The Thicket Biome is found from the west coast to KwaZulu-Natal, with most of the biome being found in the Eastern Cape. It makes up 2.5% of the area of South Africa (nearly 31 500 km2) and the vegetation ranges from shrubland to low forest. There are many evergreen and succulent trees and shrubs and many of the plants have spines or thorns. Analysis of the spider assemblages did, however, suggest that this biome overlaps considerably with other biome types, and could potentially be grouped together with the fauna of the Savanna Biome. No published surveys are known from this biome but several surveys are presently underway. To date 55 families represented by 206 genera and 464 spp. have been recorded from thicket, of which 90 are endemics and 96 near endemics. The Thomisidae with 62 spp. are the most diverse, followed by the Salticidae (38) and Araneidae (37).

#### **2.7 Agro-ecosystems**

South Africa has about 12% arable land and some of these areas were sampled to determine the spider diversity. Spiders are one of the most ubiquitous predator groups in agroecosystems (Van den Berg & Dippenaar-Schoeman, 1991a) and inventories in South Africa have provided valuable baseline information on species in agroecosystems (Table 2). As predators, spiders have a two-fold function. Not only do they feed directly on their prey, but their presence also causes indirect mortality of arthropods. The presence of spiders could disturb larvae who then drop from the plant and die. The webs spun over the surface of leaves also seems to make them less suitable for oviposition and feeding. Therefore, while considerable effort has been put into baseline surveys in agroecosystems in South Africa, there is a large scope for further experimental work on the biological control potential of the dominant agobiont spiders in each agroecosystem.

The family Salticidae has been shown to be an important component of agrobiont assemblages. In a knock-down study of three macadamia orchards in the Mpumalanga Lowveld in South Africa, 73% of the sampled spiders were salticids (jumping spiders). Salticids were also the most diverse (17 species), of which four species represented more than 61% of all spiders collected. *Thyene coccineovittata* was the most abundant species and represented 30% of all the spiders collected, followed by *T. natali* with 14%, *Viciria alba* with 9% and *Tusitala guineensis* with 8%. These four species were present throughout the year in all three orchards sampled (Dippenaar-Schoeman et al., 2001a, b).

habitats, and likely shows high levels of endemism. These biomes are thus of considerable interest biogeographically, phylogenetically and from a conservation perspective. Jenkins et al. (2011) also point to the importance of these arid areas in developing our understanding

Due to the low vegetation, fewer plant- and web-dwelling spiders are recorded, but a rich fauna of ground dwellers is present. There are a number of endemics found in ground dwelling families such as the Ammoxenidae (12 spp.) and several trapdoor spider families: Cyrtaucheniidae (8); Ctenizidae (13); Idiopidae (7) en Nemesiidae (11) (Dippenaar-Schoeman, 2002). More than 100 species of Gnaphosidae alone are known from these biomes, most of which were described by Tucker (1923) in his revision of the family in South Africa, with several endemics. Other ground dwelling spiders such as the genus *Tyrotama* (4 species) of the Hersiliidae (Foord & Dippenaar-Schoeman, 2005) and ground-dwelling Eresidae (Dippenaar-

The Thicket Biome is found from the west coast to KwaZulu-Natal, with most of the biome being found in the Eastern Cape. It makes up 2.5% of the area of South Africa (nearly 31 500 km2) and the vegetation ranges from shrubland to low forest. There are many evergreen and succulent trees and shrubs and many of the plants have spines or thorns. Analysis of the spider assemblages did, however, suggest that this biome overlaps considerably with other biome types, and could potentially be grouped together with the fauna of the Savanna Biome. No published surveys are known from this biome but several surveys are presently underway. To date 55 families represented by 206 genera and 464 spp. have been recorded from thicket, of which 90 are endemics and 96 near endemics. The Thomisidae with 62 spp.

South Africa has about 12% arable land and some of these areas were sampled to determine the spider diversity. Spiders are one of the most ubiquitous predator groups in agroecosystems (Van den Berg & Dippenaar-Schoeman, 1991a) and inventories in South Africa have provided valuable baseline information on species in agroecosystems (Table 2). As predators, spiders have a two-fold function. Not only do they feed directly on their prey, but their presence also causes indirect mortality of arthropods. The presence of spiders could disturb larvae who then drop from the plant and die. The webs spun over the surface of leaves also seems to make them less suitable for oviposition and feeding. Therefore, while considerable effort has been put into baseline surveys in agroecosystems in South Africa, there is a large scope for further experimental work on the biological control potential of the

The family Salticidae has been shown to be an important component of agrobiont assemblages. In a knock-down study of three macadamia orchards in the Mpumalanga Lowveld in South Africa, 73% of the sampled spiders were salticids (jumping spiders). Salticids were also the most diverse (17 species), of which four species represented more than 61% of all spiders collected. *Thyene coccineovittata* was the most abundant species and represented 30% of all the spiders collected, followed by *T. natali* with 14%, *Viciria alba* with 9% and *Tusitala guineensis* with 8%. These four species were present throughout the year in

Schoeman, 1989) have several species that are endemic to these arid biomes.

are the most diverse, followed by the Salticidae (38) and Araneidae (37).

of the effects of global climate change.

**2.6 Thicket, between a rock and a hard place**

dominant agobiont spiders in each agroecosystem.

all three orchards sampled (Dippenaar-Schoeman et al., 2001a, b).

**2.7 Agro-ecosystems**

Salticidae were also the dominant spider family in pistachio orchard canopies in the arid Northern Cape, although their abundance varied between the three orchards sampled: Green Valley Nuts 1 (81.2%), Remhoogte (58.8%) and Green Valley Nuts 19 (82.4%). It seems that factors such as tree canopy size, presence or absence of bark and dry leaves, climatic conditions, and orchard size may be the main influences on spider abundance (Haddad et al., 2005). The fauna of ground covers was dominated by Salticidae in Green Valley Nuts 1 (37.2%) and Remhoogte (35.0%), while Oxyopidae dominated the ground cover fauna of the youngest orchard, Green Valley Nuts 19 (31.9%) (Haddad et al., 2004a). Four families, Linyphiidae, Lycosidae, Salticidae and Gnaphosidae, dominated the ground-dwelling fauna of the orchards, although the abundance of each differed considerably between sampling methods (pitfall trapping and active searching) and also between orchards (Haddad & Dippenaar-Schoeman, 2006b).

Table 2. Crops in South Africa that have been surveyed for spiders and for which agrobiont spider species have been identified.

Preliminary data of spider prey items included Thysanoptera (Phlaeothripidae), Coleoptera (Bruchidae, Coccinellidae, Chrysomelidae), Diptera (Tephritidae, Muscidae, Cecidomyiidae), Acari (Tetranychidae), and various Lepidoptera larvae. Laboratory and field experiments on *Heliophanus pistaciae* (Salticidae), the dominant arboreal agrobiont species, indicated that it has potential as a biological control agent of the false cinch bug, *Nisius natalensis* (Hemiptera: Lygaeidae), one of the minor pests of pistachio nuts in South Africa (Haddad et al., 2004b).

South African Spider Diversity:

**4. Acknowledgments**

**5. References**

262.

*Lett* 8: 148-159.

*Ecol* 78: 573-584.

443-447.

24: 371-406.

African Perspectives on the Conservation of a Mega-Diverse Group 177

complemented by targeted studies at regional and local scales over larger temporal scales if

The authors would like to thank the Agricultural Research Council (ARC) and the South African National Biodiversity Institute (SANBI) for funding SANSA. Several volunteers and taxonomic specialists contributed to both SANSA and the National Collection of Arachnida, and their enthusiasm and assistance greatly contributed to the success of the project. We

Botes, A, McGeoch, MA, Robertson, HG, Van Niekerk, A, Davids, HP, Chown, SL. (2006).

Bursey, ML, Herbert, DG. (2004). Four new narrow-range endemic species of *Gulella* from

Butler, VP, Haddad, CR. (in press). Spider communities associated with leaf litter of three tree species in central South Africa (Arachnida: Araneae). *Afr J Ecol*.

Cardoso, P, Scharff, N, Gaspar, CS, Henriques, SS, Carvalho, R, Castro, PH, Schmidt, JB,

Chao, A, Chazdon, RL, Colwell, RK, Shen, T-J. (2005). A new statistical approach for

Coddington, JA, Agnarsson, I, Miller, JA, Kuntner, M, Hormiga, G. (2009). Undersampling

Coetzee, JH, Dippenaar-Schoeman, AS, Van den Berg, A. (1990). Spider assemblages on five

Cole, ML, Herbert, DG. (2009). Decription of four new species of *Gulella* Pfeiffer, 1856 from

De Bivort, BL, Giribet, G. (2010). A systematic revision of South African Petallidae

Dippenaar-Schoeman, AS. (1979). Spider communities in strawberry beds: seasonal changes

Dippenaar-Schoeman, AS. (1988). Annotated check list of the spiders (Araneae) of the

(Mollusca, Eupulmonata: Streptaxidae). *Zool Meded (Leiden)* 83: 547-564. Daniels, SR, Picker, MD, Cowlin, RM, Hamer, ML. (2009). Unraveling evolutionary lineages

for widespread cryptic speciation. *Biol J Linn Soc* 97: 200-216.

in numbers and species composition. *Phytophylatica* 11: 1-4.

Mountain Zebra National Park. *Koedoe* 31: 151-160.

Ants, altitude and change in the northern Cape Floristic Region. *J Biogeogr* 33: 71-90.

Eastern Cape, South Africa (Mollusca: Pulmonata: Streptaxidae). *Afr Invert* 45: 249-

Silva, I, Szuts, T, De Castro, A, Crespo, LC. (2008). Rapid biodiversity assessment of spiders (Araneae) using semi-quantitative sampling: a case study in a

assessing compositional similarity based on incidence and abundance data. *Ecol*

bias: the null hypothesis for singleton species in tropical arthropod surveys. *J Anim*

species of proteaceous plants in the fynbos biome of South Africa. *Phytophylatica* 22:

Eastern Cape, South Africa, with additional notes on two poorly known species

among South African velvet worms (Onychophora: *Peripatopsis*) provides evidence

(Arachnida: Opiliones: Cyphophthalmi) based on a combined analysis of discrete and continuous characters with the description of seven new species. *Invertebr Syst*

we are to fully understand the dynamics of spider assemblages in South Africa.

would also like to thank the data management team at the ARC.

Mediterranean forest. *Insect Cons Divers* 1: 71-84.

## **2.8 Public education, participation and awareness (SANSA)**

This constitutes another important component of the CBD and SANSA, aimed to increase awareness about arachnids to wider society through a variety of activities that includes an extensive website3 containing information on the activities of SANSA. Additional information is also available on: 1) all the arachnid orders; 2) survey results; 3) medically important spiders; 4) arachnid research; 5) virtual museum entries; and 6) copies of newsletters and reports.

Target audiences are also identified and information is disseminated in the appropriate communication medium and language through books, CD's, posters, magazine and newspaper articles, pamphlets, TV and radio (Dippenaar-Schoeman & Jocqué, 1997; Dippenaar-Schoeman, 2002; Dippenaar-Schoeman & Van den Berg, 2010; Holm & Dippenaar-Schoeman, 2010). In addition to several radio and television appearances by Dippenaar-Schoeman, the Spider Educare Programme based at the National Collection of Arachnida has conducted road shows to schools, lectures to societies and clubs, and courses presented at Universities for the past ten years.

The online Virtual Museum of South African Arachnids presently houses about 3400 images received from 80 photographers throughout the country. Not only is it a wonderful display of photographs, but it also provides valuable information on species' behaviour, colour patterns and prey. Colour patterns in particular are lost when the spiders are killed and preserved in alcohol. The Virtual Museum has provided the spider images for two full colour books that have recently been published (Dippenaar-Schoeman & Van den Berg, 2010; Holm & Dippenaar-Schoeman, 2010).

## **3. Conclusion**

Resources for surveys will always be limited and funding will have to be directed at answering targeted questions. Three biomes that require particular attention are the Thicket, Nama and Succulent Karoo Biomes. The Fynbos is also of particular significance as it seems to match the better known savanna biome in terms of spider diversity based on preliminary survey results. The hypothesized restricted distributions of species to Afromontane forests have also been rejected because of contemporary discoveries of these taxa in other biomes employing techniques that target specific microhabitats.

Two taxa, the families Theridiidae and Lycosidae, comprise the bulk of what constitutes a taxonomic impediment to a better understanding of diversity patterns in spiders. The family Salticidae is another important family, dominant in all ecosystems, that fortunately has and will continue to receive considerable taxonomic attention. The potential of this family as a surrogate of spider diversity, in conjunction with families such as Thomisidae and Gnaphosidae, could be explored, should the taxonomic impediment of taxa mentioned earlier persist.

We view the future research as a combination of optimized and standardized inventories in the degree squares of the four biomes identified. This will provide both cost efficient and comparable results. These inventories will, however, have to include protocols that target cryptic biotopes. The by-catch of other invertebrate taxa also provides the opportunity to collaborate with specialists on these taxonomic groups generating data on a "shopping basket" of taxa, as has been shown by a study that was done using the ant by-catch of one of the SANSA surveys (Schoeman & Foord, In press). These inventories have to be

3 www.arc.agric.za quick link SANSA

complemented by targeted studies at regional and local scales over larger temporal scales if we are to fully understand the dynamics of spider assemblages in South Africa.

## **4. Acknowledgments**

176 Changing Diversity in Changing Environment

This constitutes another important component of the CBD and SANSA, aimed to increase awareness about arachnids to wider society through a variety of activities that includes an extensive website3 containing information on the activities of SANSA. Additional information is also available on: 1) all the arachnid orders; 2) survey results; 3) medically important spiders; 4) arachnid research; 5) virtual museum entries; and 6) copies of

Target audiences are also identified and information is disseminated in the appropriate communication medium and language through books, CD's, posters, magazine and newspaper articles, pamphlets, TV and radio (Dippenaar-Schoeman & Jocqué, 1997; Dippenaar-Schoeman, 2002; Dippenaar-Schoeman & Van den Berg, 2010; Holm & Dippenaar-Schoeman, 2010). In addition to several radio and television appearances by Dippenaar-Schoeman, the Spider Educare Programme based at the National Collection of Arachnida has conducted road shows to schools, lectures to societies and clubs, and courses

The online Virtual Museum of South African Arachnids presently houses about 3400 images received from 80 photographers throughout the country. Not only is it a wonderful display of photographs, but it also provides valuable information on species' behaviour, colour patterns and prey. Colour patterns in particular are lost when the spiders are killed and preserved in alcohol. The Virtual Museum has provided the spider images for two full colour books that have recently been published (Dippenaar-Schoeman & Van den Berg,

Resources for surveys will always be limited and funding will have to be directed at answering targeted questions. Three biomes that require particular attention are the Thicket, Nama and Succulent Karoo Biomes. The Fynbos is also of particular significance as it seems to match the better known savanna biome in terms of spider diversity based on preliminary survey results. The hypothesized restricted distributions of species to Afromontane forests have also been rejected because of contemporary discoveries of these taxa in other biomes

Two taxa, the families Theridiidae and Lycosidae, comprise the bulk of what constitutes a taxonomic impediment to a better understanding of diversity patterns in spiders. The family Salticidae is another important family, dominant in all ecosystems, that fortunately has and will continue to receive considerable taxonomic attention. The potential of this family as a surrogate of spider diversity, in conjunction with families such as Thomisidae and Gnaphosidae, could be explored, should the taxonomic impediment of taxa mentioned

We view the future research as a combination of optimized and standardized inventories in the degree squares of the four biomes identified. This will provide both cost efficient and comparable results. These inventories will, however, have to include protocols that target cryptic biotopes. The by-catch of other invertebrate taxa also provides the opportunity to collaborate with specialists on these taxonomic groups generating data on a "shopping basket" of taxa, as has been shown by a study that was done using the ant by-catch of one of the SANSA surveys (Schoeman & Foord, In press). These inventories have to be

**2.8 Public education, participation and awareness (SANSA)**

newsletters and reports.

**3. Conclusion**

earlier persist.

3 www.arc.agric.za quick link SANSA

presented at Universities for the past ten years.

2010; Holm & Dippenaar-Schoeman, 2010).

employing techniques that target specific microhabitats.

The authors would like to thank the Agricultural Research Council (ARC) and the South African National Biodiversity Institute (SANBI) for funding SANSA. Several volunteers and taxonomic specialists contributed to both SANSA and the National Collection of Arachnida, and their enthusiasm and assistance greatly contributed to the success of the project. We would also like to thank the data management team at the ARC.

## **5. References**

South African Spider Diversity:

African Perspectives on the Conservation of a Mega-Diverse Group 179

Dippenaar-Schoeman, AS, Wassenaar, TD. (2006). A checklist of spiders from the

Dippenaar, SM, Dippenaar-Schoeman, AS, Modiba, MA, Khoza, TT (2008). A checklist of

Eeley, HC, Lawes, MJ, Reyers, B. (2001). Priority areas for the conservation of subtropical

Engelbrecht, I. (2010). Invertebrate species inventories in protected area management: are

Foord, SH, Dippenaar-Schoeman, AS. (2005). A revision of the Afrotropical species of

Foord, SH, Dippenaar-Schoeman, AS. (2006). A revision of the Afrotropical species of

Foord, SH, Dippenaar-Schoeman, AS, Van der Merwe, M. (2002). A check list of the spider

Foord, SH, Mafadza, M, Dippenaar-Schoeman, AS, Van Rensburg, BJ. (2008). Micro-scale

Govender, V, 2007. Patterns of distribution, diversity and endemism of terrestrial molluscs

Griswold, CE. (1985). A revision of the African spiders of the family Microstigmatidae

Griswold, CE. (1990). A revision and phylogenetic analysis of the spider subfamily Phyxelidinae (Araneae, Amaurobiidae). *Bull Am Mus Nat Hist* 196: 1-206. Griswold, CE. (1991a). Cladistic biogeography of Afromontane spiders. *Aust Syst Biol* 43: 73-

Griswold, CE. (1991b). A revision and phylogenetic analysis of the spider genus *Machadonia*

Griswold, CE. (1994). A revision and phylogenetic analysis of the spider genus *Phanotea* Simon (Araneae, Lycosoidea). *Ann Musee Roy Afr Centr, Sci Zool* 273: 1-83. Haddad, CR. (2006). *Spinotrachelas*, a new genus of trachelinae sac spiders from South Africa

Haddad, CR. (2010). A new species of *Poachelas* from Maputaland, South Africa (Araneae:

Haddad, CR, Dippenaar-Schoeman, AS. (2002). The influence of mound structure on the

harvester termite *Trinervitermes trinervoides*. *J Arachnol* 30: 403-408.

Corinnidae), with considerable range extension for *Poachelas striatus*. *Afr Invert* 51:

diversity of spiders (Araneae) inhabiting the abandoned mounds of the snouted

Natal, South Africa (Arachnida: Araneae). *Afr Invert* 47: 63-70.

Province, South Africa. *Koedoe* 50: 10-17.

they useful? *Afr Entomol* 18: 235-245.

(Araneae: Hersiliidae). *Afr Entomol* 13: 255-279.

*Hersilia* Audouin (Araneae: Hersiliidae). *Zootaxa*: 5-92.

(Araneae: Mygalomorphae). *Ann Nat Mus* 27: 1-37.

(Araneae: Corinnidae). *Afr Invert* 47: 85-93.

Lehtinen (Araneae, Lycosoidea). *Entomol Scand* 22: 305-351.

*Conserv* 10: 1221-1246.

35-43.

1-28.

89.

313-319.

Pietermaritzburg.

herbaceous layer of a coastal dune forest ecosystem at Richards Bay, KwaZulu-

spiders (Arachnida, Araneae) of the Polokwane Nature Reserve, Limpopo

indigenous forest in southern Africa: a case study from KwaZulu-Natal. *Biodivers*

*Hersiliola* Thorell and *Tama* Simon with the description of a new genus *Tyrotama*

fauna of the Western Soutpansberg, South Africa (Arachnida: Araneae). *Koedoe* 45:

heterogeneity of spiders (Arachnida: Araneae) in the Soutpansberg, South Africa: a comparative survey and inventory in representative habitats. *Afr Zool* 43: 156-174. Fourie, R, Haddad, CR, Jocqué, R. (2011). A revision of the purse web spider genus

*Calommata* Lucas, 1837 (Araneae: Atypidae) in the Afrotropical Region. *Zookeys* 95:

in South Africa. Unpublished M.Sc thesis, University of KwaZulu-Natal,

Dippenaar-Schoeman, AS. (1989). The African species of the sub family Penestominae

Dippenaar-Schoeman, AS. (1998) Spiders as predators of citrus pests. Agricultural Research

Dippenaar-Schoeman, AS. (2002) Baboon and Trapdoor Spiders of Southern Africa: An

Dippenaar-Schoeman, AS. (2007). South African National Survey of Arachnida: assessment and consolidation of data. *Newsl South African Nat Survey Arachnida* 3: 7-15. Dippenaar-Schoeman, AS, Craemer, C. (2000). The South African National Survey of

Dippenaar-Schoeman, AS, Jocqué, R. (1997) African spiders: An identification manual.

Dippenaar-Schoeman, AS, Leroy, A. (2003). A checklist of the spiders of the Kruger National

Dippenaar-Schoeman, AS, Leroy, A, De Jager, M, Van den Berg, A. (1999a). Spider diversity

Dippenaar-Schoeman, AS, Van den Berg, A, Prendini, L. (2009). Spiders and Scorpions

Dippenaar-Schoeman, AS, Van den Berg, AM. (2010) Spiders of the Kalahari. Agricultural

Dippenaar-Schoeman, AS, Van den Berg, AM, Van den Berg, A. (1989). Species composition

Dippenaar-Schoeman, AS, Van den Berg, AM, Van den Berg, MA. (1999b). Spiders in South

Dippenaar-Schoeman, AS, Van den Berg, AM, Van den Berg, MA, Foord, SH. (2005a).

Dippenaar-Schoeman, AS, Van den Berg, MA, Van den Berg, A. (2001b). Spiders in

Dippenaar-Schoeman, AS, Van der Walt, AE, De Jager, M, Le Roux, E, Van den Berg, A.

Dippenaar-Schoeman, AS, Wassenaar, TD. (2002). A checklist of the ground-dwelling

diversity and abundance (Arachnida: Araneae). *Afr Plant Prot* 11: 8-16. Dippenaar-Schoeman, AS, Van den Berg, MA, Van den Berg, A. (2001a). Salticid spiders in

diversity and abundance (Arachnida: Araneae). *Afr Plant Prot* 7: 39-46 Dippenaar-Schoeman, AS, Van der Merwe, M, Van den Berg, AM. (2006). Habitat

(Arachnida: Araneae, Scorpiones) of the Nylsvley Nature Reserve, South Africa.

and relative seasonal abundance of spiders from the field and tree layers of the

African cotton fields: species diversity and abundance (Arachnida: Araneae). *Afr*

Spiders in avocado orchards in the Mpumalanga Lowveld of South Africa: species

macadamia orchards in the Mpumalanga Lowveld of South Africa (Arachnida:

macadamia orchards in the Mpumalanga Lowveld of South Africa: species

preferences and seasonal activity of the Microstigmatidae from Ngome State Forest,

(2005b). The spiders of the Swartberg Nature Reserve in South Africa (Arachnida:

spiders of coastal dune forests at Richards Bay, South Africa (Arachnida: Araneae).

Identification Manual. Agricultural Research Council, Pretoria.

Park, South Africa (Arachnida: Araneae). *Koedoe* 46: 91-100.

of the Karoo National Park, South Africa. *Koedoe* 41: 31-42.

Roodeplaat Dam Nature Reserve. *Koedoe* 32: 51-60.

South Africa (Arachnida: Araneae). *Koedoe* 49: 85-89.

Council, Nelspruit.

*Koedoe* 51: 1-9.

Research Council, Pretoria.

Araneae). *Afr Plant Prot* 7: 47-51.

Araneae). *Koedoe* 49: 23-28.

*Bull Br Arachnol Soc* 12: 275-279.

*Plant Prot* 5: 93-103.

Arachnida. *Plant Prot News* 56: 11-12.

Agricultural Research Council, Pretoria.

(Araneae: Eresidae): with description of two new species. *Phytophylatica* 21: 131-134.

South African Spider Diversity:

50.

Inc., Sunderland, USA.

*Bloemfontein* 22: 113-126.

*Ann Ditsong Nat Mus Nat Hist* 1: 123-131

(Arachnida: Araneae). *Koedoe* 48: 109-115.

*Threat Taxa* 2: 815-819.

*Arachnological Colloquium:* 28.

*Phytophylatica* 23: 247-253.

Corinnidae). *Afr Invert* 51: 321-384.

African Perspectives on the Conservation of a Mega-Diverse Group 181

Lawrence, RF. (1953) The Biology of the Cryptic Fauna of Forests with Special Reference to Indigenous Forests of South Africa. Balkema Publishing Cape Town, Cape Town. Lomolino, MV, Riddle, BR, Brown, JH. (2006) Biogeography, 3rd edn. Sinauer Associates,

Lotz, LN. (2003). Afrotropical Archaeidae: 2. New species of the genera *Archaea* and *Afrarchaea* (Arachnida: Araneae). *Navors Nas Mus Bloemfontein* 19: 221-240. Lotz, LN. (2006). Afrotropical Archaeidae: 3. The female of *Eriauchenius cornutus* and new

Lotz, LN, Seaman, MT, Kok, DJ. (1991). Surface-active spiders (Araneae) of a site in semi-

Lyle, R. (2011). The male of *Fuchiba tortilis* Haddad and Lyle, 2008 and three new species of

Lyle, R, Haddad, CR. (2010). A revison of the tracheline sac spider genus *Cetonana* Strand,

Makaka, L, Little, IT, Dippenaar-Schoeman, AS, Jansen, R. (In review). Montane grassland

Mellet, MA, Schoeman, AS, Dippenaar-Schoeman, AS. (2006). The effect of Bt-cotton

Miller, JA, Griswold, CE, Haddad, CR. (2010). Taxonomic revision of the spider family

Modiba, MA, Dippenaar, SM, Dippenaar-Schoeman, AS. (2005). A check list of spiders from

Mucina, L, Rutherford, MC. (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19, South African National Biodiversity Institute, Pretoria.

Muelelwa, MI, Foord, SH, Dippenaar-Schoeman, AS, Stam, EM. (2010). Towards a

Mukherjee, A, Wilske, B, Navarro, RA, Dippenaar-Schoeman, AS, Underhill, LG. (2010).

Platnick, NI. (1991). Patterns of biodiversity: tropical vs temperate. *J Nat Hist* 25: 1083-1088.

Schoeman, CS, Foord, SH. (In press). A checklist of epigaeic ants from Marakele National Park, Limpopo Province, South Africa (Hymenoptera: Formicidae). *Koedoe*. Seshothela, SL, Dippenaar-Schoeman, AS. (2011). Termite eating spiders of the Cederberg

Van den Berg, A, Dippenaar-Schoeman, AS. (1991a). Ground-living spiders from an area

Pullin, AS. (2002) Conservation Biology. Cambridge University Press, Cambridge.

arid central South Africa. *Navors Nas Mus Bloemfontein* 7: 529-540.

South Africa (Arachnida, Araneae). *Agric Ecosyst Environ*.

Penestomidae (Araneae, Entelegynae). *Zootaxa* 2534: 1-36.

species of *Afrarchaea* (Arachnida: Araneae) from South Africa. *Navors Nas Mus*

*Spinotrachelas* Haddad, 2006 (Araneae: Corinnidae: Trachelinae) from South Africa.

1929 in the Afrotropical Region, with descriptions of two new genera (Araneae:

management and the implications on ground living spider community structure in

cultivation on spider populations in Marble Hall, South Africa. *Afr Plant Prot* 12: 40-

Sovenga Hill an inselberg in the Savanna Biome, Limpopo Province, South Africa

standardized and optimized protocol for rapid assessments: spider species richness and assemblage composition in two savanna vegetation types. *Afr Zool* 45: 273-290.

Association of spiders and lichens on Robben Island, South Africa: a case report. *J*

Wilderness Area (Araneae: Ammoxenidae). *Abstracts of the 10th African*

where the harvester termite *Hodotermes mossambicus* occurs in South Africa.

Haddad, CR, Dippenaar-Schoeman, AS. (2005). Epigaeic spiders (Arachnida: Araneae) in

Haddad, CR, Dippenaar-Schoeman, AS. (2006a). Spiders (Araneae) inhabiting abandoned

Haddad, CR, Dippenaar-Schoeman, AS. (2006b). Epigaeic spiders (Araneae) in pistachio

Haddad, CR, Dippenaar-Schoeman, AS. (2009). A checklist of the non-acarine arachnids

Haddad, CR, Dippenaar-Schoeman, AS, Wesołowska, W. (2006). A checklist of the non-

Haddad, CR, Honiball, AS, Dippenaar-Schoeman, AS, Slotow, R. (2010). Spiders as potential

Haddad, CR, Louw, SVDM, Dippenaar-Schoeman, AS. (2004a). Spiders (Araneae) in ground covers of pistachio orchards in South Africa. *Afr Plant Prot* 10: 97-107. Haddad, CR, Louw, SVDM, Dippenaar-Schoeman, AS. (2004b). An assessment of the

Hamer, ML, Slotow, RH. (2002). Conservation implications of existing data for South

Hamer, ML, Slotow, RH. (2009). A comparison and conservation assessment of the high

Herbert, DG. (2006). Rediscovery of the type species of *Euonyma* (Sabulinidae) and

Jenkins, CN, Sanders, NJ, Andersen, AN, Arnan, X, Brühl, CA, Cerda, X, Ellison, AM,

Jocqué, R. (2001). Chummidae, a new spider family (Arachnida, Araneae) from South Africa.

Jocqué, R. (2008). A new candidate for a Gondwanaland distribution in Zodariidae

Johnson, BL. (1999). The role of adaptive management as an operational approach for

two new species (Gastropoda: Eupulmonata). *J Nat Hist* 40: 1063-1081. Holm, E, Dippenaar-Schoeman, AS. (2010) Goggo Guide: The Arthropods of Southern

*Bloemfontein* 21: 1-10.

*Mus Bloemfontein* 22: 1-15.

South Africa. *Koedoe* 51: 1-9.

orchards in South Africa. *Afr Plant Prot* 12: 12-22.

Maputaland, South Africa. *Koedoe* 49: 1-22.

Maputaland, South Africa. *Afr J Ecol* 48: 446-460.

Africa (Araneae: Corinnidae). *Afr Invert* 49: 37-76.

(Araneae): *Australutica* in Africa. *Zookeys* 1: 59-66.

resource management agencies. *Conserv Ecol* 3: 8.

Drakensberg. *Soil Org* 81: 701-717.

Africa. LAPA Publishers, Pretoria.

*Distrib* 17: 652–662.

*J Zoology (Lond)* 54: 481-493.

African millipedes (Diplopoda). *Afr Entomol* 10: 29-42.

Nama Karoo grassland in the Northern Cape Province. *Navors Nas Mus*

mounds of the snouted harvester termite *Trinervitermes trinervoides* (Sjöstedt) (Isoptera: Termitidae: Nasutitermitinae) in the Free State, South Africa. *Navors Nas*

(Chelicerata: Arachnida) of the De Hoop Nature Reserve, Western Cape Province,

acarine arachnids (Chelicerata: Arachnida) of the Ndumo Game Reserve,

indicators of elephant-induced habitat changes in endemic sand forest,

biological control potential of *Heliophanus pistaciae* (Araneae: Salticidae) on *Nysius natalensis* (Hemiptera: Lygaeidae), a pest of pistachio nuts. *Biol Control* 31: 83-90. Haddad, CR, Lyle, R. (2008). Three new genera of tracheline sac spiders from southern

altitude grassland and forest millipede (Diplopoda) fauna of the South African

observations on South African species of *Gulella* (Streptaxidae), with description of

Fisher, BL, Fitzpatrick, MC, Gotelli, NJ, Gove, AD, Guénard, B, Lattke, JE, Lessard, J-P, McGlynn, TP, Menke, SB, Parr, CL, Philpott, SM, Vasconcelos, HL, Weiser, MD, Dunn, RR. (2011). Global diversity in light of climate change: the case of ants. *Divers*

**9**

*México*

**Relationships Between Bird Species Richness**

Principles implicitly addressed in most landscape level investigations of bird communities focus mainly on the arrangement of habitat patches, corridors, and matrix elements within landscapes; and patch area and isolation effects on dispersal, colonization, and local extinction (Forman, 1995). Ecologists are increasingly examining ecological patterns and processes at a scale that makes easier to understand the distribution and abundance of organisms contained within the habitat patches that compose the landscape (Forman & Gordon, 1986, Flather & Sauer, 1996, Bolger et al., 1997). Species interactions may vary for species within patches that adjoin different patch types (i.e., edge effects; Paton, 1994); for species in habitat patches of similar composition, but of differing patch sizes or distributions (i.e., habitat fragmentation effects; Robinson et al., 1995); for species requiring source-sink dynamics among patches in a landscape (i.e., metapopulations; Pulliam, 1988); and for species in habitat patches of similar composition but located within different landscape

Many studies of the effects of forest fragmentation on bird communities have been conducted in fragments surrounded by agricultural lands, and principles of island biogeography theory (MacArthur & Wilson, 1967) are usually invoked to explain patterns of species richness (Opdam, 1991). Birds are important model organisms for such studies because their taxonomy and distribution are well known, and because inventory and census methods are well developed (Ralph et al., 1995). However, in other situations, the surrounding habitat is not totally unsuitable for birds, and its characteristic determine how island-like the fragment will be (Hinsley et al., 1995, Stouffer & Bierregaard, 1995). In such cases, habitat fragmentation creates a mosaic of habitat patches of different quality, with forest fragments providing high quality habitat, and the matrix providing lower quality habitat (Wiens, 1994). For example, in North America, forest fragmentation has had an array of effects on neotropical migratory birds through habitat loss, small forest-patch size, reduced proximity of patches, more edge effect, and negative interactions with species surrounding nonforest patches (Faaborg et al., 1995, Freemark et al*.,* 1995, McGarigal &

**1. Introduction**

matrices (Renjifo, 1999).

McComb, 1995, Robinson et al., 1995).

**and Natural and Modified Habitat**

*2Museo de Zoología "Alfonso L. Herrera", Facultad de Ciencias,*

Jorge E. Ramírez-Albores1,2 and Adolfo G. Navarro-Sigüenza2 *1El Colegio de la Frontera Sur (ECOSUR). San Cristóbal de las Casas, Chiapas*

**in Southern Mexico**

*Universidad Nacional Autónoma de México*

## **Relationships Between Bird Species Richness and Natural and Modified Habitat in Southern Mexico**

Jorge E. Ramírez-Albores1,2 and Adolfo G. Navarro-Sigüenza2 *1El Colegio de la Frontera Sur (ECOSUR). San Cristóbal de las Casas, Chiapas 2Museo de Zoología "Alfonso L. Herrera", Facultad de Ciencias, Universidad Nacional Autónoma de México México*

## **1. Introduction**

182 Changing Diversity in Changing Environment

Van den Berg, AM, Dippenaar-Schoeman, AS. (1991b). Spiders, predacious insects and mites

Van den Berg, AM, Dippenaar-Schoeman, AS, Schoonbee, HJ. (1990). The effect of two pesticides on spiders in Southern African cotton fields. *Phytophylatica* 22: 435-441. Van der Merwe, M, Dippenaar-Schoeman, AS. (1996). Diversity of ground-living spiders at

Visser, D, Wright, MG, Van den Berg, A, Giliomee, JH. (1999). Species richness of arachnids

Wesołowska, W, Haddad, CR. (2009). Jumping spiders (Araneae: Salticidae) of the Ndumo

Whitmore, C, Slotow, R, Crouch, TE, Dippenaar-Schoeman, AS. (2002). Diversity of spiders

Wood, H. (2008). A revision of the assassin spiders of the *Eriauchenius gracilicollis* group, a

Game Reserve, Maputaland, South Africa. *Afr Invert* 50: 13-103.

Ngome State Forest, KwaZulu-Natal: a comparative survey in indigenous forest

associated with *Protea nitida* (Proteaceae) in the Cape Fynbos. *Afr J Ecol* 37: 334-343.

(Araneae) in a Savanna reserve, Northern Province, South Africa. *J Arachnol* 30: 344-

clade of spiders endemic to Madagascar (Araneae: Archaeidae). *Zool J Linn Soc* 152:

on South African cotton. *Phytophylatica* 23: 85-86.

and pine plantations. *Afr J Ecol* 48: 342-350.

356.

255-296.

Principles implicitly addressed in most landscape level investigations of bird communities focus mainly on the arrangement of habitat patches, corridors, and matrix elements within landscapes; and patch area and isolation effects on dispersal, colonization, and local extinction (Forman, 1995). Ecologists are increasingly examining ecological patterns and processes at a scale that makes easier to understand the distribution and abundance of organisms contained within the habitat patches that compose the landscape (Forman & Gordon, 1986, Flather & Sauer, 1996, Bolger et al., 1997). Species interactions may vary for species within patches that adjoin different patch types (i.e., edge effects; Paton, 1994); for species in habitat patches of similar composition, but of differing patch sizes or distributions (i.e., habitat fragmentation effects; Robinson et al., 1995); for species requiring source-sink dynamics among patches in a landscape (i.e., metapopulations; Pulliam, 1988); and for species in habitat patches of similar composition but located within different landscape matrices (Renjifo, 1999).

Many studies of the effects of forest fragmentation on bird communities have been conducted in fragments surrounded by agricultural lands, and principles of island biogeography theory (MacArthur & Wilson, 1967) are usually invoked to explain patterns of species richness (Opdam, 1991). Birds are important model organisms for such studies because their taxonomy and distribution are well known, and because inventory and census methods are well developed (Ralph et al., 1995). However, in other situations, the surrounding habitat is not totally unsuitable for birds, and its characteristic determine how island-like the fragment will be (Hinsley et al., 1995, Stouffer & Bierregaard, 1995). In such cases, habitat fragmentation creates a mosaic of habitat patches of different quality, with forest fragments providing high quality habitat, and the matrix providing lower quality habitat (Wiens, 1994). For example, in North America, forest fragmentation has had an array of effects on neotropical migratory birds through habitat loss, small forest-patch size, reduced proximity of patches, more edge effect, and negative interactions with species surrounding nonforest patches (Faaborg et al., 1995, Freemark et al*.,* 1995, McGarigal & McComb, 1995, Robinson et al., 1995).

Relationships Between Bird Species Richness

1975, Reyes-García & Souza, 1997).

Osumacinta

San Fernando

Tuxtla Gutiérrez

oak forest), Ur (Urban and suburban areas).

1

3

6

19

**2.2 Bird data**

and Natural and Modified Habitat in Southern Mexico 185

distinct successional stages (secondary forest), and agriculture fields, living fences, cattle pasture, shaded coffee plantations, urban and suburban areas (Miranda, 1975, FORTAM, 1984, Reyes-García & Souza, 1997). Common tree species in study area included *Pistacia mexicana*, *Cochlospermum vitifolium*, *Ceiba* sp, *Bursera bippinata, B. simaruba*, *Zuelania guidonia*, *Gyrocarpus* sp, *Acacia cornigera, A. pennatula*, *Haematoxylum* sp, *Lysiloma* sp, *Alvaradoa amorphoides*, *Swietenia humilis*, *Ficus* sp, *Fraxinus purpusii*, *Sideroxylon celastrinum* and *Heliocarpus reticulatus* (Miranda,

**Study sites Municipality Geographical coordinates Elevation (m) Habitats types**

4 16°55´N, 93°04´W 855 Tdf, Sf, C, Af, Lf 5 16°53´N, 93°07´W 630 Tdf, Tsf, Sf, Rf

7 16°52´N, 93°11´W 968 Tdf, Sf, C, Af, Lf 8 16°50´N, 93°13´W 1085 Tdf, Sf, C, Af, Lf, Ur 9 16°50´N, 93°12´W 970 Tdf, Sf, C, Af, Lf 10 16°51´N, 93°12´W 862 Tdf, Sf, C, Af, Lf, Ur 11 16°48´N, 93°10´W 835 Tdf, Tsf, Sf, To, Rf, C, Af, Lf 12 16°49´N, 93°11´W 868 Tdf, Sf, To, C, Af, Lf 13 16°49´N, 93°09´W 890 Tdf, Sf, C, Af, Lf 14 16°49´N, 93°12´W 1050 Tdf, Sf, C, Af, Lf 15 16°50´N, 93°11´W 875 C, Af, Lf, Ur 16 16°48´N, 93°11´W 883 Tdf, Sf, C 17 16°47´N, 93°10´W 710 Tdf, Sf, C 18 16°54´N, 93°10´W 995 Tdf, Tsf, Sf, Rf

20 16°45´N, 93°08´W 550 Ur 21 16°45´N, 93°05´W 508 Sf, Ur 22 16°47´N, 93°05´W 845 Tdf, Sf, C, Af, Lf 23 Chiapa de Corzo 16°42´N, 93°01´W 395 Sf, Rf, C, Af, Lf 24 16°41´N, 92°59´W 412 Sf, Rf, C, Af, Lf, Ur

Table 1. Characteristics of study sites in Chiapas Central Depression. Habitat types: Tdf (tropical deciduous forest), C (Cattle pastures), Lf (living fences), Rf (gallery forest), Tsf (Tropical semideciduous forest), Sf (secondary forest), Af (Agricultural fields), To (Tropical

Base data were collected in the field from February 2003 to November 2004 by sampling by point counts (Hutto et al., 1986, Ralph et al., 1995) that were used to asses species richness and abundance in each habitat. The number of point counts per habitat (4-8 points) was proportional to depended on the extent of different habitat types (between major coverage of the habitat sampled highest number of points). At each count station, the number of individuals of each species detected by sight and sound were recorded during a 5 min count

Chicoasén 16°57´N, 93°08´W 819 Tdf, Sf, C, Af, Lf 2 16°58´N, 93°10´W 1056 Tdf, Tsf, Sf, Rf, C, Af, Lf

16°56´N, 93°05´W 583 Tdf, Sf, C, Af, Lf

16°54´N, 93°09´W 1068 Tdf, Tsf, Sf, Af

16°45´N, 93°06´W 535 Sf, Ur

The distribution and diversity of bird communities in the tropical forests of Mexico and Central America have certainly been affected by a high degree of deforestation and therefore habitat fragmentation, but little quantitative or comparative data exist (Stiles, 1983, Flores-Villela & Geréz, 1994, Ceballos, 1995, Challenger, 1998). Some studies have identified landscape and habitat structural characteristics associated with the distribution of bird species richness in forest fragments that may be used to predict patterns of species richness in tropical deciduous forest patches (Gillespie & Walter, 2001), because different bird communities occur in response to changes in vegetation structure and species composition following logging (Morrison, 1992, Aleixo & Vielliard, 1995).

The Central Depression of Chiapas, located in southeastern Mexico, is an important area for conservation because highlights key characteristics of Middle American tropical deciduous forests: high level of endemism and the convergence of two biogeographically important migratory routes (the Gulf and the Pacific ones), thus, contains species that have migrated to the dry forest through each of these corridors. Also, there is a high turnover rate (beta diversity) between areas of tropical deciduous forest, which is also important for species conservation (Janzen, 1988, Escalante et al., 1993, Stattersfield et al., 1998). The area has also global importance for avian endemism (Stattersfield et al., 1998), and as a well-defined ecoregion (NT0211; Olson & Dinerstein, 1998, Myers et al., 2000), a Terrestrial Priority Site (Arriaga et al., 2000), an important bird area (IBAS; Arizmendi & Márquez, 2000), and the presence of some Natural Protected Areas including National Parks and Biosphere Reserve (i.e., Sumidero Canyon, El Zapotal; CONANP, 2011).

The understanding of the relationships and factors that influence bird community structure provides valuable information on the impact of habitat disturbance on populations, which is important for the conservation of these species. The goal of this contribution was to investigate differences in the species richness and composition of the bird communities in a mosaic natural and modified habitat and to evaluate how forest habitats perform to preserve species in the Central Depression of Chiapas. The results will be used to inform about appropriate strategies for the conservation of both the remnants of the original forest and the habitats created by humans with the species that inhabit them.

## **2. Material and methods**

#### **2.1 Study area**

The study area is located among the Municipalities of San Fernando, Tuxtla Gutiérrez, Chiapa de Corzo, Osumacinta and Chicoasén in the Central Depression region in Chiapas, southern Mexico (Table 1, Fig. 1). The climate is warm sub-humid with rainy summer (June to October), being May the hottest month, to moderate sub-humid in altitudes above the 1000 m (FORTAM, 1984, García, 1996). Mean annual temperature is 18-24 °C and mean annual precipitation varies between 500-2500 mm (FORTAM, 1984, INEGI, 2004, 2006). Annual precipitation shows a marked seasonality. The rainy season begins in mid-May, causing a surge of foliage and regrowth in natural vegetation areas as well as in crops and pasture grasses. This period normally lasts until the end of September. The dry season begins in December, lasting until May. The area includes a complex mixture of tropical habitats that have been classified on the basis of the physiognomic characteristics of the vegetation (Rzedowski, 1988, Reyes-García & Souza, 1997) and on the basis of land management. The natural and semi-natural habitats include tropical deciduous forest, tropical semideciduous forest, tropical oak forest, riparian forest, secondary forest, abandoned tropical forest with

The distribution and diversity of bird communities in the tropical forests of Mexico and Central America have certainly been affected by a high degree of deforestation and therefore habitat fragmentation, but little quantitative or comparative data exist (Stiles, 1983, Flores-Villela & Geréz, 1994, Ceballos, 1995, Challenger, 1998). Some studies have identified landscape and habitat structural characteristics associated with the distribution of bird species richness in forest fragments that may be used to predict patterns of species richness in tropical deciduous forest patches (Gillespie & Walter, 2001), because different bird communities occur in response to changes in vegetation structure and species composition

The Central Depression of Chiapas, located in southeastern Mexico, is an important area for conservation because highlights key characteristics of Middle American tropical deciduous forests: high level of endemism and the convergence of two biogeographically important migratory routes (the Gulf and the Pacific ones), thus, contains species that have migrated to the dry forest through each of these corridors. Also, there is a high turnover rate (beta diversity) between areas of tropical deciduous forest, which is also important for species conservation (Janzen, 1988, Escalante et al., 1993, Stattersfield et al., 1998). The area has also global importance for avian endemism (Stattersfield et al., 1998), and as a well-defined ecoregion (NT0211; Olson & Dinerstein, 1998, Myers et al., 2000), a Terrestrial Priority Site (Arriaga et al., 2000), an important bird area (IBAS; Arizmendi & Márquez, 2000), and the presence of some Natural Protected Areas including National Parks and Biosphere Reserve

The understanding of the relationships and factors that influence bird community structure provides valuable information on the impact of habitat disturbance on populations, which is important for the conservation of these species. The goal of this contribution was to investigate differences in the species richness and composition of the bird communities in a mosaic natural and modified habitat and to evaluate how forest habitats perform to preserve species in the Central Depression of Chiapas. The results will be used to inform about appropriate strategies for the conservation of both the remnants of the original forest and

The study area is located among the Municipalities of San Fernando, Tuxtla Gutiérrez, Chiapa de Corzo, Osumacinta and Chicoasén in the Central Depression region in Chiapas, southern Mexico (Table 1, Fig. 1). The climate is warm sub-humid with rainy summer (June to October), being May the hottest month, to moderate sub-humid in altitudes above the 1000 m (FORTAM, 1984, García, 1996). Mean annual temperature is 18-24 °C and mean annual precipitation varies between 500-2500 mm (FORTAM, 1984, INEGI, 2004, 2006). Annual precipitation shows a marked seasonality. The rainy season begins in mid-May, causing a surge of foliage and regrowth in natural vegetation areas as well as in crops and pasture grasses. This period normally lasts until the end of September. The dry season begins in December, lasting until May. The area includes a complex mixture of tropical habitats that have been classified on the basis of the physiognomic characteristics of the vegetation (Rzedowski, 1988, Reyes-García & Souza, 1997) and on the basis of land management. The natural and semi-natural habitats include tropical deciduous forest, tropical semideciduous forest, tropical oak forest, riparian forest, secondary forest, abandoned tropical forest with

following logging (Morrison, 1992, Aleixo & Vielliard, 1995).

(i.e., Sumidero Canyon, El Zapotal; CONANP, 2011).

**2. Material and methods**

**2.1 Study area**

the habitats created by humans with the species that inhabit them.

distinct successional stages (secondary forest), and agriculture fields, living fences, cattle pasture, shaded coffee plantations, urban and suburban areas (Miranda, 1975, FORTAM, 1984, Reyes-García & Souza, 1997). Common tree species in study area included *Pistacia mexicana*, *Cochlospermum vitifolium*, *Ceiba* sp, *Bursera bippinata, B. simaruba*, *Zuelania guidonia*, *Gyrocarpus* sp, *Acacia cornigera, A. pennatula*, *Haematoxylum* sp, *Lysiloma* sp, *Alvaradoa amorphoides*, *Swietenia humilis*, *Ficus* sp, *Fraxinus purpusii*, *Sideroxylon celastrinum* and *Heliocarpus reticulatus* (Miranda, 1975, Reyes-García & Souza, 1997).

Table 1. Characteristics of study sites in Chiapas Central Depression. Habitat types: Tdf (tropical deciduous forest), C (Cattle pastures), Lf (living fences), Rf (gallery forest), Tsf (Tropical semideciduous forest), Sf (secondary forest), Af (Agricultural fields), To (Tropical oak forest), Ur (Urban and suburban areas).

## **2.2 Bird data**

Base data were collected in the field from February 2003 to November 2004 by sampling by point counts (Hutto et al., 1986, Ralph et al., 1995) that were used to asses species richness and abundance in each habitat. The number of point counts per habitat (4-8 points) was proportional to depended on the extent of different habitat types (between major coverage of the habitat sampled highest number of points). At each count station, the number of individuals of each species detected by sight and sound were recorded during a 5 min count

Relationships Between Bird Species Richness

Arizmendi et al., 1990, Ramírez-Albores, 2010).

**2.3 Analysis**

and Natural and Modified Habitat in Southern Mexico 187

point counts conducted within a study habitat, and frequency of occurrence was based on presence or absence over all monthly samples. Relative abundances of neotropical or neartic migrants were based upon samples during months when they were present in the study area: January-April and October-January (i.e., winter visitors, summer residents). Each species was classified by a habitat guild (forest interior, generalist, and forest edge). We calculated species richness (We referred to total species richness as the total number of species per habitat) and abundance (bird abundance was obtained as the mean number of individuals detected in the total points counts per habitat) at each study site for forest interior species, generalist species, and forest edge species. Habitat guild classification was based Ehrlich et al. (1988) and also supported by other studies in fragmented forest (Brooks & Croonquist, 1990, Murcia, 1995, McIntyre, 1995, Rodewald & Yahner, 2001). Bird species were categorized into seven broad diet categories (carnivore, insectivore, nectarivore, frugivore, granivore, omnivore and aquatic) based upon primary components of the diet or subdiet obtained directly from field information and with supplemental information from literature (*i.e*., Ortiz-Pulido et al., 1995,

We used aerial photographs (scale 1:75,000; INEGI, 2001) to map land-use types of study sites and we performed direct surveys throughout the area for confirmation of site suitability. Sampling intensity was stratified among different sites based on the extent and cover proportions of different habitat on the INEGI image. Each sites was surveyed an equal number of times (sites were visited one time each month). At each site, at least 90-100% of the nonforest cover within 1 km of the study site consisted of only one disturbance type (primarily agricultural fields, cattle pastures and urbanization). We determined forest cover from classified thematic mapped imagery using ARC/INFO geographic information system software (ESRI, 1999). We calculated species richness by habitat guild: forest specialist, generalist, and early successional species, at each study site. Species richness was analyzed separately by a multiple regression analysis to assess if there was any differential response to forest disturbance characteristics, based on the species level of dependence on arboreal cover proportion. Stepwise regression analysis was performed on log-transformed total number of species, and on resident and migrant species. Bird abundance data were log (e) x + 1 transformed previous to the analyses to reduce the skewness of the data, resulting in a more interpretable analysis. An *F*-test probability value of 0.05 and 0.001 was used in all cases. Differences in species richness and guild structure of bird communities, represented as the species richness in different foraging guilds, were compared among habitat types using oneway analysis of variance (ANOVA). Tukey´s multiple range test was used for post-hoc comparisons among habitat types (Zar, 1999). The Similarity of species composition between habitats types was measured using Sorenson´s similarity index (IS=2S/N1+N2, where S is the number of common species, N1 is the number of species of habitat 1, and N2 is the number of species of habitat 2; Ravinovich, 1981). To improve the knowledge of the geographic distribution of each individual species we used a set of maps of all species of landbirds of Mexico (Navarro-Sigüenza & Peterson, 2007) constructed by ecological niche modeling (Nix, 1986, Peterson, 2001). Maps depict the potential distributions of the species using the Genetic Algorithm for Rule-set Production (GARP; Stockwell & Noble, 1992), in its PC implementation DesktopGARP (Scachetti-Pereira, 2003), using as primary source the data points contained in the Atlas of the Birds of Mexico data base (Navarro-Sigüenza et al., 2003). For generating the

period. Each count lasted for 5 minutes with a 5-minute interval between points. Birds detected at > 100 m were recorded but not used in analyses to reduce the possibility of counting the same individual twice in consecutive points. Birds detected when not conducting counts were also recorded and used to calculate total species richness. No counts were conducted on days when visibility was poor, or under windy or rainy conditions. Counts were conducted between 0700 and 1100 in the morning, and 1600 and 1900 in the afternoon (i.e., during the highest bird activity). No survey was conducted during unfavorable weather conditions (rainy, windy and mist days) because birds were less detectable under those conditions (O'Connor & Hicks, 1980, Robbins, 1981).

Fig. 1. Map of Tehuantepec Isthmus region in southern Mexico, the quadrangle depicts study region (a). Underlying map is average elevation (http://www.conabio.gob.mx). Study sites (dots) in the Central Depression of Chiapas, Mexico (b). Labels correspond to Ecoregions of World Wildlife Fund (http://www.wwf.org).

Bird species richness was calculated as the total number of species recorded in each habitat. Two estimates of relative abundance (including both visual and aural detections of both sexes) for each species were obtained for each habitat: the average number of individuals per point count, and frequency of occurrence during monthly samples. These two measurements of relative abundance assume that birds are recorded more often in areas where they are more abundant (Renjifo, 2001). The average number of individuals per point count was based on all point counts conducted within a study habitat, and frequency of occurrence was based on presence or absence over all monthly samples. Relative abundances of neotropical or neartic migrants were based upon samples during months when they were present in the study area: January-April and October-January (i.e., winter visitors, summer residents). Each species was classified by a habitat guild (forest interior, generalist, and forest edge). We calculated species richness (We referred to total species richness as the total number of species per habitat) and abundance (bird abundance was obtained as the mean number of individuals detected in the total points counts per habitat) at each study site for forest interior species, generalist species, and forest edge species. Habitat guild classification was based Ehrlich et al. (1988) and also supported by other studies in fragmented forest (Brooks & Croonquist, 1990, Murcia, 1995, McIntyre, 1995, Rodewald & Yahner, 2001). Bird species were categorized into seven broad diet categories (carnivore, insectivore, nectarivore, frugivore, granivore, omnivore and aquatic) based upon primary components of the diet or subdiet obtained directly from field information and with supplemental information from literature (*i.e*., Ortiz-Pulido et al., 1995, Arizmendi et al., 1990, Ramírez-Albores, 2010).

### **2.3 Analysis**

186 Changing Diversity in Changing Environment

period. Each count lasted for 5 minutes with a 5-minute interval between points. Birds detected at > 100 m were recorded but not used in analyses to reduce the possibility of counting the same individual twice in consecutive points. Birds detected when not conducting counts were also recorded and used to calculate total species richness. No counts were conducted on days when visibility was poor, or under windy or rainy conditions. Counts were conducted between 0700 and 1100 in the morning, and 1600 and 1900 in the afternoon (i.e., during the highest bird activity). No survey was conducted during unfavorable weather conditions (rainy, windy and mist days) because birds were less

detectable under those conditions (O'Connor & Hicks, 1980, Robbins, 1981).

Fig. 1. Map of Tehuantepec Isthmus region in southern Mexico, the quadrangle depicts study region (a). Underlying map is average elevation (http://www.conabio.gob.mx). Study sites (dots) in the Central Depression of Chiapas, Mexico (b). Labels correspond to

Bird species richness was calculated as the total number of species recorded in each habitat. Two estimates of relative abundance (including both visual and aural detections of both sexes) for each species were obtained for each habitat: the average number of individuals per point count, and frequency of occurrence during monthly samples. These two measurements of relative abundance assume that birds are recorded more often in areas where they are more abundant (Renjifo, 2001). The average number of individuals per point count was based on all

Ecoregions of World Wildlife Fund (http://www.wwf.org).

We used aerial photographs (scale 1:75,000; INEGI, 2001) to map land-use types of study sites and we performed direct surveys throughout the area for confirmation of site suitability. Sampling intensity was stratified among different sites based on the extent and cover proportions of different habitat on the INEGI image. Each sites was surveyed an equal number of times (sites were visited one time each month). At each site, at least 90-100% of the nonforest cover within 1 km of the study site consisted of only one disturbance type (primarily agricultural fields, cattle pastures and urbanization). We determined forest cover from classified thematic mapped imagery using ARC/INFO geographic information system software (ESRI, 1999). We calculated species richness by habitat guild: forest specialist, generalist, and early successional species, at each study site. Species richness was analyzed separately by a multiple regression analysis to assess if there was any differential response to forest disturbance characteristics, based on the species level of dependence on arboreal cover proportion. Stepwise regression analysis was performed on log-transformed total number of species, and on resident and migrant species. Bird abundance data were log (e) x + 1 transformed previous to the analyses to reduce the skewness of the data, resulting in a more interpretable analysis. An *F*-test probability value of 0.05 and 0.001 was used in all cases. Differences in species richness and guild structure of bird communities, represented as the species richness in different foraging guilds, were compared among habitat types using oneway analysis of variance (ANOVA). Tukey´s multiple range test was used for post-hoc comparisons among habitat types (Zar, 1999). The Similarity of species composition between habitats types was measured using Sorenson´s similarity index (IS=2S/N1+N2, where S is the number of common species, N1 is the number of species of habitat 1, and N2 is the number of species of habitat 2; Ravinovich, 1981). To improve the knowledge of the geographic distribution of each individual species we used a set of maps of all species of landbirds of Mexico (Navarro-Sigüenza & Peterson, 2007) constructed by ecological niche modeling (Nix, 1986, Peterson, 2001). Maps depict the potential distributions of the species using the Genetic Algorithm for Rule-set Production (GARP; Stockwell & Noble, 1992), in its PC implementation DesktopGARP (Scachetti-Pereira, 2003), using as primary source the data points contained in the Atlas of the Birds of Mexico data base (Navarro-Sigüenza et al., 2003). For generating the

Relationships Between Bird Species Richness

study sites.

45.3, *P<* 0.0001).

**3.2 Comparison among habitat types**

and Natural and Modified Habitat in Southern Mexico 189

Fig. 3. The relationships between the extent disturbance (%) and bird species richness at the

Tropical deciduous forest (203) had the highest number of species, whereas tropical oak forest (51) and aquatic and semiaquatic habitats (24) had the fewest species (Fig. 4). Of the total bird species recorded (278), 20 were exclusively found in tropical deciduous forest, four of tropical semideciduous forest, two of urban/suburban areas and one of cattle pastures (Appendix 1). To analyzed comparative the habitat types with bird species richness and mean abundance, and we found significant differences (F8,125 = 70.6, *P <* 0.0001, F8,125 = 106.2, *P<* 0.0001, respectively). As migration status in the different habitat types, also significant differences between residents species (F8,125 = 79.1, *P <* 0.0001) and migratory species (F8,125 =

models a set of 19 climatic variables, derived from temperature and precipitation (Hijmans et al., 2005; http://www.worldclim.org), and three topographic (Hydro1k project; http://eros.usgs.gov) was used. Individual summaries of distributions of species were summed to produce species richness maps for total species, summer resident species, winter resident species (Navarro-Sigüenza & Peterson, 2007). From the GARP maps, we derived predicted numbers of resident and migrant species using ArcView (version 3.2; ESRI, 1999). We also compared species richness values for each grid cell (resolution 0.05°) with GIS data layers summarizing Terrestrial Prioritary Regions (Arriaga et al., 2000, CONABIO, 2004) to assess whether areas recognized as priority under diverse criteria coincide with areas of greatest species richness. All statistical analyses were permormed using STATISTICA® 10 and SPSS® 19.5.

### **3. Results**

#### **3.1 Bird species composition**

A total of 279 species of 45 families was recorded from the 24 sites (Appendix 1). Of these, 193 were permanent residents and 86 were migrant species (including one occasional, two summer residents, 18 transients and 65 winter visitors). In general, the average bird richness during the study period was of 131 species/month; however, the monthly bird species richness ranged from 100 to 161 (Fig. 2). The fewest species were found in May and the most in March, April, December and January (Fig. 2). The composition of the bird community associated with percentage of disturbance in the study sites, according to habitat preferences, corresponding to 30.2% (N = 84) for forest specialists, 10.4% (N = 29) of early successional species and 39.5% (N= 110) forest generalists (Fig. 3). The distribution of each category in the sites showed greater richness of specialists and forest generalists. Forest specialist species richness (F1,22= 5.98, r= 0.46, P= 0.02) and generalist (F1,22= 17.53, r= 0.66, P= 0.0003) were negatively associated with percentage of disturbance (Fig. 3). Early succesional species richness (F1,22= 4.21, r= 0.40, P= 0.05) was slightly related to disturbance within study sites. Diet or subdiet composition of bird communities in the study sites was: 82 were insectivores, 72 insectivores/frugivores, 39 carnivores, 20 granivores/fruigivores, 14 nectarivores and 13 granivores (Appendix 1).

Fig. 2. Monthly species richness during study period. Mean and standard error for data pooled over visit and point count shown (errors bars represent 95% confidence intervals).

models a set of 19 climatic variables, derived from temperature and precipitation (Hijmans et al., 2005; http://www.worldclim.org), and three topographic (Hydro1k project; http://eros.usgs.gov) was used. Individual summaries of distributions of species were summed to produce species richness maps for total species, summer resident species, winter resident species (Navarro-Sigüenza & Peterson, 2007). From the GARP maps, we derived predicted numbers of resident and migrant species using ArcView (version 3.2; ESRI, 1999). We also compared species richness values for each grid cell (resolution 0.05°) with GIS data layers summarizing Terrestrial Prioritary Regions (Arriaga et al., 2000, CONABIO, 2004) to assess whether areas recognized as priority under diverse criteria coincide with areas of greatest species richness. All statistical analyses were permormed using STATISTICA® 10

A total of 279 species of 45 families was recorded from the 24 sites (Appendix 1). Of these, 193 were permanent residents and 86 were migrant species (including one occasional, two summer residents, 18 transients and 65 winter visitors). In general, the average bird richness during the study period was of 131 species/month; however, the monthly bird species richness ranged from 100 to 161 (Fig. 2). The fewest species were found in May and the most in March, April, December and January (Fig. 2). The composition of the bird community associated with percentage of disturbance in the study sites, according to habitat preferences, corresponding to 30.2% (N = 84) for forest specialists, 10.4% (N = 29) of early successional species and 39.5% (N= 110) forest generalists (Fig. 3). The distribution of each category in the sites showed greater richness of specialists and forest generalists. Forest specialist species richness (F1,22= 5.98, r= 0.46, P= 0.02) and generalist (F1,22= 17.53, r= 0.66, P= 0.0003) were negatively associated with percentage of disturbance (Fig. 3). Early succesional species richness (F1,22= 4.21, r= 0.40, P= 0.05) was slightly related to disturbance within study sites. Diet or subdiet composition of bird communities in the study sites was: 82 were insectivores, 72 insectivores/frugivores, 39 carnivores, 20 granivores/fruigivores, 14

Fig. 2. Monthly species richness during study period. Mean and standard error for data pooled over visit and point count shown (errors bars represent 95% confidence intervals).

and SPSS® 19.5.

**3.1 Bird species composition**

nectarivores and 13 granivores (Appendix 1).

**3. Results**

Fig. 3. The relationships between the extent disturbance (%) and bird species richness at the study sites.

#### **3.2 Comparison among habitat types**

Tropical deciduous forest (203) had the highest number of species, whereas tropical oak forest (51) and aquatic and semiaquatic habitats (24) had the fewest species (Fig. 4). Of the total bird species recorded (278), 20 were exclusively found in tropical deciduous forest, four of tropical semideciduous forest, two of urban/suburban areas and one of cattle pastures (Appendix 1). To analyzed comparative the habitat types with bird species richness and mean abundance, and we found significant differences (F8,125 = 70.6, *P <* 0.0001, F8,125 = 106.2, *P<* 0.0001, respectively). As migration status in the different habitat types, also significant differences between residents species (F8,125 = 79.1, *P <* 0.0001) and migratory species (F8,125 = 45.3, *P<* 0.0001).

Relationships Between Bird Species Richness

(aquatic), G (granivore), F (frugivore).

forest and cattle pastures (0.19; Table 2).

=38.0, *P*<0.0001).

and Natural and Modified Habitat in Southern Mexico 191

nectarivores (14), compared to the tropical oak forest, cattle pastures and agricultural fields where there were no records of these species, the differences between habitats (F8,125

**Habitat types**

Fig. 5. Diet or subdiet composition of bird communities in different habitat types of study area. Habitat types: tropical deciduous forest), C (cattle pastures), Lf (living fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af (agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq (aquatics and

subaquatics). Diet categories: I (insectivore), C (carnivore), O (omnivore), N (nectarivore), A

Similarity of species composition between habitat types indicates that the highest values were among cattle pastures and agricultural fields (0.81), followed by tropical deciduous forest and secondary forest (0.75) (Table 2). The fewest values were among gallery forest and cattle pastures (0.15), gallery forest and agricultural fields (0.19), and tropical semideciduous

Fig. 4. Species richness of birds in different habitat types at the study sites. Mean and standard error for data pooled over visit and point count shown (errors bars represent 95% confidence intervals). Habitat types: Tdf (tropical deciduous forest), C (cattle pastures), Lf (live fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af (agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq (aquatics and subaquatics).

Carnivores bird species were better represented in the tropical deciduous forest (17), agricultural fields (18) and pastures (17; Fig. 5), while the lowest numbers occurred in the tropical oak forest (5) and urban/suburban zones (4) and there were no species in living fences (F8,125 =53.9, *P*<0.0001). Insectivores-frugivores species were more abundant in the tropical deciduous forest (64) than in agricultural fields (3) and pastures (6; F8,125 =35.4, *P*<0.0001). The lowest number was recorded in living fences and pastures with one species each, presenting significant differences between habitats (F8,125 =47.2, *P*<0.0001). Tropical deciduous forest (70) had a greater number of insectivores species than gallery forest (10; F8,125 =80.1, *P*<0.0001). Tropical deciduous forest had the highest number of

Fig. 4. Species richness of birds in different habitat types at the study sites. Mean and standard error for data pooled over visit and point count shown (errors bars represent 95% confidence intervals). Habitat types: Tdf (tropical deciduous forest), C (cattle pastures), Lf (live fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af

(agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq

Carnivores bird species were better represented in the tropical deciduous forest (17), agricultural fields (18) and pastures (17; Fig. 5), while the lowest numbers occurred in the tropical oak forest (5) and urban/suburban zones (4) and there were no species in living fences (F8,125 =53.9, *P*<0.0001). Insectivores-frugivores species were more abundant in the tropical deciduous forest (64) than in agricultural fields (3) and pastures (6; F8,125 =35.4, *P*<0.0001). The lowest number was recorded in living fences and pastures with one species each, presenting significant differences between habitats (F8,125 =47.2, *P*<0.0001). Tropical deciduous forest (70) had a greater number of insectivores species than gallery forest (10; F8,125 =80.1, *P*<0.0001). Tropical deciduous forest had the highest number of

(aquatics and subaquatics).

nectarivores (14), compared to the tropical oak forest, cattle pastures and agricultural fields where there were no records of these species, the differences between habitats (F8,125 =38.0, *P*<0.0001).

Fig. 5. Diet or subdiet composition of bird communities in different habitat types of study area. Habitat types: tropical deciduous forest), C (cattle pastures), Lf (living fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af (agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq (aquatics and subaquatics). Diet categories: I (insectivore), C (carnivore), O (omnivore), N (nectarivore), A (aquatic), G (granivore), F (frugivore).

Similarity of species composition between habitat types indicates that the highest values were among cattle pastures and agricultural fields (0.81), followed by tropical deciduous forest and secondary forest (0.75) (Table 2). The fewest values were among gallery forest and cattle pastures (0.15), gallery forest and agricultural fields (0.19), and tropical semideciduous forest and cattle pastures (0.19; Table 2).

Relationships Between Bird Species Richness

Depression of Chiapas.

southern Brazil (Anjos, 2001).

needed to be discussed in detail later.

and Natural and Modified Habitat in Southern Mexico 193

Fig. 6. Maps representing of modeled species richness in the study sites at the Central

different in forest cover but diverse in habitat types associated with tropical forest are areas that contained continuous secondary forest, the same pattern found in forest fragments in

The 195 species (70%) were considered residents; as the number of resident species may be higher due to birds with less conspicuous behavior in certain periods of the year and/or difficulty in detecting those (Krügel & Anjos, 2000). Karr et al. (1982) mentioned that in some tropical environments the migratory species are capable of producing changes in the composition of bird communities. In this study migratory species (30%) played a minor role in the observed changes in the bird community. According to Arizmendi et al. (1990) and Moya-Moreno (1990), it is possible that in the study area altitudinal and latitudinal movements are correlated with fluctuations in the abundance of species. For example, some rare species are clearly features temperate environments whose populations are dispersed to other locations during times of scarcity of resources, or are migratory in passing that occur in small amounts within Chiapas. However, these seasonal changes in abundance, possibly also associated with seasonal phenology of the deciduous forest were not assessed, so that

Species richness was greater in April, October, and December, surely due to the presence of migratory species and to the beginning of reproductive activity, which make birds more detectable. On the other hand, a lower richness was found in May and August, a period in which migratory species were absent and birds were quiet, making them difficult to detect.

Table 2. Matrix similarity of bird species, based on Sorenson´s Index, among habitat types surveyed in Chiapas Central Depression.

## **3.3 General patterns of bird diversity**

Species per 1 km2 cell in the map of the region (Fig. 6) can do a high geographic consistency of the patterns. The richest areas of the study area form a strip that runs in an east-west from the eastern part of the Petén-Veracruz moist forest, following to northern part of the Chiapas Depression dry forest, and continues the Central America pine-oak forest. The richest cells within this region are precisely in the northern part of Chiapas Depression. Two cells differ with high values of richness, which are north of the Central Depression of Chiapas. We can say that there is a continuous strip of high species richness throughout the study area in east-west. In this sense, are evident two regions: the northern part of the Central Depression of Chiapas, and Gulf Coastal Plain. In fact, this latter may represent a decrease in species richness west-east (Fig. 6). The figure 6 helped to identify the species richness of areas of greatest concentration of diversity; the southern region presented the lowest concentration with a maximum of 62 species. The prediction map of migratory species richness shows the greatest number of species concentration mainly in the southern and northeastern. Most species are concentrated in the dry and moist forest, which is apparently a different distribution pattern observed in the Central America pine-oak forest.

## **4. Discussion**

Of a total of 656 bird species occurring in Chiapas according to Álvarez del Toro (1980) and Palomera-García et al. (1994), the species recorded in the study area (Central Depression of Chiapas) corresponds to 42% (279 species; Appendix 1). This high richness is a result of a complex array of habitats, convergence of two important migratory routes (of the Gulf and Pacific), as well as biogeographic (biotic provinces) and physiographic heterogeneity (Arriaga et al., 2000). Bird species richness found in study sites is similar to that in other tropical forest regions in Mexico, such as La Mancha on the coast of Veracruz (250 species; Ortiz-Pulido et al., 1995) and Chamela in Jalisco (270 species; Arizmendi et al*.,* 1990). The study sites showing greater species richness (especially in tropical deciduous forest) are

**Secondary forest**

**Secondary forest** 0.34 0.31 0.36 0.54 0.40 **Gallery forest** 0.19 0.15 0.53 0.49 **Cattle pastures** 0.81 0.24 0.37

**Living fences** 0.45 Table 2. Matrix similarity of bird species, based on Sorenson´s Index, among habitat types

Species per 1 km2 cell in the map of the region (Fig. 6) can do a high geographic consistency of the patterns. The richest areas of the study area form a strip that runs in an east-west from the eastern part of the Petén-Veracruz moist forest, following to northern part of the Chiapas Depression dry forest, and continues the Central America pine-oak forest. The richest cells within this region are precisely in the northern part of Chiapas Depression. Two cells differ with high values of richness, which are north of the Central Depression of Chiapas. We can say that there is a continuous strip of high species richness throughout the study area in east-west. In this sense, are evident two regions: the northern part of the Central Depression of Chiapas, and Gulf Coastal Plain. In fact, this latter may represent a decrease in species richness west-east (Fig. 6). The figure 6 helped to identify the species richness of areas of greatest concentration of diversity; the southern region presented the lowest concentration with a maximum of 62 species. The prediction map of migratory species richness shows the greatest number of species concentration mainly in the southern and northeastern. Most species are concentrated in the dry and moist forest, which is apparently a different

Of a total of 656 bird species occurring in Chiapas according to Álvarez del Toro (1980) and Palomera-García et al. (1994), the species recorded in the study area (Central Depression of Chiapas) corresponds to 42% (279 species; Appendix 1). This high richness is a result of a complex array of habitats, convergence of two important migratory routes (of the Gulf and Pacific), as well as biogeographic (biotic provinces) and physiographic heterogeneity (Arriaga et al., 2000). Bird species richness found in study sites is similar to that in other tropical forest regions in Mexico, such as La Mancha on the coast of Veracruz (250 species; Ortiz-Pulido et al., 1995) and Chamela in Jalisco (270 species; Arizmendi et al*.,* 1990). The study sites showing greater species richness (especially in tropical deciduous forest) are

distribution pattern observed in the Central America pine-oak forest.

**Gallery forest**

0.69 0.33 0.75 0.32 0.22 0.23 0.46 0.34

0.44 0.61 0.36 0.19 0.24 0.35 0.39

 **Cattle pastures**

0.42 0.57 0.34 0.26 0.51 0.46

**Agricul tural fields**

**Living fences**

0.22 0.32

**Urban/su burban areas**

**Tropical oak forest**

**Tropical semideciduous forest**

surveyed in Chiapas Central Depression.

**3.3 General patterns of bird diversity**

**Tropical deciduous forest**

**Tropical semideciduous forest**

**Tropical oak forest**

**Agricultural fields**

**4. Discussion**

Fig. 6. Maps representing of modeled species richness in the study sites at the Central Depression of Chiapas.

different in forest cover but diverse in habitat types associated with tropical forest are areas that contained continuous secondary forest, the same pattern found in forest fragments in southern Brazil (Anjos, 2001).

The 195 species (70%) were considered residents; as the number of resident species may be higher due to birds with less conspicuous behavior in certain periods of the year and/or difficulty in detecting those (Krügel & Anjos, 2000). Karr et al. (1982) mentioned that in some tropical environments the migratory species are capable of producing changes in the composition of bird communities. In this study migratory species (30%) played a minor role in the observed changes in the bird community. According to Arizmendi et al. (1990) and Moya-Moreno (1990), it is possible that in the study area altitudinal and latitudinal movements are correlated with fluctuations in the abundance of species. For example, some rare species are clearly features temperate environments whose populations are dispersed to other locations during times of scarcity of resources, or are migratory in passing that occur in small amounts within Chiapas. However, these seasonal changes in abundance, possibly also associated with seasonal phenology of the deciduous forest were not assessed, so that needed to be discussed in detail later.

Species richness was greater in April, October, and December, surely due to the presence of migratory species and to the beginning of reproductive activity, which make birds more detectable. On the other hand, a lower richness was found in May and August, a period in which migratory species were absent and birds were quiet, making them difficult to detect.

Relationships Between Bird Species Richness

and semideciduous forest).

and Natural and Modified Habitat in Southern Mexico 195

structure and formed by several layers of coverage are mainly species of insectivores, frugivores and nectar habits (Rappole et al., 1993). The results in this study are consistent with the above, as modified habitats had a higher proportion of species and individuals of carnivores and granivores habits compared to the original habitats (i.e., tropical deciduous

In general, the variety of habitats present in the study region seems to contribute a high proportion of species, especially considering the number of species occurring in tropical deciduous forest (203). This may be due to the structural complexity that makes an ecosystem with greater species richness in Mexico (Ceballos & García, 1995, Ceballos et al., 2010), and the fact of having a greater horizontal and vertical stratification with respect to others, thus generating increased availability of habitats and ecological niches (Blake & Loiselle, 1991, McIntyre, 1995, Villard et al., 1999), as the plant structure determines the amount and distribution of resources used by birds. The differences in diversity and richness found indicate that the tropical deciduous forest, tropical semideciduous forest, and secondary forest show a greater richness compared to other modified habitats (i.e., agricultural fields, cattle pasture). This coincides with other studies conducted in tropical environments and indicates that the original habitat loss directly affects the presence, abundance and persistence of species (Kattan et al., 1994, Laurance & Bierregaard, 1997). The results of this study suggest that species richness and diversity of habitats ranging from the study. Natural habitats (tropical deciduous forest, tropical semideciduous forest and secondary forest) appear to be attractive to a larger number of bird species, as both the richness and diversity were higher in these, which is consistent with other studies (Estrada

et al., 1997, Petit et al., 1999, Blake & Loiselle, 2001, Bojorges & Lopez-Mata, 2005).

persistence of resident and migratory species (Morales, 2002).

Modified habitats (agricultural fields, living fences, cattle pasture) had a significant contribution to the bird species richness in the study area. These habitats provide roosting sites and food resources (Lynch, 1989). This is consistent with that reported by Estrada et al. (1997), which brings a richness of 226 in the region of Los Tuxtlas (Veracruz, Mexico), finding 79% of the species found in forest areas, 80% farmland, 43% in living fences and only 5% in grassland/pasture. In addition, Petit et al. (1999) in the central area of Panama found that species richness in modified habitats (i.e., shade coffee plantations, residential areas, grasslands and pine plantations) is equal or similar to the natural habitat. However, live fences exhibited the highest species richness (77 species) from modified habitats (i.e., agricultural fields, cattle pasture), probably because their plant structure is more complex and diverse. This is consistent with other studies (i.e., Villaseñor, 1993, Villaseñor & Hutto, 1995, Morales, 2002), which state that living fences can be very attractive to a large number of individuals and species of birds, and also can support high densities as they provide food resources, roosting sites and shelter (Villaseñor & Hutto, 1995). For example, in the study area, some birds prefer to use corridors or live fences instead of open or cleared areas (Wegner & Merriam, 1979) and turnover rates are significantly more frequent along corridors connected to original habitats or with other corridors (Hass, 1995, Machtans et al., 1996). In the case where the original habitat remains, the complexity of vegetation provides alternative sites for some species, partially offsetting the fragmentation and allowing the

The similarity between the habitats types of study area indicates the existence of a high turnover of species and an apparent high connectivity between them. Suggesting that both the configuration of the environment (i.e., landscape, habitat and microhabitat) and the available amount thereof would not be equally important in the distribution of species (Karr

Seasonal variation of the avifauna in the present study was similar to that found in other tropical forest regions (Chamela region in Jalisco, Mexico and Maringá in Paraná, Brazil; Ornelas et al., 1993, Krügel & Anjos, 2000, respectively) where the species richness was greater from October to November. Our results show that species composition did not differ significantly across the 24 study sites, and similarities of different levels among the sites were common. This could suggest that most tropical forest patches still have suitable habitats that ensure availability of food, nesting sites, and protective cover for the species but are still vulnerable to persistent encroachment evident around them. In the long term this could jeopardize the ability to sustain particular bird species, especially forestdependent bird, threatened and endemic.

A considerable amount of species associated with secondary forest, open areas, clearings and forest edges remain abundant and are likely to increase in regions with small isolated forest fragments (Bierregaard & Lovejoy, 1989, Thiollay, 1992). Generalist birds, which change their diet from fruit to insects or vice-versa, are also favored in small patches (Willis, 1979). Mota (1990) found increasing, devastation of pristine areas. Although some general tendencies were observed for certain bird groups, the effects of forest fragmentation are certainly different for each species. A study in forest fragments (in Maryland) suggested that the impacts of forest fragmentation on bird communities are complex, species specific and not related only to fragment area or fragment isolation (Lynch & Whigham, 1984). The increase in species richness with fragmentation was primarily due to the addition of several migrants that were associated with edge habitats and secondary forest.

These species showed a lower frequency possibly because they were represented by few individuals, and are more sensitive to forest change and fragmentation than more widespread species, as patterns that has been shown before (Fjeldsa, 1999, Renjifo, 2001). Priority species (i.e., endemic and threatened) are important contributors to biodiversity because their restricted distributions make them globally rare and particularly vulnerable to population declines or extinction (Terborgh & Winter, 1983, Diamond 1986). Species with small ranges are also less abundant at a local scale than large-range species (Brown, 1995). The birds may demonstrate a differential response to forest fragmentation (Hobson & Bayne, 2000, Fahrig, 2003) or that probably bird species richness in the study area can be affected by other factors, such as floristic diversity, and vegetation composition and structure (Gillespie & Walter, 2001). Other effects, such as the extent and nature of the fragments edges, fragment connectivity, or fragment shape (Bierregaard et al., 1992, Laurance & Bierregaard, 1997, Cornelius et al., 2000), might be more important than forest cover in predicting the number of species found in the area (Ramírez-Albores, 2010). As for other studies, it is expected that the effect of forest cover would affect bird species richness (Kattan et al., 1994, Laurance & Bierregaard, 1997).

Diet composition was similar among habitat types, with greatest representation by insectivores and insectivores/frugivores, and decreasing representation by nectarivores and granivores. According to Petit et al. (1999) and Karr (1990), this distribution of foraging guild memberships is typical of that found in tropical forest. Tropical forest fragments resulting from human disturbance of a continuous forest are isolated more rapidly. The remaining areas suffer of progressive degradation due to isolation, which, in the long term, jeopardizes the survival of several species. In tropical environments, modified habitats are very important to a lot of carnivores, granivores and insectivores species as a temporary or permanent supply of these resources depending on their phenology and seasonality (Loiselle & Blake, 1994). On the other hand, the habitats with more complex vegetation

Seasonal variation of the avifauna in the present study was similar to that found in other tropical forest regions (Chamela region in Jalisco, Mexico and Maringá in Paraná, Brazil; Ornelas et al., 1993, Krügel & Anjos, 2000, respectively) where the species richness was greater from October to November. Our results show that species composition did not differ significantly across the 24 study sites, and similarities of different levels among the sites were common. This could suggest that most tropical forest patches still have suitable habitats that ensure availability of food, nesting sites, and protective cover for the species but are still vulnerable to persistent encroachment evident around them. In the long term this could jeopardize the ability to sustain particular bird species, especially forest-

A considerable amount of species associated with secondary forest, open areas, clearings and forest edges remain abundant and are likely to increase in regions with small isolated forest fragments (Bierregaard & Lovejoy, 1989, Thiollay, 1992). Generalist birds, which change their diet from fruit to insects or vice-versa, are also favored in small patches (Willis, 1979). Mota (1990) found increasing, devastation of pristine areas. Although some general tendencies were observed for certain bird groups, the effects of forest fragmentation are certainly different for each species. A study in forest fragments (in Maryland) suggested that the impacts of forest fragmentation on bird communities are complex, species specific and not related only to fragment area or fragment isolation (Lynch & Whigham, 1984). The increase in species richness with fragmentation was primarily due to the addition of several

These species showed a lower frequency possibly because they were represented by few individuals, and are more sensitive to forest change and fragmentation than more widespread species, as patterns that has been shown before (Fjeldsa, 1999, Renjifo, 2001). Priority species (i.e., endemic and threatened) are important contributors to biodiversity because their restricted distributions make them globally rare and particularly vulnerable to population declines or extinction (Terborgh & Winter, 1983, Diamond 1986). Species with small ranges are also less abundant at a local scale than large-range species (Brown, 1995). The birds may demonstrate a differential response to forest fragmentation (Hobson & Bayne, 2000, Fahrig, 2003) or that probably bird species richness in the study area can be affected by other factors, such as floristic diversity, and vegetation composition and structure (Gillespie & Walter, 2001). Other effects, such as the extent and nature of the fragments edges, fragment connectivity, or fragment shape (Bierregaard et al., 1992, Laurance & Bierregaard, 1997, Cornelius et al., 2000), might be more important than forest cover in predicting the number of species found in the area (Ramírez-Albores, 2010). As for other studies, it is expected that the effect of forest cover would affect bird species richness

Diet composition was similar among habitat types, with greatest representation by insectivores and insectivores/frugivores, and decreasing representation by nectarivores and granivores. According to Petit et al. (1999) and Karr (1990), this distribution of foraging guild memberships is typical of that found in tropical forest. Tropical forest fragments resulting from human disturbance of a continuous forest are isolated more rapidly. The remaining areas suffer of progressive degradation due to isolation, which, in the long term, jeopardizes the survival of several species. In tropical environments, modified habitats are very important to a lot of carnivores, granivores and insectivores species as a temporary or permanent supply of these resources depending on their phenology and seasonality (Loiselle & Blake, 1994). On the other hand, the habitats with more complex vegetation

migrants that were associated with edge habitats and secondary forest.

(Kattan et al., 1994, Laurance & Bierregaard, 1997).

dependent bird, threatened and endemic.

structure and formed by several layers of coverage are mainly species of insectivores, frugivores and nectar habits (Rappole et al., 1993). The results in this study are consistent with the above, as modified habitats had a higher proportion of species and individuals of carnivores and granivores habits compared to the original habitats (i.e., tropical deciduous and semideciduous forest).

In general, the variety of habitats present in the study region seems to contribute a high proportion of species, especially considering the number of species occurring in tropical deciduous forest (203). This may be due to the structural complexity that makes an ecosystem with greater species richness in Mexico (Ceballos & García, 1995, Ceballos et al., 2010), and the fact of having a greater horizontal and vertical stratification with respect to others, thus generating increased availability of habitats and ecological niches (Blake & Loiselle, 1991, McIntyre, 1995, Villard et al., 1999), as the plant structure determines the amount and distribution of resources used by birds. The differences in diversity and richness found indicate that the tropical deciduous forest, tropical semideciduous forest, and secondary forest show a greater richness compared to other modified habitats (i.e., agricultural fields, cattle pasture). This coincides with other studies conducted in tropical environments and indicates that the original habitat loss directly affects the presence, abundance and persistence of species (Kattan et al., 1994, Laurance & Bierregaard, 1997). The results of this study suggest that species richness and diversity of habitats ranging from the study. Natural habitats (tropical deciduous forest, tropical semideciduous forest and secondary forest) appear to be attractive to a larger number of bird species, as both the richness and diversity were higher in these, which is consistent with other studies (Estrada et al., 1997, Petit et al., 1999, Blake & Loiselle, 2001, Bojorges & Lopez-Mata, 2005).

Modified habitats (agricultural fields, living fences, cattle pasture) had a significant contribution to the bird species richness in the study area. These habitats provide roosting sites and food resources (Lynch, 1989). This is consistent with that reported by Estrada et al. (1997), which brings a richness of 226 in the region of Los Tuxtlas (Veracruz, Mexico), finding 79% of the species found in forest areas, 80% farmland, 43% in living fences and only 5% in grassland/pasture. In addition, Petit et al. (1999) in the central area of Panama found that species richness in modified habitats (i.e., shade coffee plantations, residential areas, grasslands and pine plantations) is equal or similar to the natural habitat. However, live fences exhibited the highest species richness (77 species) from modified habitats (i.e., agricultural fields, cattle pasture), probably because their plant structure is more complex and diverse. This is consistent with other studies (i.e., Villaseñor, 1993, Villaseñor & Hutto, 1995, Morales, 2002), which state that living fences can be very attractive to a large number of individuals and species of birds, and also can support high densities as they provide food resources, roosting sites and shelter (Villaseñor & Hutto, 1995). For example, in the study area, some birds prefer to use corridors or live fences instead of open or cleared areas (Wegner & Merriam, 1979) and turnover rates are significantly more frequent along corridors connected to original habitats or with other corridors (Hass, 1995, Machtans et al., 1996). In the case where the original habitat remains, the complexity of vegetation provides alternative sites for some species, partially offsetting the fragmentation and allowing the persistence of resident and migratory species (Morales, 2002).

The similarity between the habitats types of study area indicates the existence of a high turnover of species and an apparent high connectivity between them. Suggesting that both the configuration of the environment (i.e., landscape, habitat and microhabitat) and the available amount thereof would not be equally important in the distribution of species (Karr

Relationships Between Bird Species Richness

**Species Migratory**

and Natural and Modified Habitat in Southern Mexico 197

pastures), Lf (living fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af (agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq (aquatics and subaquatics). Diet categories: I (insectivore), C (carnivore), O

*Dendrocygna autumnalis* R A x *Anas discors* M A x

*Tachybaptus dominicus* R A x *Podilymbus podiceps* M A x *Pelecanus occidentalis* M C x *Phalacrocorax brasilianus* R C x *Anhinga anhinga* R C x *Ardea herodias* M C x *Ardea alba* M C x *Egretta thula* M C x *Egretta caerulea* M C x *Egretta tricolor* M C x

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

(omnivore), N (nectarivore), A (aquatic), G (granivore), F (frugivore).

**Diet or subdiet**

*Crypturellus soui* R Ox *Crypturellus cinnamomeus* R O xx

*Ortalis vetula* R GF x x x x *Penelope purpurascens* R GF x x *Colinus virginianus* R G x xx

*Bubulcus ibis* R I x

*Butorides virescens* M C x *Nycticorax nycticorax* M C x *Nyctinassa violacea* R C x *Coragyps atratus* R C x x x x x x x x *Cathartes aura* R C x x x x x x x x *Pandion haliaetus* M C x xx *Elanus leucurus* R C xx *Rostrhamus sociabilis* R Cx x *Ictinia mississippiensis* M C x x x xx *Accipiter striatus* M C x xx x x x *Accipiter cooperii* M C x x xx xx *Buteogallus anthracinus* R Cx x xx *Buteo magnirostris* R C x xx xx *Buteo nitidus* R Cx x xx *Buteo brachyurus* R Cx x *Buteo swainsoni* M Cx x x *Buteo albicaudatus* R C xx *Buteo jamaicensis* R Cx x x x x *Caracara cheriway* R C xx *Herpetotheres cachinnans* R C xx *Falco sparverius* R C x x

**status**

1990) and could confer changes in the composition of the community birds (Blake & Loiselle, 2001). Although the conservation of bird species depends on a clear understanding of their habitat requirements and the physical and biotic processes that keep (Askins, 2000), has been established that the combination of natural and modified habitats leading to new opportunities differential exploitation of space (Willson, 1974) and diversity of bird species is related to landscape diversity, so that conservation of the latter ensures the preservation of species diversity (Bôhning-Gaese, 1997).

The distribution of birds in different physiographic regions of Chiapas is highly heterogeneous (Rangel-Salazar et al*.*, 2005), and may also occur heterogeneity within each region, or even between adjacent physiographic regions (González-Domínguez, 1998). As the behavior of the birds in the Central Depression of Chiapas can be shared or influenced by other regions such as Montañas del Este, Altiplano Central, and even by the Sierra Madre de Chiapas, giving it the ability to host species of these regions (Altamirano, 2004). Biogeographic research biotic transition zones are an essential part of the study of the processes that govern the distribution and diversity of organisms (Williams et al., 1996). In this regard, species richness captures a fundamental aspect of spatial patterns of biodiversity (Koleff et al., 2003). Studying diversity patterns among the cells used in the present analysis helps to generate hypotheses about the processes that contribute to defining the current distribution patterns in the Isthmus, as the spatial turnover of species may reflect deterministic processes such as adaptation of species to different conditions, speciation, and responses to weather events or other historical effects (Condit et al., 2002).

## **5. Conclusions**

Given the continued fragmentation of natural habitats and according to the results of this study, addition and maintenance of natural and modified habitats are necessary for survival and reproduction of many species of birds in the study area. The study area, like many other regions of the country is being affected by anthropogenic factors, particularly the expansion of the agricultural frontier, forest fires, population growth and livestock, which directly affects wildlife populations wild. Studies of diversity and species richness are approximations that represent the basis to further evaluate information by monitoring the changes associated with environmental factors and especially anthropogenic. And the visualization of the biogeographic patterns over changes in species richness according to changes land permits to locate the sites that have been modified over time. This method facilitates the identification of priority areas for conservation because key to the survival of species groups threatened and endemic. Understanding the patterns of richness is closely linked the establishment of actions at the federal, regional and local levels, as they reflect as conditions of land use change are affecting populations. The need to make a stock assessment and particular requirements of each species; can support the planning, implementation and evaluation. Additional conservation actions can help to assure that these viable long-term populations are sufficient to retain species.

## **6. Appendix**

Bird species record from 24 sites of the Central Depression Chiapas. Taxonomy and order species follow AOU (2010). Migratory status: resident (R), migratory (M; including winter visitor, summer resident and transit). Habitat types: Tdf (tropical deciduous forest), C (cattle

1990) and could confer changes in the composition of the community birds (Blake & Loiselle, 2001). Although the conservation of bird species depends on a clear understanding of their habitat requirements and the physical and biotic processes that keep (Askins, 2000), has been established that the combination of natural and modified habitats leading to new opportunities differential exploitation of space (Willson, 1974) and diversity of bird species is related to landscape diversity, so that conservation of the latter ensures the preservation

The distribution of birds in different physiographic regions of Chiapas is highly heterogeneous (Rangel-Salazar et al*.*, 2005), and may also occur heterogeneity within each region, or even between adjacent physiographic regions (González-Domínguez, 1998). As the behavior of the birds in the Central Depression of Chiapas can be shared or influenced by other regions such as Montañas del Este, Altiplano Central, and even by the Sierra Madre de Chiapas, giving it the ability to host species of these regions (Altamirano, 2004). Biogeographic research biotic transition zones are an essential part of the study of the processes that govern the distribution and diversity of organisms (Williams et al., 1996). In this regard, species richness captures a fundamental aspect of spatial patterns of biodiversity (Koleff et al., 2003). Studying diversity patterns among the cells used in the present analysis helps to generate hypotheses about the processes that contribute to defining the current distribution patterns in the Isthmus, as the spatial turnover of species may reflect deterministic processes such as adaptation of species to different conditions, speciation, and

Given the continued fragmentation of natural habitats and according to the results of this study, addition and maintenance of natural and modified habitats are necessary for survival and reproduction of many species of birds in the study area. The study area, like many other regions of the country is being affected by anthropogenic factors, particularly the expansion of the agricultural frontier, forest fires, population growth and livestock, which directly affects wildlife populations wild. Studies of diversity and species richness are approximations that represent the basis to further evaluate information by monitoring the changes associated with environmental factors and especially anthropogenic. And the visualization of the biogeographic patterns over changes in species richness according to changes land permits to locate the sites that have been modified over time. This method facilitates the identification of priority areas for conservation because key to the survival of species groups threatened and endemic. Understanding the patterns of richness is closely linked the establishment of actions at the federal, regional and local levels, as they reflect as conditions of land use change are affecting populations. The need to make a stock assessment and particular requirements of each species; can support the planning, implementation and evaluation. Additional conservation actions can help to assure that

Bird species record from 24 sites of the Central Depression Chiapas. Taxonomy and order species follow AOU (2010). Migratory status: resident (R), migratory (M; including winter visitor, summer resident and transit). Habitat types: Tdf (tropical deciduous forest), C (cattle

responses to weather events or other historical effects (Condit et al., 2002).

these viable long-term populations are sufficient to retain species.

of species diversity (Bôhning-Gaese, 1997).

**5. Conclusions**

**6. Appendix**

pastures), Lf (living fences), Rf (gallery forest), Tsf (tropical semideciduous forest), Sf (secondary forest), Af (agricultural fields), To (tropical oak forest), Ur (urban and suburban areas) and Aq (aquatics and subaquatics). Diet categories: I (insectivore), C (carnivore), O (omnivore), N (nectarivore), A (aquatic), G (granivore), F (frugivore).

Relationships Between Bird Species Richness

**Species Migratory**

*Phaethornis striigularis* R N x *Florigusa mellivora* R N x x

*Archilochus colubris* M N x *Trogon melanocephalus* R IF x x *Trogon violaceus* R IF x x

**status**

*Phaethornis longirostris* R N x x

*Amazilia yucatanensis* R N x x *Amazilia viridifrons* R N x x x *Eupherusa eximia* R N x x x *Lamprolaima rhami* R N x x *Heliomaster longirostris* R N x x *Tilmatura dupontii* R N x x x

*Momotus mexicanus* R CIF x x x x *Momotus momota* R CIF x x x

*Aulacorhynchus prasinus* R GF x x *Pteroglossus torquatus* R GF x x *Ramphastos sulfuratus* R GF x x

*Sphyrapicus varius* M I x x x

*Colaptes rubiginosus* R IF x x x *Dryocopus lineatus* R I x x x

*Automolus ochrolaemus* R I x x

*Sclerurus guatemalensis* R I x *Synallaxis erythrothorax* R I x

*Dendrocincla homochroa* R I x

*Xiphorhynchus flavigaster* R I x x *Lepidocolaptes souleyetii* R I x x x

*Campephilus guatemalensis*

*Dendrocolaptes sanctithomae*

and Natural and Modified Habitat in Southern Mexico 199

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

*Panyptila sanctihieronymi* R I x x x x x x

*Colibri thalassinus* R N x x x *Chlorostilbon canivetii* R N x x x x *Amazilia beryllina* R N x x x x *Amazilia tzacatl* R N x x x x x

*Megaceryle torquata* R C x x *Megaceryle alcyon* M C x x *Chloroceryle amazona* R C x x *Chloroceryle americana* R C x x

*Melanerpes aurifrons* R IF x x x x x

*Picoides scalaris* R I x x x

R I x x x

R I x

*Taraba major* R I x x x *Thamnophilus doliatus* R I x x x x

*Charadrius vociferus* M A x *Himantopus mexicanus* R A x *Jacana spinosa* R A x *Actitis macularius* M A x *Tringa solitaria* M A x *Columba livia* R G x

*Zenaida asiatica* R GF x x x x x

*Columbina inca* R G x x x x x

*Aratinga nana* R GF x x x *Aratinga canicularis* R GF x x x x x

*Amazona autumnalis* R GF x x x

*Crotophaga sulcirostris* R GIF x x x x x

*Chordeiles acutipennis* R I x x x x *Nyctidromus albicollis* R I x x x x

*Chaetura vauxi* R I x x x x x x x

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

*Falco columbarius* M C x x x *Falco femoralis* R C x x *Falco peregrinus* M A x x

**Species Migratory**

**status**

*Patagioenas flavirostris* R GF x x x

*Claravis pretiosa* R GF x x

*Geotrygon montana* R GF x x *Aratinga holochlora* R GF x x

*Coccyzus minor* R CI x x x *Piaya cayana* R CIF x x x

*Megascops guatemalae* R C x x x

*Glaucidium brasilianum* R CI x x x *Ciccaba virgata* R C x x

*Caprimulgus ridgwayi* R I x x x

*Amazona albifrons* R GF x x

*Dromococcyx phasianellus* R CI x x *Morococcyx erythropygus* R CI x x x

*Patagioenas nigrirostris* R GF x x x

*Zenaida macroura* M GF x x x x

*Columbina minuta* R G x x x

*Columbina passerina* R G x x x

*Columbina talpacoti* R G x x x

*Leptotila verreauxi* R GF x x x x x

*Tapera naevia* R CIF x x

*Geococcyx velox* R CI x x x

*Tyto alba* R C x x x

*Pulsatrix perspicillata* R C x x x x

*Streptoprocne zonaris* R I x x x x x x

*Aeronautes saxatalis* R I x x x x x x

Relationships Between Bird Species Richness

**Species Migratory**

*Vireo leucophrys* R IF x

*Troglodytes aedon* R I x *Cistothorus platensis* R I x *Uropsila leucogastra* R I x x *Henicorchia leucophrys* R I x x *Microcerculus marginatus* R I x *Ramphocaenus melanurus* R I x x

*Catharus aurantiirostris* R IF x *Catharus minimus* M IF x *Catharus ustulatus* M IF x x *Hylocichla mustelina* M IF x

*Oreothlypis celata* M IF x

*Dendroica pensylvanica* M IF x

*Dumetella carolinensis* M IF x x

*Oreothlypis pinus* M I x x

**status**

*Vireo flavoviridis* M IF x x

*Cyanocorax morio* R O x x x x *Progne chalybea* R I x x x

*Riparia riparia* M I x x

*Catherpes mexicanus* R I x x x

and Natural and Modified Habitat in Southern Mexico 201

*Vireo philadelphicus* M I x x x x x *Vireo olivaceus* M IF x x x x x x

*Hylophilus decurtatus* R I x x x x *Cyclarhis gujanensis* R IF x x x x *Calocitta formosa* R O x x x x x x x *Cyanocorax yncas* R O x x x x x

*Tachycineta albilinea* M I x x x x x x

*Stelgidopteryx serripennis* R I x x x x x x x

*Petrochelidon pyrrhonota* M I x x x *Petrochelidon fulva* M I x x x *Hirundo rustica* M I x x x

*Thryothorus maculipectus* R I x x x x x *Thryothorus pleurostictus* R I x x x x x x x *Thryothorus modestus* R I x x x x x x

*Polioptila caerulea* R I x x x x x x *Polioptila plumbea* R I x x x x x x

*Turdus grayi* R IF x x x x x x x

*Oreothlypis ruficapilla* M IF x x x x x x *Parula americana* M I x x *Parula pitiayumi* R I x x *Dendroica petechia* M IF x x x x x

*Mimus gilvus* R IF x x x *Bombycilla cedrorum* M IF x x x x

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

R Ix x

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

*Cercomacra tyrannina* R Ix x *Grallaria guatemalensis* R I xx *Ornithion semiflavum* R Ix *Camptostoma imberbe* R Ix x x x *Myiopagis viridicata* R Ix x *Elaenia flavogaster* R IF x x

*Oncostoma cinereigulare* R I x xx *Poecilotriccus sylvia* R I xx *Rhynchocyclus brevirostris* R Ix x x *Xenotriccus callizonus* R Ix x *Contopus cooperi* M Ix x *Contopus pertinax* R Ix x x x x x *Contopus virens* M Ix x *Contopus cinereus* R Ix x x x x x *Empidonax virescens* M Ix x *Empidonax traillii* M IF x x x *Empidonax albigularis* M IF x xx x x *Empidonax minimus* M Ix x *Sayornis nigricans* R Ix x x *Pyrocephalus rubinus* R I x x *Rhytipterna holerythra* R Ix x *Myiarchus tuberculifer* R IF x x x *Myiarchus cinerascens* M IF x x x *Myiarchus nuttingi* R IF x xx x x *Myiarchus tyrannulus* R IF x x x x x x x x *Pitangus sulphuratus* R CIF x x x x x x x x *Megarhynchus pitangua* R CIF x x x x x x *Myiozetetes similis* R IF x x x x x x x x *Myiodynastes luteiventris* M IF x x x x x x x x *Legatus leucophaius* M Ix x *Tyrannus melancholicus* R IF x x x x x x x *Tyrannus vociferans* M IF x x x *Tyrannus verticalis* M IF x x x *Tyrannus tyrannus* M I x x x *Tyrannus forficatus* M IF x x x *Pachyramphus aglaiae* R IF x x *Tityra semifasciata* R IF x x x x *Vireo griseus* M IF x x x *Vireo bellii* M Ix x *Vireo solitarius* M IF x x x x x x x *Vireo huttoni* R Ix x *Vireo gilvus* M IF x x x x

**Species Migratory**

*Leptopogon amaurocephalus*  **status**

Relationships Between Bird Species Richness

**Species Migratory**

*Icterus maculialatus* R IF x

*Amblycercus holosericeus* R IF x x *Cacicus melanicterus* M IF x *Psarocolius wagleri* R GF x x *Psarocolius montezuma* R GF x x

**7. Acknowledgements**

**8. References**

**status**

and Natural and Modified Habitat in Southern Mexico 203

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

*Passerina caerulea* R GI x x x *Passerina cyanea* M GI x x x *Passerina versicolor* R GIF x x x x *Passerina ciris* M GI x x *Sturnella magna* R GI x x

*Dives dives* R GIF x x x x x x x *Quiscalus mexicanus* R O x x x x x x x x x *Molothrus aeneus* R GIF x x x x x x x *Icterus prosthemelas* R IF x x x *Icterus wagleri* R IF x x x

*Icterus spurius* M IF x x x *Icterus mesomelas* R IF x x x *Icterus pustulatus* R IF x x x x x x x *Icterus gularis* R IF x x x x x x *Icterus galbula* M IF x x x x

*Euphonia affinis* R IF x x x x x *Euphonia hirundinacea* R IF x x x x x x *Euphonia elegantissima* R IF x x x x x *Carpodacus mexicanus* R G x x x *Spinus psaltria* R GF x x x x x x x *Passer domesticus* R O x

Thanks to El Colegio de la Frontera Sur (ECOSUR-San Cristobal) and the National Council of Science and Technology in Mexico (CONACyT) for research facilities and financial support. Thanks to J. Rangel, J. León, M. Martínez and G. Escalona for comments and suggestions made during the elaboration and conclusion of the Master´s thesis. Thanks are due to the people of the local communities that we visited during fieldwork in study area. Thanks also to Museo de Zoología "Alfonso L. Herrera", Facultad de Ciencias, UNAM for

Aleixo, A. & Vielliard, M. E. (1995). Composição e dinãmica da avifauna da mata de Santa

Genebra, Campinas, São Paulo, Brasil. *Revista Brasileira de Zoología,* Vol.12, No.3,

access to data base of "Atlas de las Aves de México", especially to A. Gordillo.

(September 1995), pp. 493-511, ISSN 0101-8175.

*Cyanocompsa parellina* R G x x x x

*Dendroica magnolia* M IF x x x x x x x

*Dendroica virens* M IF x x x x x x x *Dendroica townsendi* M IF x x *Dendroica occidentalis* M IF x x x x

*Dendroica graciae* R IF x x x x *Mniotilta varia* M I x x x x x x

*Geothlypis trichas* M I x x *Geothlypis poliocephala* R I x x *Wilsonia citrina* M I x x x *Wilsonia pusilla* M IF x x x x x x x *Wilsonia canadensis* M IF x x x

*Basileupterus rufifrons* R I x x x *Icteria virens* M I x x

*Thraupis episcopus* R IF x x *Thraupis abbas* R IF x x x x x

*Saltator coerulescens* R IF x x x *Saltator atriceps* R IF x x x *Volatinia jacarina* R G x x

*Oryzoborus funereus* R G x x *Tiaris olivaceus* R G x x x x *Arremonops rufivirgatus* R GI x x x x x x x

*Aimophila botterii* R G x x *Aimophila rufescens* R GI x x

*Piranga ludoviciana* M IF x x x x x

*Pheucticus chrysopeplus* R IF x x x x x x

*Sporophila torqueola* R G x x x

*Euthlypis lachrymosa* R I x x x x

*Eucometis penicillata* R IF x x

*Piranga rubra* M IF x x x

*Habia rubica* R IF x x x *Habia fuscicauda* R IF x x x

*Pheucticus ludovicianus* M IF x x *Granatellus venustus* R IF x x x *Granatellus sallaei* R IF x x x

*Cyanerpes cyaneus* R IN x x

**Habitat types Tdf Tsf Sf To Rf Af C Lf Ur Aq**

**Diet or subdiet**

**Species Migratory**

**status**

*Dendroica coronata* M IF x

*Dendroica fusca* M I x

*Protonotaria citrea* M I x *Helmitheros vermivorum* M I x *Seiurus aurocapilla* M I x x x *Seiurus noveboracensis* M I x x x *Oporornis formosus* M I x x *Oporornis tolmiei* M I x

## **7. Acknowledgements**

Thanks to El Colegio de la Frontera Sur (ECOSUR-San Cristobal) and the National Council of Science and Technology in Mexico (CONACyT) for research facilities and financial support. Thanks to J. Rangel, J. León, M. Martínez and G. Escalona for comments and suggestions made during the elaboration and conclusion of the Master´s thesis. Thanks are due to the people of the local communities that we visited during fieldwork in study area. Thanks also to Museo de Zoología "Alfonso L. Herrera", Facultad de Ciencias, UNAM for access to data base of "Atlas de las Aves de México", especially to A. Gordillo.

## **8. References**

Aleixo, A. & Vielliard, M. E. (1995). Composição e dinãmica da avifauna da mata de Santa Genebra, Campinas, São Paulo, Brasil. *Revista Brasileira de Zoología,* Vol.12, No.3, (September 1995), pp. 493-511, ISSN 0101-8175.

Relationships Between Bird Species Richness

México, D. F., México.

970-9000-02-0, México, D.F., México.

http://www.conabio.gob.mx.

2002), pp. 666-669, ISSN 1095-9203.

ISBN:<PHONE_NUMBER>, New York, USA.

(January 1997), pp. 19-43, ISSN 1572-9710.

(January 1999), pp. 47-62, ISSN 1474-0001.

http://www.esri.com

New York, USA.

592X.

and Natural and Modified Habitat in Southern Mexico 205

Ceballos, G.; García, A.; Salazar, I. & Espinoza, E. (2010). Conservación de vertebrados de

Challenger, A. 1998. *Utilización y conservación de los ecosistemas terrestres de México. Pasado,*

CONABIO (Comisión Nacional para Uso y Conservación de la Biodiversidad). (2004).

CONANP (Comisión Nacional de Áreas Naturales Protegidas). (2011). Áreas naturales protegidas de Chiapas. 02.2011, Available in http://www.conanp.gob.mx. Condit, R.; Pitman, N.; Leigh Jr., E.G.; Chave, J.; Terborgh, J.; Foster, R. B.; Núñez, P.;

Diamond, J. (1986). The design of a nature reserve system for Indonesian New Guinea. In:

Ehrlich, P. R.; Dobkin, D. S. & Wheye, D. (1988). *The birders´ handbook: a field guide to the*

Escalante, P.; Navarro, A. G. & Peterson, A. T. (1993). A geographic, ecological and historical

ESRI. (1999). Arc view GIS ver. 3.2. Environmental Systems Research Inc., USA. Available in

Estrada, A.; Coates-Estrada, R. & Meritt, D. A. (1997). Anthropogenic landscape changes and

Faaborg, F.; Brittingham, M.; Donovan, T. & Blake, J. (1995). Habitat fragmentation in the

Fahrig, L. (2003). Effects of habitat fragmentation on biodiversity. *Annual Review of Ecology,*

Fjeldsa, J. (1999). The impact of human forest disturbance on the endemic avifauna of the

Sinauer Associates Inc., ISBN<PHONE_NUMBER>, Sunderland, USA.

University Press, ISBN 019506674X, New York, USA.

selvas secas: patrones de distribución, endemismo y vulnerabilidad. In: Diversidad, amenazas y áreas prioritarias para la conservación de las selvas secas del Pacífico de México, G. Ceballos, L. Martínez, A. García, E. Espinoza, J. Bezaury & R. Dirzo (Eds.), 369-386, Fondo de Cultura Económica-CONABIO, ISBN<PHONE_NUMBER>-31-1,

*presente y futuro*. CONABIO-Instituto de Biología, UNAM-Sierra Madre A. C., ISBN

*Regiones Terrestres Prioritarias*. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, Scale 1:1,000,000, México. 02.2011, Available in

Aguilar, S.; Valencia, R.; Villa, G.; Muller-Landau, H. C.; Losos, E. & Hubbell, S. P. (2002). Beta-Diversity in Tropical Forest Trees. *Science*, Vol.295, No.5555, (January

*Conservation biology: the science of scarcity and diversity*; M. E. Soúle (Ed.), 485-503,

*natural history of North America birds,* Simon and Shuster Adult Publishing Group,

analysis of land bird diversity in México. In: *The biological diversity of Mexico: Origins and distribution,* T. P. Ramamorthy, R. Bye, A. Lot & J. Fa (Eds.), 281-307, Oxford

avian diversity at Los Tuxtlas, Mexico. *Biodiversity and Conservation,* Vol.6, No.1,

temperate zone. In: *Ecology and management of Neotropical migratory birds,* T. E. Martin & D. M. Finch (Eds.), 357-380, Oxford University Press, ISBN<PHONE_NUMBER>,

*Evolution and Systematic,* Vol.34, No.1, (November 2003), pp. 487-515, ISSN 1543-

Udzungwa Mountains, Tanzania. *Bird Conservation International,* Vol*.*9, No.1,

AOU (American Ornithologists´ Union). (2010). *Check-list of North American birds*. In: American Ornithologists´ Union, 03. 2011, Available from http://www.aou.org. Altamirano G.O., M. A. (2004). *Obtención de la riqueza de aves y selección de especies susceptibles*

Anjos, L. Dos. (2001). Bird communities in five Atlantic forest fragments in southern Brazil. *Ornitología Neotropical,* Vol.12, No.1, (March 2001), pp. 11-27, ISSN 1075-4377. Arizmendi, M. C.; Berlanga, H.; Márquez, L.; Navarijo, L. & Ornelas, J. F. (1990). *Avifauna de*

Nacional Autónoma de México, ISBN 968-36-1571-6, México, D. F., México. Arizmendi, M.C. & Márquez, L. (2000). *Áreas de importancia para la conservación de las aves en*

Arriaga, L.; Espinoza, J. M.; Aguilar, C.; Martínez, E.; Gómez, L.; Loa, E. & Larson, J. (2000).

Askins, R. A. (2000). *Restoring North America´s birds*. Yale University Press, ISBN

Bierregaard, R. O. & Lovejoy, T. E. (1989). Effects of forest fragmentation on Amazonian

Blake, J. G. & Loiselle, B. A. (1991). Variation in resource abundance effects capture rates of

Blake, J. G. & Loiselle, B. A. (2001). Birds assemblages in second-growth and old-growth

Bôhning-Gaese, K. (1997). Determinants of avian species richness at different spatial scales. *Journal of Biogeography,* Vol.24, No.1, (January 1997), pp. 49-60, ISSN 1365-2699. Bojorges, B. J. & López-Mata, L. (2005). Riqueza y diversidad de especies de aves en una

*Mexicana (nueva serie),* Vol.21, No.1, (April 2005), pp. 1-20, ISSN 0065-1737. Bolger, D. T.; Scott, T. A. & Rotenberry, J. T. (1997). Breeding bird abundance in an

Brooks, R. P. & Croonquist, M. J. (1990). Wetland habitat and trophic response guilds for

Ceballos, G. (1995). Vertebrate diversity, ecology and conservation in neotropical dry forest,

220, Cambridge University Press, ISBN 978-0-521-11284-0, Cambridge, UK. Ceballos, G. & García, A. (1995). Conserving neotropical biodiversity: the role of dry forest

*México*. CIPAMEX, ISBN 970-18-4319-3, México, D.F., México.

Tuxtla Gutiérrez, México.

9780300093162, Nueva Haven, USA.

pp. 114-130, ISSN 0004-8038.

1353, ISSN 1523-1739.

(April 2001), pp. 304-326, ISSN 0004-8038.

No.2, (April 1997), pp. 406–421, ISSN 1523-1739.

F., México.

ISSN 0044-5967.

*de monitoreo en la zona noroeste en el estado de Chiapas*. Instituto de Historia Natural y Ecología. Informe final SNIB-CONABIO proyecto No. Y018, México, D.F., México. Álvarez del Toro, M. (1980). *Las aves de Chiapas*. (2a. ed.), Gobierno del Estado de Chiapas,

*la región de Chamela, Jalisco*. Cuadernos del Instituto de Biología No. 4, Universidad

*Regiones terrestres prioritarias de México*. CONABIO, ISBN-10<PHONE_NUMBER>, México, D.

understory bird communities. *Acta Amazonica,* Vol.19, No.1, (1989), pp. 215-241,

birds in three lowland habitats in Costa Rica. *Auk,* Vol.108, No.1, (January 1991),

forest, Costa Rica: perspectives from mist nest and point counts. *Auk,* Vol.118, No.2,

selva mediana subperennifolia en el centro de Veracruz, México. *Acta Zoológica*

urbanizing landscape in coastal southern California. *Conservation Biology,* Vol.11,

wildlife species in Pennsylvania, Journal of the Pennsylvania Academy of Science, Vol.64, No.3, (1990), pp. 93-102, ISSN 1044-6753. in a mangementBrown, J. H. (1995). *Macroecology*. University of Chicago Press, ISBN<PHONE_NUMBER>, Chicago, USA.

In: *Seasonally dry tropical forest,* S. H. Bullock, H. A. Mooney & E. Medina (Eds.), 195-

in western Mexico. *Conservation Biology,* Vol.9, No.9, (December 1995), pp. 1349-

Relationships Between Bird Species Richness

http://www.inegi.org.mx.

Washington, D. C., USA.

0225468984, Chicago, USA.

pp. 287-324, ISSN 0006-3207.

1366-1379, ISSN 1523-1739.

9780309037396, Washington, D. C., USA.

9780300054484, New Haven, USA.

1994), pp. 138-146, ISSN 1523-1739.

(December 2000), pp. 315-330, ISSN 1075-4377.

México.

2656.

and Natural and Modified Habitat in Southern Mexico 207

INEGI (Instituto Nacional de Estadística, Geográfica e Informática). (2004). *Cuaderno*

INEGI (Instituto Nacional de Estadística, Geográfica e Informática). (2006). *Cuaderno*

Janzen, D. H. (1988). Tropical dry forest: the most endangered major tropical ecosystem. In: