Zootaxa,Biodiversity, Conservation, and Hotspot Atlas of Costa Rica
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Zootaxa 1457: 1–34 (2007) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2007 · Magnolia Press ISSN 1175-5334 (online edition) Biodiversity, conservation, and hotspot atlas of Costa Rica: a dung beetle perspective (Coleoptera: Scarabaeidae: Scarabaeinae) BERT KOHLMANN1*, ÁNGEL SOLÍS2, ORTWIN ELLE3, XINIA SOTO4, & RICARDO RUSSO1 1Universidad EARTH, A.P. 4442-1000, San José, Costa Rica. *Correspondent: [email protected] 2Instituto Nacional de Biodiversidad, A.P. 22-3100, Santo Domingo de Heredia, Costa Rica. 3Abteilung Biogeographie, Am Wissenschaftspark 25-27, Universität Trier, 54296 Trier, Germany. 4GIS Consultant, A.P. 580-2070, Sabanilla de Montes de Oca, Costa Rica Abstract This paper is an analysis of the distribution of areas of high species richness and endemicity based on dung beetles living in the different Holdridge life-zones of Costa Rica by using a geographic information system (GIS). Endemism was examined in relation to whether the species were shared with Nicaragua and/or Panama, or if they were strictly Costa Rican. The species composition of dung beetle distributions in the sampling areas and life-zones was evaluated. Species distribution was also analyzed in relation to altitudinal levels. The species richness and endemicity maps served as a base for doing a gap analysis and defining four different levels of high priority conservation areas. We also investigated what percentage of these priority areas is under some type of protection or conservation scheme and which of these areas should be enlarged. Also considered is the feasibility that these areas under protection have for enlargement, considering possible problems and interactions with present land-use. We include a list of all the recorded dung beetle species for Costa Rica, as well as their presence in the different Holdridge life-zones and their endemicity status. This study clearly demonstrates the need to include insects in biodiversity-endemicity studies because different and more detailed results are obtained in relation to vertebrate and plant-based studies. Key words: Biodiversity atlas, conservation, biodiversity areas, endemism areas, gap analysis, Costa Rica, dung beetles, Scarabaeinae Resumen En el presente trabajo se realizó un análisis de la distribución de áreas de alta riqueza específica y endemismo de escarabajos del estiércol en las diferentes zonas de vida de Holdridge en Costa Rica, utilizando un sistema de información geográfica (SIG). Los endemismos fueron particularmente analizados en relación a si se compartían con Nicaragua y/o Panamá, o si eran estrictamente costarricenses. Se evaluó la representatividad de las áreas de colecta y la cobertura de zonas de vida. Se analizó también la distribución de las especies con respecto a pisos altitudinales. Los mapas de riqueza específica y endemismo sirvieron de base para realizar un análisis de tipo Gap y definir cuatro zonas de conservación de alta prioridad. Se analiza también qué porcentaje de esas áreas prioritarias se encuentran actualmente bajo algún tipo de protección, y cuáles de estas zonas deberían aumentar su cobertura al respecto. Se analiza también la factibilidad de que se logren realizar estas extensiones de protección en relación a posibles problemas e interacciones con el presente sistema de uso de suelos. Se incluye una lista de todas las especies registradas de escarabajos del estiércol de Costa Rica, así como su presencia en las diferentes zonas de vida de Holdridge y su condición de endemismo o no. Este estudio demuestra claramente la necesidad de incluir a los insectos en estudios de biodiversidad y endemismo, se pueden obtener resultados distintos y con más detalle en relación a los estudios que se basan sólo en vertebrados y plantas. Accepted by Andrew Smith: 11 Apr. 2007; published: 25 Apr. 2007 1 Introduction The distribution of biodiversity is not an even occurrence across the surface of the Earth. On the contrary, numbers vary and species composition present highs and lows in a very complex pattern. Many attempts have been undertaken to try to understand, map, and identify general spatial patterns, especially species richness and endemism of well-known groups, such as vertebrates and plants. However, very little information is available for highly speciose groups, such as insects (Gaston and Spicer 2004). At the same time, global biodiversity is decreasing precipitously with innumerable species going extinct. No one knows for sure what the final effect of this crisis will be, but one thing is certain: first we must know how this biodiversity is spatially distributed. For this reason, Morrone and Espinosa (1998) have advocated the compilation of biogeographic atlases. A biogeographic atlas represents the synthesis of the distribution patterns of taxa for a country or a biogeographic area, represented by bioclimatic predictions, panbiogeo- graphic analysis, cladistic biogeography, and areas of species richness and endemism (Nix 1989, Espinosa and Llorente 1993, Morrone et al. 1996, Lobo et al. 1997, Morrone 2000). Biogeographic atlases furnish informa- tion relevant for the identification of species richness and endemic areas, which in turn can be used for estab- lishing conservation areas and developing sustainability policies. Because the present analysis concentrates primarily on the species richness and mapping endemism it should be considered foremostly as a biodiversity atlas. Costa Rica is not a big country (Fig. 1). It has only 51,100 km2 of land surface, representing only the 0.03% of the surface of the Earth (Ministerio del Ambiente y Energía 2000). In the ranking of world biodiver- sity, it occupies the 20th place, approximately. As such, it was not considered a megadiverse country by Mit- termeier and Goettsch Mittermeier (1997), who gave this status only to the top twelve countries. However, what makes Costa Rica special is its species density (number of species per unit of area) (Obando 2002). Using this measure, Costa Rica ranks first in the world (Valerio 1999, Obando 2002). It is considered that Costa Rica possesses around 4% of the total world biodiversity and if we consider the total number of described species this number jumps then to 5.4 % (Jiménez 1995). To give a comparative idea, Costa Rica has 234.9 plant species per 1,000 km2, whereas the second most species-dense country, Colombia, has 39.4 plant species per 1,000 km2 (Obando 2002). In the same line of thought, Costa Rica has 16.9 bird species per 1,000 km2, whereas Colombia has only 1.5 bird species per 1,000 km2 (Valerio 1999, Obando 2002). Costa Rica’s extreme biodiversity is now under protection by a world-class national system of protected areas, which was initiated in the 1970’s and today protects almost 27 percent of the country (Vaughan 1994, Vaughan et al. 1998). It is precisely this biodiversity that has been attracting ecotourists to the country, making this activity the primary source for foreign currency income; surpassing in the last ten years the three traditional major exports of coffee, bananas, and beef (Damon and Vaughan 1995). This is an important and swift change for the eco- nomic paradigm of a country, especially if we consider that coffee has been the primary economic engine of Costa Rica since the mid 19th century (Hall et al. 2000). Costa Rica is considered to have a moderate number of endemics (Obando 2002), with around 1.4% of the known species (plants and vertebrates) being endemic to the country. It is estimated that around 11% of the total plant species are endemics, whereas the different vertebrate groups vary from a minimum of 0.8% in birds to a maximum of 20% for the amphibians (Obando 2002). Using plants and vertebrates, four great areas of endemism have been identified for continental Costa Rica (Fig. 15): the Central Volcanic Cordillera, the Talamanca Cordillera, the Central Pacific Region, and the Golfo Dulce Region. A fifth area has been identi- fied in Coco Island, in the Pacific Ocean. From the ecosystem point of view, cloud forests are the most endemic ecosystems (Obando 2002). This study is a first attempt to define a biodiversity atlas indicating the areas of high species richness and endemism for Costa Rica using an insect taxon (Scarabaeinae), instead of plants and vertebrates as has been 2 · Zootaxa 1457 © 2007 Magnolia Press KOHLMANN ET AL. done in the past. The analysis focuses on continental Costa Rica; Coco Island was not included because no Scarabaeinae are known from this island. A similar analysis was done by Lumaret (1978, 1990) and Lumaret and Lobo (1996) for France and the Western Palaearctic region respectively. This atlas represents a first step towards helping to define those areas most in need of conservation and sustainable use in Costa Rica using insects, as well as identifying those areas that have been undersampled in order to target future collecting efforts to these poorly-known areas. Similar analyses using plants (Araceae, Arecaceae, and Bromeliacae), fresh-water fishes, and beetles (Scarabaeidae: Dynastinae) are almost finished and will be published soon as a sequel and complement to this analysis. FIGURE 1. Map of Costa Rica showing major geographical landmarks and cities, roads, types of vegetation, and pro- tected areas (national parks, national wildlife refuges, and biological reserves; numbered 1–48) (taken from Kohlmann et al. 2002). Materials and Methods Taxon information Information regarding scarab beetle distribution was taken from the collections and electronic database