Journal of Biogeography (J. Biogeogr.
) (2004) 31, 1829–1839
Biological diversiﬁcation in a complex region: a spatial analysis of faunistic diversity and biogeography of the Andes of Colombia
´n Gustavo H. Kattan1*, Padu Franco1,2, Vladimir Rojas1,2 and Germa Morales2
´n EcoAndina/Colombia Program of Fundacio the Wildlife Conservation Society, Cali, and 2 ´a, Universidad del Departamento de Biologı Valle, Cali, Colombia
Aim Understanding large-scale patterns of beta diversity and endemism is essential for ecoregional conservation planning. We present a study of spatial patterns of faunal diversiﬁcation and biogeographical relationships in the Andean region of Colombia. This region has a great geomorphological complexity, as it is formed by several mountain ranges with different geologic origins. We hypothesize that this complexity results in a high turnover in species composition among subregions. Location The Andean region of Colombia, including the Santa Marta and Macarena mountain ranges. Methods The region was divided into subregions, represented by the eastern and western slopes of each of the three Andean cordilleras, the Cauca and Magdalena ´ , Macarena and valley bottoms, and the peripheral mountain ranges of Perija Sierra Nevada de Santa Marta. Species lists for ﬁve animal taxa (rodents, bats, birds, frogs and butterﬂies) were compiled for each subregion and similarities in species composition were determined by cluster analysis. To explore biogeographical relationships, species were classiﬁed into one of four distributional categories: endemic, tropical Andean, Andean-Central American and wide continental distribution. Results The highest species richness in the region was found in the Paciﬁc and eastern versants of the Andes, and the lowest in the Cauca and Magdalena valley bottoms. Inter-Andean slopes were intermediate in species richness. However, when species richness was calculated per unit area, the most diverse regions were the Santa Marta and Macarena ranges, the Cauca Valley watershed and the Paciﬁc slope. Although each taxonomic group had a different branching pattern, ´ -Sierra dendrograms indicated ﬁve common subregional clusterings: (1) Perija Nevada, (2) the Paciﬁc slope, (3) the eastern Andean slope, (4) the Cauca and Magdalena valley bottoms, and (5) the inter-Andean slopes. Clustering patterns of inter-Andean slopes varied among taxa. In birds, bats and rodents, grouping was by opposite slopes of the same valley, whereas frogs were grouped by mountain ranges and butterﬂies by valleys and their respective slopes. Seventyﬁve per cent of species in all taxa were found in less than ﬁve subregions. The fauna of the Magdalena and Cauca valley bottoms was composed mostly of lowland species with wide geographical distributions, whereas the cordilleran fauna was mostly restricted to the tropical Andes. Main conclusions The western and eastern versants of the Andes have the highest species richness, but are also the largest subregions. On a per unit area basis, the peripheral ranges (Santa Marta and Macarena) are the richest, followed by the western portion of the Andes (the Cauca Valley watershed and the Paciﬁc versant). Clustering patterns in dendrograms suggest two major patterns of
*Correspondence: Gustavo Kattan, Apartado ´ reo 25527, Cali, Colombia. Ae E-mail: firstname.lastname@example.org
ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi
clasiﬁcamos las especies en objectivo de explorar las relaciones biogeogra ´as de distribucio ´ n geogra ´ ﬁca: ende ´ mica. Los patrones de ´ laagrupamiento de las vertientes interandinas fueron variables. en mariposas. ecoregional planning. en los dendrogramas se destacan cinco ramas: (1) Perija ´ﬁco.G. (4) Sierra Nevada. La regio ´a de La Macarena). y (5) las vertientes interandinas. murcie ´ lagos. En cada grupo. particularly among the less vagile taxa. protected area planning. biotic diversiﬁcation. Sin embargo. relaciones biogeogra ´ n es de una gran complejidad geomorfolo ´ gica. and the third one composed by the inter-Andean slopes of the Cauca and Magdalena valleys. las vertientes opuestas de cada valle resultaron muy cercanas en ´ n de especies. 1829–1839. Colombia. Con el ´ ﬁcas. por unidad de a La Macarena y Santa Marta. y la vertiente del ´ﬁco. Three different main branches of Andean fauna can be recognized. En aves.
radiation. (2) la vertiente Pacı las planicies de los dos valles. las planicies ´as de Perija ´ y La Macarena y la de los valles del Cauca y Magdalena.
El conocimiento de los patrones de diversidad beta y endemismo a gran escala ´culo se presenta ´ n ecorregional. determinamos la similitud en composicio ´ especies por medio de analisis de agrupamiento (cluster analysis). en cambio. (3) la vertiente oriental de los Andes. Aunque el ana ´ lisis arrojo ´ diferentes patrones de agrupamiento para cada Pacı ´ ´grupo taxonomico. lista de especies para cinco grupos taxono ´ n de ranas y mariposas. murcie gos y roedores. Andes tropicales. ª 2004 Blackwell Publishing Ltd
. and the evolutionary processes that gave rise to these patterns. Biogeographical afﬁnities of the inter-Andean valley bottoms are with the lowland faunas of tropical America. Nuestra hipo ´por varias cordilleras y serranı ´ n de especies entre las tesis es que esta complejidad genera una alta composicio ´ n andina de Colombia (incluyendo la Sierra Nevada de Santa subregiones. one conﬁned to the Paciﬁc. en las ranas se agruparon primero las dos composicio ´ con sus respecvertientes de cada cordillera.
differentiation of the Andean fauna: one elevational (lowlands vs. y la ma ´ en las vertientes Pacı ´ s baja en los valles ntro del Cauca y Magdalena. La fauna de las planicies del Cauca y Magdalena esta ´ ﬁcas. andinacuatro categorı ´ n amplia. Las vertientes interandinas presentaron una riqueza ´ rea las subregiones ma ´ s diversas fueron intermedia. highlands) and one horizontal (among ranges and/or slopes). Conservation initiatives that seek to preserve representative samples of the regional biodiversity should take into account the patterns of diversiﬁcation described here. resulting in the evolution of a large number of endemic species. Para cada una de estas subregiones compilamos la ´ micos: roedores. La riqueza de especies ma ´ s alta se encocentroamericana y distribucio ´ﬁca y oriental de los Andes. cada valle se agrupo ´ compuesta tivas vertientes. Andean faunas diversiﬁed locally. In contrast. biodiversity. el valle del Cauca y sus vertientes. fue dividida en subregiones. The identiﬁcation of ﬁve main biogeographical units in the Andean region of Colombia has important implications for the conservation of the regional biota. constituidas por Marta y la serranı las vertientes oriental y occidental de cada una de las tres cordilleras. Kattan et al. speciation. ya que esta ´ conformada Esta regio ´as de diferentes orı ´genes geolo ´ gicos. principalmente por especies de tierras bajas y amplias distribuciones geogra
Journal of Biogeography 31. H. las serranı Sierra Nevada de Santa Marta. another to the eastern (Amazonian-Llanos) versant of the Andes. Keywords Andes. En este artı espacial es esencial para la planiﬁcacio ´ n de la fauna y de sus un estudio de los patrones espaciales de diversiﬁcacio ´as perife ´ ﬁcas en la regio ´ n andina de Colombia y serranı ´ ricas. aves.
1200 species of birds. 1). and natural ecosystems have been extensively ´ pez. Seventy per cent of the human population of Colombia is concentrated in the Andean region.869. Diversities of frogs and birds in the northern Andes (427 and >1400 species. planiﬁcacio
INTRODUCTION Rising at the crossroads between two continents. biodiversidad bio ´ n ecoregional. 400 species of frogs. mientras que las faunas andinas muestran una diversiﬁcacio ´ n de un gran nu ´ mero de especies ende ´ micas.000 km2. planiﬁcacio ´ n de a ´ reas protegidas.. 1996. lo americano. the connection with North America fostered a biotic interchange that enriched the autochthonous South American biota with new taxa. Considering that the area of (Fjeldsa the northern Andes (490. Duellman. the Colombian Andes harbour more than 10. 1000 species) ˚ . birds and amphibians.b). 1999). As a result of these events. one centred in the Amazon basin and another centred in the northern Andes ˚ . 1994. Central and Eastern Cordilleras). Las iniciativas de tantes implicaciones para la conservacio ´ n que buscan preservar muestras representativas de la diversidad conservacio ´ lo los patrones de diversidad descritos en este regional deben tener en cuenta no so ´culo. Magnoliaceae. two major foci of diversiﬁcation on a continental scale have been identiﬁed. with subsequent speciation and radiation events. The northern Andes reach their maximum geomorphological complexity in Colombia. cual ha resultado en la evolucio ´ viles. separated by the sedimentary basins of the Cauca and Magdalena river valleys (Fig. the northern Andes have played a key role in the diversiﬁcation of the tropical South American biota. For organisms such as vascular plants. respectively) easily surpass Amazonian diversities (305 and c. en cambio. Encompassing 11 degrees of latitude and a little over 300. La identiﬁcacio ´ ﬁcas en la regio ´ n andina de Colombia tiene imporgrandes unidades biogeogra ´ n de la biota regional.. 1994. 1993). the uplift of mountain ranges in a complex series of orogenic processes. this region is also under intense pressure. the climatic ﬂuctuations during the Pleistocene likely produced cycles of range contraction and expansion that resulted in fragmentation and isolation of populations.Biological diversiﬁcation in the Colombian Andes
´ compuesta principalmente por La fauna de las cordilleras. First. ramas en la fauna propiamente andina: una conﬁnada a la vertiente del Pacı otra a la vertiente oriental de los Andes. Gentry.000 species of plants. Duellman. 1992.000 km2. and created an incredibly diverse array of new environments that provided fertile grounds for biotic radiation and diversiﬁcation. originating almost half of all Neotropical plant species (Gentry.
particularly in the upper elevations. biodiversidad. where they are divided into three main ranges (the Western. Colombia. Several peripheral mountain ranges of different geological origins add to the physical complexity and biological diversity of the region. Andes. 1997a. y una tercera compuesta por las vertientes ´ n de cinco interandinas de los valles del Cauca y Magdalena. and 270 species of rodents and bats.000 km2). Los patrones de agrupamiento en los ´ n de la fauna dendrogramas sugieren dos patrones principales de diferenciacio ´ n vertical (tierras bajas versus tierras altas) y otro horizontal andina: un patro (entre cordilleras y entre vertientes). 1999). from the Huancabamba Depression in northern Peru to the Venezuelan Andes) is 14 times smaller than the area of the Amazon basin (6. 1982. Unfortunately. Las planicies interandinas del Cauca y ´ ﬁca con la fauna de tierras bajas del tro ´ pico Magdalena tienen aﬁnidad biogeogra ´ n local. it is evident that the region is a hotspot that requires high priority in global conservation initiatives (Myers et al. but levels of endemism also are extraordinary (Adams. 1999). 1982).e. Thirdly. Marshall & Sempere. 1986. representing up to 10% or more of the world’s biota in some taxa (Rangel. i. Uplift of the (Gentry.g. 1997). sino tambie ´ n los procesos evolutivos que originaron estos patrones. ª 2004 Blackwell Publishing Ltd
. caused innumerable vicariance events. radiacio ´ n bio ´ tica. artı Palabras clave ´ tica. Duellman. 1829–1839. esta especies restringidas a los Andes tropicales. Secondly. 1982. many of these taxa just reached the northern Andes and diversiﬁed there (e. especiacio ´ n. Fjeldsa Andes led to an explosive radiation of plant families of Gondwanan origin. 1997. The biological diversity of the northern Andes is the result of three major historical events (Gentry. Cavelier.. Lynch et al. Bibby et al. Se pueden reconocer tres grandes particularmente en los taxones menos mo ´ﬁco. spatial 1831
Journal of Biogeography 31. 2000). 1982). not only is regional diversity very high. transformed (Kattan & Alvarez-Lo The design and implementation of conservation schemes aimed at preserving a representative sample of regional biodiversity require understanding the spatial patterns of diversiﬁcation and large-scale beta diversity.
then a small number of areas will sufﬁce to ensure complete representation. faces the Amazonian lowlands in its southern half. due to interception of humid coastal winds. The eastern slope of the Eastern range. The interior-facing (inter-Andean) slopes of the Andes form the slopes of the Cauca Valley. showing the three branches of the Andes and other subregions used for the spatial analysis of diversity (only the western ´ is or Colombian portion of Serrania de Perija shown. particularly in the foothills. between the Western and Central Cordilleras. and two deep inter-Andean valleys (Fig. the eastern half of this range lies in Venezuela). then more strategically located areas will be required to cover the regional variation. the three-pronged conﬁguration and south– north axes of the mountain chains give origin to three westfacing and three east-facing slopes. But if the biota is spatially heterogeneous. which is continuous into northern Ecuador and is extremely humid. STUDY AREA At their southern end. as it intercepts the north-eastern trade winds. between the Central and Eastern Cordilleras. In addition to the subregions indicated. each branch of the Andes was divided into a western and an eastern slope. North 1832
of this point. 1829–1839. Kattan et al. The eastern Andean slope is also generally humid. on the contrary. the Colombian Andes form a single high-rising massif (the Macizo Colombiano). which is the continuation of the main axis of the Ecuadorian Andes. frogs and butterﬂies) of the Colombian Andes. Differences in geology. and Me ´a de Perija ´ . The valley bottoms tend to be dry. bats. The west-facing slope of the Western range forms the Paciﬁc versant. and the Magdalena Valley. but the range summits are humid. the Eastern range bifurcates into the ´ rida Andes. Rain shadow phenomena create subxerophytic (semi-arid) pockets.
Figure 1 Map of the Andean region of Colombia. At its northern end. Here we present results of a study of the patterns of spatial diversiﬁcation and geographical distribution of some groups of animals (rodents. so the lowlands are restricted to a narrow coastal corridor. topography and climate suggest that the history and composition of the biota might differ among ranges or even between slopes of the same range. 1). Our primary objective was to determine the degree of similarity in species composition and patterns of geographical distribution among different subregions composing the Colombian Andes.G. which extends northward the low-rising Serranı
Journal of Biogeography 31. ª 2004 Blackwell Publishing Ltd
turnover in species composition. birds. The Andes here rise abruptly from the Paciﬁc coast. If the regional biota is relatively homogeneous in its history and species composition. H. which extends north-east into Venezuela. and the llanos or savannas of Colombia and Venezuela in its northern half. Humidity patterns in the Cauca and Magdalena valleys are more variable. This information could then be used in making decisions about the areas required to obtain adequate representation of the regional biodiversity in a protected area system.
the eastern versant of the Andes and the Magdalena Valley watershed (including the slopes and valley bottom) were poor when contrasted with the Cauca Valley watershed. Lycaenidae and Riodinidae. Initial orogeny of the Santa Marta mountains can also be traced to the late Paleozoic. 1975). Also included in our analyses are two separate mountain ranges. we assigned each species to a geographical distribution category. we constructed dendrograms using cluster analyses of presence–absence matrices with the SPSS statistical package. To determine present similarities in species composition among subregions. This index equals 1 in cases of complete similarity. species that are widely distributed in tropical America or in the entire continent. ª 2004 Blackwell Publishing Ltd
. and even surpassed the eastern versant in frog richness (Table 1). Perija Marta (J ranged from 0.86) than with the rest of the 1833
Journal of Biogeography 31. Irving. Serranı Marta in a separate branch in birds. Alberico et al. we divided the Andean region of Colombia into subregions. and c the number of shared species. Per unit area. (1992). deﬁned as follows: Endemic (E). two inter-Andean valley bottoms. Species lists were compiled for each of the subregions from published national lists (Hilty & Brown. This subdivision follows. known ranges are restricted to one or a few localities). the classiﬁcation of bioge´ ndez ographical provinces of Colombia proposed by Herna Camacho et al. We used Jaccard’s coefﬁcient of similarity (J) as a distance measure. and wide distribution (WD). Pieridae (genus Catasticta).. which emerged as a relatively rich region. A species was assigned as present in a subregion if it had records in at least one locality. Although the Paciﬁc versant was still very rich. This may overestimate the geographical distribution of some species within the subregion. 1996. Species richness in the ﬁve taxonomic groups correlated with area of the subregions (Fig. and 0 if the sites have no species in common. 2). the Santa Marta massif (Sierra Nevada de ´a de Santa Marta) on the Caribbean coast. North Andean orogenesis started in the late Paleozoic. 1829–1839. where a is the number of species in site A. species that are distributed in the tropical Andes. particularly in frogs. the humid western (Paciﬁc) and eastern (Amazonian-Llanos) versants of the Andes were the richest both in total number of species and number of endemics for the ﬁve taxonomic groups. 1986. Major criteria for selecting study taxa were having a reasonable knowledge of species distributions and that lists could be obtained. species with geographical ranges restricted to <50. Dendrograms showing the clustering of subregions for the different taxa are shown in Fig. Nymphalidae [subfamilies Charaxinae. b the number of species in site B. bats and butterﬂies ´ were not available for frogs). as this valley’s bottom lies at 1000 m elevation. The Paciﬁc versant was further subdivided into an upper Paciﬁc slope (>1000 m elevation) and a lower Paciﬁc slope (200–1000 m elev. coastal biota from the analysis. Brassolinae and Satyrinae (tribe Pronophilini)]. Andean-Central American (CA). This subdivision was made to allow comparisons with the Cauca Valley slopes. The Central Cordillera was emergent but low during the Cretaceous. La from different orogenic events (Bu Macarena is part of the Guianan Shield and is the oldest range. have different geological histories and resulted ¨ rgl.Biological diversiﬁcation in the Colombian Andes (Fig. and the Serranı Santa Marta and La Macarena (Fig. butterﬂy lists were not available and were compiled de novo by three experts. but vertical uplifting to present altitudes occurred in late Tertiary time. To explore biogeographical relationships. Heliconiinae. 2000). deﬁned as the eastern and western slopes of each of the three ranges. with some modiﬁcations. Jaccard’s coefﬁcient is deﬁned as J ¼ c/(a+b)c). Orogeny of the Western Cordillera can be traced to the end of the Mesozoic. Conversely. the richest regions were the peripheral mountain ranges of Macarena and Santa Marta. although the Central Cordillera also was very rich. the lower limit on the eastern versant of the Andes was set at 500 m of elevation to exclude the lowland Amazonian biota (the Magdalena Valley lies at an average of 500 m in its mid portion.000 km2 (in many cases. RESULTS In general. The inter-Andean slopes of the Cauca and Magdalena valleys were intermediate (Table 1). Present elevations were reached during the Pliocene–Pleistocene. when the ancestral Central Cordillera was insinuated at the western margin of a miogeosyncline. Ithomiinae. Ruiz et al. 1961. and of the Eastern Cordillera to the middle Tertiary. but as we are concerned with largescale patterns of distribution.. the following common patterns of differentiation could be identiﬁed: ´a de Perija ´ grouped with Sierra Nevada de Santa 1. and descends rapidly northward). 1). a different picture emerged when species richness was tallied per unit area (Table 2). 1). Morphinae. Andean folding during the Miocene gave rise to the present conﬁguration of three Cordilleras and the two sedimentary basins of the Cauca and Magdalena valleys. The three ranges of the Colombian Andes. The lower limit of 200 m was chosen to exclude the lowland. Likewise.39 to 0. 3. Although each taxonomic group had a particular pattern of clustering. and centroid clustering. complemented by literature surveys to include new species or records that have appeared after the monograph’s publication. the ´as of Perija ´.). METHODS To determine patterns of biotic differentiation. In (separate data on Perija ´ shared a higher proportion of species with Santa general. the lowest numbers were found in the dry inter-Andean valley bottoms. In butterﬂies we included the families Papilionidae. Tropical Andean (TA). as well as the ´a de La Sierra Nevada de Santa Marta and the Serranı Macarena. and the small Serranı La Macarena east of the Andes. this does not affect our analyses. However. species that are distributed in the tropical Andes and Central American mountains.
Serranı clustered with the eastern slope of the Andes in all groups except rodents (clustering of Macarena with Santa Marta in rodents may reﬂect poor knowledge). Thus. very few species occurred in more than 10 subregions. and (5) the inter-Andean slopes (although we treat this subregion as a unit.000 60.000 80. which is extremely rich in this taxon. regressions on untransformed axes produced higher R2 values than log–log regressions. We also examined the patterns of distribution of species in each of the subregions. The proportion of species shared between the Paciﬁc and Eastern versants was intermediate in vagile groups such as birds (J ¼ 0. the Paciﬁc fauna was predominately composed of species with CA or WDs. whereas species of TA distribution were
Frogs (R = 0. there were some important differences among taxa in the clustering patterns of the different slopes.61)
Bats (R = 0. but it was as low as J ¼ 0.000 80. to explore biogeographical relationships (Fig. grouping was by opposite slopes of the same valley. Chiroptera was the only taxon with a dominance of widely distributed species. widely distributed species were also well represented in the valleys and the Eastern slope. 3. Seventy-ﬁve per cent of the species in the ﬁve taxonomic groups occurred in ﬁve or less subregions. However.
Journal of Biogeography 31.36) and bats
(J ¼ 0.13)
Rodents (R = 0. not available. Kattan et al.65) except butterﬂies (J ¼ 0.22)
20. and 36% (i. In contrast.G. ª 2004 Blackwell Publishing Ltd
Number of species
Birds (R = 0.000 40.000 80. whereas frogs grouped by mountain ranges. 1829–1839.000 40.000
Butterflies (R = 0. In all cases.36–0. In birds. The extreme outlier in the frogs plot is the Paciﬁc slope of the Andes. In the case of rodents. for which Perija Cordillera. ﬁve different faunistic subregions can be recognized: ´ -Santa Marta.
Table 1 Number of species (endemics in parenthesis) of ﬁve animal taxa in different subregions of the Colombian Andes
Subregion Butterﬂies (61) (72) (105) (55) (53) (25) (15) (29) (48) (62) (90) (73) (76) (266) Frogs 163 124 233 23 48 23 NA 12 38 82 80 52 69 464 (110) (62) (142) (10) (20) (16) Birds Rodents Bats (26) 40 (18) 26 (30) 46 (5) 18 (4) 27 (26) 21 (9) 43 (0) 20 (14) 32 (15) 42 (16) 46 (9) 53 (12) 72 (73) 107 (3) (1) (4) (1) (4) (2) (3) (0) (3) (3) (3) (4) (4) (11) 65 87 101 46 58 94 103 84 60 62 72 77 126 166
Upper Paciﬁc 210 Lower Paciﬁc 364 Paciﬁc (all) 490 Cauca Valley 212 Magdalena Valley 173 Santa Marta 161 ´ Perija 124 Macarena 138 Western east* 273 Central west* 266 Central east* 385 Eastern west* 241 Eastern east* 658 Total 1334
740 590 850 298 345 637 429 (3) 561 (22) 484 (53) 434 (45) 523 (29) 499 (32) 791 (283) 1426
NA. 2. The Cauca and Magdalena valley bottoms clustered together and were relatively similar for all groups (J ¼ 0.000
60. suggesting a shared foothills butterﬂy ´a de La Macarena fauna on both ﬂanks of the Andes.000
Figure 2 Species–area relationships for ﬁve taxonomic groups in different subregions of the Andean region of Colombia. H. and butterﬂies grouped by valley bottoms and their respective slopes (Fig. (4) the Cauca and Magdalena valley bottoms.
Eastern Cordillera (J ¼ 0.000 20. Differences in composition among subregions can be emphasized by examining how many subregions each species occurs in a measure of endemicity (Fig. it shows considerable internal heterogeneity).e. (3) the Eastern (1) Perija versant. 1225 of 3434 species in the ﬁve groups) occurred in only one subregion. (2) the Paciﬁc versant. The western (Paciﬁc) and eastern (Amazonian-Llanos) versants of the Andes branched as separate entities and were divergent in species composition. frogs and bats. 4). the only exception was butterﬂies. The upper Paciﬁc slope formed one unit with the lower Paciﬁc in birds.04 in sedentary organisms such as frogs. 5). The only exception was ´ grouped with the rest of the Eastern rodents. bats and rodents.10–0.27).62).000 60. but it was more similar to the inter-Andean slopes in butterﬂies and rodents. in which the Lower Paciﬁc region clustered with the eastern Andean slope. 3). The inter-Andean slopes of the Cauca and Magdalena watersheds tended to group together in all taxa. *Refers to the range and slope.000 80.50).73)
21 0. but TA species represent an important proportion in the Paciﬁc.38 4.42 0. in contrast to bats.91 0. Eastern and inter-Andean slopes.01 0. not available.21
better represented in the Andean and Eastern slopes.10 0.22 0.Biological diversiﬁcation in the Colombian Andes
Table 2 Area (in km2) and number of species per unit area (·10)2) for ﬁve animal taxa in different subregions of the Colombian Andes
Subregion Paciﬁc (all) Cauca Valley Magdalena Valley Santa Marta ´ Perija Macarena Western east* Central west* Central east* Eastern west* Eastern east* Area 36771 6780 19375 5998 5693 2888 14388 18995 51956 54232 60989 Butterﬂies Frogs Birds Rodents Bats 2.36 2.40 0. Butterﬂies also had a large proportion of species with wide geographical distributions in all subregions. there is a strong representation of CA and widely distributed species in all subregions. 1829–1839.35 0.15 0.18 4.52 0.57 1.30 0.25 0. and relatively few have Central American relations. our results agree with this assessment.39 1. In absolute numbers. In birds.22 0.10 0.26 0.69 0.44 1.78 1.41 0.62 7.90 1. Frogs were noteworthy for the dominance of the endemic species category in all regions.74 0. DISCUSSION The Eastern and Paciﬁc (the Chocoan biogeographical region) versants of the Andes have usually been recognized as biodiversity hotspots (Rangel.11 4. *Refers to the range and slope.78 10.09 0.81 2. but one-third to one half or more of the species were endemic or restricted to the tropical Andes.42 3.
Journal of Biogeography 31. ª 2004 Blackwell Publishing Ltd
Figure 3 Dendrograms showing similarity in species composition among subregions composing the Colombian Andes.34 0.76 0.68 0.14 0.14 0. which in general had very wide geographical distributions.43 0.13 0.53 3.54 19. However. on a per unit
NA.24 0.68 2.28 1.33 0.b).14 0.92 1.30 1.27 0.38 NA 0.12 0.90 2. 1997a.08 1.
˚ . and local radiation and subsequent dispersal of taxa. 1994. the pattern is expected from the species–area relationship. At a continental scale. Andean-Central American distribution. evidenced by the differentiation of lowland faunas in the valley bottoms and highland faunas in the Andean slopes. which tend to have wide altitudinal distributions). 1829–1839.. 1997). 1997. 1987. one major effect of the emergence of the Andes was the differentiation of CisAndean (Amazonian).
Number of subregions
Figure 4 Patterns of distribution of species in ﬁve taxonomic groups. The only exception to this pattern are butterﬂies. 1998). the TA region starting in the upper Miocene (Fjeldsa resulting in distinct avifaunas in the highlands and lowlands (Renjifo et al. wide distribution (South and Central America). tropical Andean distribution. here. with a high proportion of species of 1836
wide geographical distributions. H. whereas the highlands have autochthonous Andean biotas. 1999). endemic.
300 200 100 0 1 800 600 400 200 2–5 6–9 10–13
80 60 40 20 0 Pacific 100 80 60 40 20 0
Number of species
0 1 800 600 400 200 0 1 80 60 40 20 0 1 60 2–5 6–9 10–13 2–5 6–9 10–13 2–5 6–9 10–13
Percent of species
Pacific 80 60 40 20 0 Pacific 80 60 40 20 0 Pacific 80 60 40 20 0 Pacific
CV Slopes MV Slopes
40 20 0 1 2–5 6–9 10–13
Figure 5 Patterns of geographical distribution of species in different subregional clusters. The second one is horizontal. Andean and TransAndean (Chocoan-Central American) biotas (Lynch. In addition. the fauna and ﬂora radiated to ﬁll the new environments that were generated. but the pattern holds true for relatively well-known groups such as birds and frogs. The Cauca and Magdalena valley bottoms have typically lowland biotas. One caveat is that this may reﬂect poor knowledge of some regions (e.. Moreover. Duellman. 1982. Our results indicate that even within the Andean region. regions with a high species density (number of species per unit area) are more vulnerable. ª 2004 Blackwell Publishing Ltd
. Clustering patterns of subregions suggest two major patterns of differentiation of the Andean fauna. which are more restricted in their distribution (Renjifo et al. Kattan et al. 1994). Categories of geographical distribution are: EN. Gentry. Brown. 3. The graph shows the number of species occurring in different numbers of subregions. an important result is that by this measure the Cauca Valley watershed (including the valley’s slopes and bottom) is an extremely rich region. WD.
Journal of Biogeography 31. our results indicate that each of the ﬁve subregions identiﬁed here tend to have distinct faunas. Tyler et al. represented by the divergence of faunas among ranges or even among slopes of the same range. The diversiﬁcation of the Andean biota is likely a combination of vicariance events. In any case. CA. each valley and its corresponding slopes form a different unit (probably related to the numerical dominance of Nymphalid butterﬂies. The ﬁrst one is elevational. because many unique species could be extirpated by destroying a relatively small area. the eastern slope of the Andes). As the Andes emerged.. as deﬁned by dendrograms in Fig. Fjeldsa 1994. From a conservation perspective. there is a differentiation between Paciﬁc. Stotz. The great diversiﬁcation of the northern Andes is a result of their topographic and climatic complexity. TA. the mountain ranges of Santa Marta and Macarena turn out to be the real hotspots. in subregions composing the Colombian Andes.g.
area basis. inter-Andean and Eastern versant faunas. The Neotropical bird fauna underwent an explosive radiation in ˚ .G. 1979.
and species turnover at different localities along altitudinal ranges in the Central Andes is 30–50% (G. rodent diversity in the tropical Andes is the result of a complex history of multiple waves of invasion and radiation. the contribution of these invaders is the north. 82 (46%) were endemic to the Andes. From an oryzomine ancestor. The Andean frog fauna is dominated by the genus Eleutherodactylus. Conservation efforts should take into account these regional patterns of diversiﬁcation. Stotz. The ﬁrst wave of sigmodontine invasion from North America likely occurred as far back as the Miocene. sometime in the Eocene. 1982.. Marshall & Sempere. 1837
Journal of Biogeography 31. Patterson et al. The diversity of South American rodents. Similar patterns of differentiation have been described for some groups of butterﬂies. i. for supporting this work in various ways. Distinct elevational belts have been identiﬁed for the Andean biota. and species replacement along elevational gradients is a welldocumented phenomenon in many taxa (Terborgh. and radiated in the lowlands east of the Andes. 1971. Marı ´a Dolores Heredia. 22 to the Central Cordillera. Species turnover in frog communities may be 60% or more (Restrepo & Alberico. REFERENCES Adams.J. porcupines and allies) are hypothesized to have arrived either from Africa or North America. an akodont stock was derived and radiated in the southern Andes. Pronophiline butterﬂies. 1966. elevational ranges in a great proportion of Andean birds are 1500 m or less. Although this contribution is not considerable in birds (only 5% of the South American avifauna is composed of groups that invaded from ˚ . Of the 176 eleutherodactylines known from Colombia in 1997.. the Andes have played a key role in the radiation of the diverse sigmodontine rodents (> 250 species of Neotropical mice and rats). Zoological Journal of the Linnean Society. Probably in no other group is the effect of the Andean uplifting more dramatic as in frogs. In addition to the radiation of autochthonous South American groups.. which occurred in a series of waves during the last few million years.Biological diversiﬁcation in the Colombian Andes and the high proportion of endemisms is associated with ˚. 2002). and north into Central America. data). One example serves to illustrate this.. a group of Satyrid butterﬂies with their highest diversity in the cloud forests of the northern Andes. as part of the development of a biodiversity vision for the Northern Andes Ecoregional Complex. Although Fjeldsa alpha diversity may be low at a given Andean locality in comparison with Amazonian localities. in order to ensure representation not only of the present biological diversity. 1989. Stotz. Much of this high species turnover is due to altitudinal gradients. Some species have invaded the Andes. Eisenberg. and topographic complexity has been an important factor in this diversiﬁcation (Marshall & Sempere. 1998. We also thank several anonymous reviewers for making suggestions and pointing out literature that greatly improved this manuscript. The nonsynchronous uplifting of the three Colombian Cordilleras fragmented populations and promoted allopatric speciation (Lynch et al. 1994). 1994). but also of the evolutionary processes that gave rise to this diversity. Patterson et al. MacArthur Foundation. the spatial turnover in species composition that may occur over short distances (Gentry. show a double pattern of speciation. 1993. 27 were endemic to the Western Cordillera.. Thus. ˚ . Forty-eight of 125 species in the Colombian Andes show allopatric speciation. ª 2004 Blackwell Publishing Ltd
. whereas 61 species have parapatric distribution of close relatives at different elevational belts on the same slope (Adams. ACKNOWLEDGMENTS This work was done as part of a biodiversity analysis for the northern Andes. Hystricognath rodents (agoutis. Lucı ´a Isabel Herrera. 87. 1999). as species turnover is rapid across elevational belts. sponsored by the World Wildlife Fund. Medina et al. unpubl. which in turn dispersed back into the northern Andes (Hershkovitz. 235–320. 5). M. 1997). 1986). But at larger scales. 1998). However. 1994. 1829–1839. is the result of radiation in two different groups. but also in high levels of endemism (Fig. vicariance in regions of high recent diversiﬁcation (Fjeldsa 1994). For instance. Kattan & P. landscape diversities in the Andes may even surpass Amazonian diversities. where they presently have their highest diversity (Wood. in particular Mary Lou Higgins and Olga ´a Herna ´ ndez. with close allies at similar altitudes in different ranges. Montane faunas have further diversiﬁed by altitude. and Catherine T. since this paper. the most speciose vertebrate genus. 1986. The patterns of diversiﬁcation and regional differentiation described here have important implications for conserving the extraordinary Andean biodiversity. Luis We thank Marı ´ n Salazar for compiling lists of Miguel Constantino and Julia frogs and butterﬂies. widely divergent in their species composition and afﬁnities and in their evolutionary histories. nine were shared by these two ranges. Marshall & Sempere. Franco. and for contributing unpublished information. which then dispersed south through the Andes. This horizontal diversiﬁcation resulted not only in high diversity. For example. 1993. Our work showed the existence of at least ﬁve different faunistic subregions. differences in species composition among mountain ranges or even between slopes of the same range result in high regional diversities. Fjeldsa important in other taxonomic groups. Reig. 1997. Subsequent uplifting of the Andes during the Miocene orogenic phase set the stage for the radiation of an oryzomine stock in the northern Andes. 1998. 1985. but few are exclusively Andean. WCS’s long-term work in the Colombian Andes has been mostly supported by the John D. (1986) Pronophiline butterﬂies (Satyridae) of the three Andean Cordilleras of Colombia. Patterson. the Andean biota was enriched by the immigration of taxa from North America. Reig. Patterson. 1986. 1993). 1998. 1999).e. We thank the WWF Colombia ofﬁce. The richness of the Andean biota is due to beta diversity. many species have been added to the Colombian frog fauna). and 24 were endemic to the Eastern Cordillera (Lynch et al. for example.
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Webb). 1829–1839. and his Master’s thesis was on the biogeography of birds associated to wetlands. His interests focus on the conservation of Andean ecosystems.D. ´ n Morales recently received his MSc from Universidad del Valle. Plenum Press. H. Stehli & S. Wood. 52. and he conducts research on macroecology. His Senior thesis focused on the biogeography of Colombian mammals. Gainesville. Brown. (1985) Northern waif primates and rodents.S. The great American biotic interchange (ed. Vladimir Rojas recently received his MSc from Universidad del Valle. FL. A. His interests include the biogeography and conservation of Germa Andean fauna. (1994) Swallowtail butterﬂies of the Americas. by F. Kattan is the coordinator of the Colombia Program of the Wildlife Conservation Society. Padu Franco is a recent graduate from the biology program at Universidad del Valle.. 267–282.E. pp. His research interests include macroecology and behavioural ecology.G.Biological diversiﬁcation in the Colombian Andes ´. community ecology and population biology of endangered species. Tyler. Peru Ecology. Scientiﬁc Publishers. ª 2004 Blackwell Publishing Ltd
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BIOSKETCHES Dr Gustavo H. NY. & Wilson. patterns in the avifauna of the Cordillera Vilcabamba.
Journal of Biogeography 31.