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THE SOUTHWESTERN NATURALIST 48(4):505–525 DECEMBER 2003 BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF CALAKMUL, CAMPECHE, ME ´ XICO CARMEN POZO,* ARMANDO LUIS MARTI ´ NEZ,SANTIAGO UC TESCUM,NOEMI ´ SALAS SUA ´ REZ, AND AIXCHEL MAYA MARTI ´ NEZ Museo de Zoologı ´a-ECOSUR, El Colegio de la Frontera Sur, A.P. 424, C.P. 77000, Chetumal, Q. Roo. Me ´xico (CP, SUT, NSS, AMM) Museo de Zoologı ´a ‘‘Alfonso L. Herrera,’’ Facultad de Ciencias, Universidad Nacional Auto ´noma de Me ´xico, Ciudad Universitaria, A.P. 70-399, C.P. 04510, Me ´xico, D. F (ALM) *Correspondent: [email protected] ABSTRACT We collected 423 species of diurnal butterflies (Rhopalocera) during 250 days of observations from March 1997 through January 2000 in the region of Calakmul, Campeche, Mex- ico. We present a list of 428 species recorded for Calakmul. This is the first reported list of butterflies for the state of Campeche, and it includes 290 new records for the state, 374 new records for Calakmul, and 1 new record for Mexico: Monca telata. Museum records were found for 5 additional species: Caria melino, Cissia labe, Tmolus echion, Emesis lupina lupina, and Calephelis azteca. The accumulation curve of species estimates that 457 species should be present in Calak- mul, which suggests the list presented is 93% complete. We found that medium, tropical, semi- evergreen forest is the richest (371 species); more than a quarter of the butterflies of the Calakmul region were found exclusively in this forest. We also found the southern area slightly more abun- dant in species (82%) than the northern area (79%). Additionally, a comparison was made of the list generated for Campeche to lists from previous studies for Quintana Roo, Tikal-Guatemala National Park, and Belize. The comparison shows that 70%, 58%, and 54% of the butterfly species in Calakmul also are found in Tikal, Quintana Roo, and Belize, respectively. RESUMEN Recolectamos 423 especies de mariposas diurnas (Rhopalocera) de la regio ´n de Ca- lakmul, en el estado de Campeche, Me ´xico, desde marzo de 1997 a enero del 2000 completando 250 dı ´as de muestreo. Se presenta una lista de 428 especies registradas para Calakmul. Esta es la primera lista reportada de mariposas para el estado de Campeche y en ella se presentan 290 nuevos registros para este estado, 374 nuevos registros para la regio ´n de Calakmul y un nuevo registro para Me ´xico: Monca telata. Cinco especies ma ´s fueron encontradas en ejemplares de mu- seos: Caria melino, Cissia labe, Tmolus echion, Emesis lupina lupina y Calephelis azteca. Se presenta una curva de acumulacio ´n de especies con un nu ´ mero teo ´rico de especies esperadas de 457. De acuerdo con este ca ´lculo, la lista presentada esta ´ completa en un 93%. Encontramos que la selva tropical, mediana subperennifolia es mayor en riqueza especı ´fica (371), incluso ma ´s de un cuarto de las especies reportadas aquı ´ para Calakmul se encontro ´ exclusivamente en este tipo de vege- tacio ´n. Tambie ´n encontramos que el sur es ligeramente ma ´s rico en especies (82%) que la regio ´n del norte de Calakmul (79%). Adicionalmente, se hizo una comparacio ´n con listas de especies encontradas en el estado de Quintana Roo, en el Parque Nacional Tikal-Guatemala y en Belice, donde se muestra que el 70%, 58% y 54% de las especies de Calakmul se comparten con Tikal, Quintana Roo y Belice, respectivamente. The Calakmul Biosphere Reserve (CBR) was designated in 1989 to protect both biological and cultural heritage. With 723,185 ha, CBR is the largest protected tropical forest in Mexico. The area also includes extensive Mayan ar- chaeological sites. After the collapse of the Ma- yan culture, the area was sparsely occupied. It was not until 1940 that the logging of mahog- any (Swietenia macrophylla) and cedar (Cedrela odorata) led to the settlement of Zoh-Laguna, which is located in the center of the region. From 1960 to the present, the area has become accessible by road and many settlements have started as a consequence of the government

Transcript of BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

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THE SOUTHWESTERN NATURALIST 48(4):505–525 DECEMBER 2003

BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF CALAKMUL,CAMPECHE, MEXICO

CARMEN POZO,* ARMANDO LUIS MARTINEZ, SANTIAGO UC TESCUM, NOEMI SALAS SUAREZ,AND AIXCHEL MAYA MARTINEZ

Museo de Zoologıa-ECOSUR, El Colegio de la Frontera Sur, A.P. 424, C.P. 77000, Chetumal, Q. Roo. Mexico(CP, SUT, NSS, AMM)

Museo de Zoologıa ‘‘Alfonso L. Herrera,’’ Facultad de Ciencias, Universidad Nacional Autonoma de Mexico,Ciudad Universitaria, A.P. 70-399, C.P. 04510, Mexico, D. F (ALM)

*Correspondent: [email protected]

ABSTRACT We collected 423 species of diurnal butterflies (Rhopalocera) during 250 days ofobservations from March 1997 through January 2000 in the region of Calakmul, Campeche, Mex-ico. We present a list of 428 species recorded for Calakmul. This is the first reported list ofbutterflies for the state of Campeche, and it includes 290 new records for the state, 374 newrecords for Calakmul, and 1 new record for Mexico: Monca telata. Museum records were foundfor 5 additional species: Caria melino, Cissia labe, Tmolus echion, Emesis lupina lupina, and Calephelisazteca. The accumulation curve of species estimates that 457 species should be present in Calak-mul, which suggests the list presented is 93% complete. We found that medium, tropical, semi-evergreen forest is the richest (371 species); more than a quarter of the butterflies of the Calakmulregion were found exclusively in this forest. We also found the southern area slightly more abun-dant in species (82%) than the northern area (79%). Additionally, a comparison was made of thelist generated for Campeche to lists from previous studies for Quintana Roo, Tikal-GuatemalaNational Park, and Belize. The comparison shows that 70%, 58%, and 54% of the butterfly speciesin Calakmul also are found in Tikal, Quintana Roo, and Belize, respectively.

RESUMEN Recolectamos 423 especies de mariposas diurnas (Rhopalocera) de la region de Ca-lakmul, en el estado de Campeche, Mexico, desde marzo de 1997 a enero del 2000 completando250 dıas de muestreo. Se presenta una lista de 428 especies registradas para Calakmul. Esta es laprimera lista reportada de mariposas para el estado de Campeche y en ella se presentan 290nuevos registros para este estado, 374 nuevos registros para la region de Calakmul y un nuevoregistro para Mexico: Monca telata. Cinco especies mas fueron encontradas en ejemplares de mu-seos: Caria melino, Cissia labe, Tmolus echion, Emesis lupina lupina y Calephelis azteca. Se presenta unacurva de acumulacion de especies con un numero teorico de especies esperadas de 457. Deacuerdo con este calculo, la lista presentada esta completa en un 93%. Encontramos que la selvatropical, mediana subperennifolia es mayor en riqueza especıfica (371), incluso mas de un cuartode las especies reportadas aquı para Calakmul se encontro exclusivamente en este tipo de vege-tacion. Tambien encontramos que el sur es ligeramente mas rico en especies (82%) que la regiondel norte de Calakmul (79%). Adicionalmente, se hizo una comparacion con listas de especiesencontradas en el estado de Quintana Roo, en el Parque Nacional Tikal-Guatemala y en Belice,donde se muestra que el 70%, 58% y 54% de las especies de Calakmul se comparten con Tikal,Quintana Roo y Belice, respectivamente.

The Calakmul Biosphere Reserve (CBR) wasdesignated in 1989 to protect both biologicaland cultural heritage. With 723,185 ha, CBR isthe largest protected tropical forest in Mexico.The area also includes extensive Mayan ar-chaeological sites. After the collapse of the Ma-yan culture, the area was sparsely occupied. It

was not until 1940 that the logging of mahog-any (Swietenia macrophylla) and cedar (Cedrelaodorata) led to the settlement of Zoh-Laguna,which is located in the center of the region.From 1960 to the present, the area has becomeaccessible by road and many settlements havestarted as a consequence of the government

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strategy to encourage land settlement of thearea (Weber, 1999). The settlements are in andaround CBR, and the predominant economicactivities for this region are agriculture andforestry. Preliminary results indicate that theBiosphere Reserve supports significant biolog-ical diversity. The area is comprised of 7 trop-ical ecosystems, representing a diverse fauna ofvertebrates (Aranda, 1991; Smith et al., 2001;Calderon et al., 2003), invertebrates (Pozo yCedeno-Vazquez, 1998), and plants (Martınezet al., 2001). The growth of the human com-munity and the need for more usable land hasproduced a mosaic landscape of primary for-est, second-growth forest, and agriculturalfields (Martınez and Galindo-Leal, 2002).

There are no works reporting organized in-ventories of butterflies in Campeche. Thestudy on Mexican butterflies by Godman andSalvin (1879–1901) cited only 2 species of but-terflies for Campeche: Eurema daira eugeniaand Anaea troglodyta aidea. Eurema daira eugeniahad also been previously cited by Felder(1869). Of the few works that report speciesfor Campeche, the most important works arethose by Field (1939), who reported 20 speciesof Papilionoidea, and Hoffmann (1940, 1941),who reported 39 species of Papilionoidea and15 of Hesperiidae. This study represents thefirst organized inventory of butterflies forCampeche. The purpose of our investigationwas to monitor amphibians, reptiles, and but-terflies of the Calakmul area. In this paper, wepresent the butterfly inventory.

METHODS Study Area The Calakmul region(198159 to 178509N, 908209 to 898009W) is located inMexico in the southern part of the Yucatan Penin-sula, which extends from the Gulf of Honduras tothe Terminos Lagoon (Miranda, 1958). Calakmul isin the state of Campeche and borders the El Petenarea of Guatemala (Fig. 1). This region is character-ized by a flat lowland landscape with no importantterrain elevations (100 to 250 m). Soils are shallowand calcareous and overlie a limestone platform(Stedman-Edwards, 2000). Precipitation is highlyseasonal. A rainy season extends from June to No-vember and a dry season from December to May.Annual precipitation varies from an average of 550mm to 1,634 mm, producing dry and wet years. Al-though flooding occurs during the rainy season,once the rain stops, little water remains on the sur-face (Galindo-Leal, 2001). The mean annual tem-perature is 24.68C.

The forest in this region is transitional between

the dryer scrub forest of the north of the YucatanPeninsula to the humid tropical forest of the southin El Peten, Guatemala (Stedman-Edwards, 2000).The CBR has 50% medium tropical semi-evergreenforest, 35% low tropical semi-evergreen forest, and5% high tropical evergreen forest, with the remain-der of the habitats as grasslands and wetlands (INEy CONABIO, 1995; Garcıa-Gil et al., 2001). In thehigh tropical evergreen forest (H), located south ofthe CBR, the canopy can be higher than 25 m andsome trees reach 40 m. The trees that dominate thisforest belong to the family Sapotaceae. The mostcommon species are Manilkara zapota, M. chicle, Pou-teria sapota, P. amygdalina, P. campechiana, and P. retic-ulata (Galindo-Leal, 2001). In the medium tropicalsemi-evergreen forest (M), the canopy reaches 15 to25 m, and 25 to 50% of these trees are deciduous.The commonest species are Brosimun alicastrum, M.zapota, and Pouteria reticulata. In some areas, treeslose almost 75% of their leaves. In this case, the mostcommon species are Guaiacum sanctum, Esenbeckia(Rutaceae), Beucarnea pliabilis, and Thouinia pauci-dentata. In the low tropical semi-evergreen forest(L), the canopy never reaches more than 15 m, and75% of these trees are deciduous. This forest is dom-inated by T. paucidentata, B. pliabilis, G. sanctum, Lon-chocarpus yucatanensis, Bursera simaruba, Haematoxy-lum campechianum, Ceiba schotti, Pseudobombax ellipti-cum, and Maytenus schippi.

Literature and Collections We consulted the com-puterized database at the Museum of Zoology, Fa-cultad de Ciencias (MZFC) at the Universidad Na-cional Autonoma de Mexico. The database includesdata from Mexican butterfly specimens examined inforeign and Mexican museums and collections, aswell as data from the literature.

Sampling Protocol The inventory effort began inMarch 1997 and ended in January 2000, with a totalof 250 days in the field. This is part of a larger studyin which we conducted standardized surveys forlong-term monitoring in both disturbed and pro-tected habitats with a sampling protocol similar toAustin et al. (1996) in Tikal National Park, Guate-mala. We chose 6 localities within or near the north-ern portion of the CBR and 5 more within or nearof southern portion (Fig. 1). To attract some species,we used 35 Van Someren-Rydon traps (Rydon, 1964)baited with banana, pineapple, and beer. We alsoused hand nets and visual identification when pos-sible. During the first year, we collected in the wetand dry season, the second year we collected onlyduring the wet season, and the third year we visitedthe localities on a monthly basis. Of all the individ-uals captured, about 30% were collected and placedin glassine bags labeled with date, collector, locality,and vegetation type. For specimens with difficultidentification, we were advised by J. Llorente-Bous-quets from MZFC, A. D. Warren from Oregon State

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FIG. 1 Map of the Calakmul Biosphere Reserve, Mexico. Study sites indicated by a black circle. Longestdistance between northern sites and southern sites was 112.5 km.

University, and G. Austin from the Nevada NaturalHistory Museum. Specimens are currently stored atthe Museum of Zoology at El Colegio de la FronteraSur-Chetumal, Mexico. Duplicate specimens are de-posited at MZFC, Mexico.

Analysis We combined data from the literature,museum records, and the data we obtained in thefield to construct a table in which we specified thenew records for the Calakmul region, for the state

of Campeche, and for Mexico. We also reported thevegetation type and the area of distribution for eachspecies we recorded. The species recorded 3 or few-er times in a year were considered rare, whereascommon species were those recorded in at least 25%of the study sites and during 25% of the samplingdays (Sparrow et al., 1994). Our species list followsthe phylogenetic classification by Lamas et al. (pers.comm.). We checked the adequacy of our inventory

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FIG. 2 Species cumulative curve for butterflies inall vegetation types surveyed in Calakmul BiosphereReserve, Mexico. Asymptote calculated by Clenchmethod. Number of species expected (Se) for studyarea is 457.

FIG. 3 Species richness of butterflies found ineach type of vegetation in Calakmul Biosphere Re-serve, Mexico. Species shared among 3 forest typesreferred to as Generalist. L 5 low, tropical, semi-evergreen forest; M 5 medium, tropical, semi-ever-green forest; H 5 high, tropical, evergreen forest.See text for description of forest types.

FIG. 4 Richness of butterfly species shared by andexclusive to the low, tropical, semi-evergreen forestand medium, tropical, semi-evergreen forest by fam-ily.

by generating a species accumulation curve usingthe EstimateS 5 software (Colwell, 1997). We calcu-lated the theoretical species number for this areausing the method described by Clench (1979).

RESULTS We recorded in the field a total of65,385 individuals, representing 423 species ofbutterflies from southwestern Campeche, in-cluding 230 genera in 20 subfamilies and 5families in the superfamilies Hesperioidea andPapilionoidea. Only 5 species were not collect-ed, but were found in museums. These are Car-ia melino, Cissia labe, Tmolus echion, Emesis lupinalupina (Coleccion Nacional de Insectos. Insti-tuto de Biologıa, Universidad Nacional Auton-oma de Mexico CNIN), and Calephelis azteca(American Museum of Natural History). Ofthe 423 species collected, only 129 had beenpreviously reported or collected for Campe-che, and only 47 for the Calakmul region. Wereport 290 species in Campeche for the firsttime. For the Calakmul area, 374 species arenew records. One species, Monca telata, is a newrecord for Mexico. The theoretical speciesnumber calculated for this area was 457 species(Fig. 2). We present a list of 428 species of but-terflies from the Calakmul region (Appendix1). The list includes 231 genera in 20 subfam-ilies.

We found the medium semi-evergreen trop-ical forest the richest (371 species), represent-ing 88% of the total species recorded in theCalakmul area. We found 27% of the speciesin Calakmul only in the M forest (Fig. 3). Inthe H forest, we found 22% of the species. Sixspecies (Philaethria diatonica, Castilia ofella, Ti-gridia acesta ssp., Mechanitis menapis doryssus, Me-chanitis polymnia lycidice, and Zizula cyna cyna)are restricted to this forest type. The L forest

shared 86% of species with the M forest. Themajority of species restricted to 1 type of veg-etation were Hesperiidae (84 species) followedby lycaenid species (53 species). These were re-stricted to the L and M forest (Fig. 4).

About 60% of the species were distributedin the entire region. The remaining specieswere found exclusively in the south (21%) orthe north (18%). The south had a slightlymore abundant representation. When compar-ing our list with other studies for QuintanaRoo, Tikal-Guatemala National Park, and Be-lize, we found that 70% of the butterflies listedfor Calakmul can be also found in Tikal. Thenumbers of species shared with Quintana Rooand Belize are much less, with 58% and 54%shared, respectively (Table 1).

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TABLE 1—Distribution of species recorded fromCalakmul (CAL; this study), contiguous state ofQuintana Roo (QROO; de la Maza and Gutierrez,1992), country of Belize (BEL; Meerman, 1993,1999), and the park of Tikal, Guatemala (TIKAL;Austin et al., 1996), showing percentages of Calak-mul species unique to each region. Numbers in pa-rentheses are the number of the species exclusivelyfrom Calakmul

CAL totalCAL-

TIKALCAL-

QROOCAL-BEL

CAL-QROO-

BEL-TIKAL

428 (66)100% (15%)

29870%

24958%

23054%

15837%

DISCUSSION According to the theoreticalspecies total (SE 5 457) calculated for Calak-mul, we sampled 93% of its total butterfly fau-na. Based on these calculations, our list is al-most a complete species list of this region;however, it is important to point out that themain study effort was made in 2 types of veg-etation in the region: the L forest and M forest.There are some places in the south with patch-es of H forest, but these areas are difficult toreach in the rainy season. We think some ofthe species reported by Austin et al. (1996) forTikal might also be found in these inaccessibleH forest patches in southern Calakmul. If weare correct, this could increase our list of spe-cies to 665.

The proportion of the species shared be-tween L and M forests is similar to the propor-tion of the tree species shared between thoseforests (Perez-Salicrup, in press) in the Calak-mul region (86% versus 82%, respectively).Also, the proportion of the species endemic tothe L forest (9%) is the same for the tree spe-cies (Perez-Salicrup, in press).

Fifteen percent of the fauna recorded in Ca-lakmul has not been recorded in QuintanaRoo, the border region of Belize, or the Tikalarea in Guatemala. The species lists reportedfor Belize (Meerman, 1993, 1999) and Quin-tana Roo (de la Maza-Elvira and Gutierrez,1992) were obtained through a shorter timeand with a different survey method. We believethat increasing the sampling effort at QuintanaRoo likely would increase the number of spe-cies for Quintana Roo, thus increasing the

number of species shared between Calakmuland Quintana Roo. This is especially true forthe southern part of Quintana Roo, where thelandscape is similar to Calakmul. The case ofBelize is different, because it has habitats notpresent in Calakmul. The study by Meerman(1999) included butterflies recorded in locali-ties of the Maya Mountains. This is the richestarea for butterfly species in Belize (Meerman,1999), and it has a cloud forest that is notfound in any other area of the Yucatan Pen-insula.

Regarding the 5 species found in museumsand not observed in our field work, we knowthat Tmolus echion, found in the CNIN collec-tion for the Calakmul region, was reported inabundance in Quintana Roo (de la Maza-Elviraand Gutierrez, 1992), and it also is reportedfor Tikal (Austin et al., 1996) and Belize(Meerman, 1999). In Tikal, T. echion is foundin the L forest, the same kind of habitat thatwe surveyed in the region of Calakmul 35 kmsouth of X-Pujil. We cannot explain why we didnot find T. echion in Calakmul. Emesis lupinalupina was reported for the north and centerof Quintana Roo, and in Corozal and the MayaMountains in Belize. The species Caria melinois rare in the southern part of Quintana Roo.Cissia labe and Calephelis azteca are only foundin the Maya Mountains in Belize, making itlikely that we would not find either of these 2species in Calakmul (even considering a great-er survey effort for Quintana Roo or Belize).

The species distribution among the 5 fami-lies is similar to those reported for Tikal Parkby Austin et al. (1996): 4.2% of Papilionidae,4.9% of Pieridae, 25.7% of Nymphalidae, 25%of Lycaenidae, and 40.2% of Hesperiidae.These proportions are also similar to those re-ported by Llorente-Bousquets et al. (1996) forMexico, but are somewhat different from thosereported by Heppner (1991) for the Neotrop-ical butterfly fauna. In our list, hesperiids andpapilionids are over-represented. It is neces-sary to continue the systematic collection anddocumentation of the butterflies in all thestates of Campeche and Quintana Roo, andalso in the country of Belize.

We are grateful to the following people for theirhelp in identifying certain species in the followinggroups, Hesperiidae: A. Warren (Oregon State Uni-versity); Theclinae: R. K. Robbins (USNM); and Ly-

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caenidae: G. T. Austin (Nevada Natural History Mu-seum). We thank A. T. Novelo for his highly profes-sional and meticulous work in the field, and J. A.Padilla-Saldıvar for the elaboration of the map. Wealso thank J. Llorente-Bousquets, M. Balcazar Lara,and C. Galindo Leal for their constructive commentson the manuscript and S. Kissmann and J. J. Mor-rone Lupi for the English review. Funding for thisstudy was provided by CONABIO (projects J112 andQ049), CONACyT (32002), Pronatura Penınsula deYucatan, The Nature Conservancy, and AICAS.

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Submitted 14 December 2001. Accepted 16 December 2002.Associate Editor was Steven Goldsmith.

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AP

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IX1—

Lis

tof

butt

erfl

ies

ofC

alak

mu

l.th

elis

tin

clu

des

5fa

mili

es,4

ofth

esu

per

fam

ilyP

aplio

noi

dea

and

1of

the

Hes

per

ioid

ea.S

eete

xtfo

rd

escr

ipti

onof

vege

tati

onty

pes

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

PAP

ILIO

NID

AE

(18

spec

ies)

Bat

tus

phile

nor

acau

da(O

bert

hu

r,18

79)

Bat

tus

poly

dam

aspo

lyda

mas

(Lin

nae

us,

1758

)B

attu

sla

odam

asco

pana

e(R

eaki

rt,

1863

)Pa

ride

sse

sost

ris

zest

os(G

ary,

[185

3])

Pari

des

erith

alio

npo

lyze

lus

(C.

Feld

er&

R.

Feld

er,

1865

)Pa

ride

sip

hida

mas

iphi

dam

as(F

abri

ciu

s,17

93)

Prot

ogra

phiu

mep

idau

sep

idau

s(D

oubl

eday

,18

46)

Prot

ogra

phiu

mph

ilola

usph

ilola

us(B

oisd

uva

l,18

36)

Prot

ogra

phiu

mag

esla

usne

osila

us(H

opff

er,

1865

)

C C CK

CK

C CK

CK

CK

CK

4 4,13

4,13

13 4,13

AM

NH

CN

INA

MN

HA

MN

H,

CN

INA

MN

H

M G L-M

L-M

L-M

G L-M

L-M

G

S/r

B B S S B B B BPr

otes

ilaus

mac

rosi

laus

pent

hesi

laus

(C.

Feld

er&

R.

Feld

er,

1865

)Pr

iam

ides

roge

ri(B

oisd

uva

l,18

36)

Pria

mid

esan

chis

iade

sid

aeus

(Fab

rici

us,

1793

)T

roili

des

torq

uatu

sm

azai

(Beu

tels

pac

her

,19

77)

Her

aclid

esth

oas

auto

cles

(Rot

hsc

hild

&Jo

rdan

,19

06)

Her

aclid

escr

esph

onte

s(C

ram

er,

1777

)C

alai

des

orny

thio

nor

nyth

ion

(Boi

sdu

val,

1836

)C

alai

des

asty

alus

palla

s(G

ray,

[185

3])

Pter

ouru

sm

enat

ius

vict

orin

us(D

oubl

eday

,18

44)

C C C C CK

CK

C CK

C

3,13

3,13

4

AM

NH

AM

NH

AM

NH

M G L-M

M G G L-M

M L

B B S S/r

B B B S/r

B

PIE

RID

AE

(21

spec

ies)

Ant

eos

clor

inde

(God

art,

[182

4])

Ant

eos

mae

rula

(Fab

rici

us,

1775

)Ph

oebi

sag

arith

eag

arith

e(B

oisd

uva

l,18

36)

Phoe

bis

arga

nte

ssp

.Ph

oebi

sph

ilea

phile

a(L

inn

aeu

s,17

63)

Phoe

bis

senn

aem

arce

llina

(Cra

mer

,17

77)

C CK

CK

CK

CK

CK

3 3,21

3

CN

INSD

NH

M,

CN

INU

SNM

,C

NIN

AM

NH

,C

NIN

CN

IN

G G G G G G

B B/

cB

/c

B B/

cB

Rha

bdod

ryas

trite

ssp

.A

phri

ssa

stat

ira

stat

ira

(Cra

mer

,17

77)

Aba

eis

nici

ppe

(Cra

mer

,17

79)

Pyri

sitia

dina

wes

twoo

di(B

oisd

uva

l,18

36)

Pyri

sitia

lisa

cent

ralis

(Her

rich

-Sch

affe

r,18

65)

C CK

CK

CK

CK

21 21 3

SDN

HM

,C

NIN

CN

INU

SNM

,C

NIN

AM

NH

,C

NIN

M L-M

L-M

L-M

M

S/r

B B B N/

r

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December 2003 513Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Pyri

sitia

nise

nelp

he(R

.Fe

lder

,18

69)

Pyri

sitia

prot

erpi

apr

oter

ipa

(Fab

rici

us,

1775

)Eu

rem

aal

bula

cela

ta(R

.Fe

lder

,18

69)

Eure

ma

arbe

labo

isud

valia

na(C

.Fe

lder

&R

.Fe

lder

,18

65)

Eure

ma

dair

aeu

geni

a(W

alle

ngr

en,

1860

)K

rico

goni

aly

side

(God

art,

1819

)

CK

CK

CK

CK

CK

3 1,2,

2011

,21

AM

NH

,N

evad

a,C

NIN

CN

INA

MN

HC

NIN

CN

INA

MN

H,

CN

IN

G G G L-M

G G

B/

cB B B B B

/c

Glu

toph

riss

adr

usill

ate

nuis

(Lam

as,

1891

)Pi

erib

allia

viar

divi

ardi

(Boi

sdu

val,

1836

)A

scia

mon

uste

mon

uste

(Lin

nae

us,

1764

)G

anyr

ajo

seph

ina

jose

pha

(Sal

vin

&G

odm

an,

1868

)

CK

CK

CK

21 3 3,21

SDN

HM

AM

NH

,C

NIN

Nev

ada

AM

NH

,C

NIN

G G G L-M

B/

cB

/c

B B

NYM

PH

AL

IDA

E(1

10sp

ecie

s)

Phila

ethr

iadi

aton

ica

(Fru

hst

orfe

r,19

12)

Dio

neju

nohu

ascu

ma

(Rea

kirt

,18

66)

Agr

aulis

vani

llae

inca

rnat

a(R

iley,

1926

)D

ryad

ula

phae

tusa

(Lin

nae

us,

1758

)

C C CK

CK

3A

MN

H,

Nev

ada

AM

NH

H G G

S/r

B B BD

ryas

iulia

mod

erat

a(R

iley,

1926

)Eu

eide

sal

iphe

ragr

acili

sSt

ich

el,

1903

Euei

des

isab

ella

eva

(Fab

rici

us,

1793

)H

elic

oniu

sch

arito

nia

vazq

ueza

eW

.P.

Com

stoc

k&

F.M

.B

row

n,

1950

CK

CK

CK

CK

9

USN

M,

AM

NH

CN

INA

MN

H,

CN

INU

SNM

,A

MN

H,

CN

IN

G G G G

B/

cB B B

/c

Hel

icon

ius

erat

ope

tiver

anus

Dou

bled

ay,

1847

Eupt

oiet

acl

audi

ada

uniu

s(H

erbs

t,17

98)

Eupt

oiet

ahe

gesi

aho

ffm

anni

Stic

hel

,19

38A

nart

iaam

athe

afa

tima

(Fab

rici

us,

1793

)A

nart

iaja

trop

hae

lute

ipic

taFr

uh

stor

fer,

1907

CK

C CK

CK

3

USN

M,

AM

NH

,C

NIN

CN

IN

SDN

HM

CN

IN

G L-M

G G G

B/

cB B B

/c

B/

kSi

proe

taep

aphu

sep

aphu

s(L

atre

ille,

[181

3])

Sipr

oeta

stel

enes

bipl

agia

ta(F

ruh

stor

fer,

1907

)C C

K4

CN

INC

AS

M GN

/r

B/

cJu

noni

aev

aret

ess

p.

Chl

osyn

eer

odyl

eer

odyl

e(H

.W

.B

ates

,18

64)

CK

CK

AM

NH

,C

NIN

M-H

MB S

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514 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Chl

osyn

ega

udia

lisga

udia

lis(H

.W

.B

ates

,18

64)

Chl

osyn

eja

nais

jana

is(D

rury

,17

82)

Chl

osyn

ela

cini

ala

cini

a(G

eyer

,18

37)

The

ssal

iath

eona

Men

etri

es,

1855

Phyc

iode

sph

aon

(W.

H.

Ed

war

ds,

1864

)A

ntha

nass

aar

gent

ea(G

odm

an&

Salv

in,

1882

)A

ntha

nass

afr

isia

tulc

is(H

.W

.B

ates

,18

64)

Cas

tilia

myi

am

yia

(Hew

itso

n,

[186

4])

CK

C CK

C C C

MZ

FC

Nev

ada,

CN

INN

evad

a,C

NIN

AM

NH

L G G G L-M

L-M

G G

S B B B/

cB B

/r

B/

cB

Cas

tilia

ofel

la(H

ewit

son

,[1

864]

)H

isto

ris

odiu

sdi

ous

Lam

as,

1995

His

tori

sac

hero

nta

ache

ront

a(F

abri

ciu

s,17

75)

Smyr

nabl

omfil

dia

datis

Fru

hst

orfe

r,19

80C

olob

ura

dirc

edi

rce

(Lin

nae

us,

1758

)T

igri

dia

aces

tass

p.

C C C C C C

H G L-M

L-M

G H

S/r

B/

cB B B S/

rB

iblis

hype

ria

agan

isa

Boi

sdu

val,

1836

Mes

tra

dorc

asam

ymon

e(M

enet

ries

,18

57)

Mys

celia

cyan

iris

cyan

iris

Dou

bled

ay,

[184

8]M

ysce

liaet

husa

ethu

sa(D

oyer

e,[1

840]

)C

aton

ephe

lem

exic

ana

Jen

kin

s&

R.

G.

Maz

a,19

85C

aton

ephe

lenu

mili

aes

ite(R

.Fe

lder

,18

69)

Nes

saea

agla

ura

agla

ura

(Dou

bled

ay,

[184

8])

CK

CK

C CK

C

14 16

AM

NH

AM

NH

,N

evad

a,C

NIN

CN

IN

L-M

L-M

L-M

L L L-M

L-M

B/

cB

/c

B B/

cS/

rB B

Euni

caal

cmen

aal

cmen

a(D

oubl

eday

,[1

847]

)Eu

nica

mon

ima

(Sto

ll,17

82)

Euni

cata

tila

tatil

a(H

erri

ch-S

chaf

fer,

[185

5])

Ham

adry

asam

phin

ome

mex

ican

a(L

uca

s,18

53)

Ham

adry

asfe

brua

fere

ntin

a(G

odar

t,[1

824]

)H

amad

ryas

fero

nia

fari

nule

nta

(Fru

hst

orfe

r,19

16)

Ham

adry

asgu

atem

alen

agu

atem

alen

a(H

.W

.B

ates

,18

64)

CK

C CK

C

17,1

9

12 4,12

AM

NH

Nev

ada

AM

NH

,C

NIN

,

CN

IN

CN

IN

G L-M

G M G L-M

G

B B B/

cS/

rB

/c

B BH

amad

ryas

julit

ta(F

ruh

stor

fer,

1914

)Py

rrho

gyra

neae

rea

hyps

enor

God

man

&Sa

lvin

,18

84Py

rrho

gyra

otol

ais

otol

ais

H.

WB

ates

,18

64Te

men

isla

otho

eho

ndur

ensi

sFr

uh

stor

fer,

1907

C C CK

C16

L-M

G L-M

L-M

B/

cB N B

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December 2003 515Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Nic

afl

avill

aba

chia

na(R

.G

.M

aza

&J.

Maz

a,19

85)

Dyn

amin

edy

onis

Gey

er,

1837

Dyn

amin

eth

eseu

s(C

.Fe

lder

&R

.Fe

lder

,18

61)

Dyn

amin

epo

stve

rta

mex

ican

ad

’Alm

eid

a,19

52

C C16

AM

NH

AM

NH

,C

NIN

L-M

L-M

M G

B/

cB N

/r

BC

allic

ore

texa

titan

ia(S

alvi

n,

1869

)A

delp

haba

silo

ides

basi

loid

es(H

.W

.B

ates

,18

65)

Ade

lpha

fess

onia

fess

onia

(Hew

itso

n,

1847

)A

delp

haip

hicl

usip

hicl

eola

(H.

W.

Bat

es,

1864

)

C C CK

4A

MN

HA

MN

H

L L-M

L-M

G

S/r

B B BA

delp

hia

ixia

leuc

asFr

uh

stor

fer,

1915

Ade

lpha

naxi

aep

iphi

cla

(C.

Feld

er&

R.

Feld

er,

1867

)A

delp

haph

ylac

aph

ylac

a(H

.W

.B

ates

,18

66)

Ade

lpha

serp

am

assi

lia(C

.Fe

lder

&R

.Fe

lder

,18

67)

C C C C

L-M

L-M

L G

B B S/r

B/

cM

arpe

sia

chir

onm

ariu

s(C

ram

er,

1779

)M

arpe

sia

petr

eus

ssp

.A

rcha

eopr

epon

ade

mop

hon

cent

ralis

(Fru

hst

orfe

r,19

05)

Arc

haeo

prep

ona

dem

opho

ngu

lina

(Fru

hst

orfe

r,19

04)

Prep

ona

laer

tes

octa

via

Fru

hst

orfe

r,19

05

CK

C CK

17

AM

NH

CN

INC

NIN

CN

IN

G L-M

L-M

G L-M

B/

cB B

/c

B/

cB

Prep

ona

pyle

neph

ileta

sFr

uh

stor

fer,

1904

Zare

tisca

llidr

yas

(R.

Feld

er,

1869

)Si

dero

nega

lant

his

ssp

.A

naea

trog

lody

taai

dea

(Gu

erin

-Men

evill

e,[1

844]

)C

onsu

lel

ectr

ael

ectr

a(W

estw

ood

,18

50)

Foun

tain

eaeu

rpyl

eco

nfus

a(A

.H

all,

1929

)

C C C CK

CK

2,3,

5A

MN

H,

CN

INA

MN

HC

NIN

L-M

L-M

L-M

L-M

G L-M

B B B B/

cB B

/c

Fout

aine

agl

ycer

ium

yuca

tanu

m(W

itt,

1980

)M

emph

isar

taca

ena

(Hew

itso

n,

1869

)M

emph

isfo

rrer

i(G

odm

an&

Salv

in,

1884

)M

emph

ishe

dem

anni

(R.

Feld

er,

1869

)M

emph

isph

ilabo

isdu

vali

W.

P.C

omst

ock,

1961

Mem

phs

pith

yusa

(R.

Feld

er,

1869

)

C CK

CK

C C3

CN

INA

MN

H,

CN

IN

AM

NH

,C

NIN

L-M

M L-M

L-M

L-M

G

B B/

rB

/c

B B/

cB

Ast

eroc

ampa

idyj

aar

gus

(H.

W.

Bat

es,

1864

)D

oxoc

opa

laur

ela

ure

(Dru

ry,

1773

)D

oxoc

opa

pavo

nth

eodo

ra(L

uca

s.18

57)

Mor

pho

achi

lles

mon

tezu

ma

Gu

enee

,18

59O

psip

hane

sin

vira

efa

bric

iiB

oisd

uva

l,18

70)

C CK

CK

3,4

3

AM

NH

,SD

NH

M,

CN

IN,

Nev

ada

AM

NH

AM

NH

,C

NIN

AM

NH

,C

NIN

G G G L-M

G

B B B B/

cB

Page 12: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

516 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Oph

siph

anes

quite

ria

quir

inus

God

man

&Sa

lvin

,18

81C

alig

oat

reus

uran

usH

erri

ch-S

chaf

fer,

1850

Cal

igo

teuc

erm

emno

n(C

.Fe

lder

&R

.Fe

lder

,18

67)

Eryp

hani

sae

sacu

sae

sacu

s(H

erri

ch-S

chaf

fer,

1850

)M

anat

aria

mac

ulat

a(H

opff

er,

1874

)C

ephe

upty

chia

glau

cina

(H.

W.

Bat

es,

1864

)C

issi

ala

be(B

utl

er,

1870

)

C C C C C CK

N6

4C

NIN

L-M

G M-H

M L G

B B B B/

rS B

Cis

sia

pseu

doco

nfus

aSi

nge

r,D

eVri

es&

Eh

rlic

h,

1983

Cis

sia

sp.

Cyl

lops

issp

.H

erm

eupt

ychi

ahe

rmes

herm

es(F

abri

ciu

s,17

75)

Mag

neup

tych

ialib

ye(L

inn

aeu

s,17

67)

Pare

upty

chia

bino

cula

met

aleu

ca(B

oisd

uva

l,18

70)

Pare

upty

chia

ocir

rhoe

ssp

.

C C C

4 3

AM

NH

,C

NIN

CN

INC

NIN

AM

NH

,C

NIN

G L-M

L G L-M

G L-M

B/

cB B

/r

B/

cB B B

/c

Tayg

etis

virg

ilia

(Cra

mer

,17

76)

Tayg

etis

tham

yra

(Cra

mer

,17

79)

Vare

upty

chia

usita

tapi

eria

(C.

Feld

er&

R.

Feld

er,

1867

)Va

reup

tych

iasi

mili

s(B

utl

er,

1867

)Yp

hthi

moi

des

rena

ta(S

toll,

1780

)

C CC

NIN

CN

INC

NIN

G L-M

G G G

B/

cB B

/c

B/

cB

/c

Dan

aus

eres

imus

mon

tezu

ma

Tal

bot,

1943

Dan

aus

gilip

pus

ther

sipp

us(H

.W

.B

ates

,18

63)

Dan

aus

plex

ippu

spl

exip

pus

(Lin

nae

us,

1758

)

CK

CK

CK

7 3

USN

M,

CN

INC

NIN

AM

NH

G G G

B B BL

ycor

eaha

liaat

erga

tisD

oubl

eday

,[1

847]

Mec

hani

tism

enap

isdo

ryss

usH

.W

.B

ates

,18

64M

echa

nitis

poly

mni

aly

cidi

ceH

.W

.B

ates

,18

64Pt

eron

ymia

coty

ttoco

tytto

(Gu

erin

-Men

evill

e,[1

844]

)L

ibyt

hean

aca

rine

nta

mex

ican

aM

ich

ener

,19

43

C C C CK

CK

CN

INC

NIN

M-H

H H G G

S/r

S/r

S/r

B B

LYC

AE

NID

AE

(107

spec

ies)

Euse

lasi

ach

rysi

ppe

(H.

W.

Bat

es,

1866

)Eu

sela

sia

serg

iase

rgia

(God

man

&Sa

lvin

,18

85)

Euse

lasi

am

ystic

a(S

chau

s,19

13)

Euse

lasi

aau

rant

iaca

aura

ntia

ca(S

alvi

n&

God

man

,18

68)

Euse

lasi

asp

.M

esos

emia

tetr

ica

(Sti

chel

,19

10)

C C C C

AM

NH

,C

NIN

M L-M

M M M G

N/

rN

/r

N/

rN

/r

N B

Page 13: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

December 2003 517Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Nap

aea

umbr

aum

bra

(Boi

sdu

val,

1870

)R

hetu

sar

cius

thia

(Mor

isse

,18

38)

Rhe

tus

peri

ande

rna

evia

nus

Stic

hel

,19

10N

othe

me

erot

ass

p.

Cal

ephe

lisw

righ

tiH

olla

nd

,19

30C

alep

helis

fulm

enSt

ich

el,

1910

Cal

ephe

lisst

allin

gsi

McA

lpin

e,19

71C

alep

helis

may

aM

cAlp

ine,

1971

Cal

ephe

lisaz

teca

McA

lpin

e,19

71

C C C C C C C C NA

MN

H

L-M

G M M L-M

L-M

L-M

L-M

B B N N/

rB N S B

Cal

ephe

lisya

utep

eque

nsis

R.

G.

Maz

a&

Tu

rren

t,19

77C

alep

helis

brow

niM

cAlp

ine,

1971

Cal

ephe

listik

alA

ust

in,

1993

Cal

ephe

lissp

.C

hari

sza

ma

H.

W.

Bat

es,

1868

Car

iain

om

elic

erta

Sch

aus,

1890

Car

iast

illat

icia

Dya

r,19

12C

aria

mel

ino

Dya

r,19

12

C C C C C N

CN

IN

CN

IN

L-M

L-M

L-M

G L-M

M M

B B B B/

cB S S/

r

Car

iam

antin

eala

mpe

toG

odm

an&

Salv

in,

1886

Bae

otis

zona

tazo

nata

R.

Feld

er,

1869

Bae

otis

sulp

hure

am

acul

aria

(Boi

sdu

val,

1870

)M

elan

ispi

xepi

xe(B

oisd

uva

l,18

36)

C C C CK

CN

IN

M G M L-M

S B S/r

BM

esen

esi

lari

sG

odm

an&

Salv

in,

1878

Mes

ene

sp.

Sym

mac

hia

accu

satr

ixW

estw

ood

,[1

851]

Saro

taps

aros

psar

osG

odm

an&

Salv

in,

1886

Ant

eros

cara

usiu

sca

raus

ius

Wes

twoo

d,

[185

1]C

alyd

nast

urnu

lahe

gias

R.

Feld

er,

1869

Emes

isau

rim

na(B

oisd

uva

l,18

70)

Emes

ism

anda

nafu

ror

Bu

tler

&H

.D

ruce

,18

72

C C C C C C C

M L-M

M M L-M

L-M

L-M

G

N/

rN N

/r

N/

rS B B B

Emes

iste

nedi

ate

nedi

aC

.Fe

lder

&R

.Fe

lder

,18

61Em

esis

lupi

nalu

pina

God

man

&Sa

lvin

,18

86Em

esis

emes

ia(H

ewit

son

,18

67)

Emes

iste

gula

God

man

&Sa

lvin

,18

86A

rgyr

ogra

mm

ana

stilb

eho

lost

icta

(God

man

&Sa

lvin

,18

78)

N C C

AM

NH

,C

NIN

CN

IN

CN

IN

L-M

G L-M

M

B B B N

Page 14: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

518 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Pseu

dony

mph

idia

clea

rist

a(B

utl

er,

1871

)A

pode

mia

hypo

glau

cahy

pogl

auca

(God

man

&Sa

lvin

,18

78)

Thi

sbe

iren

eabe

lides

Stic

hel

,19

10T

hisb

ely

cori

asly

cori

as(H

ewit

son

,[1

853]

)L

emon

ias

agav

eag

ave

God

man

&Sa

lvin

,18

86Ju

dith

am

olpe

mol

pe(H

ubn

er,

[180

8])

C C C C CA

MN

H,

CN

IN

M M L-M

M M G

S N B B S/r

B/

cSy

narg

isca

lyce

myc

one

(Hew

itso

n,

1865

)Pa

ndem

osgo

dman

ii(D

ewit

z,18

77)

The

ope

virg

ilius

(Fab

rici

us,

1793

)T

heop

eeu

polis

Sch

aus,

1890

The

ope

publ

ius

inco

mpo

situ

sJ.

Hal

l,19

99

C C C CA

MN

H

L-M

L-M

M M M

B B/

rB S/

rN

/r

Cal

ocia

sma

lilin

a(B

utl

er,

1870

)L

epto

tes

cass

ius

stri

ata

(W.

H.

Ed

war

ds,

1877

)Zi

zula

cyna

cyna

(W.

H.

Ed

war

ds,

1881

)H

emia

rgus

cera

unus

Hu

bner

,[1

818]

Ever

esco

myn

tas

(God

art,

[182

4])

Cel

astr

ina

argi

olus

gozo

ra(B

oisd

uva

l,17

0)Eu

mae

usto

xea

(God

art,

1824

)B

rang

asge

tus

(Fab

rici

us,

1787

)

C C CK

C C

AM

NH

,C

NIN

USN

M,

CN

INC

NIN

USN

M,

CN

IN

L-M

G H L-M

G L G L-M

B B/

cS/

rB B N

/r

B/

cB

Even

usre

galis

(Cra

mer

,17

75)

CK

CN

INL

-MB

The

cla

hera

clid

es(G

odm

an&

Salv

in,

1887

)A

tlide

sha

lesu

s(C

ram

er,

1777

)A

tlide

sga

umer

i(G

odm

an,

1901

)A

tlide

spo

lybe

(Lin

nae

us,

1763

)A

tlide

sca

rpas

ia(h

ewit

son

,18

68)

Pseu

doly

caen

ada

mo

(H.

Dru

ce,

1875

)T

heri

tas

hem

on(C

ram

er,

1775

)C

yano

phry

sfu

sius

(God

man

&Sa

lvin

,18

87)

Cya

noph

rys

hero

dotu

s(F

abri

ciu

s,17

93)

Cya

noph

rys

long

ula

(Hew

itso

n18

68)

Rek

oam

eton

(Cra

mer

,17

79)

C C C C C C C C C C CK

CN

IN

L L-M

L-M

L-M

M G M M M G

S/r

B B B S B S/r

S/r

S/r

BR

ekoa

pale

gon

(Cra

mer

,17

80)

Rek

oam

ariu

s(L

uca

s,18

57)

Ara

wac

ussi

to(B

oisd

uva

l,18

36)

CC

NIN

CN

IN

L-M

L-M

G

B B B

Page 15: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

December 2003 519Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Ara

wac

usja

da(H

ewit

son

,18

67)

The

cla

ligur

ina

(Hew

itso

n,

1874

)T

hecl

aly

de(G

odm

an&

Salv

in,

1887

)M

agna

stig

ma

elsa

(Hew

itso

n18

77)

Chl

oros

trym

onsi

mae

this

(Dru

ry,

1773

)C

elm

iaco

nove

ria

(Sch

aus,

1902

0A

llosm

aitia

stro

phiu

s(G

odar

t,18

24)

Lam

pros

pilu

sco

lluci

a(H

ewit

son

,18

77)

The

cla

galli

ena

(Hew

itso

n,

1877

)El

ectr

ostr

ymon

sang

ala

(Hew

itso

n,

1868

)C

alyc

opis

isob

eon

(Bu

tler

&H

.D

ruce

,18

72)

C C C C C C C C C CK

CN

INC

NIN

L-M

L L M M M L M M M M

B/

rB B B N

/r

N/

rN S/

rN

/r

S/r

NSt

rym

onm

elin

us(H

ubn

er,

1813

)St

rym

onyo

joa

(Rea

kirt

,18

67)

Stry

mon

mul

ucha

(Hew

itso

n,

1867

)St

rym

once

stri

(Rea

kirt

,18

67)

Stry

mon

alea

(God

man

&Sa

lvin

,18

87)

Stry

mon

bebr

ycia

(Hew

itso

n,

1868

)St

rym

onis

tapa

(Rea

kirt

,18

67)

Stry

mon

bazo

chii

(God

art,

1824

)

C C C C C C C CK

USN

M

L G G M M M L-M

M

S/r

B S/r

S/r

B/

rS/

rB S

Stry

mon

sera

pio

(God

man

&Sa

lvin

,18

87)

Stry

mon

meg

arus

(God

art,

1824

)C C

M MB

/r

S/r

Stry

mon

ziba

(Hew

itso

n,

1868

)T

mol

usec

hion

(Lin

nae

us,

1767

)C

KN

CN

INC

NIN

MS/

r

Min

istr

ymon

azia

(Hew

itso

n,

1873

)M

inis

trym

onun

a(H

ewit

son

,18

73)

Ost

rino

tes

keila

(Hew

itso

n,

1869

)Pa

nthi

ades

bitia

s(C

ram

er,

1777

)Pa

nthi

ades

bath

ildis

(C.

Feld

er&

R.

Feld

er,

1865

)

C C C CC

NIN

L M L-M

M

N/

rN

/r

B N/

rPa

nthi

ades

phal

eros

(Lin

nae

us,

1767

)T

hecl

aec

helta

(Hew

itso

n,

1867

)Pa

rrha

sius

polib

etes

(Sto

ll,17

81)

Mic

hael

usir

a(H

ewit

son

,18

67)

Cha

lybs

jani

as(C

ram

er,

1779

)

C C C C C

L-M

M M L M

B S/r

N S N

Page 16: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

520 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

HE

SPE

RII

DA

E(1

72sp

ecie

s)

Pyrr

hopy

gem

ulle

ri(B

ell,

1934

)El

bella

scyl

la(M

enet

ries

,18

55)

Mys

celu

sam

ystis

hage

s(G

odm

an&

Salv

in,

1893

)Ph

ocid

esp

alem

onlil

ea(R

eaki

rt,

[186

7])

Phoc

ides

belu

sG

odm

an&

Salv

in,

1893

Phoc

ides

pigm

alio

npi

gmal

ion

(Cra

mer

,[1

779]

)

C C C C CK

CK

3,6

6

M M L-M

L-M

M M

B/

rS/

rB B B S

Phan

usm

arsh

allii

Kir

by,

1880

Udr

anom

iaki

kkaw

ai(W

eeks

,19

06)

Prot

eide

sm

ercu

rius

mer

curi

us(F

abri

ciu

s,17

87)

Epar

gyre

usex

adeu

scr

uza

Eva

ns,

1952

Epar

gyre

ussp

ina

spin

aE

van

s,19

52

C C C C C

L-M

L L-M

L-M

L-M

B S/r

B N BEp

argy

reus

sp.

Poly

gonu

sm

anue

lim

anue

liB

ell

&W

.P.

Com

stoc

k,19

48C

K8

M L-M

B B/

cC

hioi

des

catil

lus

albo

fasc

iatu

s(H

ewit

son

,18

67)

Chi

oide

szi

lpa

(Bu

tler

,18

72)

Agu

naas

ande

ras

ande

r(H

ewit

son

,18

67)

Agu

nacl

axon

Eva

ns,

1952

Agu

naau

runc

ehy

pozo

nius

(Plo

tz,

1880

)A

guna

met

ophi

s(L

atre

ille,

[182

4])

Agu

naco

eloi

des

Au

stin

&M

ielk

e,19

97T

yphe

danu

sun

udla

tus

(Hew

itso

n18

67)

Typ

heda

nus

ampy

x(G

odm

an&

Salv

in,

1893

)T

yphe

danu

ssa

las

H.

A.

Free

man

,19

77

C C C C C C C C C C

M M L-M

L M L-M

L M L-M

L-M

S S/r

B/

rS N

/r

B S/r

N/

rB B

Poly

thri

xoc

tom

acul

ata

(Sep

p,

1848

)Po

lyth

rix

asin

e(H

ewit

son

,18

67)

Cod

atra

ctus

caro

los

Eva

ns,

1952

Cod

atra

ctus

alca

eus

(Hew

itso

n,

1867

)C

odat

ract

usyu

cata

nus

H.

A.

Free

man

,19

77C

oadt

ract

usm

elon

(God

man

&Sa

lvin

,18

93)

C C C C C C

L-M

L-M

M L-M

L-M

M

B/

rB S/

rB S S/

r

Page 17: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

December 2003 521Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Rid

ens

ally

niH

.A

.Fr

eem

an,

1979

Urb

anus

prot

eus

prot

eus

(Lin

nae

us,

1758

)U

rban

usvi

terb

oana

(Eh

rman

n,

1907

)U

rban

usbe

lli(H

ayw

ard

,19

35)

Urb

anus

esm

eral

dus

(Bu

tler

,18

77)

Urb

anus

evon

aE

van

s,19

52

C C C C C C

M L-M

L-M

M L L-M

B/

rB N

/r

N/

rB

/r

BU

rban

usdo

rant

esdo

rant

es(S

toll,

[179

0])

Urb

anus

tele

us(H

ubn

er,

1821

)U

rban

usta

nna

Eva

ns,

1952

Urb

anus

sim

plic

ius

(Sto

ll,[1

790]

)U

rban

uspr

ocne

(Plo

tz,

1881

)U

rban

usdo

ryss

usdo

ryss

us(S

wai

nso

n,

1831

)

C C C C C C

L-M

L-M

L-M

M L-M

L-M

B B B N/

rB

/r

BU

rban

usal

bim

argo

albi

mar

go(M

abill

e,18

75)

Ast

rapt

esfu

lger

ator

azul

(Rea

kirt

,[1

867]

)A

stra

ptes

egre

gius

(Bu

tler

,18

70)

Ast

rapt

esen

otru

s(S

toll,

[178

1])

Ast

rapt

esal

ardu

sla

tiaE

van

s,19

52A

stra

ptes

alec

tor

hopf

feri

(Plo

tz,

1882

)

C C CK

C C C

6

L L-M

L-M

L-M

L L-M

N B B S/r

S/r

BA

stra

ptes

anap

hus

anne

ttaE

van

s,19

52N

arco

sius

pari

sihe

len

(Eva

ns,

1952

)N

arco

sius

sp.

Cal

liade

sze

utus

(Mos

chle

r,18

79)

Aut

ocht

onlo

ngip

enis

(Plo

tz,

1882

)A

utoc

hton

zare

x(H

ubn

er,

[181

])

C C C C C

L-M

L-M

L-M

MM

M

B S S B/

rN N

/r

Aut

ocht

onsp

p.

The

ssia

jala

pus

(Plo

tz,

1882

)A

chal

arus

albo

cilia

tus

albo

cilia

tus

(Mab

ille,

1877

)A

chal

arus

toxe

us(P

lotz

,18

82)

Cab

ares

potr

illo

potr

illo

(Lu

cas,

1857

)N

ascu

sph

ocus

(Cra

mer

,[1

777]

)

C C C C C

M M M L-M

L-M

M

N B N/

rB B N

/r

Page 18: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

522 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Ocy

baca

lath

ana

cala

nus

(God

man

&Sa

lvin

,18

94)

Cel

aeno

rrhi

nus

stol

aE

van

s,19

52Sa

pthi

lepi

acl

oniu

s(C

ram

er,

[177

5])

Cog

iaca

lcha

s(H

erri

ch-S

chaf

fer,

1869

)A

rteu

rotia

trac

tipen

nis

trac

tipen

nis

Bu

tler

&H

.D

ruce

,18

72Po

lyct

orcl

eta

Eva

ns,

1953

C C C C CK

CK

6,15

6,10

L-M

L-M

L-M

L-M

M M

B B/

rB B S/

rN

/r

Nis

onia

des

godm

aE

van

s,19

53N

ison

iade

sru

besc

ens

(Mos

chle

r,18

77)

Nis

onia

des

sp.

Pelli

cia

arin

aE

van

s,19

53Pe

llici

adi

mid

iata

Her

rich

-Sch

affe

r,18

70Pa

chyn

euri

asp

p.

C C C C C

L-M

L L-M

L-M

L L-M

B N/

rB B N B

Stap

hylu

svu

lgat

a(M

osch

ler,

1879

)St

aphy

lus

leni

sSt

ein

hau

ser,

1989

Gor

gyth

ion

begg

apy

ralin

a(M

osch

ler,

1877

)G

orgy

thio

nvo

xE

van

s,19

53Ze

rasp

p.

Qua

drus

ceri

alis

(Sto

ll,[1

782]

)

C C C C C C

M L-M

L-M

M L-M

M

N/

rB

/r

B N/

rB N

Qua

drus

cont

uber

nalis

(Mab

ille,

1883

)Q

uadr

uslu

gubr

islu

gubr

is(R

.Fe

lder

,18

69)

Sost

rata

nord

ica

Eva

ns,

1953

Pach

eslo

xus

zonu

la(M

abill

e,18

89)

Ata

rnes

salle

i(C

.Fe

lder

&R

.Fe

lder

,18

67)

Myl

onm

enip

pus

(Fab

rici

us,

1777

)M

ylon

pelo

pida

s(F

abri

ciu

s,17

93)

C C CK

CK

C C CK

6 6 6

M L-M

L-M

M M L-M

M

N/

rB B N

/r

N/

rB N

Car

rhen

esca

nesc

ens

cane

scen

s(R

.Fe

lder

,18

69)

Xen

opha

nes

tryx

us(S

toll,

[178

0])

Ant

igon

usne

arch

us(L

atre

ille,

[181

7])

Ant

igon

user

osus

(Hu

bner

,[1

812]

)Sy

stas

eapu

lver

ulen

ta(R

.E

lder

,18

69)

Aet

hilla

lavo

chre

aB

utl

er,

1872

C C C C C C

M M M L-M

L-M

L-M

S/r

B/

rS/

rB S B

Page 19: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

December 2003 523Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Ach

lyod

esbu

siru

she

ros

Eh

rman

n,

1909

Eant

isth

raso

(Hu

bner

,[1

807]

)Ea

ntis

tam

enun

d(W

.H

.E

dw

ard

s,[1

871]

)G

rais

stig

mat

icus

stig

mat

icus

(Mab

ille,

1883

)T

imoc

hare

str

ifasc

iata

trifa

scia

ta(H

ewit

son

,[1

868]

)

CK

C C C CK

6 6

L-M

M M L G

N/

rN

/r

S S/r

BT

imoc

hare

sru

ptifa

scia

tus

rupt

ifasc

iatu

s(P

lotz

,18

84)

Ana

stru

sse

mpi

tern

usse

mpi

tern

us(B

utl

er&

H.

Dru

ce,

1872

)A

nast

rus

near

eris

(Mos

chle

r,18

79)

Ebri

etas

sp.

Ebri

etas

anac

reon

(Stu

din

ger,

1876

)C

yclo

glyp

hath

rasi

bulu

s(F

abri

ciu

s,17

93)

C C C C C

M M M M M M

S/r

N/

rS/

rS S/

rN

/r

Hel

ias

cam

aE

van

s,19

53C

ampt

ople

ura

ther

amen

esM

abill

e,18

77C

hiom

ara

geor

gina

geor

gina

(Rea

kirt

,18

6)C

iom

ara

mith

rax

(Mos

chle

r,18

79)

C C C CK

6

M M M L

S/r

S N/

rS/

rG

esta

invi

sus

(Bu

tler

&H

.D

ruce

,18

72)

Pryg

usad

epta

Plo

tz,

1884

Pyrg

usph

ileta

sW

.H

Ed

war

ds,

1881

Pyrg

usoi

leus

(Lin

nae

us,

1767

)H

elio

pete

sm

acai

ra(R

eaki

rt,

[186

7])

Hel

iope

tes

arsa

lte(L

inn

aeu

s,17

58)

C C C C C CK

3

L G M G L-M

G

S/r

B N/

rB B B

Hel

iope

tes

alan

a(R

eaki

rt,

1868

)Sy

napt

epe

cta

Ean

s,19

55Za

rias

pes

mys

(Hu

bner

,[1

808]

)A

ntho

ptus

insi

gnis

(Plo

tz,

1882

)A

ntho

ptus

epic

tetu

s(F

abri

ciu

s,17

93)

Cor

ticea

cort

icea

(Plo

tz,

1883

)

C C C C C C

L-M

L-M

L-M

M M L-M

B B B N/

rN B

/r

Cor

ticea

lysi

as(P

lotz

,18

83)

Vin

ius

tryh

ana

(Kay

e,19

14)

Cal

limor

mus

juve

ntus

Scu

dd

er,

1872

Cal

limor

mus

satu

rnus

(Her

rich

-Sch

affe

r,18

69)

Vir

gavi

rgin

ius

(Mos

chle

r,18

83)

Vir

gacl

ench

iL

.D

.M

iller

,19

70

C C C C C C

L-M

M L-M

L-M

M L

B B B B S/r

S

Page 20: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

524 vol. 48, no. 4The Southwestern Naturalist

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Mna

sicl

esge

taG

odm

an,

1901

Mna

sicl

eshi

ceta

onG

odm

an,

1901

Met

hion

opsi

sin

a(P

lotz

,18

82)

Mna

seas

bico

lor

(Mab

ille,

1889

)Ph

anes

alet

es(G

eyer

,[1

832]

)M

onca

tela

ta(H

erri

ch-S

chaf

fer,

1869

)M

onca

tyrt

aeus

(Plo

tz,

1883

)N

astr

ale

ucon

ele

ucon

e(G

odm

an,

1900

)

C C C C C M C C

L-M

L-M

L-M

L-M

L M L-M

M

B B B B/

rS/

rS/

rB

/r

N/

rC

ymae

nes

theo

geni

s(C

apro

nn

ier,

1874

)C

ymae

nes

treb

ius

(Mab

ille,

1891

)C

ymae

nes

frau

s(G

odm

an,

1900

)Ve

hiliu

sin

ca(S

cud

der

,18

72)

Vehi

lius

illud

ens

(Mab

ille,

1891

)

C C C C CK

6

L L-M

M L-M

L-M

S/r

B N/

rB B

Rem

ella

rem

us(F

abri

ciu

s,17

98)

Rem

ella

spp

.L

erem

aac

cius

(J.

E.

Smit

h,

1797

)L

erem

alir

isE

van

s,19

55L

erem

alo

chiu

s(P

lotz

,18

83)

Mor

ysva

leri

usva

lda

Eva

ns,

1955

Mor

ysge

isa

lyde

God

man

,19

00Ve

ttius

fant

asos

(Sto

ll,[1

780]

)

C C C C C C CK

15,1

8

L L-M

L-M

L-M

L L-M

M L-M

N/

rB B B S/

rB N B

Vetti

uson

aca

Eva

ns,

1955

Vetti

uste

rtia

nus

(Her

rich

-Sch

affe

r,18

69)

Tro

mba

xant

hura

(God

man

,19

01)

C C CK

6

M M L-M

N/

rN

/r

B/

rSy

nale

cyna

xa(H

ewit

son

,18

67)

Car

ystu

sph

orcu

s(C

ram

er,

[177

7])

Dam

ascl

avus

(Her

rich

-Sch

affe

r,18

69)

Car

ysto

ides

sp.

Peri

char

esph

ilete

sad

ela

(Hew

itso

n,

[186

7])

Ors

escy

nisc

a(S

wai

nso

n,

1821

)

C C C C C C

M L M L L-M

N/

rN

/r

N/

r

S/r

S/r

Page 21: BUTTERFLIES (PAPILIONOIDEA AND HESPERIOIDEA) OF ... - Bio …

December 2003 525Pozo et al.—Butterflies of Calakmul, Campeche, Mexico

AP

PE

ND

IX1—

Con

tin

ued

.

Tax

onN

ewre

cord

s1L

iter

atu

rere

cord

s2M

use

um

s3

Vege

ta-

tion

typ

e4A

rea5

Rhi

ntho

nos

ca(P

lotz

,18

83)

Con

gach

ydae

a(B

utl

er,

1877

)C

opae

odes

min

ima

(W.

H.

Ed

war

ds,

1870

)H

ylep

hila

phyl

eus

phyl

eus

(Dru

ry,

[177

3])

Ata

lope

des

cam

pest

ris

cam

pest

ris

(Boi

sdu

val,

1852

)Po

lites

vibe

xpr

aece

ps(S

cud

der

,18

72)

C C C C C C

M L-M

M L-M

L-M

L-M

N/

rS/

rN

/r

B B B/

rW

alle

ngre

nia

otho

otho

(J.

E.

Smit

h,

1797

)Po

mpe

ius

pom

peiu

s(L

atre

ille,

[182

4])

Am

blys

cirt

esto

lteca

tolte

caSc

ud

der

,18

72L

erod

eaeu

fala

(W.

H.

Ed

war

ds,

1869

)L

erod

eaar

abus

(W.

H.

Ed

war

ds,

1882

)Pa

noqu

ina

ocol

a(W

.H

.E

dw

ard

s18

63)

Pano

quin

ahe

cebo

lus

(Scu

dd

er,

1872

)Pa

noqu

ina

leuc

as(F

abri

ciu

s,17

93)

C C C C C C C C

L L-M

M L L L L-M

L

S/r

B N/

rS/

rS/

rS/

rB

/r

BPa

noqu

ina

paup

er(M

abill

e,18

78)

Pano

quin

aev

aden

s(S

toll,

[178

1])

Vace

rra

litan

a(H

ewit

son

,[1

866]

)A

ides

brill

a(H

.A

.Fr

eem

an,

1970

)N

eoxe

niad

eslu

da(H

ewit

son

,18

77)

C C C C C

M L-M

L M L-M

S/r

N S/r

N/

rB

1N

ewre

cord

s;C

K5

Cal

akm

ul;

C5

Cam

pec

he;

M5

Mex

ico.

2L

iter

atu

rere

cord

s:1

5Fe

lder

,18

69;

25

God

man

&Sa

lvin

,18

78–1

901;

35

Fiel

d,

1939

;4

5H

offm

ann

,19

40;

55

Joh

nso

nan

dC

omst

ock,

1941

;6

5H

offm

ann

,19

41;

75

Forb

es,

1943

;8

5B

ell

and

Com

stoc

k,19

48;

95

Com

stoc

kan

dB

row

n,

1950

;10

5Fr

eem

an,

1967

;11

5W

ellin

g,19

73;

125

Jen

kin

s,19

83;

135

Beu

tels

pac

her

and

How

e,19

84;

145

Jen

kin

s,19

84;

155

Ken

dal

lan

dM

cGu

ire,

1984

;16

5d

ela

Maz

aan

dT

urr

ent,

1985

;17

5d

ela

Maz

a,19

87;

185

Bal

caza

r,19

88;

195

Jen

kin

s,19

90;

205

Beu

tels

pac

her

,19

91;

215

Llo

ren

te-B

ousq

uet

set

al.,

1997

.3

CN

IN5

Col

lecc

ion

Nac

ion

ald

eIn

sect

os.

Inst

itu

tod

eB

iolo

gia,

Un

iers

idad

Nac

ion

alA

uto

nom

ad

eM

exic

o.4

Vege

tati

onty

pe:

M5

med

ium

,tr

opic

al,

sem

i-eve

rgre

enfo

rest

;L

5lo

w,

trop

ical

,se

mi-e

verg

reen

fore

st;

H5

hig

h,

trop

ical

,se

mi-e

verg

reen

fore

st,

G5

gen

eral

ist.

5A

rea

ofd

istr

ibu

tion

:N

5n

orth

;S

5so

uth

;B

5br

oad

;r

5ra

re;

c5

com

mon

.6

Not

fou

nd

inou

rfi

eld

wor

k.