FOREST MANAGEMENT AND BIODIVERSITY: A STUDY OF AN INDICATOR GROUP IN SOUTHERN

Yariely del Rocío Balam-Ballote and Jorge L. León-Cortés

SUMMARY

Previous studies on succesional dynamics of insect commu- use and geographic range for 63 was detected nities have revealed important changes along succesional sta- (χ²= 13.26, df= 1, p<0.005). Species with narrow geographic ges among community structures and insect life histories. This ranges were usually grouped in unmodified habitats, whereas work evaluates the changes and diversity of a butterfly fauna in widespread species were much more likely to make some use a forest-managed landscape in Southern Mexico. One-hundred of human-modified habitats. A declining proportion of species and forty three species were collected belonging to 102 genera restricted to unmodified, or primary, habitats with increasing in five families. was the most diverse family with geographic range was quantified. Forest-management practices 54 species (38%). Rarefaction and rank abundance curves re- on local habitats as well as differences among species’ habi- corded in each forest stand suggested important differences for tat preferences and geographical distributions can account for species richness values at comparable levels of sampling effort. an increasing proportion of widespread species in the study Species abundances were positively correlated with their lands- landscape and comparable sites. Monitoring of insect indicator cape distribution sizes. Based on field observations and on li- groups in extractive landscapes is needed to establish guidelines terature review, a designation of habitat preferences for a set for ecologically, economically, and socially sustainable forest of species was given. A significant association between habitat zoning and use regimes.

Introduction adequate levels of species rich- are considered to toration schemes due to their ness under some semi-intense be adequate responders to quick reaction to changes as Succesional stages represent forest management could be habitat change (Brown and regards to habitat quality and an ecologically important case achieved through the mainte- Freitas, 2000; Caballero et quantity (Thomas, 1984; Ty- of change in diversity through nance of a patchwork that ide- al., 2009). Their sensitivity to ler et al., 1994; Wood and time. Former studies on suc- ally includes areas of primary disturbance and change allow Gillman, 1998). However, the cesional dynamics of insect forest, and a network of for- them to be good models for response of some butterfly communities reveal important est patches with a given man- the study in environmental species to habitat disturbance changes along succesional stag- agement regime (Wood and gradients, as is the case for depends upon the characteris- es among community structures Gillman, 1998). In addition, in dragonflies, beetles, moths, tics of particular organisms, and insect life histories, namely order to be considered a genu- and ants (Tyler et such as body size, dispersal host plant specificity, migra- inely sustainable method of for- al., 1994; Brown and Freitas, ability, life history, population tions, and body size, among est management, selective log- 2000; Arellano et al., 2008). size or rarity, because in some others. For example, excessive ging needs to be economically Butterflies in particular are cases, narrowly-distributed extraction of native tree spe- viable, provides a sustainable useful responders in environ- species seem to experience cies from forests can affect the yield of timber, and preserves mental studies because they no adverse effects of forest structure of communities, im- biodiversity (Bawa and Seidler, could decrease in abundance disturbance. Increasing habi- pacting the pattern of coloniza- 1998). However, in tropical sys- or even disappear due to (even tat fragmentation could even tion and diversity of insects in tems the costs of forest distur- subtle) changes in habitat benefit generalists, which have fragmented systems (Wood and bance, mostly caused by selec- quality, namely, forest cover recorded important population Gillman, 1998; Shahabuddin tive logging, upon biodiversity and vegetation composition increases as habitat becomes and Ponte, 2004; Waltz and have been poorly measured. and structure (León Cortés et more heterogeneous (Brandle Covington, 2004; Ohwaki et Can significant effects of forest al., 2004). Additionally, but- et al., 2002; León-Cortés et al., 2007). It is thought that an management upon biodiversity terflies have been proposed al., 2004; Waltz and Coving- optimal strategy to safeguard elements be expected? as useful responders in res- ton, 2004).

Keywords / Butterfly / Chiapas / Forestry Development / Forest Management / Indicator Species / Received: 06/02/2009. Modified: 05/17/2010. Accepted: 06/02/2010.

Yariely del Rocío Balam-Bal- Jorge L. León-Cortés. Ph.D. in Panamericana y Av. Periférico lote. MSc, El Colegio de la Ecology) University of Leeds, Sur S/N. San Cristóbal de las Frontera Sur (ECOSUR), Uni- UK. Researcher, ECOSUR, Casas, Chiapas, México. e- dad San Cristóbal de las Ca- México. Address: Departamen- mail: [email protected] sas, Chiapas, México. e-mail: to de Ecología y Sistemática [email protected] Terrestres, ECOSUR, Carr.

526 0378-1844/10/07/526-08 $ 3.00/0 JUL 2010, VOL. 35 Nº 7 MANEJO FORESTAL Y BIODIVERSIDAD: ESTUDIO DE UN GRUPO INDICADOR DE INSECTOS EN EL SUR DE MÉXICO Yariely del Rocío Balam-Ballote y Jorge L. León-Cortés RESUMEN

Estudios previos sobre dinámica sucesional de comunidades ciación significativa entre uso de hábitat y extensión geográfi- de insectos revelan cambios importantes a lo largo de los esta- ca para 63 especies de mariposas (χ²= 13,26, dl= 1, p<0,005). dios sucesionales en la estructura de las comunidades y la his- Las especies con extensiones geográficas reducidas se agruparon toria de vida de los insectos. Se evaluaron cambios y diversidad usualmente en hábitat no modificados, mientras que las de dis- de las mariposas en un paisaje con manejo forestal del sur de tribución amplia tendían a hacer uso de hábitat intervenidos. Se México, recolectándose 143 especies pertenecientes a 102 géne- registró una baja proporción de especies restringidas a hábitat ros de cinco familias, siendo Nymphalidae la más diversa (54 no modificados en relación con el incremento de su distribución especies; 38%). Las curvas de rarefacción y los niveles de abun- geográfica. Las prácticas locales de manejo forestal así como las dancia registradas en cada rodal sugirieron diferencias importan- diferencias en la preferencia de hábitats y distribución geográfica tes en riqueza de especies para niveles de esfuerzo de captura de las especies conllevan una creciente proporción de especies de comparables. La abundancia de especies estuvo correlacionada distribución amplia en la zona de estudio y sitios comparables. positivamente con el tamaño de su distribución. Con base en ob- Se requiere del monitoreo de grupos indicadores de insectos en servaciones de campo y revisión de la literatura, se designaron zonas de extracción para establecer pautas para la subdivisión y preferencias de hábitat para las especies. Se detectó una aso- régimen de uso ecológico, económica y socialmente sostenibles.

MANEJO DE SELVAS E BIODIVERSIDADE: ESTUDO DE UM GRUPO INDICADOR DE INSETOS NO SUL DO MÉXICO Yariely del Rocío Balam-Ballote e Jorge L. León-Cortés RESUMO

Estudos prévios sobre dinâmica sucesional de comunidades tat e extensão geográfica para 63 espécies de borboletas (χ²= de insetos têm revelado mudanças importantes ao longo dos 13,26, dl= 1, p<0,005). As espécies com extensões geográficas estadios sucessionais na estrutura das comunidades e a história reduzidas agruparam-se usualmente em hábitats não modifica- de vida dos insetos. Foram avaliadas mudanças e diversidad e dos, enquanto que as dispersas tendiam a fazer uso de hábitats das borboletas em uma selva protegida do sul do México,se re- interditados. Foi achada uma proporção em descenso de espé- colhendo 143 espécies pertencentes a 102 gêneros de cinco fa- cies restringidas a hábitats não modificados (primários) que au- milias, sendo Nymphalidae a mais diversa (54 espécies; 38%). mentavam sua extensão geográfica. As práticas de manejo flo- A rarefação e curvas de níveis de abundância registradas em restal local e as diferenças na preferência de distribuição das cada stand sugerem diferenças importantes em riqueza de espé- espécies levam a uma crescente proporção de espécies de ampla cies para níveis de esforço de captura comparáveis. A abundân- distribuição na área do estudo e locais comparáveis. Requere-se cia de espécies se correlacionou positivamente com a extensão de monitoramento de insetos indicadores em zonas de extração de sua distribuição. Com base em observações de campo e re- para estabelecer pautas para a subdivisão e regime de uso eco- visão da literatura, se designaram preferências de hábitat para lógico, económica e socialmente sustentáveis. as espécies. Houve associação significativa entre uso de hábi-

The objectives of this pa- mented in the landscape con- mer rains between June and ging and in lesser proportion per are 1) to assess the local sidered in the present study is October. Roughly 75% of the from farming and livestock (forest stand) and landscape as ecologically sustainable as total area of Coapilla is char- activities. In particular, logged diversity of butterflies in a it claims to be. acterized by steep slopes where areas through the Coapilla forest-managed landscape, 2) forest stands are imbibed; the landscape include a one-year to examine changes in spe- Methods remaining 25% of land consists intensive rotation scheme in cies composition as regards of flat areas located around which four selective cuttings to species habitat preferences Study site the Coapilla’s urban settlement (with different intensity) are and geographical distributions, (Gerez-Fernández et al., 2003). expected to allow regeneration and 3) to compare the overall The ejido Coapilla (hereafter The original Coapilla of forest stands over a 10-year butterfly diversity and species “Coapilla”, 17º05'10'', 17º11'02''N; landscape included mostly period. Very few biodiver- composition patterns of the 93º05'45'', 93º12'25''W; Figure 1) habitats, but an sity assessments have been study landscape as regards to is located in northern Chiapas, important proportion (16%, carried out in this area, but other sites for which butterfly México, and entitles a total area ~1000ha) of this landscape recent insect surveys suggest diversity assessments have of 13571ha. The area receives has now been converted into the area of Coapilla to be a been carried out. Ultimately, it an average annual rainfall of forest stands that include dif- site holding unusual numbers is discussed whether the type 1800mm. The climate is warm ferent successional stages, of endemic and narrowly-dis- of forest management imple- and humid with heavy sum- which mostly result from log- tributed species for beetles

JUL 2010, VOL. 35 Nº 7 527 and ants (Robert S. other (Magurran, 1988). Anderson and John T. In addition, the slopes of Longino, personal com- the equations of the lin- munication). No pre- ear regressions for each vious butterfly assess- rank-abundance distribu- ments had been carried tion were calculated and out in Coapilla. compared among forest stands. When the slope Butterfly sampling is zero, distribution of abundance values of the Before carrying out species is approximate- butterfly transects, a ly equivalent (Tokeshi, six-month pilot study 1993). was set up for identify- ing butterflies at sight. Butterfly abundance- Previous experience distribution in this and other sites relationship in Chiapas allowed us to identify specimens The abundance-dis- at sight (León-Cortés tribution relationship et al., 2004; Molina- for Coapilla's butterflies Martínez and León- was examined at the Cortés 2006; Marín et landscape scale. But- al., 2009). Nonethe- terfly distributions were less, certain species Figure 1. The location of the Coapilla landscape in Chiapas, Mexico proportions of forest complexes were caught stands occupied, where- using aerial nets; e.g., as butterfly abundances the representatives of tribes Tyler et al, (1994), and Gar- lation curves failed to reach were recorded from transect Achlyodidini, Pyrrhopy- wood and Lehman (2005), an asymptote, sample-based counts at forest stands. Lin- gini, Erynnini, , as well as the entomological rarefaction curves scaled to ear regressions between abun- and Polyommatinae. Caught collections of El Colegio de individuals and samples were dance and distribution were specimens were prepared and la Frontera Sur, the Zoology standardized and truncated to calculated using the ordinary identified in the Entomologi- Museum at Faculty of Science, the value of the lesser entry least squares method (mod- cal Collection of ECOSUR UNAM, and their curators, for of the total sample, and hence el 1; Gaston et al., 2000). (voucher specimens have been butterfly identifications. allowing for a comparison of Abundance was used as the deposited therein). relative richness values among dependent variable. Before In April 2007, fixed tran- Accumulation and dominance forest stands. In doing so it statistical analyses were con- sects of ~500m in length (Ta- curves was attempted to differentiate ducted, both abundance val- ble I) were established along under-sampling effects from ues and distributions were each of the forest stands. From To examine local diversity real differences among forest transformed so as to homog- May to November 2007, tran- (the recorded diversity at each stand values. enize variance. Abundance sect routes were walked be- forest stand), rarefaction curves In addition, for each forest values were log10 transformed, tween 9:00 and 15:00, un- of butterfly species were built stand, rank-abundance curves whilst distributions were Arc- der ideal conditions to record up for each forest stand in were constructed with the val- sine transformed (distribu- butterflies (Pollard and Yates, Coapilla, using the program ues grouped in descending tional data are proportions of 1993; Tyler et al., 1994). We EstimateS (Colwell, 1997). order, and abundance values all forest stands occupied). followed DeVries (1987, 1997), Since one or more accumu- were compared among each Habitat use and geographic Table I range categories The number of transects and habitat features for each forest stand in Coapilla, during 2007 Habitat preferences and geographic range data were Forest Fixed Transect Total Butterfly Number of Vegetation Number of analyzed for Papilionidae, stand number length number diversity flowering height tree species and Nymphalidae, of transects (m) of transects plants (cm) recorded based on the information pro- 1 2 500 13 59 (391) 33 59 12 vided by DeVries (1987), Llor- 2 1 800 7 44 (213) 15 61 4 ente-Bousquets et al. (2006), 3 1 800 9 55 (310) 30 91 12 Warren et al. (2010) and Tyler 4 2 500 11 42 (219) 45 126 10 et al. (1994). Only 63 of the 5 1 800 9 52 (248) 29 119 9 143 species recorded in Coap- 6 1 1500 11 79 (1070) 35 83 6 illa were included in the anal- 7 1 800 11 57 (6134) 39 107 2 ysis. Following Thomas (1991) 8 1 1500 9 58 (312) 40 124 13 and Léon-Cortés et al. (2003) 9 1 800 11 55 (427) 29 65 3 butterflies were classified as Forest 1 500 11 28 (108) 22 74 14 occupying unmodified habitat Total 12 102 (when no mention was made

528 JUL 2010, VOL. 35 Nº 7 sites for which butterfly diver- “Forest” recorded the highest sity assessments have been car- and poorest butterfly richness, ried out were compared using respectively. ANOVA tests (Sokal and Rohlf, In addition, rarefaction 1997). Comparisons were re- curves per forest stand sug- stricted to sites that exhibited gested important differences similar vegetation patterns, for species richness values mostly associated to moun- at comparable levels of sam- tainous habitats in southern pling effort (Figure 2). Levels Mexico, and for which adequate of species richness remained inventories are known to have comparable when changing been carried out (Armando sample units. Forest stand 6 Luis-Martínez, personal com- recorded the highest diversity munication). of butterflies, whereas the most pristine forest stand remained Results as the relatively poorest condi- tion in Coapilla. Butterfly diversity Abundance ranks Figure 2. Rarefaction curves per forest stand (numbers on the right). The A total of 9432 butter- distributions discontinuous vertical arrow indicates standardized sample values at which fly individuals was recorded forest stands were compared. and grouped in 143 species Abundance ranks distribu- and five families. tions differed among forest that they occupy sec- Fifty four species stands studied (Figure 3). Less ondary or agricul- were recorded in disturbed forest stands showed tural habitats), or as the Nymphalidae abundance curves with gentler occupying secondary, family (38%), slopes in comparison to most agricultural, or oth- 32 in Hesperi- disturbed habitats. er human-modified idae (22%), 27 in habitats. In addition, Pieridae (19%), Abundance distribution each species was 20 in relationship assigned to a given (14%), seven in geographical range Papilionidae (5%) A positive relationship be- category. Geograph- and three in Rio- tween abundance and distri- ic range categories dinidae (2%). The bution was identified at the were grouped as: most abundant landscape scale. Species’ abun- species record- dances were positively cor- 1- Restricted to Figure 3. Rank-abundance curves (log scale in Y axis) for two ed were Eunica related with their landscape southeast Mexico representative forest stands in Coapilla, during 2007. monima (6522 distribution sizes (F = 603.7, and Central America 1,146 records; 69%), P<0.001, r=0.89; Figure 4). but not further south Hermeuptychia hermes (554 than . records; 6%), and Eurema Habitat use categories and 3- Widespread in the Neo- mexicana mexicana (271 re- geographical distribution 2- North America to Cen- tropics (at least reaching Bra- cords; 3%). Table I shows but- tral America and/or south- zil or ). ern Mexico to northern South terfly total richness recorded A significant association America. Because the sample size per forest stand. Stand 6 and between habitat use and geo- for each family was rather small, family samples were col- lated to examine a possible overall as- sociation between habitat use and geographical range using χ2 tests.

Species composition in Coapilla and comparable sites

The proportion Figure 5. The proportion of butterfly species using unmodified (a) and modi- fied (b) habitats, when increasing geographic range; geographic distribution of species grouped categories, 1: restricted to southeast Mexico and Central America but not for each geographic further south than Panama, 2: North America to Central America, and/or Figure 4. Abundance distribution- relationship for range categories for southern Mexico to northern , and 3: widespread in the Neo- butterflies in Coapilla, Chiapas. Coapi­lla and other tropics. (Geographic categories modified from Thomas, 1991).

JUL 2010, VOL. 35 Nº 7 529 graphic range for 63 species dition). Previous results have might seek for several pieces and silvicultural mosaics with (χ²= 13.26, df= 1, p<0.005; suggested that diversity often of habitat to complete what >30% conversion, including Figure 5) was detected in Co- increases with disturbance it has been called “functional selective logging, show shifts apilla. Seventeen species were near or below natural levels habitat” (Dennis et al., 2003). in species composition with classified as using unmodified (perhaps the condition of for- In addition, species abun- irreversible loss of many com- habitats, and 46 species were est stand 6), although some dance and identity showed ponents of butterfly communi- placed as mostly occupying sensitive species might be important responses to in- ties, and otherwise indicating modified habitats. Species with eliminated at even low levels creased disturbance and dif- non-sustainable land and re- narrow geographic ranges were of interference. For example, ferent forest-use regimes. source use (Brown, 1997). On commonly grouped in unmodi- some spatially-restricted spe- Typically, forest-affiliated the other hand, Janzen (1988) fied habitats; whilst widespread cies, namely Satyrotayget- species were associated to has suggested that a resulting distributed species were as- is satyrina, Enantia jethys, forest habitats (Table II). mosaic of habitats and suc- sociated to human-modified Dismorphia crisia virgo and Sedentary species that persist cessional stages of vegetation habitats. Jonaspyge jonas were almost within relatively small forest can support more species of exclusively found in forest areas were found rarely and than a pristine Species composition between stands with important levels in low densities. In contrast, tropical forest could realisti- Coapilla and other some common spe- cally support. The analysis sites cies that occur in of habitat preferences and the surrounding re- geographic distributions for a The proportion of gion were extremely set of species from Coapilla species grouped in abundant, such as showed that few forest species distinct geographical E. monima, result- that possess relatively narrow distribution categories ing in numerous and geographic ranges were clas- was significantly dif- widespread distri- sified as making use of modi- ferent among a sam- bution records. At fied habitats. Geographically ple of sites for which a landscape scale, restricted or endemic butterfly butterfly diversity had sedentary spe- species are often regarded as been thoroughly as- cies have greater biotope specialists, although sessed (F4 = 11.615, Figure 6. The proportion of species recorded in five different potential to be lo- they can be among the most p<0.001; Figure 6). sites in Mexico as regards to their geographical distribution calized within the abundant species at sites Less disturbed sites, category. Geographic range is R1: restricted to southeast study area (Cowley where they occur (León-Cor- such as Santiago Co- Mexico and Central America but not further south than et al., 1999) and, tés et al., 2000). Some bio- Panama, R2: North America to Central America, and/or maltepec and Los southern Mexico to northern South America; and R3: wide- in contrast, mobile tope specialists might occupy Mazos (sites 4 and 5, spread in the Neotropics. Sites are 1: Coapilla (this work), 2: species sometimes disturbed habitats, but in most respectively; in Figure Los Altos Chiapas (Miss-Barrera et al., 2010), 3: Sierra de have a higher prob- cases the latter are used very 6) showed a relatively Atoyac, Guerrero (Vargas-Fernández et al., 1991), 4: Santiago ability of being re- rarely by these species, and even proportion of spe- Comaltepec, Oaxaca (Luis-Martínez et al., 1991), 5: Los corded away from usually only when modified cies distributed among Mazos, Jalisco (Vargas-Fernández et al., 1999). their main breeding and unmodified habitats are distinct geographical habitats, resulting in adjacent (Thomas, 1991). For categories. In contrast, heav- of forest cover as is the case widespread distributions. In instance, many savannah and ily disturbed sites, such as for “Forest”, and forest stands general, the densities and tropical dry forest butterflies Los Altos de Chiapas, exhib- 4, 1 and 2. In contrast, widely distributions of Coapilla but- appear to congregate in mesic ited a poor representation of and common species such as terflies were positively re- habitats, such as gallery for- narrowly-distributed species. Eunica monima, sen- lated at the landscape scale ests, or migrate to higher ele- nae and Danaus gilippus were (Figure 4). These results vations in dry seasons (DeVr- Discussion recorded in open heavily-used are broadly consistent with ies, 1987). Furthermore, many forest stands, as is the case previous studies on other species may be specialists on The 143 species of butter- for forest stands 6, 8 and 9. taxonomic groups that have naturally-occurring succes- flies registered in forest stands These species have been able demonstrated positive rela- sional habitats, and so be able of Coapilla represent ~12% of to locate scattered resources tionships between density to occupy human-modified the total number of species throughout the landscape, and and distribution regardless of areas that provide the same recorded for Chiapas (Luis- although moderate levels of the spatial scale considered successional conditions. At Martínez et al., 2003). Even disturbance generate a patchy (Niemelä and Spence, 1994; this point, and for the species considering these important structure of habitats, this Cowley et al., 2001). whose habitat preferences and levels of butterfly diversity for could be beneficial for an im- What changes in species geographic distributions could a single landscape, butterfly portant set of species. In line composition can be perceived be determined, there seems to distribution and structure have with this idea, species such as a result of forest manage- be an increasing proportion of experienced important shifts as Adelpha iphicleola iphi- ment? Depending on its scale rather “weedy” species (>25% at the local (forest stand) and cleola, Altinote ozomene nox, and frequency, habitat modifi- Coapilla butterflies have been landscape scales. There was nimbice nimbice, cation may provoke a variety regarded as widespread and almost a three-fold increase in Dircenna klugii klugii and Pa- of responses from the mem- common species) in this for- butterfly diversity between the rides photinus exhibited a mix bers of a given Neotropical ested landscape. Furthermore, most evenly-forested site (what of preferences (i.e. occurring butterfly community (Raguso comparisons for species com- we recognized as the “pristine in a variety of habitats, from and Llorente-Bousquets, 1991). position of Coapilla butter- condition”) and forest stand 6 open to relatively preserved Previous assessments have flies and other sites in Mexico (the most heterogeneous con- conditions). These butterflies suggested that agricultural yielded significant differences.

530 JUL 2010, VOL. 35 Nº 7 TABLE II CHECK-LIST OF BUTTERFLIES FROM COAPILLA, CHIAPAS, MEXICO a, b Taxon Abundance Range Habitat Taxon Abundance Range Habitat Abaeis nicippe (Cramer) 4 2 2 Hermeuptychia hermes (Fabricius) 554 NA NA Achlyodes pallida (Felder) 8 NA NA Hypanartia godmanii (Bates) 6 2 1 Adelpha iphicleola iphicleola (Bates) 12 1 2 Hypoleria lavinia cassotis (Bates) 31 1 1 Adelpha leuceria leuceria (Druce) 9 1 1 Jonaspyge jonas (Felder & Felder) 17 NA NA Adelpha pithys (Bates) 13 1 1 Laothus barajo (Reakirt) 1 NA NA Agraulis vallinae incarnata (Riley) 1 3 2 Leptophobia aripa elodia (Boisduval) 20 2 1 Altinote ozomene nox (Bates) 3 1 2 Leptotes marina (Reakirt) 2 NA NA Anartia fatima fatima (Fabricius) 1 NA NA Lerema accius accius (Smith) 3 NA NA Ancyloxypha numitor (Fabricius) 3 NA NA Lerema lumina (Herrich-Schäffer) 20 NA NA Anteos clorinde (Godart) 2 3 2 Librita librita (Plötz) 1 NA NA Anteos maerula (Fabricius) 2 3 2 nemesis atthis (Doubleday) 1 NA NA ardys ardys (Hewitson) 2 2 2 chiron marius (Cramer) 6 3 2 Anthanassa drusilla leeles (Bates) 5 3 2 Marpesia petreus (Cramer) 13 3 2 Anthanassa otanes otanes (Hewitson) 2 NA NA Marpesia zerynthia dentigera (Fruhstorfer) 1 NA NA Anthanassa aff. ptolyca ptolyca (Bates) 2 NA NA Mechanitis polymnia lycidice Bates 3 3 2 Anthanassa texana texana (Edwards) 2 NA NA pixe pixe (Boisduval) 1 NA NA Aphrissa statira statira (Cramer) 1 3 2 Mestra dorcas amymone (Ménétriés) 17 2 2 Apuecla maeonis (Godman & Salvin) 1 NA NA Methion aff. melas Godman 27 NA NA Astraptes fulgerator (Walch) 4 NA NA Morpho helenor montezuma Guenée 2 2 2 (Boisduval & Leconte) 24 NA NA Nathalis iole Boisduval 9 NA NA Autochton vectilucis (Butler) 69 NA NA Niconiades sp. 2 NA NA Autochton sp. 52 NA NA Noctuana lactifera bipuncta (Plötz) 1 NA NA Biblis hyperia aganisa (Boisduval) 2 3 2 Noctuana stator (Godman) 1 NA NA Calephelis sp. 31 NA NA Oxeoschistus tauropolis tauropolis (Westwood) 1 NA NA Caligo uranus Herrich-Schäffer 2 1 2 Panthiades bathildis (Felder) 1 NA NA Calycopis clarina (Hewitson) 1 NA NA Papias aff. dictys (Godman) 9 NA NA Calycopis isobeon (Butler & Druce) 21 NA NA Papilio polyxenes asterius Stoll 1 2 2 Catasticta flisa flisa (Herrich-Schäffer) 12 2 1 Parides photinus (Doubleday) 114 1 1 Catasticta nimbice nimbice (Boisduval) 28 1 2 Parrhasius moctezuma (Clench) 1 NA NA Catasticta nimbice ochracea (Bates) 1 NA NA Pereute charops nigricans Joicey & Talbot 13 NA NA Catasticta teutila flavifaciata Beutelspacher 1 2 1 Phocides urania urania Godman & Salvin 1 NA NA Celastrina argiolus gozora (Boisduval) 20 NA NA Phoebis agarithe agarithe (Boisduval) 64 3 1 Chlosyne erodyle erodyle (Bates) 17 1 2 Phoebis neocypris virgo (Butler) 9 NA NA Chlosyne janais janais (Drury) 14 2 2 Phoebis philea phílea (Linnaeus) 19 3 2 Chlosyne aff. marina (Geyer) 2 NA NA Phoebis sennae marcellina (Cramer) 9 3 2 Cissia pompilia (Felder & Felder) 2 NA NA Poanes inimica (Butler & Druce) 12 NA NA Consul excellens genini (Le Cerf) 2 NA NA Poanes melane (Edwards) 32 NA NA Cupido comyntas (Godart) 58 NA NA Poanes zabulon (Boisduval & Leconte) 40 NA NA Cyllopsis hedemanni hedemanni Felder 27 1 2 Pompeius pompeius (Latreille) 4 NA NA Cyllopsis hilaria (Godman) 3 NA NA epidaus epidaus (Doubleday) 4 NA NA Danaus gilippus thersippus (Bates) 13 2 2 Pterourus garamas electryon (Bates) 4 1 1 Danaus plexippus plexippus (Linnaeus) 6 3 2 Pterourus menatius victorinus (Doubleday) 1 NA NA Denivia augustinula (Goodson) 2 NA NA Pyrgus communis (Grote) 4 NA NA anna anna (Guérin-Ménéville) 28 NA NA Pyrgus oileus (Linnaeus) 25 NA NA Diaethria pandama (Doubleday) 1 NA NA Pyrisitia dina westwoodi (Boisduval) 3 2 2 Dione juno huascuma (Reakirt) 18 3 2 Pyrisitia nise nelphe (Felder) 107 3 3 Dione moneta poeyii Butler 86 3 2 Pyrisitia proterpia (Fabricius) 31 3 2 Dircenna klugii klugii (Geyer) 38 1 2 Quasimellana sp. 2 NA NA Dismorphia crisia virgo (Bates) 27 1 1 Quinta cannae (Herrich-Schäffer) 1 NA NA Doberes anticus (Plötz) 1 NA NA Rekoa menton (Cramer) 1 NA NA Dryas iulia moderata (Riley) 10 3 2 Remella sp. 8 NA NA Echinargus isola (Reakirt) 2 NA NA Satyrotaygetis satyrina (Bates) 57 1 1 Electrostrymon sangala (Hewitson) 1 NA NA Siproeta epaphus epaphus (Latreille) 33 2 2 Enantia jethys (Boisduval) 10 NA NA Siproeta stelenes biplagiata (Fruhstorfer) 1 3 2 Epargyreus aspina Evans 6 NA NA Smyrna blomfildia datis Fruhstorfer 5 2 1 Erynnis tristis tatius (Edwards) 3 NA NA Strymon cestri (Reakirt) 1 NA NA Eunica monima (Stoll) 6522 3 2 Strymon melinus (Hübner) 1 NA NA Euptoieta claudia daunius (Herbst) 3 NA NA Synapte salenus (Mabille) 1 NA NA Euptoieta hegesia meridiania Stichel 32 2 2 anieta cluvia (Godman & Salvin) 10 NA NA Eurema albula celata (Felder) 2 3 2 aegides (Herrich-Schäffer) 1 NA NA Eurema daira eugenia (Wallengren) 42 3 2 Thecla maeonis (Godman & Salvin) 1 NA NA Eurema mexicana mexicana (Boisduval) 271 3 2 Tmolus crolinus Butler & Druce 1 NA NA Eurema salome jamapa (Reakirt) 244 2 1 Urbanus prodicus Bell 46 NA NA Fountainea glycerium glycerium (Doubleday) 1 NA NA Urbanus simplicius (Stoll) 6 NA NA Ganyra josephina josepha (Salvin & Godman) 1 NA NA Urbanus teleus (Hübner) 7 NA NA Greta annette annette (Guérin-Ménéville) 28 1 1 Vanessa annabella (Field) 4 NA NA Hamadryas februa ferentina (Godart) 8 3 1 Vanessa virginiensis (Drury) 20 2 2 Heliconius charithonia vazquezae (Comstock & 25 3 2 Vehilius inca (Scudder) 2 NA NA Brown) Yphthimoides renata (Stoll) 3 NA NA Heliconius hortense (Guérin-Ménéville) 20 NA NA Zerene cesonia cesonia (Stoll) 2 NA NA Hemiargus hanno antibubastus Hübner 2 NA NA Ziegleria ceromia (Hewitson) 2 NA NA Heraclides thoas autocles (Rothschild & Jordan) 4 2 2 Zizula cyna (Edwards) 27 NA NA a The author of each taxon is given. The order is alphabetic, first by , then by species, subspecies (taxonomic authorities after Llorente-Bousquets et al., 2006, and Warren et al., 2010). b Abundance denotes the total number of individuals recorded for a given species. Range indicates the geographic range categories assigned for a given species: 1) restricted to southeast Mexico and Central America but not further south than Panama; 2) North America to Central America, and/or southern Mexico to northern South America; and 3) widespread in the Neotropics. Habitat indicates the habitat use recorded for a given species: 1) species making use of unmodified habitat; 2) species making use of modified habitat. NA: habitat use or geographic range information not available.

JUL 2010, VOL. 35 Nº 7 531 Within the bounds of geo- vegetation, soil biota, and received a M.Sc. scholarship DeVries PJ (1987) The Butterflies graphical or historical factors indicator insect communities. from the National Council of Costa Rica and their Natu- ral History: Vol. I: Papilioni- that might produce inherent Mexico is a megadiverse for Science and Technolo- dae, Pieridae y Nymphalidae. differences among butterfly country, but it has high rates gy in Mexico (CONACYT, Princeton University Press. faunas, it is concluded that of deforestation and many 203016). The project was Princeton, NJ, USA. 327 pp. current differences in spe- ecological-impoverished ar- supported by ECOSUR’s re- DeVries PJ (1997) Butterflies of cies composition might be eas resulting from supposed search line “Dinámica de Po- Costa Rica and their natural a result of landscape condi- sustainable forest-managing blaciones y Comunidades de history: Vol. II (). Princeton University Press. tions; and hence, changes in practices. Mapping α and β Insectos.” Princeton, NJ, USA.. 288 pp. the proportions of widespread diversity can contribute to Garwood K, Lehman R (2005) or spatially-restricted species the conservation of natural REFeRENCES Butterflies of Northeastern might potentially differ due resources by helping to iden- Mexico. Nuevo Leon, San Luis Potosi, Tamaulipas. A Photo- to fundamental changes in tify species-rich hotspots and Arellano L, León-Cortés JL, Ovas- the landscape conditions. The areas that include as many graphic Checklist. Eye Scry. kainen O (2008) Patterns of McAllen, TX, USA 188 pp. immediate effect that expan- species (Gentry, 1992). 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E, Boege-Schmidt E (2003) tinue to be lost, local (alpha) pilla landscape seems to hold Informe de Evaluación de Brown KSJr (1997) Diversity, dis- Certificación SmartwWood diversity may not noticeably important levels of diversity turbance, and sustainable use para Ejido Coapilla Chiapas, decrease, and may sometimes for butterflies. The result- of Neotropical forests: insects México. Smartwood, USA. increase (e.g. Raguso and ing affiliations of the organ- as indicators for conservation 116 pp. Llorente-Bousquets, 1991) but isms that persist or colonize monitoring. J. Insect Cons. 1: Janzen DH (1988) Ecological char- 25-42. regional (beta) diversity will these forest-managed areas acterization of a Costa Rican decline (Thomas, 1991). should be a matter of concern Brown KSJr, Freitas AL (2000) At- dry forest caterpillar fauna. lantic forest butterflies: indica- Biotropica 20: 120-135. 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KJ (2001) Density-distribution Anim Cons. 7: 241-250. 1997). Indeed, the usefulness and Sergio Villafuerte helped relationships in British but- Llorente-Bousquets J, Luis-Mar- terflies. I. The effect of mobil- of forest systems to humans in the field. Andrew Warren tínez A, Vargas-Fernández I ity and spatial scale. J. Anim. is just one aspect of their and Armando Luis-Martínez (2006) Apéndice general de Ecol. 70: 410-425. overall complex functioning, confirmed butterfly identifica- Papilionoidea: lista sistemática, just as likely to be modified tions. Henry Castañeda and Dennis RLH, Shreeve TG, Van distribución estatal y provin- by a change in forest struc- Miguel Icó helped identifying Dyck H (2003) Towards a cias biogeográficas. In Mor- functional resource-based rone JJ, Llorente-Bousquets J ture (such as removal of large plant species from Coapilla. concept for habitat: a butter- (Eds.) 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