, W. D., C. Ulloa U., A. Pool & O. M. Montiel (Eds). 2001. Flora de Nicaragua. Monographs in Systematic Botany from the Missouri Botanical Garden. 85 Ulloa, J. B. & E. A. Melgar. 1983. Estudio preliminar de las poblaciones del ―pez ejote‖ (Melaniris guija) en el Lago de Ilopango, El Salvador. CENDEPESCA. MAG. El Salvador. 11 pp. Ulloa, J. B. & R. Castro. 1985a. Situación actual de la pesca en la Laguna de Metapán, algunas relaciones técnicas para su manejo. CENDEPESCA. MAG. El Salvador. 8 pp. Ulloa, J. B. & R. Castro. 1985b. Diagnóstico de la pesca en el Lago de Gϋija, relacionada con el empleo de diversos métodos y aperos de pesca. CENDEPESCA. MAG. El Salvador. 23 pp. Ulloa, J. B., N. Avalos, & T. Sorto. 1988. Informe sobre la producción pesquera del embalse Cerrón Grande 1986-1987. CENDEPESCA. MAG. El Salvador. 23 pp. Ulloa, J. B., E. A. Hernández, J. A González & W. E. Ramírez. 2003. Evaluación de Algunos Aspectos Biológicos de la Captura de Peces y Camarones Obtenida con Redes de Enmalle Número Nueve en El Estero de Jaltepeque, CENDEPESCA, MAG, El Salvador, 24 pp. Vásquez, M. J. 1993. Ictiofauna del Estero Barra de Santiago, Publicaciones ocasionales, Museo de Historia Natural de El Salvador. 40 pp. Villa, J. 1971. Sinopsis de los peces de nicaragua. 3ª edición, Departamento de Biología, Universidad Nacional Autónoma de Nicaragua, Managua, 132 pp. Villa, J. 1982. Peces nicaragüenses de agua dulce. Colección Cultural Serie Geografía y Naturaleza. 253 pp. Villacorta, R. 2000. Estudio preliminar de la flora del Complejo de San Marcelino. Asociación Salvadoreña para la Conservación del Medio Ambiente (ASACMA). Waid, R. M., R.L. Raesly, K.R. McKaye & J.K. McCrary. 1999. Zoogeografía íctica de lagunas cratéricas de Nicaragua. Encuentro (Revista de la Universidad Centroamericana, Nicaragua) 51: 65–80. Baseline inventory of flora and fauna 243 Yáñez-Arancibia, A. 1975. Sobre los estudios de peces en lagunas costeras: Nota Científica. An. Inst. Cienc. del Mar y Limnol. Univ. Nal. Autón. México. 8 pp. Zapata F.A & R.Robertson. 2006. How many species of shore fishes are there in the Tropical Eastern Pacific?. Journal of Biogeography. 14 pp. 244 Baseline inventory of flora and fauna