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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
506 vol. 48, no. 4The Southwestern Naturalist
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
December 2003 507Pozo et al.—Butterflies of Calakmul, Campeche, Mexico
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
508 vol. 48, no. 4The Southwestern Naturalist
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).
December 2003 509Pozo et al.—Butterflies of Calakmul, Campeche, Mexico
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-
510 vol. 48, no. 4The Southwestern Naturalist
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.
512 vol. 48, no. 4The Southwestern Naturalist
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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
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
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
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
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
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
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
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
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
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
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
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
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
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.