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Journal of

HE LEPIDOPTERISTS' SOCIETY

Volume 58                                                                         2004                                                                     Number 3

Journal of the Lepidopterists' Society 58(3), 2004, 125-142

PAPILIONOIDEA OF THE EVERGREEN TROPICAL FORESTS OF MEXICO

Jose L. Salinas-Gutierrez1, Armando Luis-Martinez2 And Jorge Llorente-Bousquets3

Museo de Zoologia, Facultad de Ciencias, UNAM, Apdo. Postal 70-399, Mexico, 04510 D. E, MEXICO

ABSTRACT. The diurnal butterflies in 11 geographical units of evergreen tropical forest in Mexico were studied, giving in a total list of 683 species of Papilionoidea (excluding Hesperiidae). This is the first list of Mexican butterflies which covers a specific type of vegetation. The species richness in this zone makes evident the need for adequate conservation strategies for these ecosystems, whose extent is rapidly decreasing. The results are compared with other areas of Neotropical rain forest.

Additional key words: butterflies, distribution, evergreen tropical forest, Neotropical region, richness.

RESUMEN. Se efectuo un trabajo de mariposas diurnas en 11 unidades geograficas con bosque tropical perennifolio en Mexico, registrandose una lista total de 683 especies de Papilionoidea (sin incluir Hesperiidae). Esta es la primera lista de mariposas en Mexico que involucra la cobertura de un tipo de vegetacion especifico. La riqueza de especies presentes en la zona hace patente la necesidad de crear estrategias de conservacion en este ecosistema que esta disminuyendo su extension rapidamente. Los resultados se comparan con otros sitios de bosque tropical del Neotropico.

Palabras claves: mariposas, distribucion, bosque tropical perennifolio, region Neotropical, riqueza.

The rain forests, or evergreen tropical forest (ETF) ecosystems are considered top priority for world conservation. This is because these forests shelter approximately 50% of the species of the planet Earth. In addition, their rate of decrease is one of the fastest (Wilson 1988, Dirzo & Garcia 1992). The Neotropical region contains approximately 20% of the species on the Earth (Myers 1988); within this region, the ETF has the largest extent of all the regions.

The northernmost distribution of the ETF in America is in Mexico. Originally, the ETF covered 13% of the nation, but Granillo (1985) and Toledo (1988) remark that at present it covers from 10 to a

1  sgjl@att.net.mx

2 alm@hp.fciencias.unam.mx

3 Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Santa Fe de Bogota, Columbia, email:jlb@hp.fciencias.unam.mx

15% of the original area, being replaced by pastures and other agroecosystems. An annual deforestation rate of 4% was registered for the last 25 years by Dirzo & Garcia (1992) in the ETF of the Los Tuxtlas region in Veracruz, Mexico.

The ETF was once distributed in Mexico from southern San Luis Potosi and northern Veracruz, in the north, through parts of the states of Hidalgo, Puebla, Oaxaca, southern Veracruz, to the north and northeast of Chiapas and in some parts of Tabasco, Campeche and Quintana Roo (Fig. 1). Along the Pacific slope, it occupied the southernmost area of Sierra Madre de Chiapas and the Tapachula-Mapastepec (low Soconusco region), isolated by the Sierra Madre, the Tehuantepec isthmus and the Central Depression of Chiapas (Rzedowski 1978). Rzedowski (1996) estimated that the plant species in the ETF include over a third of the total flora of the country.

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Journal of the Lepidopterists' Society

Fig. 1. Distribution in Mexico of the geographical units having evergreen tropical forests.

Although Papilionoidea have been the subject of many studies, the knowledge of this lepidopteran superfamily is still very incomplete. Robbins & Opler (1997) estimated its approximate richness as 17,500 species; furthermore, they pointed out that this is one of the most studied groups of insects, with 90% of its species being known. These authors assumed that the greatest richness of this group is within the Neotropical region; similar results were obtained by Heppner (1991). There are not enough faunistic studies in the region and many of the studies were based on sporadic collections in extensive, ecologically heterogeneous areas. The situation becomes more critical in the ETF, whose understanding is based on very few studies, most of which were made during this century in a few areas, such as Los Tuxtlas, Veracruz and Sierra de Juarez, Oaxaca in Mexico.

The studies made by Lamas et al. (1991, 1996), Brown (1984), Emmel & Austin (1990), and Austin et al. (1996), among others, show the need for making more intensive systematic collections in these communities, especially in areas located close to mountains (increasing the diversity; G. Lamas pers. com.). Because of the inaccessibility of the areas occupied by the ETF and the consequent logistic and financial problems for systematic studies there, methods have been proposed to estimate the potential number of species living in each community in a quick and accurate way (Soberon & Llorente 1993, Colwell & Coddington 1994).

The large diversity of butterflies living in the ETF of Mexico has attracted attention of interested students since the 19th century; nevertheless, only about 10 faunistic studies are available which can be compared

with some accuracy, such as those made by Hoffmann (1933), Ross (1964, 1976-1977), Routledge (1977), de la Maza & de la Maza (1985a, b), de la Maza & White (1990), Luis et al. (1991, 1995), Raguso & Llorente (1991, 1997), Martinez (1994), and Villegas (1998).

Materials And Methods

The first step to obtain a list of the butterflies species inhabiting the ETF in Mexico was to compile, systematize and summarize the studies made in these communities. A specialized bibliography was consulted for five states: Tabasco, Routledge (1977), Martinez (1994) and Villegas (1998); Chiapas, Hoffmann (1933), de la Maza & de la Maza (1985a, 1985b), and unpublished data of faunistic surveys made in Yaxchilan by members of the Museo de Zoologia de la Facultad de Ciencias, UN AM; Oaxaca, the data available for the Chimalapas region and Luis et al. (1991); Veracruz, Ross (1964, 1976-1977), Raguso & Llorente (1991, 1997) and Luis et al. (1995); for San Luis Potosi, de la Maza & White (1990). The bibliography was used to obtain comparable lists, in order to be able to tabulate the data by region and by locality. Only records with vouchers collected in ETF were considered.

Once the areas in each report were analyzed, "geographical units" were defined for each collection site, giving 11 units (Fig. 1): in Tabasco, Cerro del Cocona (CC), Agua Blanca (AB) and Tenosique (TEN); in Chiapas, Chajul (CHJ), Yaxchilan (YAX) and Soconusco (SOC); in Oaxaca, La Gringa (LG), Chalchijapa (CHA) and Sierra de Juarez (SJ); in Veracruz, Los Tuxtlas (LT); and in San Luis Potosi, Huichihuayan (SLP).

For the geographical units LT, SJ and SLP, only the localities having ETF were included, since the original studies were made in several plant communities. As these studies included sites whose altitudes do not correspond to the distribution of the ETF, a detailed revision was made so as not to overestimate the richness of the fauna by inclusion of montane species or those of semideciduous tropical forest.

Results And Discussion

Based on the above defined 11 units, 683 species living in the ETF of Mexico were registered, in 272 genera, 18 subfamilies and 4 families, about 53% of the total butterfly fauna of the country (assuming a total of 1,295 species of Papilionoidea and excluding the Hesperiidae). The percentage of species in each family is similar to that reported in other regions of Mexico (Sierra de Manantlan, Jalisco-Colima: Vargas et al. 1999;

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Sierra de Atoyac de Alvarez, Guerrero: Vargas et al. 1992): Papilionidae 5.4%, Pieridae 7.7%, Nymphalidae 45.1%, and Lycaenidae 41.7%.

Even though the studies chosen for this analysis used different methods, time periods and objectives, a general comparison is presented for each of the 11 geographical units by species in each family and subfamily, together with the estimated total number of species inhabiting the ETF in Mexico (Table 1).

According to Llorente et al. (1993) and Luis et al. (2000) the butterfly fauna of Mexico represents about 10% of the wold total, thus indicating that Mexico is a megadiversity country. In this paper only a single plant community was analyzed, covering 13% of the total land surface; due to deforestation over the last 50 years, this is now reduced to less than 2%. The percentage of species living in this plant community is surprisingly high, representing 50% of the Mexican butterfly fauna. Heppner (1991) gave a total of 7,927 species of butterflies in the Neotropical region, including the family Hesperiidae, which was excluded in the present paper; even so the ETF included 8.6% of this fauna. If only the 4,800 species of Papilionoidea are considered, the ETF in Mexico holds 14.2% of this figure.

Table 1 also includes the percentage of the Mexican total in each subfamily. This table shows that LT is the most diverse geographical unit, with 482 species

(70.6%), followed by SJ with 355 (52%), CHJ with 352 (51.5%), and SLP with 300 (43.9%). The latter geographical unit is the northernmost ETF site in Mexico, followed by LT and SJ. Considering that Mexico is in a transitional biogeographical zone between the Nearctic and the Neotropical regions, and that the predominance of these elements varies with latitude and altitude, this suggest an increase of species with clear Nearctic affinity, which do not belong to the ETF in a strict sense.

The geographical units LT, SJ and CHJ have similar patterns of number of species per subfamily, with the exception of the Theclinae in LT, which include 54.78% of the species mentioned for ETF, or 50% of those for all of Mexico. This may be due to the fact that, for many years, amateur collectors have paid local people for hairstreak specimens (L. Gonzalez-Cota pers. com.). This collecting effort is clearly seen in the percent representation (Table 1).

The geographical unit LT has the highest representation for 8 out of 18 subfamilies collected. Heppner (1991) mentioned the importance of the Ithomiinae, Morphinae and Brassolinae, which have maximum number of species in the following geographical units: Ithomiinae in SJ with 21 species; Morphinae in CHJ, CHA, SJ and LT with three species; and Brassolinae in CHJ and LT with 11 species.

TABLE 1. Species richness of Papilionoidea by geographical unit. See 'Methods' for abbreviations.

CC

AB

TEN

CHJ

YAX

SOC

LG

CHA

SJ

LT

SLP

MEX

Papilionidae

Papilioninae

6

15(18.29)

11(13.41)

7 14(17.07)

9

24(29.27)

8

15(18.29)

4

9(10.98)

8

11

10

12

16(19.51) 12(14.63) 29(35.37) 30(36.59) 20(24.39) 37(45.12) 82

Pieridae

Dismorphiinae

Coliadinae

Pierinae

12

2(9.52)

14(40)

7(11.29)

8

1(4.76) 13(37.14) 1(1-61)

11

0 17(48.57) 4(6.45)

19

4(19.05) 17(48.57) 10(16.13)

IS

2(9.529 16(45.71) 6(9.68)

14

2(9.52)

13(37.14)

5(8.06)

10

2(9.52)

12(34.29)

5(8.06)

12

25

26

21

27

1(4.76) 4(19.05) 14(40) 21(60) 4(6.45) 16(25.81)

7(33.33) 4(19.05) 10(47.62) 22(62.86) 20(57.14) 24(68.57) 16(25.81) 11(17.74) 19(30.65)

21 35 62

NOTE: The numbers in family rows correspond to the total genera in each geographical unit and, in parentheses, the percentage of species per subfamily in relation to the total in Mexico. ETF: represents the total for Mexico ETF MEX: represents the total for all of Mexico

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Journal of the Lepidopterists' Society

Table 1 also details the total number of species in the ETF of Mexico for Papilionidae (37), Pieridae (53), Nymphalidae (308), and Lycaenidae (285), and the total

0.30

0,36

0.42

0.48

054

~CC

-YAX

LG

CHA

AB

TEN

CHJ

-c

LLT -SLP

-soc

Fig. 2. UPGMA dendrogram of the geographical units based on

THE JACCARD SIMILARITY COEFFICIENT

number of genera per family for each geographical unit. The geographical units having the highest number of exclusive species are LT (84), CHJ (40), and SJ (23). When the geographical units were gruped by state, the three in Tabasco (CC-AB-TEN) shared 78 species; in Oaxaca (LG-CHA-SJ), 105; and in Chiapas (CHJ-YAX-SOC) 84. In Chiapas, the units CHJ-SOC share 114 species, while CHJ-YAX share 170; this makes sense since the latter two geographical units belong to the same biogeographical "island" of ETF, while SOC, on the Pacific slope, belongs to a different biogeographical area.

TABLE 2. Synthetic matrix of species shared by the geographical units.

The geographical units SLP and LT, the northernmost localities of ETF, share 243 species, this large figure suggests that both belong to the same biogeographical "island." If the four geographical units having that largest number of species are grouped (LT-SJ-CHJ-SLP), these share only 138 species, again emphasizing the importance of the geographical history of such units, with mixture of species in the northern units having influence over the total number of species.

The UPGMA (unweighted pair-group method using arithmetic averages) dendrogram of all geographical units (Fig. 2) was prepared, based on the Jaccard coefficient to assess the degree of similarity. This similarity between the geographical units was based in part on the synthetic data matrix (Table 2) that groups the species shared by each unit.

As mentioned above, SOC belongs to a distinct geographical "island" of ETF, a fact seen in the dendrogram where it appears as the most dissimilar from all other geographical units. An important group is that whose geographical units have the largest collection effort (SJ-LT-CHJ) and number of species. The remaining group whose geographical units have the lowest number of species, may need more collecting effort. The similarities between LG and CHA, and that between AB and TEN are probably due to their geographical proximity (Fig. 1).

Table 3 compares the numbers of species in various areas of ETF in the Neotropical region, such as Tikal, Guatemala (Austin et al. 1996); Jaru, Manaus, Campinas and Rondonia in Brazil (Brown 1984, Emmel & Austin 1990); Pakitza, Tambopata and Rio Napo in Peru

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TABLE 3. Butterfly species richness in some localities in the Neotropical region.

NOTE: Asterisks indicate missing data.

ABBREVIATIONS: AL = Atlantic lowland; PE = Pacific evergreen (DeVries 1997).

(Lamas et al. 1991, 1996); several localities in Costa Rica (DeVries 1997); and Misahualli in Ecuador (Racheli & Racheli 1998). When these data are compared with those from Mexico (Table 1), the species richness of geographical units such as LT or S J becomes obvious, comparable with that of areas such as Tikal (328 species) or Campinas (368). LT and SJ have the largest know numbers of Pieridae (45 and 41, respectively), for Mexico (and also the Neotropical region).

Conclusions

The ETF of Mexico show a total of 683 species of Papilionoidea (excluding Hesperiidae). Robbins & Opler (1997) state that the higher diversity of butterflies follows the amount of rainfall. The data in this paper include collections made in sites having 1,500 mm minimum average annual rainfall, supporting the empirical observation that wet sites have a large species diversity. However, other parameters must be considered to explain and compare the diversity and richness of species, such as area and topographical or ecological heterogeneity.

Emmel & Austin (1990) discussed the role played by the great "microheterogeneity" present in the locality of Jaru. Environmental heterogeneity is also relevant in our observations; for example SLP and LT are more heterogeneous, giving mixture of species from montane and lowland areas, due to altitude (montane effect) in LT and latitude in SLP.

Acknowledgments

The authors thank Dr. Juan Jose Morrone Lupi for reviewing the manuscript. Also, we thank the "Consejo Nacional para la Ciencia y la Tecnologia" (CONACYT) project 32002, Direccion General de Asuntos del Pesonal Academico (DGAPA-IN) project 209900 and the "Comision para el Conocimiento y Uso de la Biodiversidad" (CONABIO) for financial aid for field work in the regions of Chimalapas, Oaxaca and Yaxchilan, Chiapas.

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Received for publication I November 2001; revised and accepted 5 September 2003

Appendix 1: Preliminary list of species. Data for each species are represented in the following format first column is the species name and subsequent columns mention the sites where the butterflies were collected.Asterisks (Vindicate doubtful data.

Taxon

Papilionidae

Battus philenor philenor

B. polydamas polydamas

B. laodamas copanae

B. ingenuus

B. lycidas

Parides photinus

P. montezuma

P. eurimedes mylotes

P. sesostris zestos

P. panares panares

P. panares lycimenes

P. erithalion polyzelus

P. iphidamas iphidamas

Protographium epidaus epidaus

P. philolaus philolaus

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP X                                                                           XXX

xxxxxxxxxxx xxxx xxxx

X                                         XX

XX                        XX

X                        XX                        XXX

X                                 XX

XXXXXXXXXX

xxxxx               xxxx

XX X XXX                                         X

XXXXXXXXXX X XXXX XXX XXXX                                         XXX

XXXX                                 XXX

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Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

P. agesilaus neosilaus

P. dioxippus lacandones

P. calliste calliste

P. thyastes marchandi

Eurytides salvini

Protesilaus macrosilaus penthesilaus

Mimoides thymbraeus thymbraeus

M. ilus branchus

M. phaon phaon

Priamides pharnaces

P. anchisiades idaeus

P. erostratus erostratinus

P. erostratus erostratus

Troilides torquatus tolus

Calaides ornythion ornythion

C. astyalus pallas

C. androgens epidaurus

Heraclides thoas autocles

H. cresphontes

Papilio polyxenes asterius

Pyrrhosticta victorinus victorinus

P. abderus abderus

X X

X X

X X

X X X X

X X X

Pieridae

Enantia Una marion

E. albania albania

E. jethys

E. mazai mazai

Lieinix nemesis atthis

Dismorphia amphiona praxinoe

Dismorphia crisia virgo

D. eunoe eunoe

D. eunoe popoluca

D.  theucharila fortunata Zerene cesonia cesonia Anteos clorinde nivifera A. maerula lacordairei Phoebis agarithe agarithe P. argante argante

P. neocypris virgo

P. philea philea

P. sennae marcellina

Rhabdodryas trite ssp.

Aphrissa statirajada

Abaeis nicippe

Pyrisitia dina westwoodi

P. lisa centralis

P. nise nelphe

P. proterpia proteryria

Eurema agave millerorum

E.  albula celata E. boisduvaliana E. daira

E. mexicana mexicana

E. salome jarnapa

E. xantochlora xantochlora

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Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SQC LG CHA S] LT SLP

Nathalis iole Kricogonia lyside

Hesperocharis costaricensis pasion Archonias brassolis aproximata Charonias theano nigrescens Catastictaflisa flisa Catasticta nimbice nimbice C. ochracea ochracea Pereute charops charops Melete hjcimnia isandra M. polyhymnia florinda Glutophrissa drusilla tenuis Pontia protodice Leptophobia aripa elodia Itaballia demophile centralis I. pandosia kicaha Pieriballia viardi viardi Perrhybris pamela chajulensis Ascia monuste monuste Ganyra josephina josepha G. phaloe tiburtia

X X X

Nymphalidae

Altinote ozomene nox Actinote guatemalena veraecrusis A. guatemalena guatemalena A. thalia anteas Philaethria diatonica Dionejuno huascuma

D.  moneta poeyii Agraulis vanillae incarnata Dryadula phaetusa Dryas iulia moderata Eueides aliphera gracilis

E.  isabella eva E. lineata

E. procula asidia

E. vibilia vialis

Laparus doris viridis

Heliconius charitonia vazquezae

Heliconius cydno galanthus

H. erato petiveranus

H. hecale zuleika

H. hecale fornarina

H. hecalesia octavia

H. hortense

H. ismenius telchinia

H. sapho leuce

H. sara veraepacis

Euptoieta claudia daunius

E. hegesia hoffmanni

Vanessa atalanta rubria

Cynthia cardui

C. virginiensis

C. anabella

Nymphalis antiopa antiopa

X X

X X

X X X X X X X X X X

X

X

X X

X X

X

X X X

X X

X X X

X

X X

X X

X

X X

X

X X

X X

X X X X

X X

X X X X X

X X X

X X

X X X

X

X

X X

X X X X

X X X X X X X

X X

X

X X

X X X X X X

X X X

X X

X X

X X

X X X X X X X X X X

X X

X X

X X X X X X X

X X

X X X X X X X X X

X X X

X X X

X X X X

X X X X X

X X

X X

X X X X X X X

Volume 58, Number 3

133

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Polygonia interrogations

Hypanartia dione

H. godmanii

H. lethe

H. kefersteini

Anartia amathea venusta

A. jatrophae luteipicta

Siproeta epaphus epaphus

S. stelenes biplagiata

S. superba superba

Junonia coenia

]. evarete

Chlosyne erodyle erodyle

C. gaudialis gaudialis

C. hippodrome hippodrome

C. janais

C. lacinia lacinia

C marina marina

C. rosita browni

Thessalia theona theona

T. theona thekla

Texola elada elada

T. elada ulrica

Microtia elva horni

Phyciodes mylitta mexicana

P. vesta graphica

P. phaon

Phyciodes tharos tharos

Anthanassa ardys ardys

A. argentea

A. atronia sydra

A. atronia atronia

A. drusilla lelex

A. frisia tulcis

A. ptolyca ptolyca

A. texana texana

A. annulata

Tegosa anieta cluvia

T. anieta luka

T. guatemalena

T. similis

Eresia clara clara

K. phillyra phillyra

Costilla eranites mejicana

C. myia myia

C. ofella ofella

Historis odius dious

Coea acheronta acheronta

Baeotus beotus beotus

Smyrna blomfildia datis

S. karwinskii

Colobura dirce dirce

Tigridia acesta ssp.

Biblis hyperia aganisa

Mestra dorcas amymone

Myscelia cyananthe cyananthe

134

Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Myscelia cyaniris cyaniris M. ethusa ethusa Catonephele mexicana

C.  numilia esite Catonephele cortesi Nessaea aglaura aglaura Eunica alcmena

E. alpais excelsa

Eunica sydonia caresa

E. malvina alhida

E. monima

E. mygdonia omoa

E. olympias augusta

E. venusia

E. tatila tatila

Hamadryas amphinome mexicana

H. februa ferentina

H. feronia farinulenta

H. fornax fornacalia

H. glauconome glauconome

H. guatemalena marmarice

H. guatemalena guatenwlena

H. iphthime joannae

H. laodamia saurites

Ectima erycinoides ssp.

Pyrrhogyra edocla edocla

P. neaerea hypsenor

P. otolais otolais

Temenis laothoe hondurensis

Epiphile adrasta adrasta

E. hermosa

E. orea plutonia

Bolhoneura sylphis veracruzana

B. sylphis lacandona Nicaflavilla hachiana Dynamine artemisia glauce

D.  ate

D. dyonis

D. postverta mexicana D. theseus Diaethria anna D. astala astala Cyclogramma bacchis Cyclogramma pandama Callicore astarte casta Callicore astarte patelina

C.  lyca lyca

C. texa grijalva

C. texa titania

C. tolima tehuana

C. tolima pacifica

C. pitheas

Adelpha basiloides basiloides

A. baeotia milleri

A. baeotia oberthurii

A. celerio diademata

Volume 58, Number 3

135

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Adelpha bredowii eulalia

A. cytherea marcia

A. escalantei

A. felderi j arias

A. iphiclus iphicleola

A. diazi

A. ixia leucas

A. leuceria leuceria

A. leucerioides leucerioides

A. lycorias rnelanthe

A. naxia epiphicla

A. paroeca emathia

A. phylaca phylaca

Adelpha pithys

A. salmoneus salmonides

A. serpa massilia

A. zalmona sophax

A. fessonia fessonia

Basilarchia archippus hoffmanni

Marpesia chiron marius

M. corita corita

M. harmonia

M. petreus tethys

M. zerynthia dentigera

Archaeoprepona amphimachus amphiktion

A. demophon centralis

A. demophoon gulina

A. meander phoehus

A. phaedra aelia

Prepona deiphile brooksiana

P. deiphile escalantiana

P. dexamenes medinai

P. laertes octavia

P. pylene philetas

Agrias aedon rodriguezi

A. amydon oaxacata

Zaretis callidryas

Z. itus ellops

Siderone galanthis ssp.

S. syntiche syntiche

Anaea troglodyta aidea

Consul electra electra

C. fabius cecrops

Fountainea eurypyle confusa

F. glycerium glycerium

F. halice martinezi

F. ryphea ryphea

Memphis artacaena

M. aureola

M. dia ssp.

M. forreri

M. hedemanni

M. herbacea

M. mora orthesia

M. phila boisduvali

M. neidhoeferi

X X

X X X

X X

X X X X X

X X

X X

X X

X X

X X X

X X X

X X X X

X X X X X

X X

X X X

X X X

X X

X X

X X X X

X X X

X X X X X X X

X X

X X

X X X

X X X

X X X

X X

X X

X X X X

X X

X X

X X X X X X

X X

X X

X X

X X

X X X X X X X X

X X X

X X

X X X X X X X X X X X X X

X

X X X X

X X

X X X X X

X X

X

X

X X

X X X

X X

X X

X X X

X

X X

X X X X X

X X

136

Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Memphis oenomais

M. philumena xenica

M. pithyusa

M. proserpina

M. xenocles Carolina

M. perenna perenna

Asterocampa idyja argus

Doxocopa cyane mexicana

D. laure laure

D. laurentia cherubina

D. pavon theodora

Morpho achilles montezuma

M. achilles octavia

Pessonia luna luna

Iphimedeia telemachus justitiae

I. telemachus ssp.

I. telemachus oaxacensis

Vynastor darius stygianus

D. macrosiris strix

Opsiphanes boisduvalii

O. cassiae

O. tamarindi

O. invirae fabricii

O. quiteria quirinus

Caligo atreus uranus

C. oileus scamander

C. prometheus memnon

Erijphanis aesacus aesacus

Narope cyllastros testacea

Pierella luna rubecula

Manataria maculata

Cepheuptychia glaucina

Chloreuptychia sericeella

Cissia confusa

C. labe

C. terrestris

Cyllopsis hedemanni hedemanni

C. hedemanni tamaulipensis

C. hilaria

C. suivalens escalantei

C. pyracmon

C. gemma, freemani

C. dospassosi

Euptychia mollina

Hermeuptychia hermes

H. alcinoe

Oxeoschistus hilara hilara

Magneuptychia libye

Megeuptychia antonoe

Pareuptychia hinocula metaleuca

P. interjecta

P. ocirrhoe

Pseudodebis zimri

Satyrotaygetis satyrina

Splendeuptychia kendalli

Paramacera xicaque xicaque

X X X X

X X

X X X

X X

X X

X X X

X X

X X X X

X X

X X X

X X X X X X X X X X X X

X X

X X X X

X X X

X X

X X

X X X X X X X

X X X X X X

X X X

X X X

X X X X X X X

X X X

X X X X X X X X X X X X X X X X X

X X X X X

X X

Volume 58, Number 3

137

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Taygetis kerea kerea

T. leuctra

T. rnermeria excavata

T. uncinata

T. virgilia

T. weymeri

T. thamyra

Vareuptychia usitata pieria

V. themis

V. similis

V. undina

Yphthimoides renata disaffecta

Dioriste tauropolis tauropolis

Pedaliodes dejecta circumducta

Danaus eresimus montezuma

D. gilippus thersippus

D. plexippus plexippus

Lycorea halia atergatis

L. ilione albescens

Anetia thirza thirza

Tithorea harmonia hippothous

T. tarricina duenna

Aeria eurimedea pacifica

Olyras crathis theon

Melinaea litis flavicans

M. lilis irnitata

Mechanitis lysimnia utemaia

M. menapis doryssus

M. polymnia lycidice

Hyposcada virginiana virginiana

Oleria paula

Napeogenes tolosa tolosa

Hypothyris euclea valora

H. lycaste dionaea

Ithomia leila

I. patilla patilla

Callithomia hezia hedila

C.  hezia wellingi Dircenna dero ssp.

D. jemina ssp.

D.  klugii klugii Episcada salvinia salvinia Pteronymia artena artena P. cotytto

P. simplex fenochioi

Godyris zavaleta sosunga

Hypomenitis annette annette

Greta morgane oto

G. nero nero

Hypoleria lavinia cassotis

Libytheana carinenta mexicana

Lycaenidae

Euselasia cataleuca

E.  chrysippe

E. regipennis regipennis E. sergia sergia

138

Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Euselasia procula

E. hieronymi hieronymi

E. inconspicua

E. pusilla

E. eubule eubule

E. aurantiaca aurantiaca

Hades noctula

Perophthalma tullius lasus

Leucochimona vestalis vestalis

L. lepida nivalis

Mesosemia telegone telegone

M. gaudiolum

M. gemina

Eurybia patrona persona

E. lycisca

E. halimede elvina

Hermathena oweni

Diophtalma lagora iphias

Napaea eucharila picina

N. theages theages

N. umbra umbra

Cremna actoris

C. thasus subrutila

Lyropteryx lyra cleadas

Ancyluris jurgensenii montezuma

A.  inca mora Rhetus arcius thia

R. periander naevianus

Isapis agyrtus hera

Brachyglenis dodone

Notheme erota diadema

Lepricornis melanchroia

Calephelis nemesis nemesis

C. mexicana

C. fulmen

C. stallingsi

C. huasteca

C. acapulcoensis

C. yucatana

C. perditalis perditalis

C. montezuma

C. laverna laverna

Charis gynaea zama

C. velutina

Chalodeta chaonitis

Carta ino melicerta

C. domitianus vejento

C. rhacotis rhacotis

C. lampeto

Baeotis zonata simbla

B.  sulphurea macularia B. sulphurea sulphurea Lasaia meris

L. agesilas callaina

L. sessilis

L. maria anna

X X

X X X X X X X

X

X X

X X

X X X X X X X X X X X

X X

X X

X X

X X

X X

X X

Volume 58, Number 3

139

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

L. narses*

L. sula peninsularis

Melanis pixe pixe

M. cephise cephise

M. cephise huasteca

Mesene croceella

M. margaretta margaretta

Xenandra caeruleata

Chimastrum argenteum argenteum

Symmachia rubina rubina

S. accusatrix

Symmachia probetor championi

S. tricolor hedemanni

Pterographium sagaris tyriotes

Sarota gamelia

S. acanthoides myrtea

S. chrysus dematria

Anteros formosus micon

A. carausius carausius

Calydna lusca venusta

C. sturnida hegias

C. sinuata

Emesis aurimna

E. saturata

E. liodes

E. mandana furor

E. vulpina

E. fatimella nobilata

E. tenedia tenedia

E. lupina

E. ocypore aethalia

E. zela zela

E. emesia emesia

X X X X X X

X X X X X

E. cypria)

E. tegula

E. zela cleis

Argyrogrammana holosticta

Pseudonymphidia clearista

Pachythone gigas

Apodemia multiplaga

A. hypoglauca hypoglauca

A. walkeri

Thisbe irenea helides

T. lycorias lycorias

Lemonias caliginea

L. agave

Juditha molpe

Synargis calyce mycone

S. ethelinda nymphidioides

S. nycteus

Menander menander purpurata

Pandemos godmanii

Calospila pelarge

C. sudias

Theope pedias isia

T. virgilius virgilius

X X

X X

X X X X

X X

X X X X

X X

140

Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Theope eupolis

T. cratylus

T. publius

T. eleutho

T. mania

T. diores

Calociasrria lilina

Nymphidium ascolia ascolides

Brephidium exilis exilis

Leptotes marina

L. cassius striata

Zizula cyna cyna

Hemiargus ceraunus

H. isola isola

H. huntingtoni hannoides

Everes comijntas

E. amyntula amyntula

Celastrina argiolus gozora

Eumaeus childrenae

E. toxea

Paiwarria antigonus

Theorema eumenia

"Thecla" (busa group) busa

Evenus regalis

E. coronata

E. batesii

"Thecla" (gibberosa group) barajo

"Thecla" (eunus group) eunus

Allosmaitia strophius

Vseudolycaena damo

Areas imperialis

A. cypria

Theritas mavors

"Thecla" (hemon group) augustinula

"Thecla" (hemon group) theocritus

"Thecla" (hemon group) hemon

"Thecla" (hemon group) lisus

Atlides gaumeri

A. polybe

A. inachus

A. carpasia

A. halesus

A. caranus*

Radissima umbratus

"Thecla" (ligurina group) ligurina

"Thecla" (ligurina group) lyde

Denivia theocritus

Contrafacia ahola

C. imma

Thereus cithonius

T. oppia

T. ortalus

Arawacus togarna

A. sito

A. jada

Rekoa meton

X X X

X X

X X X

X X

X X X

X X X

Volume 58, Number 3

141

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Rekoa palegon

R. zebina

R. marius

R. stagira

Ocaria petelina

O. thales

O. ocrisia

Magnastigrna elsa

Chlorostrymon simaethis

C. telea

Cyanophrys goochoni

C. arnyntor

C. Justus

C. herodotus

C. miserabilis

C. longula

Panthiades bitias

P. ochus

P. bathildis

P. phaleros

Oenornaus ortygnus

0.  atesa

Parrhasius polibetes P. orgia

P. moctezuma Michaelus jebus M. thordesa M. hecate M. vibidia M. zenaida Ignata gadira

1.  nr. gadira I. norax

Strymon melinus S. albata

S. alea

S. bazochii

S. mulucha

S. yojoa

S. cestri

S. astiocha

S. istapa

S. ziba

S. nr. megarus #7

S. serapio

S. basalides0

Lamprospilus collucia

"Thecla" (anuria group) gailiena

Kisutam syllis

K. hesperitis

K. ceromia

K. denarius

K. guzanta

K. sethon

"Thecla" (camissa group) vespasianus

Electrostryrnon mathewi

X

X

X

X X

X X

X X

X X

142

Journal of the Lepidopterists' Society

Appendix 1. Continued

Taxon

CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP

Electrostrymon sangala

E. canus

Calycopis calus

C. demonassa

C. atnius

C. clarina

C. isobeon

C. susanna

C. drusilla

C. trehula

Tmolus echion

T. crolinus

T. cydrara

Crimsinota phobe

"Tmolus" (heraldica group) heraldica

"Thecla" (empusa group) halciones

"Thecla" (keila group) keila

Siderus philinna

S. gargophia

S. caninius

S. thoria

Aubergina paetus

"Thecla" (my con group) my con

"Thecla" (tephraeus group) tephraeus

"Thecla" (tephraeus group) syedra

"Thecla" (tephraeus group) ambrax

Ministrymon clytie

M. arola

M. una

M. inoa

M. phrutus

M. azia

Janthecla janthodonia

J. janthina

Ipidecla schausi

Brangas neora

B. getus

B. coccineifrons

B.  carthaea

"Thecla" (cupentus group) cupentus Chalybs janias

C.  hassan

"Thecla" (theia group) theia

Hypostrymon critola

Iaspis nr castitas

I. temesa

Nesiostrymon calchinia

N. celona

N. dodava

Erora carta

E. opisena

E. muridosca

"Thecla" (ares group) semones

Caerofethra carnica

Celmia celmus

"Thecla" (color group) conoveria______

X X X X

X X

X X

X X

X X

X X

X X

X X

X X X X

X X X

X X X X X

X X X

X X X

X X X X X X X X X X X X X

X X X

X X X X X X X X X X