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Remanentes de bosque tropical como refugios de la diversidad de aves dentro de una matriz de desarrollo turistico en la Peninsula de Yucatan, Mexico.

Tropical forest remnants as shelters of avian diversity within a tourism development matrix in Yucatan Peninsula, Mexico

Tourism development is an important driver of forest fragmentation in some countries in tropical areas, in addition to the historic clearing for cattle and agriculture (Bierregaard & Stouffer, 1997; Lambin, Geist, & Lepers, 2003). Construction of tourism developments and associated infrastructure (golf courses, residential zones, recreational parks, roads, etc.) result in fragmentation of forest habitats (Fahrig, 1997; Christ, Hillel, Matus, & Sweeting, 2003; White et al., 2012), leaving many different shapes and sizes of forest remnants. Further, selective extraction of native vegetation and introduction of exotic species to increase the value of tourism complexes (Chettri, Chandra, Sharma, & Jackson, 2005; Schlaepfer, Sax, & Olden, 2011), modify plant species composition, and forest structure and complexity (vertical stratification and plant species composition). Altogether, these environmental modifications reduce the availability of habitats with suitable attributes (e.g., food resources and shelter) to forest-dependent wild fauna, including bird communities (McGarigal & McComb, 1995; Newsome, Moore, & Dowling, 2002; Buckley, 2004).

In addition, if the number of remnants increases, distance between them increases and the exposed edge becomes larger (Fahrig, 1997; Sodhi, 2002; Sekercioglu, 2007), resulting as plausible scenario a higher mortality of bird species by high nest predation as well lower food availability near to the edge of remnants with respect to their interior (e.g., Whyte, Didham, & Briskie, 2005; Newmark & Stanley, 2011). However, these effects depend on the attributes of avian community such as: migratory status, feeding guilds, species richness, and abundance (Stouffer & Bierregaard, 1995; Bierregaard & Stouffer, 1997); as well as forest type and the local threats facing each of them. Nevertheless, some bird species are able to use forest remnants surrounded by secondary growth, in a matrix with pasture and crops and other land uses, with stable population sizes and even experiencing significant increases in their populations (Hughes, Daily, & Ehrlich, 2002; Sekercioglu, Loarie, Oviedo, Ehrlich, & Daily 2007). Thus, this biodiversity corresponds to species generalists or species associated with anthropogenic activities (Krauss et al., 2010). Forest-interior bird species (i.e., specialist species) abilities to use the matrix of modified habitats surrounding forest fragments may affect their vulnerability in fragmented landscapes i.e. species that avoid the matrix tend to decline or disappear in fragments, while those (i.e., generalist bird species) that tolerate or exploit the matrix often remain stable or increase. However, it is not known what happens in a tourism development where forest remnants are interspersed by residential buildings and tourism activities, which are increasing across the tropical forest in Latin American.

During the period 2000-2010, world tropical forest deforestation was 62 % (Keenan et al., 2015), resulting in 6.5 million hectares lost per year. However, in Mexico showed the largest deforestation rates, with 197 651 hectares lost from the 2001 to 2015 period (see details in http://www.globalforestwatch. org/country/ MEX). Tropical forest originally covered about 8 % of the country, being considered a world "hotspot" because of its high biodiversity and endemism (Myers, Mittermeier, Mittermeier, Da Fonseca, & Kent, 2000). Unfortunately, this ecosystem has experienced high deforestation rates, particularly since the early 1970's, because of conversion to pastures and crops, and the establishment of tourism development. Nonetheless, it is still possible to find considerable amount of tropical forest in the Yucatan Peninsula. However, these tropical forest area consist of forest remnants surrounded by mosaics of agricultural land, tourism development and secondary growth. Therefore, it is very important to know the characteristics and extension of these remnants of tropical forest and evaluate if it possible to conserve bird diversity and richness compared to other areas with different land uses. In order to know if the tropical forest remnants are functioning as bird diversity shelters within a matrix dominated by tourism development in one of the most important tourism area in Mexico, our goal was to better understand differences in bird species richness among natural and modified habitats in Akumal region in Quintana Roo, Mexico. In addition, to investigate the role of the different habitat types in a matrix dominated by tourism development. This study aims to provide a general understanding of how bird communities are affected by tourism development. We expected to find a lower species richness and a distinctive bird species composition in modified environments compared with natural environments (mature tropical forest and tropical forest remnants).

MATERIALS AND METHODS

Study area: The present study was carried out in Akumal, an area with several tourism developments (covering approximately 143 [km.sup.2]) located in the Yucatan Peninsula between 20[degrees]30' N - 87[degrees]12' W & 20[degrees]10' N - 87[degrees]26' W (Fig. 1), at the municipality of Tulum in Quintana Roo, Mexico. This site ranges in elevation from 0 to 20 masl, climate of warm subhumid type with abundant rainfalls in summer. Annual average temperature ranges from 25 to 28 [degrees]C, and annual precipitation between 1 300 and 1 500 mm. Dominant natural vegetation in the area is tropical semideciduous forest, tropical deciduous forest, and tropical flooded forest associated with secondary growth; as well as relicts of dunes coast vegetation and mangrove. Common tree species in the study area included Brosimum alicastrum, Bursera simaruba, Manilkara zapota, Talisia olivaeformis, Metopium brownei, Caesalpinia gaumeri, Thrinax radiata, Coccothrinax readi and Pseudophoenix sargentu. The mangrove relicts founded are dominated by Rhizophora mangle and Laguncularia racemose. From the tide line, where the sand accumulates and the soil is very unstable plants, are established Sesuvium portulacastrum, trailblazers like Ambrosia hispida, Salicornia and Hymenocallis littoralis bigelavii. This vegetation is the limit to stable dunes where there is a thicket forming shrub species complex as Cocoloba uvifera, Ipomoea pescaprae, Camavelia rosea, Sophora tomentosa, and Ernodea littoralis, among others (Miranda, 1959; Rzedowski, 1978).

Habitat classification: Habitat classification was based on main vegetation cover, land uses, and the pattern of utilization by settlements as follow: (a) mature tropical forest: tropical semideciduous and tropical deciduous forest >2 ha with mature trees >10 years with canopy height 8 to 15 m and, diameters >20 cm; (b) tropical forest remnants: tropical semideciduous and tropical deciduous forest remnants <2 ha with mature trees <10 years, with canopy height 4 to 8 m and, diameters <20 cm within golf courses and residential zones; (c) modified environments by tourism developments, that include golf course and artificial water bodies in golf course; hotels zones and residential with natural and introduced vegetation; (e) modified environments by urban developments, crops and livestock, that include urban zone with natural and introduced vegetation, cattle pastures and agricultural fields; (f) coast dunes, beach zone and small remnants mangrove.

Bird surveys: Point counts surveys were conducted along transects in the different habitat types (see above; Hutto, Pletschet, & Hendricks, 1986) from April 2009 to November 2010, for a total of 412 point counts in 96 days. Points were randomly selected to represent different types of natural vegetation and land uses in the area (107 [km.sup.2]). Distance between sampling points were at least 250 m to avoid double-counting of highly local species (Hutto et al., 1986; Ralph, Saber, & Droege, 1995). Observation time by point was 20 min, as proposed for tropical environments (Vielliard, 2000). Points were located in both edge and interior of the forest remnants. Sampling was conducted monthly mostly in the morning (06:00 to 11:30 h) and in the afternoon (15:30 to 20:00 h), additional to nocturnal observations. Birds were identified by sight and sound (mostly), excluding birds that overflew the sampling points. Sampling was avoided on rainy days. Species richness was expressed as the total number of species recorded in each habitat, because effort was approximately equal at all habitats (21 days of sampling effort per habitat, with exception of coast dunes, beach zone and small remnants mangrove, which was 12 days). For species identification, Peterson and Chalif (1989), and Howell and Webb (1995) guides were used, and nomenclature and taxonomic status followed AOU (2017), as well as some supplements.

Bird attributes: Birds were categorized as resident or migratory species according to their presence during the study period and complemented with Howell and Webb (1995). Feeding habits were categorized according to which the species was feeding most frequently, which was complemented with literature sources (Peterson & Chalif, 1989; Howell & Webb, 1995) and field observations: omnivores, nectarivores, carnivores, frugivores, granivores, and insectivores (included aquatic invertebrates as well as bark insectivores, aerial insectivores, trunk insectivores, generalist insectivores, ground insectivores, and leaf insectivores). Habitat use preferences were categorized based on Blair (1996), and based on main cover vegetation of the land uses (see above).

Statistical analyses: Species richness was calculated as the cumulative number of species observed in the study area. EstimateS v.9 was used to compute species accumulation curves for the species detected by survey (number of sampling days) (Colwell, 2013). Species accumulation curves estimate the number of species expected in the study area and to compare qualitatively avian richness among habitat types, based on randomized re-sampling from all pooled samples. Asymptote from species accumulation curves was constructed by Michaelis-Menton species richness estimation function using EstimateS v.9 (Colwell & Coddington, 1994). This method estimates of total species richness based on successively larger numbers of samples from the data set. Non-parametric estimator Jackknife 2 was selected based for having the slightest bias in the accuracy data (Walther & Moore, 2005; Hortal, Borges, & Gaspar, 2006). The Shannon diversity index (H') and Simpson's index (D) were obtained to estimate diversity among habitats (Krebs, 2000). Point Abundance Index (PAI) was calculated by dividing the number of detections for each species by the total number of point's sampled (Blondel, Ferry, & Frochot, 1970). To understand how community composition differs, and what species are present and how the habitats differ in the mix of species they have, we conducted a hybrid multidimensional scaling ordination (HMDS), using the Bray-Curtis dissimilarity index on untransformed species abundance. The hybrid MDS was introduced by Faith, Michin and Belbin (1987) and combines both the PCoA (principal coordinate analysis or classical MDS) and the non-metric MDS (NMDS). It has the advantage of assuming a linear relationship between the ecological distances obtained by the ordination and the dissimilarity measures where it is most often straight (the PCoA part), and only monotonicity where ecological distances (in the ordination space) are too high to be accurately measured (the NMDS part; Faith et al., 1987). Differences between natural environments and modified environments (see above) were tested using a permutation multivariate analysis of variance (PERMANOVA; Anderson, 2005). Data of the coast dunes, beach zone and mangrove were not included given the low number of sampling points made in those areas. All analyses were conducted using Minitab (see details http:// www.minitab.com/).

RESULTS

We recorded a total of 1 914 bird sightings during the study period, with a bird density of 54.3 individuals/observation-hour. A total of 160 species and 50 families was recorded, from which only five species are considered endemic, and 10 species were most frequently recorded (Appendix). Accumulation curves for sampling by census reached an asymptote (Fig. 2A) in the value of 170 species. In this context, Jackknife's 2 estimator resulted in a value of 177 species, indicating that the probability of encountering more species increasing sampling effort is very low (Fig. 2B). From all detected bird species, 99 were permanent residents, 47 were winter visitors, 11 were transients, and three introduced (Appendix). The avian community in the study have a predominance of insectivore species (N = 97, see Appendix).

Bird community attributes by habitats: A total of 96 species were found in mature tropical forest, 92 in tropical forest remnants, 79 in modified environments by tourism developments and, 40 in modified environments by urban developments, crops and livestock (Appendix). Accumulation curves showed that the expected species richness present in mature tropical forest had the highest bird richness (Jackknife 2 = 114), followed by tropical forest remnants with expected species (Jackknife 2 = 110), modified environments by tourism developments (Jackknife 2 = 91), and (Jackknife 2 = 57; Fig. 2B). Only 17 species were exclusively found in mature tropical forest, three in tropical forest remnants, and the rest was shared, and four in modified environments by tourism developments and two in modified environments by urban development, crops and livestock, while the rest were found in two to four habitat types (Appendix). The species richness and diversity values were highest in mature tropical forest (96 species, H' = 3.78 [+ or -] 0.006, D= 0.93 [+ or -] 0.010) and tropical forest remnants (94 species, H'= 3.32 [+ or -] 0.008, D= 0.90 [+ or -] 0.010); while, modified environments by tourism developments (72 species, H'= 2.89 [+ or -] 0.014, D= 0.73 [+ or -] 0.030), and modified environments by urban developments, crops and livestock (40 species, H'= 2.73 [+ or -] 0.012, D= 0.69 [+ or -] 0.029) presented the lowest species richness and diversity values (Fig. 3). Bird species richness and diversity values (H', D) varied significantly among habitats (Fig. 3; P< 0.001), with few species detected in modified environments compared with mature tropical forest and tropical forest remnants. This was supported also by the HMDS ordination explained 55 % of the variation in species composition among habitats. Clear gradients in community composition were observed along both axes, with the centroids for mature tropical forest sites and tropical forest remnants having negative values on both axes and the centroids for modified environments (by tourism developments and by urban developments, crops and livestock) having positive values (Fig. 4). Mature tropical forest and tropical forest remnants sites were clearly separated from modified environments, and grouping of same land-use sites was significant (F2 = 28.632, [R.sup.2] = 0.412, P< 0.05) showing that bird richness differ significantly between the four analyzed habitats (Fig. 4). Similarly, with migratory status, the highest species richness of resident and migratory species was recorded in mature tropical forest (69 resident species and 26 migratory species) and tropical forest remnants (62 resident species and 35 migratory species) while modified environments by tourism developments (45 resident species, 26 migratory species, and one introduced species) and modified environments by urban developments, crops and livestock (30 resident species, eight migratory species, and two introduced species) recorded the lowest species richness; but not significant difference depending on migratory status, both migratory and resident species respond the same way. Insectivore species were better represented in the mature tropical forest, and tropical forest remnants (>16). Frugivores and nectarivores species were slightly higher and abundant in mature tropical forest and, tropical forest remnants (with six species in each habitat). Carnivores (18), granivores (10) and omnivores (10) species were better represented in modified environments (particularly in cattle pastures and agricultural fields), and insectivore species were better represented in mature tropical forest (65 species) and, tropical forest remnants (59; Appendix). Results in this study are consistent with respect to that modified environments (i.e., agricultural and livestock areas) had a higher proportion of carnivores and granivores species in comparison with tropical forest.

DISCUSSION

Our study revealed that mature and tropical forest remnants in Akumal region had higher bird species diversity that the modified environments, which is expected because modified environments lack suitable vegetative remnants, shrubs and canopy cover that limits food density and diversity, nest placement, and predator avoidance. Above mentioned reveals the importance of tropical forest remnants for bird diversity conservation in a tourism area, as an important shelters to the bird community. According with our results, Bennet and Saunders (2010) mentioned that the forest remnants are very important in terms of shelter, feeding and nesting areas, particularly to birds that depend on native vegetation.

Bird species recorded accounted for 32 % of all species reported for Quintana Roo by Correa and MacKinnon (2011), being the order Passeriformes the most representatives with 52 % (83 species) from the total recorded. Abundance index values (PAI) showed a large number of species with low PAI, as well as few species with intermediate to high PAI compared to the pattern observed in other surveys (Aleixo & Vielliard, 1995; Lyra-Neves, Martins, Mendes, Rodrigues, & Lacerda, 2004). Bird species richness in the study was similar to other tropical forest areas with a predominance of insectivore species (e.g., Estrada, Coates-Estrada, & Meritt, 1997; Blake & Loiselle, 2001). Omnivore species abundance can be directly related to the variety of available resources for change in land use and declining native resources like fruits. However, the presence of frugivore species, also some bark insectivore species indicate that the study area is relatively well conserved (Blake & Loiselle, 2015). Others signs of relative adequate habitat conditions included the occurrence of mixed-species flocks (Stotz, Fitzpatrick, Parker, & Moskovits, 1996), and trunk insectivores. Frequency and structure of mixed-species flocks also suggests habitat conditions at the study area were adequate for many common in tropical forest bird species according to Stotz et al. (1996). Most bird species recorded in this study were dependent on forest edge, these results suggest that the sensitivities of bird species to vegetation are associated with their dependence of food resources as availability of native fruit (Hasiu, Gomes, & Silva, 2007).

The differences in the species richness and diversity found in this study indicated that the mature tropical forest and tropical forest remnants present greater diversity and richness compared with modified environments. This accords with other studies in tropical environments, and indicates that the loss of original habitats directly influences the presence, abundance and persistence of species (Kattan, Alvarez-Lopez, & Giraldo, 1994; Laurance & Bierregaard, 1997; Rocha, Virtanen, & Cabeza, 2015). The higher avian diversity found in tropical forest may be due to high numbers of individuals and mature vegetation that provide many different microhabitats, which promote varieties of bird species compared with habitats with different land covers (e.g., with human infrastructure or tourism development). However, others studies have found highest richness in modified environments than natural environments (Petit, Petit, Christian, & Powell, 1999; Martin, Viano, Ratsimisetra, Laloe, & Carriere, 2012), but this may be due to the environmental heterogeneity that can get to present the area.

Tropical forest remnants had a significant contribution to the bird species richness and diversity in the study area which is consistent with those reported by Estrada et al. (1997) in Los Tuxtlas region in Veracruz, Mexico. On the other hand, bird composition in terms of the feeding guilds is related to vegetation structure (Laurance & Bierregaard, 1997). Different groups of bird species were found that respond differently to the conversion of forest to modified environments. Not surprisingly, tropical forest assemblages were characterized by a high proportion of forest-associated species, whereas modified habitats were dominated by generalists and open habitat specialists. However, modified environments by urban developments, crops and livestock are very important to a lot of carnivores, granivores and insectivores species because of temporarily or permanently provide such resources depending on their phenology and seasonality (Loiselle & Blake, 1994).

In general, the tropical forest remnants that presents the study area appears to contribute to the relatively high species richness, especially considering the number of species occurring in mature tropical forest. Results of this study showed evidence that tropical forest remnants are significantly important in tourism zones as an available habitat for birds. The continuing expansion of tourism complex, particularly large-scale, will likely result in the simplification and loss of bird diversity. That is particularly important in tourism zones from Quintana Roo because these remnants representing shelters, feeding or nesting areas for birds dependent from natural environments; as well as responsible for maintaining an important proportion of regional avian diversity. The importance of tropical forest remnants provides important habitats for many species of resident and migrant birds in Yucatan Peninsula. Our results confirm the great need for conservation (preserved areas), restoration with native vegetation, and ecological studies of tropical forests remnants, because represent the first step to take actions for conservation of regional avian diversity in the Yucatan Peninsula subjected to anthropogenic activities. An added potential value to this tourist area to attract other tourism type (as birdwatchers) as an alternative to preserve and promote ecological tourism. Furthermore, create incentives for protection and preservation on natural areas and, native biota, which allow preserve these tropical forest remnants.

ACKNOWLEDGMENTS

Thanks to C. Vazquez and L. del Villar for the help and support in the field work. To Akumal municipally authorities, Villas Jade Beach, Golf and Spa, Naj K'aax Residential, Bahia Principe Residences and Golf, Bahia Principe Hotels and Resorts properties for the facilities during the field work.

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APPENDIX

Bird species recorded were classified according to their migratory
status and feeding habits in each habitat type in Akumal region,
Quintana Roo, Mexico

Species                       Migratory status     Feeding       PAI
                                                   habits *

Dendrocygna autumnalis            Resident            I        0.9856
Anas discors                   Winter visitor         I        0.6312
Nomonyx dominicus                 Resident            I        0.9856
Ortalis vetula                    Resident            Fr       1.8874
Phoenicopterus ruber              Resident            I        0.1322
Podilymbus podiceps            Winter visitor         I        0.6312
Columba livia                    Introduced           Om       2.2480
Patagioenas flavirostris          Resident            Fr       0.8523
Streptopelia decaocto            Introduced           Om       0.8523
Columbina passerina               Resident            Gr       1.2340
Zenaida asiatica                  Resident            Gr       1.2340
Zenaida aurita                    Resident            Gr       1.2003
Coccyzus minor                    Resident          Fr, I      0.0045
Chordeiles acutipennis            Resident            I        0.0987
Nyctidromus albicollis            Resident            I        0.0846
Chaetura vauxi                    Resident            I        1.0084
Anthracothorax prevostii          Resident            Ne       0.0012
Archilochus colubris           Winter visitor         Ne       0.0458
Chlorostilbon canivetii           Resident            Ne       0.0683
Amazilia yucatanensis             Resident            Ne       0.0879
Amazilia rutila                   Resident            Ne       0.0879
Gallinula chloropus            Winter visitor         I        0.0875
Fulica americana               Winter visitor         I        0.0875
Himantopus mexicanus             Transient            I        0.0987
Pluvialis squatorola           Winter visitor         I        0.0784
Pluvialis dominica               Transient            I        0.0012
Charadrius semipalmatus        Winter visitor         I        0.5489
Charadrius vociferus           Winter visitor         I        0.6231
Jacana spinosa                    Resident            I        0.0023
Actitis macularius             Winter visitor         I        0.0987
Tringa solitaria               Winter visitor         I        0.0846
Arenaria interpres             Winter visitor         I        0.0846
Calidris minutilla             Winter visitor         I        0.0846
Calidris pusilla                 Transient            I        0.0784
Leucophaeus atricilla          Winter visitor         Ca       0.0784
Hydroprogne caspia             Winter visitor         Ca       0.0458
Chlidonias niger                 Transient          I, Ca      0.0030
Thalasseus elegans             Winter visitor         Ca       0.0458
Thalasseus maximus             Winter visitor         Ca       0.0458
Fregata magnificens               Resident            Ca       0.6307
Sula leucogaster                  Resident            Ca       0.5543
Phalacrocorax brasilianus         Resident            Ca       0.9936
Anhinga anhinga                   Resident            Ca       0.9701
Pelecanus occidentalis            Resident            Ca       0.6111
Ardea herodias                 Winter visitor         Ca       0.0224
Ardea alba                        Resident            Ca       0.0458
Egretta thula                     Resident          I, Ca      0.1322
Egretta caerulea               Winter visitor       I, Ca      0.1322
Egretta tricolor               Winter visitor       I, Ca      0.0112
Bubulcus ibis                     Resident            I        0.7789
Butorides virescens               Resident          I, Ca      0.0112
Euducimus albus                   Resident            I        0.6803
Coragyps atratus                  Resident            Ca       1.9635
Cathartes aura                    Resident            Ca       1.9648
Pandion haliaetus              Winter visitor         Ca       0.0112
Buteogallus anthracinus           Resident            Ca       0.0157
Rupornis magnirostris             Resident            Ca       0.1002
Buteo nitidus                     Resident            Ca       0.0875
Tyto alba                         Resident            Ca       0.0045
Megascops guatemalae              Resident            Ca       0.0012
Glaucidium brasilianum            Resident            Ca       0.0012
Trogon melanocephalus             Resident            Fr       0.0085
Trogon caligatus                  Resident            Fr       0.0088
Momotus coeruliceps               Resident            Om       0.0654
Eumomota superciliosa             Resident            Om       0.0879
Megaceryle alcyon              Winter visitor         Ca       0.0085
Chloroceryle americana            Resident            Ca       0.0084
Melanerpes pygmaeus               Resident            I        0.0879
Melanerpes aurifrons              Resident            I        0.6321
Picoides scalaris                 Resident            I        0.6004
Campephilus guatemalensis         Resident            I        0.0081
Herpetotheres cachinans           Resident            Ca       0.0879
Falco sparverius               Winter visitor         Ca       0.0701
Falco columbarius              Winter visitor         Ca       0.0556
Eupsittula nana                   Resident            Fr       0.7540
Amazona xantholora                Resident            Fr       0.6412
Sittasomus griseicapillus         Resident            I        0.0683
Xiphorhynchus flavigaster         Resident            I        0.0023
Synallaxis erythrothorax          Resident            I        0.0023
Camptostoma imberbe               Resident            I        0.0023
Myiopagis viridicata              Resident            I        0.0245
Elaenia flavogaster               Resident            I        0.0023
Oncostoma cinereigulare           Resident            I        0.0245
Todirostrum cinereum              Resident            I        0.0023
Rhynchocyclus brevirostris        Resident            I        0.0023
Contopus virens                  Transient            I        0.0245
Contopus cinereus                 Resident            I        0.0335
Attila spadiceus                  Resident            I        0.0335
Myiarchus yucatanensis            Resident            I        0.0335
Myiarchus tuberculifer            Resident            I        0.0278
Myiarchus tyrannulus              Resident            I        0.1150
Pitangus sulphuratus              Resident            Om       0.2369
Myiozetetes similis               Resident            I        0.4481
Myiodynastes luteiventris         Resident            I        0.1150
Tyrannus melancholicus            Resident            I        0.2369
Tyrannus couchii                  Resident            I        0.2369
Tyrannus tyrannus                Transient            I        0.1150
Tityra semifasciata               Resident          Fr, I      0.2369
Pachyramphus aglaiae              Resident            I        0.1123
Vireo pallens                     Resident            I        0.2369
Vireo philadelphicus           Winter visitor         I        0.2369
Vireo magister                    Resident            I        0.1123
Psilorhinus morio                 Resident            Om       1.0523
Cyanocorax yucatanicus            Resident            Om       1.1238
Stelgidopteryx serripennis     Winter visitor         I        1.0523
Riparia riparia                  Transient            I        1.0035
Petrochelidon fulva               Resident            I        1.0523
Hirundo rustica                  Transient            I        1.0035
Thryothorus maculipectus          Resident            I        0.1123
Thryothorus ludovicianus          Resident            I        0.1122
Uropsila leucogastra              Resident            I        0.2369
Polioptila caerulea               Resident            I        0.2035
Catharus ustulatus               Transient            I        0.1122
Turdus grayi                      Resident            Om       0.2369
Hylocichla mustelina           Winter visitor         I        0.1122
Dumetella carolinensis            Resident            I        0.2369
Melanoptila glabirostris          Resident            I        0.1123
Mimus gilvus                      Resident          Fr, I      0.2369
Arremonops rufivirgatus           Resident            Gr       0.1123
Euphonia hirundinacea             Resident            Fr       0.2369
Dives dives                       Resident            Om       0.9856
Quiscalus mexicanus               Resident            Om       3.4522
Molothrus aeneus                  Resident            Gr       0.9856
Icterus prosthemelas              Resident            I        0.2369
Icterus cucullatus                Resident            Om       0.2568
Icterus chrysater                 Resident            I        0.1148
Icterus auratus                   Resident            I        0.1148
Icterus galbula                Winter visitor         Om       0.1148
Seiurus aurocapilla            Winter visitor         I        0.1123
Helmitheros vermivorum         Winter visitor         I        0.1123
Parkesia noveboracensis        Winter visitor         I        1.0035
Mniotilta varia                Winter visitor         I        0.1123
Protonotaria citrea              Transient            I        0.0041
Oreothlypis peregrina            Transient            I        0.0041
Oreothlypis ruficapilla        Winter visitor         I        0.1148
Geothlypis poliocephala           Resident            I        0.0245
Geothlypis trichas             Winter visitor         I        0.0041
Setophaga citrina              Winter visitor         I        0.0041
Setophaga ruticilla            Winter visitor         I        0.0041
Setophaga americana            Winter visitor         I        0.0041
Setophaga magnolia             Winter visitor         I        0.1123
Setophaga petechia             Winter visitor         I        0.1148
Setophaga caerulescens         Winter visitor         I        0.0041
Setophaga virens               Winter visitor         I        0.0041
Cardellina canadensis          Winter visitor         I        0.0041
Cardellina pusilla             Winter visitor         I        0.0245
Icteria virens                 Winter visitor         I        0.0041
Thraupis abbas                    Resident          Fr, I      0.1148
Piranga roseogularis              Resident            I        0.2035
Piranga rubra                  Winter visitor         I        0.2369
Cardinalis cardinalis             Resident            Gr       0.1148
Pheucticus ludovicianus        Winter visitor       I, Gr      0.2035
Cyanocompsa parellina             Resident            Gr       0.2369
Passerina caerulea             Winter visitor         Gr       0.2568
Passerina cyanea               Winter visitor         Gr       0.1148
Volatinia jacarina                Resident            Gr       0.1123
Cyanerpes cyaneus                 Resident            Ne       0.1123
Sporophila torqueola              Resident            Gr       0.1148
Saltator atriceps                 Resident            Gr       0.2035
Saltator coerulescens             Resident            Gr       0.2369

Species                          Habitat use preferences **

Dendrocygna autumnalis                     Td (Gc)
Anas discors                               Td (Gc)
Nomonyx dominicus                          Td (Gc)
Ortalis vetula                          Td (Gc), Cbm
Phoenicopterus ruber                         Cbm
Podilymbus podiceps                     Td (Gc), Cbm
Columba livia                              Ca (Us)
Patagioenas flavirostris                     Mt
Streptopelia decaocto                 Td (Rh), Ca (Us)
Columbina passerina                     Tfr, Td (Rh)
Zenaida asiatica              Mt, Tfr, Td (Gc, Rh), Ca (Cp, Us)
Zenaida aurita                      Td (Rh), Ca (Cp, Us)
Coccyzus minor                             Mt, Tfr
Chordeiles acutipennis                Mt, Tfr, Td (Gc)
Nyctidromus albicollis                       Mt
Chaetura vauxi                Mt, Tfr, Td (Gc, Rh), Ca (Cp, Us)
Anthracothorax prevostii                     Mt
Archilochus colubris                       Mt, Tfr
Chlorostilbon canivetii                    Mt, Tfr
Amazilia yucatanensis                      Mt, Tfr
Amazilia rutila                   Mt, Tfr, Td (Rh), Ca (Us)
Gallinula chloropus                     Td (Gc), Cbm
Fulica americana                        Td (Gc), Cbm
Himantopus mexicanus                       Td (Gc)
Pluvialis squatorola                         Cbm
Pluvialis dominica                      Td (Gc), Cbm
Charadrius semipalmatus                      Cbm
Charadrius vociferus                    Td (Gc), Cbm
Jacana spinosa                          Td (Gc), Cbm
Actitis macularius                      Td (Gc), Cbm
Tringa solitaria                        Td (Gc), Cbm
Arenaria interpres                           Cbm
Calidris minutilla                           Cbm
Calidris pusilla                             Cbm
Leucophaeus atricilla                        Cbm
Hydroprogne caspia                           Cbm
Chlidonias niger                             Cbm
Thalasseus elegans                           Cbm
Thalasseus maximus                           Cbm
Fregata magnificens                     Td (Gc), Cbm
Sula leucogaster                        Td (Gc), Cbm
Phalacrocorax brasilianus               Td (Gc), Cbm
Anhinga anhinga                         Td (Gc), Cbm
Pelecanus occidentalis                  Td (Gc), Cbm
Ardea herodias                          Td (Gc), Cbm
Ardea alba                              Td (Gc), Cbm
Egretta thula                           Td (Gc), Cbm
Egretta caerulea                        Td (Gc), Cbm
Egretta tricolor                        Td (Gc), Cbm
Bubulcus ibis                              Ca (Cp)
Butorides virescens                     Td (Gc), Cbm
Euducimus albus                         Td (Gc), Cbm
Coragyps atratus              Mt, Tfr, Td (Gc, Rh), Ca (Cp, Us)
Cathartes aura                Mt, Tfr, Td (Gc, Rh), Ca (Cp, Us)
Pandion haliaetus                            Cbm
Buteogallus anthracinus                   Tfr, Cbm
Rupornis magnirostris               Mt, Tfr, Ca (Cp, Us)
Buteo nitidus                         Mt, Tfr, Ca (Cp)
Tyto alba                           Td (Rh), Ca (Cp, Us)
Megascops guatemalae                       Mt, Tfr
Glaucidium brasilianum                       Mt
Trogon melanocephalus                      Mt, Tfr
Trogon caligatus                             Mt,
Momotus coeruliceps                          Mt
Eumomota superciliosa                      Mt, Tfr
Megaceryle alcyon                       Td (Gc), Cbm
Chloroceryle americana                  Td (Gc), Cbm
Melanerpes pygmaeus                        Mt, Tfr
Melanerpes aurifrons              Mt, Tfr, Td (Rh), Ca (Us)
Picoides scalaris                   Tfr, Td (Rh), Ca (Us)
Campephilus guatemalensis                  Mt, Tfr
Herpetotheres cachinans                 Tfr, Ca (Cp)
Falco sparverius                        Tfr, Ca (Cp)
Falco columbarius                 Mt, Tfr, Td (Gc), Ca (Cp)
Eupsittula nana                 Mt, Tfr, Td (Rh), Ca (Cp, Us)
Amazona xantholora                           Mt
Sittasomus griseicapillus                    Mt
Xiphorhynchus flavigaster                    Mt
Synallaxis erythrothorax                     Mt
Camptostoma imberbe                        Mt, Tfr
Myiopagis viridicata                       Mt, Tfr
Elaenia flavogaster                        Mt, Tfr
Oncostoma cinereigulare                      Mt
Todirostrum cinereum                         Mt
Rhynchocyclus brevirostris                   Mt
Contopus virens                            Mt, Tfr
Contopus cinereus                 Mt, Tfr, Td (Rh), Ca (Cp)
Attila spadiceus                           Mt, Tfr
Myiarchus yucatanensis                       Tfr
Myiarchus tuberculifer          Mt, Tfr, Td (Rh), Ca (Cp, Us)
Myiarchus tyrannulus            Mt, Tfr, Td (Rh), Ca (Cp, Us)
Pitangus sulphuratus            Mt, Tfr, Td (Rh, Gc), Ca (Us)
Myiozetetes similis               Mt, Tfr, Td (Rh), Ca (Us)
Myiodynastes luteiventris                  Mt, Tfr
Tyrannus melancholicus          Tfr, Td (Gc, Rh), Ca (Cp, Us)
Tyrannus couchii                  Mt, Tfr, Td (Gc), Ca (Cp)
Tyrannus tyrannus                 Mt, Tfr, Td (Rh), Ca (Cp)
Tityra semifasciata                        Mt, Tfr
Pachyramphus aglaiae                       Mt, Tfr
Vireo pallens                                Mt
Vireo philadelphicus                       Mt, Tfr
Vireo magister                               Mt
Psilorhinus morio                 Mt, Tfr, Td (Rh), Ca(Cp)
Cyanocorax yucatanicus                Mt, Tfr, Td (Rh)
Stelgidopteryx serripennis    Mt, Tfr, Td (Gc, Rh), Ca (Cp, Us)
Riparia riparia                         Td (Gc), Cbm
Petrochelidon fulva                 Td (Gc), Cbm, Ca (Cp)
Hirundo rustica                   Td (Rh, Gc), Ca (Cp, Us)
Thryothorus maculipectus                   Mt, Tfr
Thryothorus ludovicianus                   Mt, Tfr
Uropsila leucogastra                       Mt, Tfr
Polioptila caerulea               Mt, Tfr, Td (Rh), Ca (Us)
Catharus ustulatus                         Mt, Tfr
Turdus grayi                          Mt, Tfr, Td (Rh)
Hylocichla mustelina                       Mt, Tfr
Dumetella carolinensis                     Mt, Tfr
Melanoptila glabirostris                   Mt, Tfr
Mimus gilvus                      Mt, Tfr, Td (Rh), Ca (Us)
Arremonops rufivirgatus                    Mt, Tfr
Euphonia hirundinacea                        Mt
Dives dives                     Mt, Tfr, Td (Gc, Rh), Ca (Us)
Quiscalus mexicanus                 Td (Gc, Rh), Ca (Us)
Molothrus aeneus                    Td (Gc, Rh), Ca (Us)
Icterus prosthemelas                       Mt, Tfr
Icterus cucullatus                    Mt, Tfr, Td (Rh)
Icterus chrysater                          Mt, Tfr
Icterus auratus                            Mt, Tfr
Icterus galbula                         Tfr, Td (Rh)
Seiurus aurocapilla                        Mt, Tfr
Helmitheros vermivorum                     Mt, Tfr
Parkesia noveboracensis                    Mt, Tfr
Mniotilta varia                       Mt, Tfr, Td (Rh)
Protonotaria citrea                        Mt, Tfr
Oreothlypis peregrina                      Mt, Tfr
Oreothlypis ruficapilla               Mt, Tfr, Td (Rh)
Geothlypis poliocephala                    Mt, Tfr
Geothlypis trichas                           Tfr
Setophaga citrina                          Mt, Tfr
Setophaga ruticilla                        Mt, Tfr
Setophaga americana                        Mt, Tfr
Setophaga magnolia                         Mt, Tfr
Setophaga petechia                    Mt, Tfr, Td (Rh)
Setophaga caerulescens                     Mt, Tfr
Setophaga virens                           Mt, Tfr
Cardellina canadensis                      Mt, Tfr
Cardellina pusilla                  Tfr, Td (Rh), Ca (Us)
Icteria virens                        Mt, Tfr, Td (Rh)
Thraupis abbas                             Mt, Tfr
Piranga roseogularis                         Mt
Piranga rubra                              Mt, Tfr
Cardinalis cardinalis                        Tfr
Pheucticus ludovicianus               Mt, Tfr, Td (Rh)
Cyanocompsa parellina                 Mt, Tfr, Td (Rh)
Passerina caerulea                Tfr, Td (Rh), Ca (Us, Cp)
Passerina cyanea                        Tfr, Ca (Cp)
Volatinia jacarina                      Tfr, Ca (Cp)
Cyanerpes cyaneus                          Mt, Tfr
Sporophila torqueola                    Tfr, Ca (Cp)
Saltator atriceps                          Mt, Tfr
Saltator coerulescens                      Mt, Tfr

* Feeding habits: Omnivores (Om); Nectarivores (Ne); Carnivores (Ca);
Frugivores (Fr); Granivores (Gr); Invertebrates (I, included aquatic
invertebrates, bark insectivores aerial insectivores, trunk
insectivores, generalist insectivores, ground insectivores, and leaf
insectivores.

** Habitat use preferences: Mature tropical forest (Mt); Tropical
forest remnants (Tfr), Modified environments by tourism developments
(Td): Golf course and artificial water bodies in golf course (Gc),
Hotel and residential zones with native and introduced vegetation
(Rh); Modified environments by urban development, crops and livestock
(Ca): Urban zone with native and introduced vegetation (Us), Cattle
pastures and agricultural fields (Cp); and, Coast dunes, beach and
mangrove zones (Cbm).


Jorge E. Ramirez-Albores (1,2) & Marlin Perez-Suarez (1)*

(1.) Instituto de Ciencias Agropecuarias y Rurales (ICAR), Universidad Autonoma del Estado de Mexico (UAEMex). Carr. El Cerrillo-Piedras Blancas s/n. C.P. 50090, Toluca de Lerdo, Estado de Mexico, Mexico; mperezs@uaem.mx*

(2.) Museo de Zoologia "Alfonso L. Herrera", Departamento de Biologia Evolutiva, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico. Apdo. Postal 70-399. Ciudad de Mexico, C.P. 04510, Mexico; jorgeramirez22@hotmail.com

Received 24-XI-2017. Corrected 08-II-2018. Accepted 07-III-2018.

Caption: Fig. 1. Map show the location of study area in Yucatan Peninsula, Mexico. The location of study area is found in the eastern Yucatan Peninsula, and it's delimited by a square that it's shown in detail in the right panel.

Caption: Fig. 2. Species accumulation curve for birds sampled by census in the study area. Observed species richness (a) and expected (b) in the study area and habitat type: Mature tropical forest (Mt), Tropical forest remnants (Tfr), Modified environments by tourism developments (Td), and Modified environments by urban developments, crops and livestock (Ca).

Caption: Fig. 3. Boxplots of mean species richness (a), mean abundance (b), (c) Shannon diversity index, and (d) Simpson's diversity index of bird species at Akumal region, Quintana Roo, Mexico: Mature tropical forest (Mt), Tropical forest remnants (Tfr), Modified environments by tourism developments (Td), and Modified environments by urban developments, crops and livestock (Ca). Lines represent minimum, first quartile, median, third quartile, and maximum.

Caption: Fig. 4. Ordination plot of HMDS axes showing gradients in bird composition in mature tropical forest ([??]), tropical forest remnants ([??]), modified environments by tourism developments ([??]) and modified environments by urban developments, crops and livestock ([??]).
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Author:Ramirez-Albores, Jorge E.; Perez-Suarez, Marlin
Publication:Revista de Biologia Tropical
Date:Jun 1, 2018
Words:6950
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