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Biodivers Conserv DOI 10.

1007/s10531-012-0259-5 ORIGINAL PAPER

Cross-taxon congruence of a and b diversity among ve leaf litter arthropod groups in Colombia
Jimmy Cabra-Garca Christian Bermudez-Rivas Ana Milena Osorio Patricia Chacon

Received: 13 January 2011 / Accepted: 17 February 2012 Springer Science+Business Media B.V. 2012

Abstract In this study a and b diversity patterns of ve leaf litter arthropod groups (ants, predatory ants, oribatid mites, spiders and other arachnids) were described and compared in 39 sampling patches of a transformed landscape in southwestern Colombia, that represented ve vegetation types: secondary forest, riparian forest, giant bamboo forest, pasture and sugarcane crop. It was also assessed whether some taxa could be used as diversity surrogates. A total of 6,765 individuals grouped in 290 morphospecies were collected. Species richness in all groups was lower in highly transformed vegetation types (pasture, sugarcane crop) than in native ones (forests). In contrast, there were no clear tendencies of b diversity among vegetation types. Considering sampling patches, 0.142% of the variation in a diversity of one taxonomic group could be explained from the a diversity of another, and 0.233% of the variation of b diversity of a given taxon was explained by that in other groups. Contrary to recent ndings, we concluded that patterns of a diversity are more congruent than patterns of b diversity. This fact could be attributed to a sampling effect that promotes congruence in a diversity and to a lack of a clear regional ecological gradient that could promote congruent patterns of b diversity. We did
Electronic supplementary material The online version of this article (doi:10.1007/s10531-012-0259-5) contains supplementary material, which is available to authorized users. J. Cabra-Garca (&) C. Bermudez-Rivas A. M. Osorio P. Chacon Grupo de investigacion en Biologa, Ecologa y Manejo de Hormigas, Seccion de Entomologa, Departamento de Biologa, Universidad del Valle, A. A. 25360 Cali, Colombia e-mail: jimjacag@gmail.com C. Bermudez-Rivas e-mail: christianax@gmail.com A. M. Osorio e-mail: anamilenaosorio@gmail.com P. Chacon e-mail: patricia.chacon@correounivalle.edu.co Present Address: J. Cabra-Garca Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo CEP: 05508-090, Brazil

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not nd evidence for an ideal diversity surrogate although diversity patterns of predatory ants had the greatest congruencies. These results support earlier multi-taxon evaluations in that conservation planning should not be based on only one leaf litter arthropod group. Keywords Ants Arachnids Cross-taxon congruence Diversity indicators Multi-taxa inventories Southwestern Colombia Surrogate taxa Tropical dry forest

Introduction Shortage of knowledge on biodiversity, especially in tropical environments (Lawton et al. 1998; Wrigth 2005), coupled with rapid rates of species loss induced by human activities, is a major challenge for conservation biology (Negi and Gadgil 2002; Brooks et al. 2002; McGeoch 2007). This challenge can be faced effectively only by obtaining as much information as possible with minimum requirements in time and resources (McGeoch 2007). In this context the assessment of biodiversity indicators on multiple scales has gained great momentum over the last decade (Caro and Odoherty 1999; Duelli and Obrist 2003; Wolters et al. 2006; McGeoch 2007). In particular, different arthropod groups have attracted interest as potential indicators of biodiversity because of their signicant contribution to global species richness, biomass and a large variety of ecological functions and lifestyles (Kotze and Samways 1999; Moritz et al. 2001; Sauberer et al. 2004; Schulze et al. 2004; Lovell et al. 2007; Basset et al. 2008a, b; Kessler et al. 2009; Maleque et al. 2009; Uehara-Prado et al. 2009; Finch and Lofer 2010; Leal et al. 2010; Vasquez-Velez et al. 2010). The use of biodiversity indicators is based on the premise that across large areas the number of species in one well-studied taxon correlates well with the number of species in other lesser known taxa (Schall and Pianka 1978; Prendergast and Eversham 1997). It is assumed that this taxon will positively reect the patterns and processes of biological diversity in the landscape (Kotze and Samways 1999). According to Gaston (1996a, b), there are four possible explanations for richness correlation between different taxa: (1) random coincidence, (2) interactions between taxa, (3) similar responses to common factors and (4) responses to different environmental factors that are spatially covariant. Ants and arachnids, especially spiders, have been among the most commonly used groups to evaluate the effect of environmental changes (fragmentation, pollution and habitat loss) on the diversity of arthropod communities (Marc et al. 1999; Alonso 2000; Maleque et al. 2009). However, the species richness patterns of these groups have not been consistent as there have been congruent responses in some scenarios (Weaver 1995; Niemela et al. 1996; Sauberer et al. 2004; Leal et al. 2010), while in others no statistically signicant correlation has been detected (Oliver and Beattie 1996; Kotze and Samways 1999). Moreover, this trend does not apply only for these taxa; various studies have not found consistent patterns of variation in diverse plants, vertebrates and invertebrates (Lawton et al. 1998; Oliver et al. 1998; Kati et al. 2004; Heino et al. 2005; Barlow et al. 2007; Kessler et al. 2009; Heino et al. 2009; Heino 2010). Nevertheless, the search for indicator groups and their assessment across different environments and spatial scales remains an important issue because of the potential for simplifying data requirements for conservation planning (Larsen et al. 2009; Caro 2010; Lewandowski et al. 2010; Toranza and Arim 2010). In the Tropics, assessments of congruency across invertebrate taxa to identify potential surrogates have focused mainly on humid tropical forests (Lawton et al. 1998; Barlow et al.

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2007; Basset et al. 2008a; Uehara-Prado et al. 2009; Leal et al. 2010). In contrast, investigations in dry ecosystems are scarce. The tropical dry forest is one of the most endangered ecosystems in the Neotropics (Janzen 1988). Given the fertility of its soils, it has undergone intense transformation for beneting urban populations in the form of agriculture and livestock (Alvarez et al. 1998). Recent fragmentation studies in the upper Cauca River basin (southwestern Colombia), showed that forests (10,716 ha) represent only 1.76% of the landscape, and other native elements such as giant bamboo forests account for barely 1%; whereas crops (52%) and pastures (36%) dominate (Arcila 2007). Because of the current context, the upper Cauca River basin provides a framework suitable for assessing potential biodiversity indicators. According to Kotze and Samways (1999), epigeic invertebrates are appropriate in multitaxa conservation-oriented research because they (1) include two of the most studied terrestrial taxa (Carabidae and Formicidae); (2) are associated with one of the most complex habitat systems, i.e. the soil; and (3) are easily captured using standard techniques such as pitfall trapping. In this paper we describe and compare a and b diversity of ve leaf litter arthropod groups (ants, predatory ants, oribatid mites, spiders and other arachnids) in 39 sampling patches of the upper basin of the Cauca River valley. In addition, we assess the congruence of a and b diversities among groups, explore the effect of sampling completeness in congruence analyses, and evaluate whether some taxa can be used as diversity surrogates.

Materials and methods Study area The study was carried out in the upper basin of the Cauca River Valley, which comprises three provinces in southwestern Colombia (from the northern Cauca Province to central Risaralda Province) (Fig. 1a). This valley is 220 km long and 1020 km wide with a total area of 420,000 ha. According to Holdridges life zone classication system, this region belongs to the tropical dry forest life zone. It is located at altitudes from 900 to 1,100 masl, the average annual temperature is above 24C, and the average annual rainfall ranges from 1,000 to 2,000 mm (CVC 1990). Ten sites with one tropical dry forest fragment were selected across the basin (Fig. 1a). These forests are secondary (originated from clear-cut harvesting followed by a natural tree regeneration) with relatively well-structured canopies approximately 25 m in height, dominated by tree species such as Anacardium excelsum, Xilopia ligustrifolia, Laetia americana, Ficus spp., Cecropia sp. and Ceiba petandra (CVC 1990; Armbrecht and Ulloa-Chacon 2003). At each site sampling patches were identied within a radius of 1,500 m from the centroid of the dry forest fragment (Fig. 1b). These patches were grouped into the following vegetation types: sugarcane crop, grassland, giant bamboo forest and riparian forest, comprising a total of 39 sampling patches (Supplementary material 1). Sampling methods Fieldwork was conducted between October and December 2005 and FebruaryMay 2006, covering rainy and dry seasons. We applied the sampling protocol used to characterize the diversity of ants and staphylinids in rural landscapes of the Andean region (Jimenez et al. 2008; Abada et al. 2010; Vasquez-Velez et al. 2010) and the Cauca River geographic

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Fig. 1 Study area. a Geographic site of the upper Cauca River basin, southwestern Colombia. b An example of a sampling site: A fragment of tropical dry forest and surrounding sampling patches located within a radius of 1,500 m from the centroid of the forest fragment

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valley (Arcila-Cardona et al. 2008). For each sampling patch two 50-m linear transects were marked and separated by a distance of no less than 50 m. Both transects included six sampling units, set up every 10 m and consisting of two methods of capture: mini-Winkler sacks and pitfall traps, placed parallel to the transect and separated by 10 m. In total, the sampling effort included 39 sampling patches, 78 transects and 468 sampling units. Samples collected with each method of capture in each sampling unit, were pooled for data analysis due to the great complementarity in the species composition collected with each method and the enhanced sampling completeness. Processing samples Ants and spiders were separated from the rest of arthropods collected and identied to the species level, using all available keys and consulting specialists. Specimens that could not be identied to a specic category were grouped into morphospecies (unnamed species diagnosed with standard taxonomic techniques) (Basset et al. 2008b). The arachnid orders used in the analysis were: Araneae, Schizomida, Sarcoptiformes: Oribatida, Opiliones, Scorpiones and Pseudoscorpiones. Only adults were considered for data analyses because juvenile identication is difcult and ambiguous in many cases due to marked variations during ontogeny (Coddington et al. 1996; Foelix 1996; Norton and Behan-Pelletier 2009). The vouchers were deposited in the Museo de Entomologa de la Universidad del Valle (MUSENUV). Data analysis Taxa that were used in the analyses were ants (excluding predatory ants), predatory ants (subfamilies Ponerinae, Proceratiinae, Ectatomminae and Amblyoponinae), oribatid mites, spiders and other arachnids. The last group comprises pseudoscorpions, scorpions, harvestmen and schizomids due to their low sampling frequency. Given the social behavior of ants, we used presenceabsence data as a measure of abundance as Longino (2000) suggests, rather than the number of individuals. a Diversity Observed richness (aobs) and estimated species richness (aest) per sampling patch, were used to analyze patterns of a diversity of each group across the study area. Estimated richness (aest) was calculated by using the average of four incidence-based estimators (ICE, Chao 2, Jack 1 and Jack 2) with the EstimateS 8.2 program (Colwell 2010). In order to compare a and b diversity more directly (Kessler et al. 2009), we also calculated differences in species richness between each sampling patch pair (all possible combinations of the 39 sampling patches). This was done for the observed (Daobs) and the estimated (Daest) richness values. We grouped the sampling patches into ve contrasting vegetation types (secondary forests, riparian forests, giant bamboo forests, sugarcane crops and pastures), in order to visually compare patterns of a diversity for each group. KruskalWallis tests and a posteriori multiple comparisons with the Statistica 7.0 program (Statsoft 2004) were used to compare richness values across vegetation types.

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b Diversity Similarities in the species composition of sampling patches pairs (all possible combinations of the 39 sampling patches) were quantied with the quantitative Srensen similarity index (also known as BrayCurtis index) (bobs) according to Kessler et al. (2009). To correct for incomplete sampling, the similarity index of Chao et al. (2005) was used, which is based on the foregoing index but includes an estimate of incompleteness (best). Raw abundance values were transformed to the fourth root in order to standardize abundance between taxa (Su et al. 2004); in the case of ants these values correspond to presence/ absence data as stated above. Similarities in the species composition among vegetation types were also evaluated. Correlation analyses In our analyses a diversity correspond to the diversity within sampling patches and b diversity corresponds to the diversity among sampling patches. We used the Spearman rank correlation (due to the absence of normality or homoscedasticity in some variables) to test for cross-taxon a diversity correlations (aobs, aest) and the Mantel test for cross-taxon b diversity correlations (bobs, best) and for cross-taxon Da diversity correlations (Daobs, Daest). Because of a possible spatial effect on the correlation coefcient values (distance between sampling patches), we tested spatial autocorrelation using the Mantel test to correlate a matrix of geographical distance between sampling patches and matrices of differences in a diversity (Da) and b diversity. This multivariate measure evaluates the null hypothesis of no relationship between two similarity or dissimilarity matrices (Su et al. 2004; Kessler et al. 2009). Probabilities are assigned by repeatedly randomizing the arrangement of similarity matrices, recalculating correlation coefcients each time and comparing the observed correlation value to the randomly generated ones (Kessler et al. 2009). All Mantel tests were conducted with the XLSTAT program (Addinsoft 2008), using 9,999 randomizations. Mantel analyses were also used to assess the relationship between Da diversity and b diversity. We calculated mean sampling completeness (i.e. the ratio between observed and mean estimated richness) for each taxon. To assess the potential impact of mean sampling completeness in correlation analyses, we correlated this variable with the correlation coefcients obtained in each combination (e.g. predatory ants/spiders), using the lower of the two values for mean sampling completeness (Kessler et al. 2009). Based on Leal et al. (2010) a surrogate was dened as being reasonable if it explained[60% of the variation in total species richness, good if it explained [70%, and excellent if it explained [80%.

Results A total of 6,765 individuals grouped into 290 morphospecies were collected (Supplementary material 2). The most abundant taxa were ants and oribatid mites, which accounted for 89.66% of all captures. The groups with the highest number of morphospecies were ants and spiders with 54.13 and 22.41%, respectively, of the total registered (Table 1). The regional mean sampling completeness ranged from 55% for other arachnids to 87% for predatory ants (Table 1).

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Biodivers Conserv Table 1 Richness, abundance and mean sampling completeness of ants and arachnids in 39 sampling patches of the upper Cauca River basin Taxon Ants Predatory ants Oribatid mites Spiders Other arachnids Abundance 3,107 569 2,390 504 195 aobs 135 22 41 65 27 aest 169.36 25.31 58.17 117.55 49.04 Regional MSC (%) 79.71 86.92 70.48 55.30 55.06 RSC (%) 3389 42100 33100 41100 43100

MSC mean sampling completeness, RSC range of mean sampling completeness for individual sampling patches

Diversity patterns across vegetation types In general richness patterns were consistent in all taxa; species richness tended to decrease in highly intervened vegetation types (pastures and sugarcane crops) in relation to native ones (forests) (Fig. 2). This trend was statistically signicant in ants and other arachnids (Fig. 2). All taxa evaluated showed the highest value of species richness in secondary forest and the lowest value in sugarcane crops, except for other arachnids, which had the lowest richness in pastures. The average observed similarity (bobs) between vegetation types ranged from 13% (other arachnids) to 54% (ants); average estimated similarity (best) ranged from 27% (other arachnids) to 94% (ants). Correlation analysis a Diversity Correlation coefcients of aobs diversity varied considerably (Table 2). Four of the 10 pairwise comparisons were signicant and all were positive, with ants and predatory ants most strongly correlated (q = 0.65; P \ 0.0001). Similar results were obtained with Daobs diversity although the number of signicant correlations was higher in relation to aobs (Table 3). Correlation coefcients for aest and Daest tended to decrease in relation to aobs and Daobs (Fig. 3); likewise, the number of signicant correlations decreased from 4 to 2 (aobs vs. aest) and from 9 to 8 (Daobs vs. Daest) (Tables 2, 3). No signicant correlations were found between mean sampling completeness and correlation coefcients in any diversity measure (aobs q = 0.25, P = 0.49; aest q = -0.25, P = 0.48; Daobs q = 0.31, P = 0.37; Daest q = 0.16, P = 0.66). Geographical distance between sampling patches was not correlated with any a diversity measure (Table 4). b Diversity Eight of the 10 pairwise comparisons of bobs were signicant, all of which were positive. Correlation coefcients of bobs diversity ranged from 0.05 to 0.49 (Table 5), similar results were obtained with best diversity although the correlation values were somewhat lower. Geographical distance between sampling patches was correlated with beta diversity measures across all taxa, except for predatory ants. All except one of the signicant correlations were negative (Table 4).

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Fig. 2 Species richness (aobs) of taxa evaluated across vegetation types. The box plot shows median (black square), 50% (box limits) and 95% of all values (brackets). Different letters denote signicant differences (P \ 0.05). Patterns of estimated richness (aest) show roughly similar patterns and are therefore not shown. SF secondary forest, BF giant bamboo forest, RF riparian forest, SC sugarcane crop, P pasture

No matter which diversity measure was used (a or b), no individual taxon reached any surrogate threshold ([60, [70, 80% [of the variation explained). In general cross-taxon a-diversity correlations were higher than cross-taxon b-diversity correlations (Fig. 3).

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Biodivers Conserv Table 2 Correlations (P values) of a-diversity (species richness per sampling patch) among the ve study groups Ants Ants Predatory ants Oribatid mites Spiders Other arachnids 0.65*** 0.41* 0.28 0.58** Predatory ants 0.3 0.31 -0.03 0.43* Oribatid mites 0.18 0.25 0.15 0.13 Spiders 0.21 -0.12 0.08 0.18 Other arachnids 0.41* 0.38* 0.19 0.25

Values above the diagonal were based on the estimated richness values (aest); values below, on the observed richness values (aobs) * P \ 0.05, ** P \ 0.001, *** P \ 0.0001 Table 3 Correlations (P values) of Da-diversity (differences in species richness per sampling patch) among the ve study groups Ants Ants Predatory ants Oribatid mites Spiders Other arachnids 0.68*** 0.4** 0.25** 0.47*** Predatory ants 0.35** 0.25** -0.03 0.4** Oribatid mites 0.16* 0.22** 0.14* 0.12* Spiders 0.11* -0.19* 0.03 0.10* Other arachnids 0.31** 0.30** 0.08 0.12*

Values above the diagonal were based on the estimated richness values (Daest); values below, on the observed richness values (Daobs) * P \ 0.05, ** P \ 0.001, *** P \ 0.0001

a Diversity versus b diversity All comparisons of patterns of a and b diversity resulted in low correlation values, ranging from q = -0.16 to q = 0.06, averaging q = -0.01 for Daobs versus bobs and from q = -0.16 to q = 0.09, averaging q = -0.02 for Daest versus best.

Discussion a Diversity Contrary to the results reported by other authors (Oliver and Beattie 1996; Kotze and Samways 1999), a diversity patterns across taxa were in general consistent (Fig. 2). Vegetation types with canopy layer such as forests had the highest richness values in relation to homogeneous and highly intervened vegetation types such as pastures and sugar cane crops. Variations in a diversity are primarily related to local environmental factors and interactions between populations (Halffter and Moreno 2005). In this context the so-called habitat heterogeneity hypothesis (Tews et al. 2004), becomes important in explaining the diversity patterns observed in this study. This hypothesis assumes that the structural complexity of habitats may provide more niches and diverse ways of exploiting the environmental resources, thereby increasing species diversity (Tews et al. 2004). Thus

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Fig. 3 Correlation (P) values of a diversity (the number of species per sampling patches), Da diversity (differences in species numbers between sampling patches), and b diversity (similarities in the species composition between sampling patches) of each of the ve study groups relative to the other 4 groups. The box plots show medians (black square), 50% (the box limits) and 95% of all values (brackets). A ants, PA predatory ants, OM oribatid mites, S spiders, OA other arachnids

similarity in richness patterns (Fig. 2) may reect a common trend of the evaluated taxa against an increase in habitat heterogeneity, from the highly intervened vegetation types to the native ones. Vegetation of a given environment determines certain characteristics of the leaf litter layer such as nutritional quality, depth and heterogeneity (Facelli and Pickett 1991; Cornelissen et al. 1999, Koivula et al. 1999), which directly affect the epigeic arthropod assemblages analyzed (Uetz 1979; Bultman and Uetz 1982; Bultman et al. 1982; Hansen 2000; McGlynn et al. 2009) by reducing or increasing factors such as competitive exclusion between species that inhabit this microenvironment (Wagner et al. 2003; Achury

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Biodivers Conserv Table 4 Mantel correlations (P values) of diversity measures and geographical distance between sampling patches Taxon Ants Predatory ants Oribatid mites Spiders Other arachnids * P \ 0.05, ** P \ 0.001 Daobs -0.056 -0.049 -0.061 -0.033 -0.005 Daest -0.031 -0.044 -0.059 -0.026 -0.006 bobs -0.118* 0.004 -0.09* -0.17* 0.26** best -0.09* 0.005 -0.12* -0.17* -0.13*

Table 5 Mantel correlations (P values) of b-diversity (differences in species composition between sampling patches) among the ve study groups Ants Ants Predatory ants Oribatid mites Spiders Other arachnids 0.58** 0.44** 0.16* 0.32** Predatory ants 0.37** 0.28** 0.06 0.25** Oribatid mites 0.14* 0.25** 0.10* 0.10* Spiders 0.18* 0.04 0.12* 0.05 Other arachnids 0.12* 0.03 0.02 0.17*

Values above the diagonal were based on Chaos Srensen similarity estimator (best); values below, on Srensens quantitative (BrayCurtis) similarity index (bobs) * P \ 0.05, ** P \ 0.001

et al. 2008). Thus, further analyses of these properties in the leaf litter are needed to establish to what extent it contributes to the richness patterns of the groups evaluated. In accordance with Kessler et al. (2009), patterns of species richness were fairly similar, regardless of the measure used (aobs, aest, Daobs, Daest). Predatory ants and the rest of the ant assemblage had the strongest signicant correlations in almost all cases (Tables 4, 5). Despite the variation of the species richness (aobs) of ants that can be explained by the richness of predatory ants is under our dened ranges for a diversity surrogate (\60%), both taxa appear to respond similarly to the conditions of the landscape in terms of species richness. Two investigations evaluated the potential use of ants as biodiversity indicators in the upper Cauca River basin: Armbrecht and Ulloa-Chacon (2003) reported the little re ant Wasmannia auropunctata (Roger 1863) as a negative indicator of ant diversity in 14 dry forest fragments, and Chacon et al. (2008) suggested predatory ants as positive indicators of ant diversity in 10 dry forest fragments. The latter authors suggest that congruence in richness patterns of these taxa could be related to the fact that the two groups share similar microhabitat requirements for nesting. Our study is the second investigation that reports a signicant and positive relationship between predatory ant species richness and ant species richness in the basin; nevertheless the group did not reach the established surrogate thresholds. We found different degrees of correlations (Table 2), which might be associated with some of the explanations suggested by Gaston (see Introduction section); however a

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detailed assessment of environmental factors is necessary to explore mechanisms behind of the congruence and/or inconsistencies between the patterns, and further research is required on the natural history of the groups. First of all, the distance between sampling patches had no signicant effect on the variation in the richness of the taxa (Table 4) so the spatial factor in terms of site and distance between sampling patches can be eliminated as an explanation for the observed patterns. Another factor that can contribute to consistency in richness patterns is a similar response of each taxon to an increment in area (the speciesarea relationship); nevertheless a preliminary exploration of this subject did not suggest a linear relationship of aobs with patch area (Bermudez-Rivas unpublished data). In our comparisons, the range of the variation of species richness of a given group that could be explained by the richness of another was 0.142%. These values are within the range reported by Schulze et al. (2004) for ve groups in Indonesia, Barlow et al. (2007) for 15 groups in Brazilian Amazonia and Leal et al. (2010) for nine groups in Brazilian Atlantic forest and Caatinga, and are higher than values reported in other studies (Lawton et al. 1998; Kessler et al. 2009). Some factors that can inuence the absence of consensus in the assessment of surrogate taxa are variation in (1) area in which correlations are assessed (Weaver 1995; Negi and Gadgil 2002; Sauberer et al. 2004; Wolters et al. 2006; Hess et al. 2006), (2) sampling protocols (Sauberer et al. 2004; Leal et al. 2010) and (3) measures for the consistency of diversity patterns (Su et al. 2004; Barlow et al. 2007; Lovell et al. 2007; Basset et al. 2008a; Leal et al. 2010; Grantham et al. 2010). It is interesting to note that the correlation coefcients obtained with aest tended to be lower in relation to that of aobs which is consistent with some of the comparisons reported by Kessler et al. (2009). This fact has a close relationship with mean sampling completeness, because when we compare groups with distinct sampling completeness, Daobs will be necessarily different in relation with Daest which changes correlation patterns. b Diversity Cross-taxon b-diversity correlations were lower than cross-taxon a-diversity correlations (Fig. 3). All correlations were positive as occurred in earlier studies (Barlow et al. 2007; Kessler et al. 2009). In our comparisons, only 0.2 (spiders vs. other arachnids)33% (ants vs. predatory ants) of the variance of similarities in the species composition of a given group could be explained by the similarity of another. These values are in the range reported by Barlow et al. (2007) and Uehara-Prado et al. (2009). Kessler et al. (2009) state that the b diversity of all taxa along an ecological gradient (land use, elevation, climate, soil fertility, etc.) will tend to shift more strongly when the ecological conditions are more divergent. Nevertheless, the highly modied area (upper Cauca River basin) where we evaluated correlations is a complex landscape with no clear regional ecological gradient. Each sampling patch and its surrounding matrix are a complex and unique mosaic landscape where interaction between species occurs. This could be a reason for the lower cross-taxon correlations in b diversity, which is contrary to results in recent publications (Su et al. 2004; Kessler et al. 2009). A pattern that seems to be consistent across all taxa is that b diversity contributes in a large proportion to gamma diversity (Bermudez-Rivas unpublished data), a pattern that have been found for spiders and predatory ants (Arcila Cardona et al. 2008; Cabra-Garca et al. 2010). Our results suggest that distance between sampling patches (Table 4) explain a very low percentage (0.0066.760%) of the variation of b diversity across the sampling patches.

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a versus b diversity Our results suggest that, on average, only 0.015% of b diversity variation of a given group could be explained by the patterns of a diversity of another group. This result is congruent with that reported by Kessler et al. (2009). Contrary to recent ndings (Barlow et al. 2007; Kessler et al. 2009; Uehara-Prado et al. 2009), cross-taxon a-diversity correlations were higher than cross-taxon b-diversity correlations. As proposed by Leal et al. (2010), the fact that the groups evaluated in this study were collected by the same method of capture could promote congruence in a-diversity measures given that several studies using different collection methods at different spatial scales tended not to detect correlations between groups (Leal et al. 2010). It is likely that taxa living and operating in the same microhabitat have more congruent diversity patterns than those from different microhabitats (Leal et al. 2010). To our knowledge, this is the rst approach in Colombia that aims to assess congruence in a and b diversity patterns of different leaf litter arthropods (multi-taxa approach) in a tropical dry forest landscape. Therefore this research can serve as a baseline for comparing diversity patterns from other arthropod taxa in order to identify robust surrogates of diversity across different spatial scales. Ideally, conservation planning would be based on detailed surveys, including a thorough knowledge of the affected species life histories, distributions and interactions with other species and the physical environment (Favreau et al. 2006). In the Colombian context, however, detailed data are costly to accumulate and cannot be obtained in the timeframe within which landscape-altering decisions are made. For this reason assessment of diversity congruence indicators is a very important tool for guiding conservation planning.

Conclusions Although no group reached the surrogate thresholds, we found a signicant and positive relationship between predatory ants and ants species richness. Future research needs to evaluate whether this pattern is consistent regardless of scale, habitat, sampling method and diversity measures used. Our results suggest that conservation planning in the upper Cauca River basin should not be based on only a single leaf litter arthropod taxon given that each group appears to be affected by different factors to a different extent, which could be related to the lack of strong congruencies (Caro 2010).
Acknowledgments Biologists Angela Arcila, Nestor Zuniga, Paloma Vejarano, Pilar Lopez and Rafael Achury participated in the eldwork. Researchers Fernando Fernandez, Eduardo Florez, David Luna, Jairo Moreno, Osvaldo Villareal, Lillan Sendales, Alexandra Sierra, Wojciech Niedbaa and Jari Niemela contributed with taxonomic determination and provided literature. Three anonymous reviewers and the Asso ciate Editor Jari Niemela provided detailed comments and important suggestions on an earlier version of the paper. The program Joven Investigadores e Innovadores Virginia Gutierrez de Pineda from Colciencias and Universidad del Valle funded the participation of J. Cabra-Garca and A. M. Osorio. The Instituto de Investigacion de Recursos Biologicos Alexander von Humboldt (IAvH) funded part of this project (Agreement No. 05-01-24843-054CE).

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