Animal Conservation (2004) 7, 169–179 C 2004 The Zoological Society of London. Printed in the United Kingdom DOI:10.1017/S1367943004001258

Bird responses to shade coffee production

Cesar´ Tejeda-Cruz and William J. Sutherland Centre for Ecology, Evolution and Conservation, School of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, UK (Received 2 June 2003; accepted 16 October 2003)

Abstract It has been documented that certain types of shade coffee plantations have both biodiversity levels similar to natural forest and high concentrations of wintering migratory . These findings have triggered a campaign to promote shade coffee as a means of protecting Neotropical migratory . Bird censuses conducted in the El Triunfo Biosphere reserve in southern have confirmed that shade coffee plantations may have bird diversity levels similar to, or higher than, natural forest. However, coffee and forest differed in species composition. Species with a high sensitivity to disturbance were significantly more diverse and abundant in primary ecosystems. Neotropical migratory birds, granivorous and omnivorous species were more abundant in disturbed habitats. Insectivorous bird species were less abundant only in shaded monoculture. Foraging generalists and species that prefer the upper foraging stratum were more abundant in disturbed habitats, while a decline in low and middle strata foragers was found there. Findings suggests that shade coffee may be beneficial for generalist species (including several migratory species), but poor for forest specialists. Although shade coffee plantations may play an important role in maintaining local biodiversity, and as buffer areas for forest patches, promotion of shade coffee may lead to the transformation of forest into shade coffee, with the consequent loss of forest species.

INTRODUCTION Sterling, 1997b; Estrada et al., 1998; Megchun´ Guerrero, 1999; Moguel & Toledo, 1999; Petit et al., 1999; Roberts, Coffee is one of the world’s most traded commodities Cooper & Petit, 2000a), the species composition is and the major source of foreign exchange earnings for generally dominated by generalist and opportunistic many developing countries. It is estimated that over species. There is also evidence that differences in species 125 million people worldwide are dependent on coffee richness and composition between native forest and shade for their livelihoods (International Coffee Council, 2001). coffee patches are a function of ecosystem structural Coffee in northern Latin America covers around 44% complexity (Gallina, Mandujano & Gonzalez-Romero, of the total arable cropland (Rice & Ward, 1996). This ´ 1996; Parrish & Petit, 1996; Greenberg et al., 1997b; region has 3.1 million hectares of coffee production and Calvo & Blake, 1998; Megchun Guerrero, 1999), or a generates 34% of the world’s production (Perfecto & ´ function of distance from forest (Parrish & Petit, 1996; Armbrecht, 2002). Roberts, Cooper & Petit, 2000b). In terms of avifauna, In the Neotropics, coffee is an important conservation studies have reported that shade coffee plantations issue because it is often produced in countries identified may provide suitable habitat for several bird species as megadiverse (Perfecto & Armbrecht, 2002). Shade (Greenberg et al., 1997a,b; Wunderle & Latta, 2000). In coffee has higher biodiversity levels than unshaded sun fact, public and scientific interest in coffee increased after coffee (Nestel, Dickschen & Altieri, 1993; Perfecto & studies revealed that a great number of Neotropical Vandermeer, 1994, 1996; Perfecto & Snelling, 1995; migratory bird species winter in shade coffee plantations Wunderle & Latta, 1996; Greenberg et al., 1997a). (Vaninni, 1994; Greenberg, 1996; Wexler, 2002). How- Although some studies have found similar or higher levels ever, shade coffee has been found to be detrimental for of bird species richness in shade coffee in comparison forest associated species (Greenberg et al., 1997b; Roberts to natural forest (Aguilar-Ortiz, 1982; Estrada, Coates- et al., 2000a; Armbrecht & Perfecto, 2003). Estrada & Meritt, 1993, 1994, 1997; Parrish & Petit, In recent years much traditional shade coffee has 1996; Wunderle & Latta, 1996; Greenberg, Bichier & been converted into sun coffee in Latin America. Sun coffee grow without shade and insecticides are All correspondence to: Cesar´ Tejeda-Cruz. Fax: +44(0) 1603 extensively used. The combination of insecticides with 592250; E-mail: [email protected] the elimination of the shade overstory may result in 170 C. TEJEDA-CRUZ &W.J.SUTHERLAND reduced biodiversity (Vaninni, 1994; Gallina et al., 1996; Table 1. Sampling effort and altitudinal range of treatments Wunderle & Latta, 1996; Greenberg et al., 1997a). Given the replacement of shade coffee by sun coffee, there have Habitat No. of Altitude been urgent demands to stop the transformation of shade Habitat code samples (m a.s.l.) coffee into sun coffee, with shade coffee being linked in Cloud forest CF 47 1700–2200 public awareness campaigns to forest conservation and Montane rain forest MR 43 1100–1699 the protection of migratory songbirds (Rice & McLean, Lower montane rain forest LM 41 450–1099 1999). Since 1995, conservation organisations have been Rustic coffee RC 46 500–1200 promoting ‘shade coffee’ over ‘sun coffee’ as a way to stop Shaded monoculture SM 55 1000–1600 this trend (National Audubon Society, 1998; Sherry, 2000; Sun coffee SC 4 950–1050 Conservation International, 2001; Wexler, 2002; Rappole, Total samples 236 King & Vega Rivera, 2003). However, to divide coffee production systems into ‘traditional’ versus ‘modern’ or m a.s.l., metres above sea level. ‘shade’ versus ‘sun’ does not reflect the real complexity of these production systems. In practice, they represent Chiapas, the southernmost Mexican State. It covers almost the extreme ends of a continuum of intensification, with 120 000 ha, comprising a buffer zone of about 93 500 ha several stages between the two extremes. Subsequently, (79%) and five core zones totalling 25 700 ha (21%). The shade coffee conservation value is related to the degree core zones are pristine forest where economic activities of intensification of the system used (Rice & Ward, 1996; are not allowed. The buffer zone is private land, ‘ejidos’ Calvo & Blake, 1998). The usefulness of the campaign (land communally owned) and individual farms. Roughly to promote shade coffee production is debatable since: 14 000 people live in the buffer zone. Approximately (1) biodiversity found in shade coffee is impoverished by 60% of the buffer zone (56 000 ha) is densely forested, the presence of generalistic and opportunistic species and with the remaining 40% (approximately 37 400 ha) under a decrease in forest associated species (Canaday, 1996; agricultural production with coffee as the main cash crop Greenberg et al., 1997b; Petit et al., 1999; Roberts et al., as well as small scale maize and cattle ranching, mainly 2000a; Rappole et al., 2003) and (2) this marketing for local consumption (IHN, 1995; INE, 1999). strategy may inadvertently encourage an increase in shade Coffee production systems in El Triunfo can be coffee area at the expense of natural forest (Rappole et al., classified as: (1) rustic coffee, in which the original 2003). tree layer is maintained and coffee bushes substitute for Like most tropical areas, southern Mexico is currently the original understory plants; (2) traditional polyculture, under increasing land use pressure by local people. Our where some original trees are maintained, but coffee is study area, El Triunfo Biosphere Reserve, in Chiapas, grown alongside many useful introduced plants and trees; has one of the most important cloud forest remnants in (3) shaded monoculture, where planted leguminous trees Mexico. This area is considered to be an Important Bird (mainly species of the genus Inga) are almost exclusively Area (Arizmendi & Marquez,´ 2001) and an Endemic Bird used to provide shade for coffee bushes and (4) unshaded Area (Stattersfield et al., 1998) because of the quality monoculture (sun coffee), where coffee is a monoculture and uniqueness of its bird fauna. Coffee production is planted with no tree shade at all (Moguel & Toledo, the main economic activity. It has been estimated that 1999). more than 80% of the buffer zone inhabitants work in In our study area, rustic coffee was present mainly activities related to coffee production. However, coffee in the form of small production units in ‘ejido’ land. expansion has been identified as the main threat to Shaded monoculture was found mainly in the coffee primary ecosystems (IHN, 1995; Tejeda Cruz et al., 1997). ‘fincas’ (large, single-owned coffee plantations). Finally, Moreover, Conservation International is currently running sun coffee was very localised in small stands in some of the its flagship project, namely ‘Conservation Coffee’, with biggest plantations. The original natural forest from which small coffee growers in El Triunfo’s buffer and influence the coffee plantations were derived can be separated into areas. Hence there is an urgent need to investigate the cloud forest, montane rain forest and lower montane rain effects of different coffee production systems on biota forest (Breedlove, 1981). There is an important altitudinal at a local scale. Our objectives were to analyse how variable that may have an effect on the results. Table 1 different shade coffee systems affect bird community shows the altitudinal ranges for the treatments studied. structure in relation to natural forest. Specifically we Lower montane rain forest, located at elevations examined how coffee system intensity affected bio- below 1100 m above sea level (a.s.l.), where human diversity, species composition, feeding guild, foraging density is greater, has been historically more exposed to strata and the abundances of forest associated species, anthropogenic disturbance. As a result, it has a fragmented migratory species and generalist species. distribution embedded in a mosaic of coffee plantations, maize fields and pasture lands. These fragments show various levels of disturbance. Therefore, lower montane rain forest will be considered to be disturbed rain forest. STUDY AREA Cloud forest in El Triunfo is dense, consisting of broad- El Triunfo Biosphere Reserve (15◦ 09 10 N, 15◦ 57 leaved evergreen trees with a mean canopy height of 19.81 02 N and 92◦ 34 04 W, 9 3 ◦ 12 2 W) is located in (± 0.68) m and a canopy cover of 91%. Cloud forest Birds in shade coffee plantations 171 dominant tree species include: Hedyosmum mexicanum, per habitat by means of a rarefaction analysis using a Matudea trinervia, Conostegia volcanalis, Heliocarpus Monte Carlo simulation procedure based on the number donell-smithii, Glossostipula concinna, Amphitecna mon- of individuals found in the habitat with the lowest number tana and Oreopanax xalapensis. Montane rainforest is of individuals (Gotelli & Entsminger, 2001). Simulations characterised by the presence of a well-developed under- were run 1000 times and the mean expected number of story of small trees and shrubs, a mean canopy height of species per treatment was compared on the basis of the 24.9 (± 1.24) m and a canopy cover of 92%. Dominant 95% confidence interval (CI), where non-overlapping CI trees include: Ficus cooki, Coccoloba matudae, Dipholis bars indicate a significant difference between means. minutiflora, Roupala montana, Liquidambar styraciflua, Species were assigned to functional groups. Foraging Pinus oocarpa and Quercus spp (Heath & Long, 1991). strata preferences were defined following Stotz et al. The lower montane rain forest tree layer is dominated by (1996), but the strata classifications were simplified to Astronium graveolens, Rheedia edulis, Bursera simaruba, improve the response resolution. Feeding guilds were Manilkara achras, Casearia sp, Croton guatemalensis, defined using information from Stiles & Skutch (1989), Erythroxylum mexicanum and Randia armata. Its mean Terborgh et al. (1990), Howell & Webb (1995) and canopy height is 23.05 (± 0.95) m and its canopy cover Greenberg et al. (1997b). Species response to disturbance is 84%. Rustic coffee has a tree layer dominated by was categorised by their sensitivity to disturbance Trichilia moschata ssp matudai, Lonchocarpus sp, Inga according to an independent assessment (Parker, Stotz & laurina, Inga rodrigueziana and mombin; its Fitzpatrick, 1996). Differences between treatments were canopy cover is 60% and its mean canopy height is compared using one-way analysis of variance (ANOVA) or 16.31 (± 0.67) m. Finally, shaded monoculture has a mean Kruskal-Wallis test depending upon data normality (Zar, canopy height of 11.19 (± 0.64) m and a canopy cover of 1996). 41%. In shaded monoculture tree species of the genus Inga Differences in species composition among treatments are predominant, principally I. micheliana and I. paterna. were graphically analysed using non-metric multidimen- Also important are Trema micrantha, Alchornea latifolia, sional scaling (NMDS) methods. This method graphically Cedrela sp, Montanoa sp and Ulmus mexicanum.(C.T.-C., shows the similarity between samples in terms of species unpubl. results). composition and abundances. Distance between points indicates degree of similarity. We favoured the use of NMDS because it does not rely on any assumptions about data distribution (Clarke & Warwick, 1994). Analyses METHODS were performed using PRIMER v.5 software. Similarity Bird survey matrices for bird species were built using the Bray–Curtis similarity index from untransformed data. Differences in Data were collected following the same methods and at species composition were examined using the analysis of the same points in May–July 2000 and 2001 for overall similitude test (ANOSIM: Clarke & Warwick, 1994). avifauna, and in January 1997 for migratory birds, except that in 1997 montane rain forest and sun coffee were not sampled. To establish distribution and relative abundance of bird species, we conducted bird censuses using 10-min RESULTS fixed-radius point counts (Hutto, Pletschet & Hendricks, We recorded a total of 182 bird species (4617 individuals) 1986). To mitigate possible effects of differences in of which 33 were migratory bird species (500 individuals: detectability between habitats we recorded all bird species see Appendix). Randomised species accumulation curves detected and numbers of individuals within and beyond the for each of the habitat types are shown in Fig. 1(a). Most 25 m radius. All censuses were carried out between 15 min habitat types had similar species richness, with the after sunrise and 10:00 am when bird activity decreases. exception of cloud forest, which had a significantly lower Point counts were placed systematically within treatments diversity according to the results of rarefaction ana- and at least 150 m apart, to avoid double counting of lysis based on 1000 simulations (Fig. 1(b). We found individuals. To avoid edge effects, all samples were placed the following number of bird species per habitat (with at least 50 m from any boundary with neighbouring the estimated number of species derived from rarefaction vegetation. analysis in parentheses): cloud forest, 51 (46); montane rain forest, 85 (78), lower montane rain forest, 86 (77), rustic coffee, 80 (70), shaded monoculture, 79 (69) and Data analysis sun coffee, 6 (insufficient data). All analyses were carried out using 2000–2001 data, Species composition varied between habitats (Fig. 2). except the analysis of migratory birds, which was carried NMDS scores showed that cloud forest and montane rain out using 1997 data. Species richness was compared forest are distinctly different from the other treatments. graphically using species accumulation curves const- However, rustic coffee and lower montane rain forest ructed from the means of 100 randomisations of sample clustered together, showing similar species composition. order (Colwell & Coddington, 1997; Colwell, 2000). Finally, shaded monoculture and sun coffee each clustered Sample effort varied among treatments (see Table 1). apart. Analysis of similitude test results were significant Therefore, we calculated the expected number of species for all pairs of habitats (Table 2). Differences between 172 C. TEJEDA-CRUZ &W.J.SUTHERLAND

90 (a) Table 2. Results from the analysis of similitude test and the average dissimilarity for all of the habitats sampled and for all habitat pairs 80 Habitats Mean dissimilarity 70 compared R P (%) 60 LM, RC 0.126 0.001 75.27 50 SM, RC 0.545 0.001 85.47 40 MR, LM 0.473 0.001 86.27 SM, SC 0.621 0.001 88.91 Cloud forest Number of species 30 CF, MR 0.650 0.001 88.99 Montane rain forest SM, LM 0.723 0.001 89.13 20 Lower montane rain forest Rustic coffee MR, RC 0.644 0.001 90.28 10 Shaded monoculture MR, SM 0.715 0.001 92.91 Sun coffee SC, RC 0.843 0.001 93.62 0 0 102030405060SC, LM 0.995 0.001 97.02 MR, SC 0.811 0.001 97.35 Number of samples CF, SM 0.955 0.001 98.41 CF, LM 0.995 0.001 98.88 100 (b) CF, RC 0.987 0.001 99.27 90 CF, SC 0.997 0.001 100 80 Global 0.710 0.001 70 60 CF, cloud forest; MR, montane rain forest; LM, lower montane rain forest; RC, rustic coffee; SM, shaded monoculture; SC, sun coffee. 50 40 30 rustic coffee and lower montane rain forest showed smaller 20 differences than other habitat pairs (Table 2, Fig. 2). 10 Neotropical migratory species occurred in rustic coffee and shaded monoculture in similar numbers to lower Expected number of species (95% CI) 0 Cloud Montane Lower Rustic Shaded montane rain forest. In contrast, cloud forest held forest rain forest montane coffee monoculture lower numbers (Fig. 3(a)). However, migratory species rain forest abundance was significantly higher in rustic coffee and Fig. 1. (a) Species accumulation curves for bird species in all the shaded monoculture than in either montane rain forest habitat types. Species richness varied among habitats: cloud forest or cloud forest (H = 19.332, d.f. = 3, P < 0.001). The (51), montane rain forest (85), lower montane rain forest (86), rustic migratory bird abundance in shade coffee is three times coffee (80), shaded monoculture (79) and sun coffee (4). (b) Mean that of cloud forest and nearly double that of montane rain expected number of species based on 1000 simulations in samples forest. of 500 observations. Means are considered to be different if the Despite having a species richness that was comparable 95% Confidence Intervals (CI; bars shown) do not overlap. to, or greater than, natural ecosystems, shade coffee plantations contained fewer species and fewer individuals of species with a high sensitivity to habitat disturbance (H = 128.3, d.f. = 4, P < 0.001: Fig. 3(b)). Montane rain Cloud forest Shaded monoculture forest and cloud forest presented 12 and 11 sensitive 1.5 Montane rain forest Rustic coffee Lower montane rain forest Sun coffee species, respectively; while lower montane rain forest showed an intermediate level with seven sensitive species. 1 In contrast, only four sensitive species were recorded in each of the coffee plantations, except in sun coffee which 0.5 had none. 0

−0.5 Feeding and strata guilds

−1 Abundances of species grouped in feeding guilds dif- stress 0.18 fered between treatments (Fig. 4(a)). Insectivorous and −1.5 frugivorous species were less common in shaded mono- −2 −1.5 −1 −0.5 0 0.5 1 1.5 2 2.5 culture, while the other treatments showed no differences = = < Fig. 2. Non-metric multi-dimensional scaling ordination of the (insectivores: H 64.282, d.f. 4, P 0.001; frugivores: abundances of species showing differences in species composition H = 18.994, d.f. = 4, P = 0.001). Granivorous species among habitats. The stress value indicates a fair goodness-of-fit of were more abundant in disturbed sites (lower montane the configuration. rain forest, rustic coffee and shaded monoculture), when Birds in shade coffee plantations 173

40 2.0 compared to cloud forest and montane rain forest (H = (a) 20.523, d.f. = 4, P < 0.001). The opposite pattern was shown by nectarivorous species, which were less abundant 30 1.5 in disturbed sites (H = 2.029, d.f. = 4, P < 0.001). There is a gradual increase in omnivorous species along the 20 1.0 disturbance gradient. At one extreme, cloud forest had the lowest abundance of omnivorous species, while at the other end, shaded monoculture possessed the highest 10 0.5 abundance (H = 24.899, d.f. = 4, P < 0.001). Coffee plantations and lower montane rain forest had = 0 0.0 higher abundances of upper stratum species (H 36.057, CF LM RC SM d.f. = 4, P < 0.001) and species with no preference = = < 14 6 for a particular strata (H 95.871, d.f. 4, P 0.001: (b) Fig. 5(b)). However, rustic coffee and shaded monoculture

Number of species 12 5 held lower abundances of species that prefer the middle strata (H = 106.656, d.f. = 4, P < 0.001) and low level 10 4 Mean number of individuals strata (H = 14.901, d.f. = 4, P = 0.005). Disturbed rain 8 3 forest, like both coffee systems, showed a high abundance 6 2 of upper strata species, but abundances of species with a preference for middle, low and variable strata were similar 4 1 to those found in natural ecosystems. 2 0

0 DISCUSSION CF MR LM RC SM Species richness Fig. 3. Number of species (columns) and mean number of in- dividuals per treatment (points with error bars) for Neotropical The results suggest that shade coffee plantations may migrants (a) and species with high sensitivity to disturbance (b). CF, sustain a high bird abundance and species richness. In cloud forest; MR, montane rain forest; LM, lower montane rain particular, shade coffee is important for wintering forest; RC, rustic coffee; SM, shaded monoculture; SC, sun coffee. Neotropical migrants. Although montane rain forest was not evaluated during winter, we would expect it to have (a) a lower level of Neotropical migrants to those found 12 Cloud forest Montane rain forest Lower montane rain forest in shade coffee systems and disturbed forest. This is 10 Rustic coffee because secondary habitats are well known for supporting Shaded monoculture a large number of migratory species populations during 8 winter, sometimes surpassing primary habitats (Terborgh, 1989; Rappole, 1995; Stotz et al., 1996). However, 6 differences in species composition between forest and shade coffee were significant. Differentiation in feeding 4 guild composition, preferred foraging strata composition 2 and sensitivity to disturbance suggests that shade coffee may be beneficial for generalist species, but negatively 0 affect forest specialists. Insectivore Frugivore Granivore Nectarivore Omnivore Although the sun coffee sample size was very small, the results suggest that its species richness is very low 10 (b) (Fig. 1(a)). Other studies have reported the low bird diversity of unshaded coffee (Wunderle & Latta, 1996; 8 Greenberg et al., 1997a). Mean number of individuals The elevation gradient in our study may confound 6 some of the comparisons between habitats because higher elevations have lower bird diversity (Stotz, 1998). This 4 problem has also been acknowledged in previous studies (Wunderle & Latta, 1996; Greenberg et al., 1997a). Species richness in shade coffee systems is similar to 2 montane rain forest and lower montane rain forest and is even higher than that found in cloud forest. Species 0 composition and richness in lower montane rain forest is High Middle Low Variable more similar to rustic coffee than to montane rain forest. Fig. 4. Bird composition across habitats. (a) mean abundance per Lower montane rain forest is located at lower elevations, feeding guild and (b) preferred foraging strata. where the landscape is a mosaic of forest patches in a 174 C. TEJEDA-CRUZ &W.J.SUTHERLAND matrix of varying levels of disturbance including shade pattern suggests that shade coffee may have limited coffee, maize crops and pasture lands. resources during part of the breeding season for resident The high bird species richness found in rustic coffee and species (from March to July for several species). Genera- shaded monoculture may be explained by the structural list species can ‘spread the risk’ of extinction by being complexity of these systems. It is known that habitat polyphagous as opposed to monophagous food specialists structural heterogeneity is often positively related to (Vazquez & Simberloff, 2002). Thus, a reduction in diversity (MacArthur & MacArthur, 1961). In addition, the abundance of frugivorous and insectivorous species coffee plantations are located near to large patches of in shaded monoculture is probably related to seasonal natural forest. Distance to the forest in our samples changes in resource availability, coupled with impoveri- averaged 52 m for rustic coffee and 257 m for shaded shed tree diversity. Fewer tree species means a reduced monoculture (C.T.-C., unpubl. results). Proximity to chance of finding a fruiting tree at some times of the adjacent extensive forest may enhance the habitat value year. In shaded monoculture, shade is dominated by a of shade coffee for many forest associated species (Petit few species of Inga trees (2.4 tree species per sample: et al., 1999; Roberts et al., 2000a,b). Parrish & Petit C.T.-C., unpubl. results). The same reasoning may be used (1996) found that the further the plantation was from to explain the decline of nectarivores in disturbed habitats. the continuous forest, the lower the number of forest The herbaceous stratum is more developed in coffee species shared with forest interior and forest edges. In plantations and disturbed forest than it is in undisturbed addition, Mart´ınez & Peters (cited by Moguel & Toledo, forest systems since canopy cover is more open and more 1999) presented empirical evidence on the effect of sunlight is available. This observation may explain the plantation isolation. They found more species in - higher abundance of granivorous species found in coffee ations adjacent to forest patches and significantly systems. fewer in isolated plantations. However, when coffee In shade coffee plantations, the original bush layer plantations are adjacent to primary forest, it is difficult has been replaced by coffee bushes. Few bird species to determine the ability of the plantation alone to prefer to use coffee bushes for foraging and none feed support breeding bird populations (Moguel & Toledo, on coffee beans (Greenberg et al., 1997a; our pers. obs.). 1999). Thus, abundance reduction in birds that prefer to forage Our results support the notion that shade coffee systems in the lower and middle strata may be explained by the are an important habitat for wintering migratory species. substitution of native bushes by coffee plants and the This finding is endorsed by several previous studies reduction in leaf litter because of more open canopy. (Vaninni, 1994; Wunderle & Latta, 1996; Greenberg et al., However, this bird reduction in the mid and lower strata 1997a,b; Megchun´ Guerrero, 1999; Petit et al., 1999). was compensated for by an increase in the abundance of Nevertheless, several Neotropical migratory species are canopy and strata generalist species. known to use secondary habitats in the wintering grounds (Terborgh, 1989; Rappole, 1995). Moreover, Vaninni (1994) and Greenberg et al. (1997b) noticed that shaded High sensitivity species monoculture may provide a dry season refuge for several bird species. The dry season in our study area occurs from Despite having the lowest species richness, cloud forest November to April, hence this may partially explain the held the highest number and abundance of high sensitivity high abundance of Neotropical migratory species in shade species. Montane rain forest also possessed high diversity coffee sites. and abundance of high sensitivity species. The low richness and abundance of high sensitivity species that was found in lower montane rain forest pinpoints disturbance associated with this habitat. Both shade coffee habitats Ecological specialisation showed numbers of high sensitivity species that were It is widely believed that generalist species would benefit similar to those found in lower montane rain forest. from disturbance, while specialists may be negatively Nevertheless, some of the forest species (including affected (McKinney & Lockwood, 1999; Vazquez & some high sensitivity species) were present in shade Simberloff, 2002). In our study, we considered omni- coffee systems. However, bird use does not necessarily vorous species to be less specialised than species with a correspond to reproductive success (Van Horne, 1983). narrower feeding niche and species with no preference These species’ populations can respond to shade coffee for a particular foraging stratum more generalist than disturbance in different ways: (1) they benefit through birds that prefer one. The results suggest that the a reduction in the number of competitors, so they greater abundance and richness of omnivorous and strata successfully breed in coffee systems, (2) they make a generalists in shade coffee systems may, thus, be positively facultative use of coffee systems resources, but breed related to disturbance. in nearby primary forest or, (3) coffee systems are Fruit and nectar are more available in coffee plantations a ‘sink’ habitat for the surplus individuals produced than other habitats during the dry season (Vaninni, 1994), in neighbouring primary forest (Pulliam, 1988). More in particular Inga dominated shade coffee provides large research is necessary to assess how different forest species numbers of arthropods (Johnson, 2000) and abundant respond to the perturbation caused by coffee produc- nectar resources (Greenberg et al., 1997a). This seasonal tion. Birds in shade coffee plantations 175

Shade coffee conservation value Arizmendi,M.D.C.&Marquez,´ L. (Eds) (2001). Areas´ de importancia para la conservacion´ de las aves en Mexico´ .Mexico,D.F.: Much of the case for the current campaign for shade CONABIO. coffee conservation is based on the fact that shade coffee Armbrecht, I. & Perfecto, I. (2003). Litter-twig dwelling ant species shelters biodiversity levels that are similar to forest richness and predation potential within a forest fragment and ecosystems and provides habitat for several migratory neighbouring coffee plantations of contrasting habitat quality in birds. However, diversity indices alone do not provide Mexico. Agr. Ecosyst. Environ. 97: 107–115. reliable conservation criteria since communities with Breedlove, D. E. (1981). Introduction to the Flora of Chiapas.San Francisco, CA: California Academy of Sciences, USA. similar biodiversity can have very different ecological Calvo, L. & Blake, J. (1998). Bird diversity and abundance on two compositions (Rappole et al., 2003). This criterion alone different shade coffee plantations in Guatemala. Bird Consersv. Int. can lead to actions that might result in forest trans- 8: 297–308. formation. If current campaigns are successful and criteria Canaday, C. (1996). Loss of insectivorous birds along a gradient of enforcement is weak, local producers may choose to human impact in Amazonia. Biol. Conserv. 77: 63–77. transform forested areas into shade coffee. Clarke, K. R. & Warwick, R. M. (1994). Change in marine communities: The value of shade coffee to bird conservation remains an approach to statistical analysis and interpretation. Plymouth: Plymouth Marine Laboratory. incompletely understood. Shade coffee, like many other Colwell, R. K. (2000). EstimateS: statistical estimation of species anthropogenic perturbations, is detrimental for forest richness and shared species from samples. http://viceroy.eeb.uconn. associated species. Therefore, the transformation of forest edu/EstimateS. into shade coffee could be of limited conservation value. Colwell, R. K. & Coddington, J. A. (1997). Estimating terrestrial However, shade coffee provides habitat for several biodiversity through extrapolation. Phil. Trans. R. Soc. Lond. Ser. resident and migratory birds. Undoubtedly shade coffee B 345: 101–118. Conservation International (2001). Conservation coffee. http://www. is preferable to the drastic perturbation that results from celb.org/pressreleases/conservationcoffeefactsheet5-22.pdf transformation into intensive monoculture production (accessed 25-May-2003). systems. In other words, to transform sun coffee or other Estrada, A., Coates-Estrada, R., Anzures Dadda, A. & Cammarano, P. monocultures into shade coffee would increase bird (1998). Dung and carrion beetles in tropical rain forest fragments diversity and be beneficial to some Neotropical migrants and agricultural habitats at Los Tuxtlas, Mexico. J. Trop. 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Appendix. Occurrence of birds in the habitats surveyed in El Triunfo Biosphere Reserve, Chiapas, Mexico. Occurrence is based on the average number of detections per sample (x100) in fixed-radius point counts during May–July 2000 and 2001 for resident species and January 1997 for migratory birds (MR and SC not surveyed in 1997). Numbers in parentheses indicate winter occurrence of birds detected in both periods. An asterisk after the common name indicates bird species with a high sensitivity to habitat perturbation. Status code: r, resident; m, migratory. Diet code: i, insectivore; f, frugivore; n, nectarivore; g, granivore; o, omnivore; c, carnivore. Strata code: L, lower; M, middle; U, upper. Habitat code: CF, cloud forest; MR, montane rain forest; LM, lower montane rain forest; RC, rustic coffee; SM, shaded monoculture; SC, sun coffee.

Habitat Common name Scientific name Status Diet Stratum CF MR LM RC SM SC

Thicket Tinamou Crypturellus cinnamomeus r o L 0.0 0.0 7.3 17.4 3.6 0.0 Gray Hawk Asturina nitida r c U 0.0 0.0 0.0 4.3 0.0 0.0 Barred Forest-Falcon Micrastur ruficollis r c M 0.0 2.3 4.9 0.0 0.0 0.0 Collared Forest-Falcon Micrastur semitorquatus r c U 0.0 0.0 2.4 2.2 0.0 0.0 Laughing Falcon Herpetotheres cachinnans r c U 0.0 0.0 0.0 4.3 0.0 0.0 Bat Falcon Falco rufigularis r c U 0.0 0.0 0.0 6.5 0.0 0.0 Plain Chachalaca Ortalis vetula r f G 0.0 0.0 14.6 6.5 3.6 0.0 Crested Guan Penelope purpurascens r f G 0.0 0.0 2.4 6.5 0.0 0.0 Highland Guan∗ Penelopina nigra r f U 4.3 14.0 17.1 10.9 1.8 0.0 Horned Guan∗ Oreophasis derbianus r f G 4.3 0.0 0.0 0.0 0.0 0.0 Crax rubra r f L 0.0 0.0 2.4 0.0 0.0 0.0 Spotted Wood-Quail∗ Odontophorus guttatus r g L 0.0 2.3 0.0 0.0 0.0 0.0 Singing Quail∗ Dactylortyx thoracicus r g L 19.1 30.2 0.0 0.0 9.1 0.0 Red-billed Pigeon Columba flavirostris r f U 0.0 4.7 24.4 19.6 47.3 0.0 Band-tailed Pigeon Columba fasciata r f U 0.0 0.0 0.0 0.0 7.3 0.0 White-winged Dove Zenaida asiatica r g G 0.0 0.0 0.0 0.0 16.4 0.0 Inca Dove Columbina inca r g L 0.0 0.0 0.0 0.0 3.6 0.0 White-tipped Dove Leptotila verreauxi r g L 0.0 18.6 126.8 108.7 20.0 0.0 White-faced Quail-Dove∗ Geotrygon albifacies r g L 42.6 7.0 0.0 0.0 1.8 0.0 Green Parakeet Aratinga holochlora r f U 0.0 37.2 0.0 0.0 0.0 0.0 Orange-fronted Parakeet Aratinga canicularis r f U 0.0 0.0 29.3 10.9 0.0 0.0 White-crowned Parrot Pionus senilis r f U 0.0 37.2 0.0 0.0 56.4 0.0 White-fronted Parrot Amazona albifrons r f U 0.0 0.0 24.4 0.0 0.0 0.0 Squirrel Cuckoo Piaya cayana r I U 0.0 9.3 22.0 30.4 3.6 0.0 Striped Cuckoo Tapera naevia r I L 0.0 0.0 14.6 26.1 0.0 0.0 Pheasant Cuckoo Dromococcyx phasianellus r I L 2.1 2.3 0.0 0.0 3.6 0.0 Groove-billed Ani Crotophaga sulcirostris r I G 0.0 0.0 0.0 0.0 1.8 0.0 Spectacled Owl Pulsatrix perspicillata r I U 0.0 0.0 2.4 0.0 0.0 0.0 Ferruginous Pygmy-Owl Glaucidium brasilianum r c U 0.0 0.0 0.0 0.0 0.0 25.0 Rufous Sabrewing Campylopterus rufus r n M 2.1 20.9 0.0 0.0 0.0 0.0 Violet Sabrewing Campylopterus hemileucurus r n M 17.0 7.0 0.0 4.3 0.0 0.0 Emerald-chinned Hummingbird Abeillia abeillei r n M 4.3 7.0 0.0 0.0 0.0 0.0 Black-crested Coquette Lophornis helenae r n U 2.1 0.0 0.0 0.0 0.0 0.0 White-eared Hummingbird Hylocharis leucotis r n G 0.0 2.3 0.0 0.0 3.6 0.0 Berylline Hummingbird Amazilia beryllina r n U 0.0 2.3 12.2 15.2 5.5 0.0 Cinnamon Hummingbird Amazilia rutila r n G 0.0 0.0 0.0 2.2 0.0 0.0 Green-throated Mountain-gem Lampornis viridipallens r n M 55.3 2.3 0.0 0.0 0.0 0.0 Amethyst-throated Hummingbird Lampornis amethystinus r n M 0.0 4.7 0.0 0.0 0.0 0.0 Blue-throated Hummingbird Lampornis clemenciae r n M 2.1 0.0 0.0 0.0 0.0 0.0 Long-billed Starthroat Heliomaster longirostris r n U 0.0 0.0 0.0 2.2 0.0 0.0 Ruby-throated Hummingbird Archilochus colubris m n M 0.0 – 4.8 17.4 0.0 – Violaceous Trogon Trogon violaceus r o U 0.0 20.9 39.0 28.3 18.2 0.0 Mountain Trogon Trogon mexicanus r o U 2.1 9.3 0.0 0.0 3.6 0.0 Collared Trogon Trogon collaris r o U 21.3 25.6 43.9 30.4 0.0 0.0 Resplendent Quetzal Pharomachrus mocinno r o M 8.5 0.0 0.0 0.0 0.0 0.0 Tody Motmot∗ Hylomanes momotula r I L 0.0 2.3 2.4 0.0 0.0 0.0 Blue-throated Motmot∗ Aspatha gularis r I L 8.5 14.0 0.0 0.0 0.0 0.0 Blue-crowned Motmot Momotus momota r o M 0.0 55.8 73.2 63.0 20.0 0.0 Turquoise-browed Motmot Eumomota superciliosa r I M 0.0 0.0 19.5 26.1 0.0 0.0 Emerald Toucanet Aulacorhynchus prasinus r f U 42.6 25.6 0.0 0.0 10.9 0.0 Collared Aracari Pteroglossus torquatus r f U 0.0 2.3 14.6 19.6 0.0 0.0 Acorn Woodpecker Melanerpes formicivorus r I U 0.0 2.3 0.0 0.0 9.1 0.0 Golden-fronted Woodpecker Melanerpes aurifrons r o U 0.0 4.7 43.9 69.6 25.5 0.0 Yellow-bellied Sapsucker Sphyrapicus varius m I U 4.2 – 0.0 0.0 0.0 – Hairy Woodpecker Picoides villosus r I U 2.1 11.6 0.0 0.0 0.0 0.0 Golden-olive Woodpecker Piculus rubiginosus r I U 0.0 7.0 12.2 4.3 3.6 0.0 Lineated Woodpecker Dryocopus lineatus r o U 0.0 0.0 7.3 0.0 0.0 0.0 178 C. TEJEDA-CRUZ &W.J.SUTHERLAND

Appendix. Cont.

Habitat Common name Scientific name Status Diet Stratum CF MR LM RC SM SC

Scaly-throated Foliage-gleaner∗ Anabacerthia variegaticeps r I M 34.0 30.2 0.0 0.0 3.6 0.0 Ruddy Foliage-gleaner Automolus rubiginosus r I L 38.3 14.0 4.9 0.0 3.6 0.0 Tawny-throated Leaftosser∗ Sclerurus mexicanus r I L 0.0 16.3 2.4 0.0 0.0 0.0 Ruddy Woodcreeper∗ Dendrocincla homochroa r I L 4.3 4.7 9.8 4.3 0.0 0.0 Olivaceous Woodcreeper Sittasomus griseicapillus r I M 0.0 7.0 0.0 0.0 0.0 0.0 Northern Barred-Woodcreeper∗ Dendrocolaptes sanctithomae r I M 0.0 0.0 4.9 0.0 0.0 0.0 Ivory-billed Woodcreeper Xiphorhynchus flavigaster r I M 0.0 0.0 58.5 32.6 7.3 0.0 Spotted Woodcreeper Xiphorhynchus erythropygius r I M 10.6 0.0 12.2 4.3 1.8 0.0 Streak-headed Woodcreeper Lepidocolaptes souleyetii r I M 0.0 0.0 0.0 2.2 0.0 0.0 Spot-crowned Woodcreeper Lepidocolaptes affinis r I M 21.3 9.3 0.0 0.0 1.8 0.0 Barred Antshrike Thamnophilus doliatus r I M 0.0 0.0 17.1 21.7 1.8 0.0 Greenish Elaenia Myiopagis viridicata r o U 0.0 0.0 2.4 0.0 0.0 0.0 Ochre-bellied Flycatcher Mionectes oleagineus r f G 0.0 4.7 2.4 0.0 0.0 0.0 Paltry Tyrannulet Zimmerius vilissimus r o U 34.0 23.3 0.0 2.2 0.0 0.0 Northern Bentbill Oncostoma cinereigulare r I M 0.0 9.3 0.0 0.0 0.0 0.0 Eye-ringed Flatbill Rhynchocyclus brevirostris r o M 0.0 14.0 2.4 2.2 0.0 0.0 Yellow-olive Flycatcher Tolmomyias sulphurescens r o U 0.0 11.6 87.8 69.6 52.7 0.0 Stub-tailed Spadebill Platyrinchus cancrominus r I L 0.0 0.0 9.8 2.2 0.0 0.0 Tufted Flycatcher Mitrephanes phaeocercus r I M 10.6 0.0 0.0 0.0 0.0 0.0 Greater Pewee Contopus pertinax r I U 0.0 0.0 0.0 15.2 0.0 0.0 Tropical Pewee Contopus cinereus m I G 0.0 0.0 0.0 0.0 9.1 0.0 Least Flycatcher minimus m I U 0.0(0.0) 0.0 2.4(0.0) 2.2(8.7) 0.0(0.0) 0.0 Yellowish Flycatcher Empidonax flavescens r I M 42.6 30.2 0.0 0.0 0.0 0.0 Black Phoebe Sayornis nigricans r I G 0.0 0.0 0.0 0.0 1.8 0.0 Bright-rumped Attila Attila spadiceus r o U 0.0 4.7 26.8 21.7 0.0 0.0 Dusky-capped Flycatcher Myiarchus tuberculifer r o U 2.1 23.3 51.2 37.0 38.2 0.0 Great Kiskadee Pitangus sulphuratus r o G 0.0 2.3 0.0 0.0 1.8 0.0 Boat-billed Flycatcher Megarynchus pitangua r o U 0.0 4.7 0.0 0.0 5.5 0.0 Social Flycatcher Myiozetetes similis r o U 0.0 7.0 34.1 45.7 41.8 25.0 Sulphur-bellied Flycatcher Myiodynastes luteiventris t o U 0.0 7.0 12.2 32.6 50.9 0.0 Tropical Kingbird Tyrannus melancholicus r o U 0.0 2.3 2.4 2.2 9.1 0.0 Rose-throated Becard Pachyramphus aglaiae r o U 0.0 2.3 4.9 0.0 5.5 0.0 Masked Tityra Tityra semifasciata r o U 0.0 9.3 4.9 0.0 50.9 0.0 Long-tailed Manakin Chiroxiphia linearis r f M 0.0 53.5 161.0 80.4 0.0 0.0 Yellow-throated Vireo Vireo flavifrons m I U 0.0(0.0) 2.3 82.9(14.3) 123.9(21.7) 0.0(0.0) 0.0 Blue-headed Vireo Vireo solitarius m o U 0.0 – 0.0 0.0 12.9 – Warbling Vireo Vireo gilvus m o U 0.0 0.0 0.0 8.7 0.0 0.0 Brown-capped Vireo Vireo leucophrys m o U 14.9 2.3 0.0 0.0 1.8 0.0 Philadelphia Vireo Vireo philadelphicus m o U 0.0 – 0.0 0.0 6.5 – Lesser Greenlet Hylophilus decurtatus r o U 0.0 0.0 12.2 0.0 1.8 0.0 Green Shrike-Vireo Vireolanius pulchellus r o U 0.0 16.3 14.6 2.2 0.0 0.0 Rufous-browed Peppershrike Cyclarhis gujanensis r I U 0.0 0.0 14.6 15.2 34.5 0.0 Green Jay Cyanocorax yncas r o U 0.0 130.2 70.7 58.7 54.5 0.0 Black-throated Jay∗ Cyanolyca pumilo r o U 42.6 0.0 0.0 0.0 0.0 0.0 Band-backed Wren Campylorhynchus zonatus r I U 0.0 0.0 0.0 0.0 14.5 0.0 Spot-breasted Wren Thryothorus maculipectus r I M 0.0 2.3 29.3 4.3 12.7 0.0 Rufous-and-white Wren Thryothorus rufalbus r I L 2.1 32.6 43.9 32.6 0.0 0.0 Banded Wren Thryothorus pleurostictus r I L 0.0 14.0 0.0 2.2 0.0 0.0 Plain Wren Thryothorus modestus r I L 0.0 2.3 29.3 47.8 1.8 0.0 Southern House Wren Troglodytes musculus r I L 0.0 7.0 2.4 21.7 100.0 50.0 Rufous-browed Wren Troglodytes rufociliatus r I M 34.0 0.0 0.0 0.0 0.0 0.0 White-breasted Wood-Wren Henicorhina leucosticta r I L 4.3 125.6 36.6 6.5 5.5 0.0 Gray-breasted Wood-Wren Henicorhina leucophrys r I L 193.6 7.0 0.0 0.0 0.0 0.0 Long-billed Gnatwren Ramphocaenus melanurus r I M 0.0 0.0 7.3 0.0 0.0 0.0 Brown-backed Solitaire∗ Myadestes occidentalis r f U 134.0 39.5 2.4 2.2 18.2 0.0 Orange-billed Nightingale- Catharus aurantiirostris r o L 21.3 34.9 4.9 2.2 7.3 0.0 Thrush Ruddy-capped Nightingale- Catharus frantzii r o L 27.7 0.0 0.0 0.0 0.0 0.0 Thrush Spotted Nightingale- Catharus dryas r o L 114.9 23.3 0.0 0.0 0.0 0.0 Thrush∗ Swainson’s Thrush Catharus ustulatus m f M 0.0(0.0) 0.0 17.1(4.8) 37.0(21.7) 0.0(25.8) 0.0 Hermit Thrush Catharus guttatus m o M 4.2 – 23.8 60.9 0.0 – Black Robin∗ Turdus infuscatus r o U 57.4 0.0 0.0 0.0 0.0 0.0 Birds in shade coffee plantations 179

Appendix. Cont.

Habitat Common name Scientific name Status Diet Stratum CF MR LM RC SM SC

Mountain Robin Turdus plebejus r o G 36.2 0.0 0.0 0.0 0.0 0.0 Clay-colored Robin Turdus grayi r o G 0.0 18.6 17.1 69.6 180.0 25.0 White-throated Robin Turdus assimilis r o M 0.0 60.5 63.4 82.6 56.4 0.0 Gray Catbird Dumetella carolinensis m o L 0.0 0.0 0.0 2.2 0.0 0.0 Blue-and-white Mockingbird Melanotis hypoleucus r o L 0.0 0.0 0.0 0.0 9.1 0.0 Cedar Waxwing Bombycilla cedrorum m f U 0.0 – 0.0 0.0 38.7 – Gray Silky-flycatcher Ptilogonys cinereus r f U 0.0 0.0 0.0 0.0 0.0 0.0 Tennessee Warbler Vermivora peregrina m o U 0.0(0.0) 0.0 17.1(90.5) 0.0(43.5) 0.0(103.2) 0.0 Nashville Warbler Vermivora ruficapilla m o G 0.0(0.0) 0.0 0.0(14.3) 2.2(13.0) 0.0(3.2) 0.0 Crescent-chested Warbler Parula superciliosa r I U 42.6 0.0 0.0 0.0 0.0 0.0 Yellow Warbler Dendroica petechia m I U 0.0 0.0 0.0 2.2 0.0 0.0 Magnolia Warbler Dendroica magnolia m I U 0.0 0.0 12.2 4.3 0.0 0.0 Black-throated Green Warbler Dendroica virens m I U 0.0(0.0) 0.0 2.4(0.0) 0.0(4.3) 0.0(29.0) 0.0 Townsend’s Warbler Dendroica townsendi m I U 4.2 – 14.3 0.0 6.5 – Hermit Warbler Dendroica occidentalis m I U 0.0 – 0.0 0.0 19.4 – Grace’s Warbler Dendroica graciae m I U 0.0 – 4.8 0.0 0.0 – Black-and-white Warbler Mniotilta varia m I U 16.7 – 0.0 4.3 3.2 – American Redstart Setophaga ruticilla m I U 0.0(0.0) 0.0 0.0(4.8) 4.3(4.3) 0.0(0.0) 0.0 Seiurus aurocapillus m I L 4.2 – 0.0 0.0 0.0 – MacGillivray’s Warbler Oporornis tolmiei m I L 0.0 – 0.0 0.0 9.7 – Wilson’s Warbler Wilsonia pusilla m I M 10.6(95.8) 0.0 0.0(28.6) 2.2(39.1) 0.0(77.4) 0.0 Canada Warbler Wilsonia canadensis m I M 2.1 0.0 0.0 0.0 0.0 0.0 Slate-throated Redstart Myioborus miniatus r I U 97.9 41.9 0.0 0.0 1.8 0.0 Fan-tailed Warbler Euthlypis lachrymosa r I L 0.0 0.0 12.2 2.2 3.6 0.0 Golden-crowned Warbler Basileuterus culicivorus r o M 23.4 125.6 75.6 17.4 0.0 0.0 Rufous-capped Warbler Basileuterus rufifrons r o L 0.0 23.3 36.6 47.8 60.0 75.0 Golden-browed Warbler Basileuterus belli r o L 63.8 7.0 0.0 0.0 0.0 0.0 Common Bush-Tanager Chlorospingus ophthalmicus r o M 234.0 62.8 0.0 0.0 0.0 0.0 Red-crowned Ant-Tanager∗ Habia rubica r o M 0.0 7.0 9.8 8.7 0.0 0.0 Red-throated Ant-Tanager Habia fuscicauda r o L 0.0 4.7 63.4 10.9 0.0 0.0 Summer Tanager Piranga rubra m o U 0.0(0.0) 0.0 0.0(9.5) 0.0(4.3) 1.8(0.0) 0.0 Western Tanager Piranga ludoviciana m o U 0.0(0.0) 0.0 48.8(28.6) 21.7(21.7) 0.0(0.0) 0.0 Flame-colored Tanager Piranga bidentata r o U 12.8 25.6 0.0 0.0 34.5 0.0 White-winged Tanager Piranga leucoptera r o U 0.0 4.7 0.0 13.0 9.1 0.0 Blue-gray Tanager Thraupis episcopus r o U 0.0 0.0 0.0 0.0 1.8 0.0 Yellow-winged Tanager Thraupis abbas r o U 0.0 9.3 14.6 39.1 116.4 0.0 Scrub Euphonia Euphonia affinis r f U 0.0 7.0 0.0 4.3 3.6 0.0 Yellow-throated Euphonia Euphonia hirundinacea r f U 0.0 0.0 17.1 34.8 0.0 0.0 Blue-crowned Chlorophonia Chlorophonia occipitalis r f U 78.7 39.5 0.0 0.0 3.6 0.0 Azure-rumped Tanager Tangara cabanisi r f U 0.0 46.5 0.0 0.0 0.0 0.0 Blue-black Grassquit Volatinia jacarina r g L 0.0 0.0 0.0 0.0 18.2 0.0 Cinnamon-bellied Flowerpiercer Diglossa baritula r o G 2.1 0.0 2.4 0.0 0.0 0.0 White-naped Brush-Finch Atlapetes albinucha r o L 12.8 0.0 0.0 0.0 0.0 0.0 Chestnut-capped Brush-Finch Buarremon brunneinucha r o L 59.6 0.0 0.0 0.0 0.0 0.0 Prevost’s Ground-Sparrow Melozone biarcuatum r o L 0.0 0.0 0.0 4.3 14.5 0.0 White-eared Ground-Sparrow Melozone leucotis r o L 0.0 39.5 12.2 10.9 12.7 0.0 Rusty Sparrow Aimophila rufescens r o L 0.0 0.0 0.0 0.0 3.6 0.0 Rufous-collared Sparrow Zonotrichia capensis r o L 0.0 0.0 0.0 0.0 5.5 0.0 Black-headed Saltator Saltator atriceps r o G 0.0 9.3 36.6 100.0 47.3 100.0 Yellow Grosbeak Pheucticus chrysopeplus r o U 19.1 0.0 0.0 0.0 10.9 0.0 Rose-breasted Grosbeak Pheucticus ludovicianus m o U 0.0(0.0) 0.0 4.9(28.6) 6.5(8.7) 0.0(3.2) 0.0 Painted Bunting Passerina ciris m g G 0.0(0.0) 0.0 2.4(14.3) 0.0(0.0) 0.0(3.2) 0.0 Melodious Blackbird Dives dives r o G 0.0 2.3 4.9 21.7 83.6 0.0 Great-tailed Grackle Quiscalus mexicanus r o G 0.0 0.0 0.0 10.9 50.9 0.0 Molothrus aeneus r o L 0.0 0.0 0.0 0.0 47.3 0.0 Black-vented Oriole Icterus wagleri r o U 0.0 23.3 0.0 0.0 12.7 0.0 Orchard Oriole Icterus spurius m o U 0.0 – 9.5 0.0 0.0 – Yellow-backed Oriole Icterus chrysater r o U 0.0 0.0 0.0 0.0 10.9 0.0 Spot-breasted Oriole Icterus pectoralis r o U 0.0 0.0 2.4 0.0 0.0 0.0 Altamira Oriole Icterus gularis r o U 0.0 0.0 19.5 8.7 0.0 0.0 Baltimore Oriole Icterus galbula m o U 0.0 – 0.0 4.3 0.0 – Yellow-billed Cacique Amblycercus holosericeus r I L 0.0 0.0 4.9 0.0 3.6 0.0 Black-headed Siskin Carduelis notata r g U 0.0 0.0 0.0 0.0 10.9 0.0 Hooded Grosbeak Coccothraustes abeillei r g U 0.0 0.0 0.0 0.0 5.5 0.0