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Full text of "Journal of the Lepidopterists' Society"

OL, 

Erf 



\ NOV 1 2004 

LiBRAK^ 



Volume 58 Number 3 
27 October 2004 
ISSN 0024-0966 



Journal of the 

Lepidopterists' Society 




Published quarterly by The Lepidopterists' Society 



THE LEPIDOPTERISTS' SOCIETY 

Executive Coi n< a 

James K. Adams, President Niklas Wahlberg, Vice President 

Susan J. Weller, Immediate Past President Ernest II. Williams, Secretary 

Gary G. Anweiler, Vice President 
Marc Epstein, Vice President 



Kelly M. Richers, Treasurer 



William E. Conner 
Rebecca Simmons 
Charles V. Covell Jr. 



Robert M. Pyle 
John A. Shuey 
Andrew D. Warren 



Members at large: 
Akito Kawahara 
Jane M. Ruffin 
Erik B. Runquist 

Editorial Board 

John W Brown (Chairman) 

Michael E. Toliver (Journal) 

Lawrence F. Gall (Memoirs) 

Phillip J. Schappert (Neios) 

John A. Snyder (Website) 

Carla M. Penz (at large) 

Honorary Life Members of the Society' 

Charles L. Remington (1966), E. G. Munroe (1973), Ian F. B. Common (1987), 
John G. Franclemont (1988), Lincoln P. B rower (1990), Douglas C. Ferguson (1990), 
Hon. Miriam Rothschild (1991), Claude Lemaire (1992), Frederick H. Rindge (1997) 



The object of The Lepidopterists' Society, which was formed in May 1947 and formally constituted in December 1950, is "to pro- 
mote the science of lepidopterology in all its branches, ... to issue a periodical and other publications on Lepidoptera, to facilitate 
the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea- 
sures" directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the 
News of The Lepidopterists' Society. Prospective members should send to the Assistant Treasurer full dues for the current year, to- 
gether with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is is- 
sued, with addresses and special interests. 

Active members — annual dues $45.00 within the U.S., $50.00 outside the U.S. 
Affiliated members — annual dues $10.00 within the U.S., $15.00 outside the U.S. 
Student members — annual dues $20.00 within die U.S., $25.00 outside the U.S. 
Sustaining members — annual dues $60.00 within the U.S., $65.00 outside the U.S. 
Life members — single sum $1,800.00 
Institutional subscriptions — annual $60.00 
Airmail postage for the News $15.00 

Send remittances, payable to The Lepidopterists' Society, to: Kelly M. Richers, Treasurer, 9417 Carvalho Court, Bakersfield, CA 
93311; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 
For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To 
order back issues of the Memoirs, write for availability and prices to Kenneth R. Bliss, 28 DuPont Avenue, Piscataway, NJ 08854. 



The additional cost for members outside the U.S. is to cover mailing costs. 



Journal of The Lepidopterists' Society (ISSN 0024-0966) is published quarterly by The Lepidopterists' Society, % Los Angeles 
County Museum of Natural Histoiy, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA 
and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists' Society, % Natural Histoiy Museum, 
900 Exposition Blvd., Los Angeles, CA 90007-4057. 



Cover illustration: Painted Ladv, Vanessa cardui L., from the 2003 influx to central Illinois, USA. Photo by Mike Toliver 



Journal of 

The Lepidopterists' Society 



Volume 58 



2004 



Number 3 



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



PAPILIONOIDEA OF THE EVERGREEN TROPICAL FORESTS OF MEXICO 



Jose L. Salinas-Gutierrez 1 , Armando Luis-Martinez 2 And Jorge Llorente-Rousquets 3 

Museo de Zoologfa, Facultad de Ciencias, UNAM, Apdo. Postal 70-399, Mexico, 04510 D. F„ MEXICO 

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

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

RESUMEN. Se efectuo un trabajo de mariposas diurnas en 1 1 unidades geograficas con bosque tropical perennifolio en Mexico, 
registrandose una lista total de 6S3 especies de Papilionoidea (sin inclnir Hesperiidae). Esta es la primera lista de mariposas en Mex- 
ico que involucra la cobertura de un tipo de vegetacion especifico. La riqueza de especies presentes en la zona hace patente la 
necesidad de crear estrategias de conservation en este ecosistema que esta disminuyendo su extension rapidamente. Los resultados 
se comparan con otros sitios de bosque tropical del Neotropico. 

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



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

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

1 sgjl@att.net.mx 

2 alm@hp.fciencias.unam.mx 

3 Institute de Ciencias Naturales, Universidad Nacional de Colom- 
bia, Santa Fe de Bogota, Columbia, email:jlb@hp.fciencias.unam.mx 



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

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



126 



Journal of the Lepidopterists' Society 



1 


CERRO COCONA 


cc 


2 


AGUA BLANCA 


AB 


3 


TENOSK3UE 


TEN 


4 


CHAJUL 


CHJ 


5 


YAXCHILAN 


VAX 


6 


SOCONUSCO 


SOC 


7 


LA GRINGA 


LG 


8 


CHALCHUAPA 


CHA 


S 


SERRA BE JUAREZ 


SJ 


10 


LOS TUXTLAS 


Lt 


11 


HUICHMUAYAN 


SLP 




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

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

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

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



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

Materials And Methods 

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

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

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

Results And Discussion 

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



Volume 58, Number 3 



127 



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

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

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

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



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

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

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



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







CC 


AB 


TEN 


CHJ 


VAX 


soc 


LG 


CHA 


SJ 


LT 


SLP 


ETF 


MEX 


Papilionidae 


6 


5 


7 


9 


8 


4 


8 


5 


11 


10 


9 


12 




Papilionmae 


15(18.29) 


11(13.41) 


14(1707) 


24(29.27) 


15(1829) 


9(10.98) 


16(19.51) 


12(14.63) 


29(35.37) 


30(36.59) 


20(24.39) 


37(45.12) 


S2 


Pieridae 


12 


8 


11 


19 


15 


14 


10 


12 


25 


26 


21 


27 




Dismorphiinae 


2(9.52) 


1(4.76) 





4(19.05) 


2(9.529 


2(9.52) 


2(9.52) 


1(4.76) 


4(19.05) 


7(33.33) 


4(19.05) 


10(47.62) 


21 


Coliadinae 


14(40) 


13(37.14) 


17(48.57) 


17(48.57) 


16(45.71) 


13(37.14) 


12(34.29) 


14(40) 


21(60) 


22(62.86) 


20(57.14) 


24(68.57) 


35 


Pierinae 


7(11.29) 


1(1.61) 


4(6.45) 


10(16.13) 


6(9.68) 


5(8.06) 


5(8.06) 


4(6.45) 


16(25.81) 


16(25.81) 


11(17.74) 


19(30.65) 


62 


Nynphalidae 


69 


59 


51 


94 


70 


62 


64 


68 


94 


98 


78 


115 




Hehconinae 


19(46.34) 


17(41.46) 


13(31.71) 


19(46.34) 


12(29.27) 


13(31.71) 


14(34.15) 


16(39.02) 


23(56.10) 


19(46.34) 


13(31.71) 


28(68.29) 


41 


Nymphalinae 


16(10.88) 


14(9.52) 


14(9.52) 


21(14.29) 


17(11.56) 


18(12.24) 


11(7.48) 


15(10.20) 


27(18.37) 


27(18.37) 


38(25.85) 


51(34.69) 


147 


Limenitidinae 


37(25.87) 


28(19.58) 


30(20.98) 


56(39.16) 


37(25.87) 


33(23.08) 


31(21.68) 


32(22.38) 


56(39.16) 


59(41.26) 


40(27.97) 


90(62.94) 


143 


Charaxinae 


14(21.88) 


4(6.25) 


3(4.69) 


22(34.38) 


11(17.19) 


7(10.94) 


17(26.56) 


10(15.63) 


26(40.63) 


23(35.94) 


14(21.88) 


38(59.38) 


64 


Apaturinae 


3(27.27) 


1(9.09) 


1(9.09) 


4(36.36) 


2(18.18) 


3(27.27) 


1(9.09) 





5(45.45) 


4(36.36) 


406.36) 


5(45.45) 


11 


Morphinae 


1(10) 


1(10) 


1(10) 


3(30) 


1(10) 


1(10) 


1(10) 


3(30) 


3(30) 


3(30) 


1(10) 


6(60) 


10 


Brassolinae 


4(22.22) 


3(16.67) 


6(33.33) 


11(61.11) 


6(33.33) 


5(27.78) 


7(38.89) 


6(33.33) 


9(50) 


11(61.11) 


7(38.89) 


12(66.67) 


IS 


Satyrinae 


8(7.41) 


6(5.56) 


5(4.63) 


20(18.52) 


15(13.89) 


9(8.33) 


11(10.19) 


8(7.41) 


21(19.44) 


25(23.15) 


16(14.81) 


41(37.96) 


108 


Danainae 


4(52.14) 


4(57.14) 


2(28.57) 


4(57.14) 


3(42.86) 


2(28.57) 


4(57.14) 


4(57.14) 


4(57.14) 


6(85.71) 


4(57.14) 


6(85.71) 


7 


Ithomiinae 


11(23.40) 


12(25.53) 


6(12.77) 


20(42.55) 


11(23.40) 


12(25.53) 


15(31.91) 


17(36.17) 


21(44.68) 


20(42.55) 


8(17.02) 


30(63.83) 


47 


Libythemae 


1(50) 


1(50) 


1(50) 


1(50) 


1(50) 





1(50) 


1(50) 


1(50) 


1(50) 


1(50) 


1(50) 


2 


Lycaenidae 


36 


9 


12 


71 


43 


13 


11 


23 


55 


97 


58 


116 




Riodininae 


18(7.96) 


6(2.65) 


8(3.54) 


64(28.32) 


21(9.29) 


8(3.54) 





1(0.44) 


54(23.89) 


76(33.63) 


42(18.58) 


125(55.31 


226 


Theclinae 


29(12.61) 


6(2.61) 


5(2.17) 


49(21.30) 


33(14.35) 


5(2.17) 


8(3.48) 


28(12.17) 


30(13.04) 


126(54.78 


50(21.74) 


150(65.22 


230 


Poryommatmae 


4(9.76) 








3(7.32) 


4(9.76) 





4(9.76) 


1(2.44) 


5(12.20) 


7(17.07) 


7(17.07) 


10(24.39) 


41 


TOTAL 


207 


129 


130 


352 


213 


145 


160 


173 


355 


482 


300 


683 


1295 



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



128 



Journal of the Lepidopterists' Society 



Table 1 also details the total number of species in the 
ETF of Mexico for Papilionidae (37), Pieridae (53), 
Nymphalidae (30S), and Lycaenidae (285), and the total 



0.30 



0.36 



0.42 



0.48 



■c 



054 

— cc 

—YAX 

LG 

CHA 

AB 

TEN 

CHJ 

SJ 

LT 

SLP 

S0C 



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

THE JACCARD SIMILARITY COEFFICIENT. 

number of genera per family for each geographical unit. 
The geographical units having the highest number oi 
exclusive species are LT (84), CHJ (40), and SJ (23). 
When the geographical units were gruped by state, the 
three in Tabasco (CC-AB-TEN) shared 78 species; in 
Oaxaca (LG-CHA-SJ), 105; and in Chiapas (CHJ-YAX- 
SOC) 84. In Chiapas, the units CHJ-SOC share 114 
species, while CHJ-YAX share 170; this makes sense 
since the latter two geographical units belong to the 
same biogeographical "island" of ETF, while SOC, on 
the Pacific slope, belongs to a different biogeographical 
area. 
TABLE 2. Synthetic matrix of species shared by the geographical units. 



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

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

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

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







CC 


AB 


TEN 


CHJ 


YAX 


SOC 


LG 


CHA 


SJ 


LT 


SLP 


CC 

























AB 


105 























TEN 


105 


85 





















CHJ 


164 


110 


114 



















YAX 


135 


89 


98 


170 

















SOC 


91 


66 


70 


114 


90 















LG 


118 


83 


82 


139 


110 


75 













CHA 


118 


84 


77 


137 


105 


75 


115 











SJ 


172 


116 


119 


243 


160 


121 


139 


142 









LT 


185 


114 


122 


273 


185 


119 


143 


157 


293 







SLP 


130 


81 


91 


168 


132 


93 


102 


99 


198 


243 






Volume 58, Number 3 



129 



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















LOCALITIES 










FAMILIES 


GUATEMALA 






BRASIL 






PERU 




COSTA RICA 


ECUADOR 




Tikal 


Jam Manaus 


i Campinas 


Rondonia Pakitza 


Tambopata 


Rio Napo 


AL 


PE 


Misahualli 


Papikonidae 


18 


23 


7 


17 


18 


25 


26 


26 


16 


17 


36 


Pieridae 


23 


26 


7 


29 


29 


31 


27 


23 


26 


26 


34 


Nymphalidae 


141 


343 


137 


208 


275 


371 


341 


238 


219 


174 


317 


Lycaemdae 


98 


89 


50 


54 


87 


181 


172 


68 


* 


* 


* 


Rjodiiudae 


48 


196 


111 


60 


203 


251 


242 


153 


97 


79 


* 


TOTAL 


328 


677 


312 


368 


612 


859 


808 


508 


358 


296 


387 



NOTE: Asterisks indicate missing data. 

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



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

Conclusions 

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

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

Acknowledgments 

The authors thank Dr. Juan Jose Morrone Lupi for reviewing 
the manuscript. Also, we thank the "Consejo Nacional para la 
Ciencia y la Tecnologfa" (CONACYT) project 32002, Direction 
General de Asuntos del Pesonal Academico (DGAPA-IN) pro- 
ject 209900 and the "Comisi6n para el Conocimiento y Uso de la 
Biodiversidad" (CONABIO) for financial aid for field work in 
the regions of Chimalapas, Oaxaca and Yaxchilan, Chiapas. 



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Received for publication 1 November 2001; 
September 2003 



wised and accepted 5 



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



Taxon 



Papilionidae 

Battus philcnor pliilenor 

B. polydamas polydamas 

B. laodamas copanae 

B. ingenuus 

B. Ii/cidas 

Parides photinus 

P. montezuma 

P. eurimedes mylotes 

P. sesostris zestos 

P. panares panares 

P. panares lydmenes 

P. eritlialion polyzelus 

P. iphidamas iphidamas 

Protographium epidaus epidaus 

P. philolaus philolaus 



2C 


AB 


TEN 


CHJ 


YAX 


SOC 


LG 


CHA 


sj 


LT 


SLP 




X 














X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 
X 
X 




X 


X 


X 


X 
X 
X 


X 
X 
X 




X 






X 




X 
X 






X 


X 
X 


X 
X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 




X 
X 


X 
X 


X 


X 
X 




X 


X 




X 










X 






X 


X 


X 


X 


X 


X 


X 


X 


X 


X 




X 




X 


X 


X 


X 




X 


X 


X 




X 


X 


X 


X 










X 


X 


X 




X 


X 


X 


X 








X 


X 


X 



Volume 58, Number 3 



131 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



P. agesilaus neosilaus 

P. dioxippus lacandones 

P. calliste calhste 

P. thyastes marchandi 

Eurytides salvini 

Protesilaus macrosilaus penthesilaus 

Mimoides thymbraeus thymbraeus 

M. ilus branchus 

M. phaon phaon 

Priamides pharnaces 

P. anchisiades idaeus 

P. erostratus erostratimis 

P. erostratus erostratus 

Troilides torquatus tolus 

Calaides ornythion ornythion 

C. astyalus pallas 

C. androgeus epidaurus 

Heraclides thoas autocles 

H. cresphontes 

Papilio polyxenes asterius 

Pi/rrhosticta victorinus victorinus 

P. abderus abderus 



X 




X 
X 


X 
X 

X 
X 
X 


X 
X 




X 

X 
X 


X 


X 

X 
X 

X 
X 


X 

X 
X 

X 


X 

X 


X 


X 


X 


X 






X 


X 


X 


X 




X 




X 


X 


X 




X 


X 


X 
X 


X 
X 


X 
X 


X 




X 


X 


X 
X 




X 


X 


X 
X 


X 
X 


X 

X 

X 








X 


X 




X 




X 


X 


X 






X 


X 


X 


X 


X 




X 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 


X 






X 




X 


X 


X 


X 
X 
X 


X 
X 
X 


X 
X 
X 
X 



Fieri dae 

Enantia Una motion 

E. albania albania 

E.jethys 

E. mazai mazai 

Lieinix nemesis atthis 

LHsmorphia amphiona praxinoe 

Dismorphia crista virgo 

D. eunoe eunoe 

D. eunoe popoluea 

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

P. neocypris virgo 

P. philea philea 

P. sennae marcellina 

PJiabdodryas trite ssp. 

Aphrissa statirajada 

Abaeis nicippe 

Pyrisitia dina westwoodi 

P. lisa centralis 

P. nise nelphe 

P. proterpia proterpia 

Eurema agave millerorum 

E. albula celata 
E. boisduvaliana 
E. daira 

E. mexicana mexicana 

E. salome janmpa 

E. xantochlora xantochlora 









X 










X 


X 
X 

X 


X 
X 

X 


X 


X 




X 


X 


X 


X 




X 


X 
X 

X 


X 


X 






X 






X 


X 


X 


X 






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X 










X 


X 


X 


X 




X 


X 


X 


X 






X 


X 


X 


X 


X 


X 


X 


X 


X 






X 


X 


X 


X 


X 


X 


X 


X 


X 






X 


X 


X 


X 


X 




X 


X 


X 


X 


X 
X 


X 
X 


X 
X 


X 
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X 
X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 






X 


X 


X 


X 




X 


X 


X 




X 


X 


X 


X 


X 


X 
X 


X 


X 


X 
X 


X 
X 


X 
X 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 




X 




X 


X 








X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 






















X 
X 


X 


X 
X 


X 

X 


X 
X 


X 


X 


X 


X 
X 


X 
X 


X 
X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 




X 


X 


X 




X 


X 
X 




X 
X 
X 


X 

X 


X 
X 



132 



Journal of the Lepidopterists" Society 



Appendix 1. Continued 



T;i\on 



cc 



AB TEN C11J VAX SOC LG CHA SJ LT SLP 



Nathalis iole 

Kricogonia lyside 

Hesperocharis costaricensis pasion 

Archonias brassolis aproximata 

Charonias theano nigrescens 

Catasticta flisa flisa 

Catasticta nimbice nimbice 

C. ochracea ochracea 
Pereute champs champs 
Melete lycimnia isandra 
M. polyhymniaflorinda 
Clutophrissa drusilla tenuis 
Pontic protodlce 
Leptophobia aripa elodia 
Itaballia demophile centralis 
I. pandosia kicaha 
Pieribtdlia viardi viardi 
Perrhybris pamela chajulensis 
Ascia monuste monuste 
Gaiu/ra josephina josepha 

G. phaloe tiburtia 

Nymphalidae 

Altinote ozvmene nox 

Actinote guatemalena veraecrusis 

A. guatemalena guatemalena 

A. thalia anteas 

Philaethria diatonica 

Dime juno huascuma 

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

E. Isabella eva 
E. lineata 

E. procula asidia 

E. vibilia vialis 

Laparus doris viridis 

Heliconius charitonia vazquezae 

Hcliconius cydno galanthus 

H. erato petiveranus 

H. hecale zuleika 

H. hecale fornarina 

H. hecalesia octavia 

H. hortense 

H. ismenius telchinia 

H. sapho leuce 

H. sara veraepacis 

Euptoieta claudia daunius 

E. hegesia hoffmanni 

Vanessa atalanta rubria 

Cynthia cardui 

C. virginiensis 

C. anabella 

Nymphalis antiopa antiopa 









X 


X 


X 






X 


X 


X 








X 




X 






X 


X 
X 










X 






X 




X 


X 










X 


X 








X 
X 

X 


X 
X 
X 
X 
X 


X 

X 

X 


X 




X 


X 
X 




X 






X 
X 


X 


X 


X 


X 


X 


X 
X 


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X 


X 


X 


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X 
X 


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X 
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X 
X 
X 


X 






X 




X 


X 


X 


X 
X 


X 
X 


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X 


X 
X 




X 


X 


X 
X 


X 


X 


X 




X 


X 


X 


X 


X 


X 


X 


X 


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X 




X 








X 


X 


X 


X 






















X 










X 




X 


X 


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X 


X 
X 


X 


X 


X 


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X 






X 










X 


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X 


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X 


X 


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X 
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X 
X 


X 
X 


X 
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X 


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X 
X 
X 


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X 
X 
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Volume 58, Number 3 



133 



Appendix 1. Continued 



Taxon 



CC 



AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Polygonia interroeationis 

Hypanartia dione 

H. godmanii 

H. lethe 

H. kefersteini 

Anartia amathea venusta 

A.jatrophae luteipicta 

Siproeta epaphus cpaphus 

S. stelenes biplagiata 

S. superba superba 

Junonia coenia 

]. evarete 

Chlosyne erodyle erodyle 

C. gaudialis gaudialis 

C. hippodrome hippodrome 

C. janais 

C. lacinia lacinia 

C. marina marina 

C. rosita browni 

Thessalia theona theona 

T. theona thekla 

Texola elada elada 

T. elada ulrica 

Microtia elva homi 

Phyciodes mylitta mexicana 

P. vesta graphica 

P. phaon 

Phyciodes tharos tharos 

Anthanassa ardys ardys 

A. argentea 

A. atronia sydra 

A. atronia atronia 

A. drusilla lelex 

A. frisia tulcis 

A. ptolyca ptolyca 

A. texana texana 

A. annulata 

Tegosa anieta cluvia 

T. anieta luka 

T. guatemalena 

T. similis 

Eresia clara clara 

E. phillyra phillyra 

Catfilia eranites mejicana 

C. myia myia 

C. ofella ofella 

Hkioris odius dious 

Coea acheronta acheronta 

Baeotus beotus beotus 

Smyrna blomfildia datbs 

S. karwinskii 

Colobura dirce dirce 

Tigridia acesta ssp. 

Biblis hyperia aganisa 

Mestra dorcas amymone 

Myscelia cyananthe cyananthe 













X 






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134 



Journal of the Lepidopterists" Society 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Myscelia cyaniris cyaniris 
M. ethusa ethusa 
Catonephele mexicana 

C. numilia esite 
Catonephele cortesi 
Nessaea aglaura aglaura 
Eunica alcmena 

E. alpais excelsa 

Eunica sijdonia caresa 

E. malvina albida 

E. monima 

E. mygdonia omoa 

E. olympias augusta 

E. venusia 

E. tatila tatila 

Hamadnjas amphinome mexicana 

H. febma ferentina 

H. feronia farinulenta 

H. fomax fornacalia 

H. glauconome glauconome 

H. guatemalena marmarice 

H. guatemalena guatemalena 

H. iplitliime joannae 

H. laodamia saurites 

Ectima erycinoides ssp. 

Pyrrhogyra edocla edocla 

P. neaerea hypsenor 

P. otolais otolais 

Temenis laothoe hondurensis 

Epiphile adrasta adrasta 

E. hermosa 

E. area plutonia 

Bolboneura sylphis veracruzana 

B. sylphis lacandona 
NicaflamUa hachiana 
Dijnamine artemisia glauce 

D. ate 

D. dyonis 

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

C. lyca lyca 

C. texa grijalva 

C. texa titania 

C. tolitna tehuana 

C. tolima pacifica 

C. pitheas 

Adclpha basiloides basiloides 

A. baeotia milleri 

A. baeotia oberthurii 

A. celerio diademata 



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Volume 58, Number 3 



135 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Adelpha bredowii eulalia 

A. cytherea marcia 

A. escalantei 

A. felderi jarias 

A. iphiclus iphicleola 

A. diazi 

A. ixia leucas 

A. leuceria leuceria 

A. leucerioides leucerioides 

A. lycorias melanthe 

A. naxia epiphicla 

A. paroeca emathia 

A. phylaca phylaca 

Adelpha pithys 

A. salmoneus salmonides 

A. serpa massilia 

A. zahnona sophax 

A. fessonia fessonia 

Basilarchia archippus hoffmanni 

Marpesia chiron marius 

M. corita corita 

M. harmonia 

M. petreus tethys 

M. zerynthia dentigera 

Archaeoprepona amphimachus amphiktion 

A. denwphon centralis 

A. demophoon gulina 

A. meander phoebus 

A. phaedra aelia 

Prepona deiphile brooksiana 

P. deiphile escalantiana 

P. dexamenes medinai 

P. laertes octavia 

P. pijlene philetas 

Agrias aedon rodriguezi 

A. amydon oaxacata 

Zarette callidryas 

Z. itus ellops 

Siderone galanthis ssp. - 

S. syntiche syntiche 

Anaea troglodyta aidea 

Consul electra electra 

C. fabius cecrops 

Fountainea eurypyle confusa 

F. glycerium glyceriutn 

F. halice martinezi 

F. ryphea ryphea 

Memphis artacaena 

M. aureola 

M. dia ssp. 

M. forreri 

M. hedemanni 

M. herbacea 

M. mora orthesia 

M. phila boisduvali 

M. neidhoeferi 









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136 



Journal of the Lepidopterists' Society' 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Memphis oenomais 

M. philumena xenica 

M. pithyusa 

M. proserpina 

M. xenocles Carolina 

M. perenna perenna 

Asterocampa idyja argus 

Doxocopa cyane mexicana 

D. laure laure 

D. laurentia cherubina 

D. pavon theodora 

Morpho achilles montezuma 

M. achilles octavia 

Pessonia luna luna 

Ipliimedeia telemachus justitiae 

1. telemachus ssp. 

I. telemachus oaxacensis 

Vynastor darius stygianus 

D. macrosiris strix 

Opsiphanes boisduvalii 

O. cassias 

O. tamarindi 

O. invirae fabricii 

O. quiteria quirinus 

Caligo atreus uranus 

C. oileus scamander 

C. prometheus memnon 

Eryphanis aesacus aesacus 

Narope cyllastros testacea 

Pierella luna rubecula 

Manataria maculata 

Cepheuptychia glaucina 

Chloreuptychia sericeella 

Cissia confusa 

C. lobe 

C. terrestris 

Cyllopsis hedemanni hedemanni 

C. hedemanni tamaulipensis 

C. hilaria 

C. suiualens escalantei 

C. pijracmon 

C. gemma freemani 

C. dospassosi 

Euptychia mollina 

Hermeuptt/chia hermes 

H. alcinoe 

Oxeoschistus hilara hilara 

Magneuptychia libye 

Megeuptychia antonoe 

Pareuptijchia binocula metaleuca 

P. interjecta 

P. ocirrhoe 

Pseudodebis zimri 

Satyrotaygetis satyrina 

Splendeuptychia kendalli 

Paramacera xicaque xicaque 









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Volume 58, Number 3 



137 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Taijgetis kerea kerea 

T. leuctra 

T. mermeria excavata 

T. uncinate 

T. virgilia 

T. weymeri 

T. thamyra 

Vareuptychia usitata pieria 

V. themis 

V. similis 

V. undina 

Yphthimoides renata disaffecta 

Dioriste tauropolis tauropolis 

Pedaliodes dejecta circumducta 

Danaus eresimus montezuma 

D. gilippus thcrsippus 

D. plexippus plexippns 

Lycorea halia atergatis 

L. ilione albescens 

Anetia thirza thirza 

Tithorea harmonia luppothous 

T. tarricina duenna 

Aeria eurimedea pacifica 

Olyras crathis theon 

Melinaea lilis flnmcans 

M. lilis imitata 

Mechanitis lysimnia utemaia 

M. menapis doryssus 

M. pohjmnia lycidice 

Hyposcada virginiana virginiana 

Oleria paula 

Napeogenes tolosa tolosa 

Hypothyris euclea vaiora 

H. lycaste dionaea 

Ithomia leila 

I. patilla patilla 

Calhthomia hezia hedila 

C. hezia wettingi 
Dircenna dero ssp. 
D.jemina ssp. 

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

P. simplex fenochioi 

Godyris zavaleta sosunga 

Hypomenitis annette annette 

Greta morgane oto 

G. nero nero 

Hypoleria lavinia cassotis 

Libytheana carinenta mexicana 

Lycaenidae 

Euselasia cataleuca 

E. chrysippe 

E. regipennis regipennvs 
E. sergia sergia 







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138 



Journal of the Lepidopterists' Society 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Euselasia procula 

E. hieronymi hieronymi 

E. inconsplcua 

E. pusilla 

E. eubule eubule 

E. aurantiaca aurantiaca 

Hades noctula 

Perophthalma tullius lasus 

Leucochimona vestalis vestalis 

L. lepida nivalis 

Mesosemia tclegone telegone 

M. gaudiolum 

M. gemina 

Eurybia patrona persona 

E. lycisca 

E. halimede elvina 

Hermathena oweni 

Diophtahna lagpra iphias 

Napaea eucharila picina 

N. theages theages 

N. umbra umbra 

Cremna actoris 

C. thasus subrutila 

Lyropteryx hjra cleadas 

Ancijluris jurgensenii montezuma 

A. inca mora 
Rhetus arcius thia 

R. periander naevianus 

Isapis agi/rtus hera 

Brachyglenis dodone 

Notheme erota diadema 

Lepricornis melanchroia 

Calephelis nemesis nemesis 

C. mexicana 

C. fulmen 

C. stallingsi 

C. huasteca 

C. acapulcoensis 

C. yucatana 

C. perditalis perditalis 

C. montezuma 

C. laverna laverna 

Charts gynaea zaina 

C. velutina 

Chalodeta chaonitis 

Caria ino melicerta 

C. domitianus vejento 

C. rhacotis rhacotis 

C. lampeto 

Baeotis zonata simbla 

B. sulphurea macularia 
B. sulphurea sulphurea 
Lasaia meris 

L. agesilas callaina 

L. sessilis 

L. maria anna 



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Volume 58, Number 3 



139 



Appendix 1. Continued 



Tiixon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



L. narses' 

L. sula peninsularis 

Melanis pixe pixe 

M. cephise cephise 

M. cephise huasteca 

ilesene croceella 

M. margaretta margaretta 

Xenandra caendeata 

Chimastrum argenteum argenteum 

Symmachia rubina rubina 

S. accusatrix 

Symmachia probetor championi 

S. tricolor hedemanni 

Pterographium Sagarin tyriotes 

Sarota gamelia 

S. acanthoides myrtea 

S. chrysus dematria 

Anteros formosus micon 

A. carausius carausius 

Calydna lusca venusta 

C. stumula hegias 

C. sinuata 

Emesis aurimna 

E. saturata 

E. liodes 

E. mandana furor 

E. vulpina 

E.fatimella nobilata 

E. tenedia tenedia 

E. lupina 

E. ocypore aethalia 

E. zela zela 

E. emesia emesia 

E. cypria paphia 

E. tegula 

E. zela cleis 

Argyrogrammana holosticta 

Pseudonymphidia clearista 

Pachythone gigas 

Apodemia multiplaga 

A. hypoglauca hypoglauca 

A. walkeri 

Thisbe irenea belides 

T. lycorias lycorias 

Lemonias caliginea 

L. agave 

Juditha molpe 

Synargis calyce mycone 

S. ethelinda nymphidioides 

S. nycteus 

Menander menander purpurata 

Pandemos godmanii 

Calospila pelarge 

C. sudias 

Theope pedias tiia 

T. virgilius virgilius 



X X X X X X 



X 
X 
X 
X 
X 



X 
X 



X 
X 



X 
X 
X 
X 



X 
X 



X X 

X 

X 
X X 

X X X X 
X 

X X X X X 

X XXX 



X 

X 
X 
X 







X 


X 


X 

X 


X 
X 


X 


X 


X 




X 


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X 




X 


X 


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X 


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X 


X 
X 
X 






X 


X 


X 


X 


X 


X 


X 
X 


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X 


X 




X 






X 




X 


X 


X 


X 


X 


X 


X 




X 


X 


X 
X 


X 


X 








X 


X 


X 


X 


X 


X 


X 


X 
X 




X 






X 


X 


X 


X 




X 


X 
X 


X 


X 


X 


X 


X 


X 




X 


X 
X 
X 




X 
X 


X 
X 


X 


X 


X 




X 




X 



140 



Journal of the Lepidopterists' Society 



Appendix 1. Continued 



Taxon 



CC 



AB TEN CHJ YAX SOC LG CHA S] LT SLP 



Theope eupolis 

T. cratylus 

T. publius 

T. eleutho 

T. mania 

T. diorcs 

Calociasma lilina 

Nymphidium ascolia ascolides 

Brephidium exilis exilis 

Leptotes marina 

L. cassius striata 

Zizula cyna cyna 

Hemiargus ceraunus 

H. isola isola 

H. huntingtoni hannoides 

Everes comyntas 

E. amyntula amijntula 

Celastrina argiolus gozora 

Eumaeus childrenae 

E. toxea 

Paiwarria antigonus 

Theorema eumenia 

"Thecla" (busa group) busa 

Evenus regalis 

E. coronata 

E. batesii 

"Thecla" (gibberosa group) barajo 

"Thecla" (eunus group) eunus 

Allosmaitia strophius 

Pseudolycaena damo 

Areas imperialis 

A. cypria 

Theritas mavors 

"Thecla" (hemon group) augustinula 

"Thecla " (hemon group) theocritus 

"Thecla" (hemon group) hemon 

"Thecla" (hemon group) lisus 

Atlides gaumeri 

A. pohjbe 

A. inachus 

A. earpasia 

A. halesus 

A. caranus° 

Radissima umbratus 

"Thecla" (ligurina group) ligurina 

"Thecla" (ligurina group) lude 

Denivia theocritus 

Contrafacia ahola 

C. imma 

Thereus cithonius 

T. oppia 

T. ortalus 

Arawacus togarna 

A. sito 

A. jada 

Rekoa meton 



X 
X 
X 



X 
X 



X 
X 
X 



X 
X 
X 



X 


X 


X 


X 




X 


X 




X 




X 


X 


X 





X X X X 
X 

X X 
X X 

X 
X 



X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 










X 






X 


X 





X 






X 






X 


X 




X 


X 




X 


X 




X 


X 


X 


X 


X 


X 




X 


X X 


X 


X 


X 


X 




X 


X 
X 


X 




X 


X 


X 


X 




X 


X 


X 


X 


X 




X 


X 






X 


X 




X 




X X 


X 






X 






X 


X 


X 


X 


X 


X 






X 


X 


X 


X 


X 


X 


X 


X 

X 
X 






X 


X 




X 






X 


X 
X 
X 




X 






X 






X 


X 




X 






X 


X 




X 






X 






X 




X X 


X 




X X 


X 


X 
X 


X 


X 


X 



Volume 58, Number 3 



141 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Rekoa palegon 

R. zebina 

R. marius 

R. stagira 

Ocaria petelina 

O. thales 

O. ocrisia 

Magnastigma elsa 

Chlorostrymon simaethis 

C. telea 

Cyanophrys goodsoni 

C. amyntor 

C. fusius 

C. herodotus 

C. miserabilis 

C. longula 

Panthiades bitias 

P. ochus 

P. bathildis 

P. phaleros 

Oenomaus ortygnus 

0. atesa 

Parrhasius polibetes 
P. orgia 

P. moctezuma 
Mich aelus jebus 
M. thordesa 
M. hecate 
M. uibidia 
M. zenaida 
lgnata gadira 

1. nr. gadira 
I. norax 

Strymon melinus 
S. albata 

S. alea 

S. bazochii 

S. mulucha 

S. yojoa 

S. cestui 

S. astiocha 

S. istapa 

S. ziba 

S. nr. megarus #1 

S. serapio 

S. basalides' 

Lamprospilus collucia 

"Thecla" (aruma group) gattiena 

Kisutam syllis 

K. hesperitis 

K ceromia 

K. denarius 

K. guzanta 

K. sethon 

"Thecla" (carrussa group) cespasianus 

Electrostrymon mathewi 



X 
X 



X X 



X X 



X 
X 



X 
X 



X 


X 


X 




X 


X 


X 


X 


X 


X 


X 




X 




X 




X 




X 





X 

X 



X 

X 

X 
X 



X 
X X 



X 


X 


X 
X 
X 
X 
X 

X 
X 


X 

X 






X 


X 






X 


X 






X 


X 






X 


X 


X 




X 
X 


X 
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X 


X 


X 


X 


X 


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X 


X 
X 


X 






X 


X 




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X 


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X 






X 


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X 






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X 








X 
X 


X 
X 




X 










X 


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X 




X 








X 


X 


X 




X 


X 






X 




X 




X 
X 
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X 


X 
X 


X 
X 




X 
X 






X 


X 






X 


X 


X 




X 










X 


X 



142 



Journal of the Lepidopterists' Society 



Appendix 1. Continued 



Taxon 



CC AB TEN CHJ YAX SOC LG CHA SJ LT SLP 



Electrostryinon sangala 

E. canus 

Calycopis calus 

C. demonassa 

C. atnius 

C. clarina 

C. isobeon 

C. susanna 

C. drusilla 

C. trebula 

Tinolus echion 

T. crollnus 

T. cijdrara 

Criinsinota phobe 

"Tmolus" (heraldica group) heraldica 

"Thecla" (empusa group) halciones 

"Thecla" (keila group) keila 

Siderus philinna 

S. gargophia 

S. caninius 

S. thoria 

Auberglna paetus 

"Thecla" (mycon group) mycon 

"Thecla" (tephraeus group) tepliraeus 

"Thecla" (tephraeus group) suedra 

"Thecla" (tephraeus group) ambrax 

Ministrymon clytie 

M. arola 

M. una 

M. inoa 

M. phrutus 

M. azia 

Jantheclajanthodonia 

J.janthina 

lpidecla schausi 

Brangas neora 

B. getus 

B. coccineifrons 

B. carthaea 

"Thecla" (cupentus group) cupentus 
Chalybs janias 

C. hassan 

"Thecla" (theia group) theia 

Hypostrymon critola 

Iaspis nr castitai 

I. teraesa 

Nesiostrymon calch in ia 

N. celona 

N. dodava 

Erora carlo 

E. opisena 

E. muridosca 

"Thecla" (ares group) semones 

Caerofethra carnica 

Celmia celmus 

"Thecla" (color group) conoveria 



X 
X 
X 
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Journal of the Lepidopterists' Society 
58(3), 2004, 143- 16S 

HISTOIRE GENERALE ET ICONOGRAPHIE DES LEP1DOPTERES ET DES CHENILLES DE 

L'AMERIQUE SEPTENTRIONALE BY BOISDUVAL & LE CONTE (1829-[1837]): ORIGINAL DRAWINGS 

USED FOR THE ENGRAVED PLATES AND THE TRUE IDENTITIES OF FOUR FIGURED TAXA 

John V. Calhoun 1 

977 Wicks Drive, Palm Harbor, Florida 34684-4656, USA 

ABSTRACT. A set of 149 Lepidoptera drawings is currently deposited in the Thomas Cooper Library, University of South Car- 
olina. Forty of the 55 butterfly drawings include figures that were copied for the engraved plates in Histoire Generate et Iconogra- 
phie des h&pidopteres et des Chenilles de I'Amerique Septentrionale (Boisduval & Le Conte 1829-[1837]). Identifications are pro- 
vided herein for the 80 butterfly species in these drawings, as well as manuscript notes that likely accompanied 21 other drawings 
reproduced in Histoire Gene'rale. Pieris cleomes Boisduval & Le Conte is shown to be synonymous with Ascia monuste phileta 
(Fabricius). The figures identified as Thecla favonius (J. E. Smith) represent both Satyrium favonius and Strymon melinus (Hiib- 
ner). The figures identified as Libythea motya (Hiibner) represent Libytheana motya and Libytheana carinenta bachmanii (Kirt- 
land). Also included are remarks about the identity of Melitaea ismeria Boisduval & Le Conte and the validity of the Boisduval type 
specimens of North American Lepidoptera. 



Additional key words: John Abbot, Emile Blanchard, Paul Dumenil, Georgia, South Carolina, type locality. 



I recently traced the history of a set of 149 
Lepidoptera drawings now deposited in the Thomas 
Cooper Library, University of South Carolina 
(Calhoun 2003). These were rendered in graphite and 
watercolor by at least four artists: English naturalist 
John Abbot (1751-ca.l840), who resided in Virginia 
and Georgia from 1773 until his death; French 
zoologist Emile (or Charles Emile) Blanchard (1819- 
1900); French engraver and publisher Paul C. R. C. 
Dumenil (1779-?); and probably American naturalist 
John E. Le Conte, Jr. (1784-1860). My analysis of 
these drawings confirmed the claim of art historian 
Vivian Rogers-Price (1983) that some were copied for 
plates in Histoire Generate et Iconographie des 
Lepidopteres et des Chenilles de I'Amerique 
Septentrionale [General History and Iconography of 
the Lepidoptera and the Caterpillars of Northern 
America] by Jean B. A. D. de Boisduval and J. E. Le 
Conte, Jr., published in 26 livraisons from 1829 to 
1837 (usually cited as [1833]). After the publication of 
Histoire Generale, Boisduval retained these drawings 
for many years and they eventually passed into the 
hands of lepidopterist Charles M. Oberthur. Oberthiir 
died in 1924 and the drawings have not been 
examined by another lepidopterist since that time. 

These 149 drawings are of great relevance to 
American lepidopterists. The Thomas Cooper Library 
has digitized all 149 drawings and made them available 
for viewing on the Internet (USC 2003). As part of 
their study, I was afforded the opportunity to offer 
identifications and other pertinent information. Due 

1 Research Associate, Florida State Collection of Arthropods, DPI, 
FDACS, Gainesville, Florida 32614, USA 



to the ephemeral nature of Internet web sites, I have 
decided to also present this data in print and 
incorporate many additional details. I also pro\ide 
information on three sets of surviving manuscript 
notes by John Abbot that contain entries relevant to 
published plates in Boisduval & Le Conte (1829- 
[1837]). Evidence from this research clarifies the 
origin of specimens figured in Boisduval & Le Conte 
(1829-[1837]) and resolves the status of Pieris cleomes 
Boisduval & Le Conte, as well as the butterflies 
figured as Thecla favonius (J. E. Smith) and Libythea 
motya (Hiibner). Further evidence regarding die 
identity of Melitaea ismeria Boisduval & Le Conte 
augments Calhoun (2003). 

Materials and Methods 
The butterfly drawings at the University of South 
Carolina were examined (in person and through digital 
scans) and compared with the published plates in 
Boisduval & Le Conte (1829-[1837]). Also consulted 
were the plates in Smith & Abbot (1797), as well as 
original drawings and manuscript notes bv John Abbot 
deposited in the Alexander Turnbull Library 
(Wellington, New Zealand), the Houghton Library 
(Harvard University), and The Natural History 
Museum, London. Inscriptions on the drawings were 
compared with the known handwriting of Abbot, 
Boisduval, and Le Conte. The National Museum of 
Natural Histoiy (Washington, D. C.) and The Natural 
History Museum, London, were searched for relevant 
specimens from Boisduvals collection. 



RESULTS 
Original butterfly drawings. 



Fifty-five of die 



144 Journal of the Lepidopterists' Society 

Table 1. Butterfly species depicted in original drawings at the Thomas Cooper Library, University of South Carolina. B&L = Boisduval & 
Le Conte (1S29-[1837]). Figures: D = dorsal, V = ventral, m = male, f = female, L = larva, P = pupa. 

Figures Artist 

B&L copied credited on 

No. Artist Species depicted Figures plate for B&L B&L plate B&L name 

1 Abbot Papilio cresphontes Cramer Df,Vf,L,P 12,13 Df,Vf,L,P Abbot Papilio Thoas 
NOTES: drawing has a penciled reference to B&L Plates 12 and 13. 

2 Abbot Eurytides marcellus (Cramer) Dm,Vm,L,P 2 Dm,Vm,L,P Abbot Papilio Marcellus 
NOTES: depicts the summer form "lecontei" (Rothschild & Jordan), named in honor of B&L coauthor J. E Le Conte. Includes Abbot's 
inscription, "Autumnal Ajax." 

3 Abbot Eurytides marcellus (Cramer) Dm,Df,Vm,L,P 1 Dm,Vm,L,P Abbot Papilio Ajax 
NOTES: depicts the spring form "marcellus" and has a penciled reference to B&L Plate 1. Includes Abbot's inscription, "Spring Ajax." 
Drawing was figured by Rogers-Price (1983). 

4 Dumenil? Eurytides celadon (Lucas) Df, Vf 3 Df.Vf Abbot Papilio Sinon 
NOTES: the style seems consistent with drawing 37 that Boisduval attributed to Dumenil. 

5 Abbot Papilio polyxenes (Fabricius) Df 
NOTES: includes Abbot's inscription "Ni Female." 

6 Abbot Papilio polyxenes (Fabricius) Dm,Vm,L,P 4 Dm,Vm,L,P Abbot Papilio Asterias 
NOTES: includes a penciled reference to B&L Plate 4. Includes Abbot's inscription "Ni Male Troilus." 

7 Abbot Ascia monuste (Linnaeus) Dm,Df,Vm,L,P 16 Dm,Df,Vf,L,P Abbot Pieris Cleonies 
NOTES: includes a penciled reference to B&L Plate 16, which was copied from this drawing and another trom Abbot with a hostplant (see 
text). Figures in this drawing were probably used for the original description of P. cleomes. See Figs 10, 11. 

8 Abbot Phoebis sennae (Linnaeus) Df,Dm,Vf,L,P 24 Df,Dm,Vf,L,P Abbot Callidnjas Eubule 
NOTES: includes a penciled reference to B&L Plate 24 and Abbot's inscription, "Eubule." The ventral female on B&L Plate 24 was 
misidentified as a male. Scudder (1888-1889, PI. 76. fig. 2) copied the larva. Holland (1898, PL 2, fig. 2) and Klots (19.51, Pi. 5, fig. 25) 
reproduced Scudder's larva. 

9 Abbot Zerene cesonia (Stoll) Dm.Df,Vf,L,P 22 Dm,Df,Vf,L,P Abbot Colias Coesonia 
NOTES: includes a penciled reference to Plate 22, which was probably copied from this drawing and another from Abbot with duplicate 
figures and a hostplant. Although the ventral figure is identified on the B&L plate as a male, die corresponding figure in this drawing appears 
to be a female. 

10 Abbot a. Parrhasius m-album (B&L) Dm,Df,Vm,L,P 27 Dm,Df,Vm,L,P Abbot Thecla Psyche 

b. Strymon melinus (Hiibner) Dm,Vm,L,P 

NOTES: the figures of P. m-album were probably consulted for the original description of T. psyche. Includes a penciled reference to B&L 
Plate 27 and figure numbers used on the plate. The hostplant "smilax" is written below the figures of S. melinus. Although Oberthiir (1920) 
believed these figures of S. melinus were copied for B&L plate 28 of T. hyperici, they are not analogous. 

11 Abbot a. Callophrys niphon (Hiibner) Df.Vf.L.P 33 Df,Vf,L,P Abbot Thecla Niphon 

b. Satyrium titus (Fabricius) Dm,Df,Vf,L,P 34 Dm,Df,Vf,L,P Abbot Thecla Mopsus 

c. Callophrys gryneus (Hiibner) Dm,Vf,L,P 33 Dm,Vf,L,P Abbot Thecla Smilacis 
NOTES: the figures of C. gryneus were probably consulted for die original description of T. smilacis. Adult figures of S. titus from this 
drawing and the ventral male of Dumenil's drawing 13 were used to illustrate the species on B&L Plate 34. Le Conte wrote "Pine" below C. 
niphon. "Oak, Eupatorium coelestinum" below S. titus, and "Smilax" below C. gryneus. These plants were reported as hosts for these 
butterflies in B&L. Scudder (1888-1889) copied die pupa of C. niphon (PI. 84, fig. 40), the larva and pupa of S. titus (PI. 75, fig. 35; PI. 84, 
fig. 37), and the larva and pupa of C. gryneus (PI. 75, fig. 31; PI. 84, fig. 30). Holland (1898, PI. 5, figs. 30, 37 & 40) and Klots (1951, PI. 5, fig. 
14) reproduced some of Scudder's figures. See Fig. 4. 

12 Abbot a. Satyrium liparops (Le Conte) Df,Vf,L,P 31 Df,Vf,L,P Abbot Thecla Liparops 

b. Callophrys henrici (Grote & Robinson) Dm,Df,Vf,L,P 31 Df,Vf,L,P Abbot Thecla I rus 

NOTES: the figures of S. liparops were used for die original description of Thecla liparops. Le Conte wrote "Oak, Chestnut" below the 
figures of S. liparops (mentioned bv Scudder (1888-1889) and in his notes at Harvard). "Vaccinium" is written below the figures of C. henrici, 
which corresponds to the mention o("vaccinium" as a hostplant of this species in B&L. C. henrici was not described until 1867 and was 
often confused with C. irus in earlier literature. Scudder ( 1888-1889, PI. 75, fig. 28) copied the larva of C. henrici (as "Incisalia irus"). See 
Figs. 3, 6. 

13 Dumenil? a. Satyrium titus (Fabricius) Dm.Vm 34 Vm Abbot Thecla Mopsus 

b. Strymon melinus (Hiibner) Dm,Vm — 

c. Paectes pygmaea (Hiibner) Dm 

NOTES: the ventral male of S. titus from this drawing and the adults of Abbot's drawing 1 1 were used to illustrate the species on B&L Plate 34. 



Volume 58, Number 3 



145 



Table 1. Continued 



No. 



Artist 



Species depicted 



Figures 





Figures 


Artist 




B&L 


copied 


credited on 




plate 


for B&L 


B&L plate 


B&L name 



14 Blanehard a. Calycopis cecrops (Fabricius) Dm,Df,Vm,Vf 35 Dm,Df,Vm,Vf Abbot Thecla Poeas 

b. Hemiargus ceraunus (Fabricius) Dm.Df.Vf 35 Dm,Df,Vf Abbot Argus Pseudoptiletes 

NOTES: entire drawing was copied for B&L Plate 35 in the same layout. Figures of H. ceraunus accompanied the original description of A 
pseudoptiletes and are consistent with the Floridian H. c. antibubastus (Hiibner). 

15 Blanehard a. Lycaena lujllus (Cramer) Dm,Df,Vm 38 Dm,Df,Vm Abbot Polyommatus Thoe 

b. Lijcaena epixanlhe (B&L) Df,Vm 38 Df.Vm Abbot Polyoimnatus Epixanthe 

NOTES: entire drawing was copied for B&L Plate 38 in the same layout. Figures of L. epixanthe accompanied die original description of P. 
epixanthe. 

16 Abbot a. Celastrina ladon (Cramer) Dm.Df.Vm.L.P 36 Dm,Df,Vm,L.P Abbot Argus Pseudargiolus 

b. Cupido comyntas (Godart) Dm,Df,Vm,L,P 36 Dm, Df.Vm, L,P Abbot Argus Comyntas 

NOTES: the figures of C. ladon were consulted for the original description of A. pseudargiolus. Includes Abbot's inscription of "Argiolus." 
Scudder (1888-1889. PL 75, figs. 29, 44: PI. 84, figs. 42, 43) copied all die larvae and pupae diat he said were "formerly used in Boisduval and 
LeConte's iconography." Holland (1898, PI. 5, figs. 42, 43) and Klots (1951, PL 6, fig. 16) reproduced some of Scudder's figures. Rogers- 
Price (19S3) figured die entire drawing. 

17 Blanehard Calephelis virginiensis (Guerin-Meneville) Dm,Vm 37 Dm,Vm Abbot Nymphidia Pumila 
NOTES: figures accompanied die original description of iV. pumila and were combined with Abbot's figures of Feniseca tarquinius 
(Fabricius) on B&L Plate 37. 

18 Abbot Danaus plexippus (Linnaeus) Dm,Vm,L,P 40 Dm.Vm.L.P Abbot Danais Archippus 
NOTES: Rogers-Price (1983) figured the entire drawing. 

19 Abbot Danaus gilippus (Cramer) Df,Vf,L,P 39 Df,Vf,L.P Abbot Danais Berenice 
NOTES: includes Abbot's inscription of "Gillippus." 

20 Blanehard Heliconius charithonia (Linnaeus) Dm.Vm 41 Dm.Vm Blanehard Heliconia Charitonia 
NOTES: entire drawing was copied for B&L Plate 41 in the same layout. Based in the width of the yellow bands, the figures probably 
represent die Floridian subspecies H. c. tuckeri W. P. Comstock & F. M. Brown. 

21 Abbot Agraulis vanillae (Linnaeus) Dm,Df,Vf,L,P 42 Df,Vf,L,P Abbot Agraulis Vanillae 
NOTES: drawing has penciled reference to B&L Plate 42, as well as figure legends used on the published plate. It has an attribution to 
Abbot in Boisduval's hand ("abbot Pinxit") and includes Abbot's inscription of "Passifora," probably in reference to die name Papilio 
passifiorae of Smidi & Abbot (1797). This name was crossed out and Le Conte wrote "vanillae" below it. Rogers-Price (1983) figured the 
entire drawing. 

22 Abbot Speyeria idalia (Drury) Df.Vf 43 Df,Vf "Leconte" Argynnis Idaho 
NOTES: style is consistent with Abbot, but credited to Le Conte on B&L Plate 43. Abbot is known to have illustrated diis species at least 
one other time; a single male that was "Met with by Mr. Elliot [Stephen Elliott] in his journey to die mountains" (drawing and notes in The 
Natural History Museum, London). Strangely, the text in B&L did not refer to the occurrence of the species in Virginia or Georgia. 
References to New York and Jamaica were probably derived from Cramer ([1775]). The paper used for this drawing differs slighdv from 
other Abbot drawings in diis set, suggesting that it was original rendered for a different set. 

23 Blanehard a. Boloria ■ selene (Denis & Schiffermuller) Df.Vf 45 Df.Vf Blanehard Argynnis Myrina 

b. Speyeria cybele (Fabricius) Df.Vf 45 Df.Vf Blanehard Argynnis Cybele 

c. Boloria bellona (Fabricius) DfVf 45 Df.Vf Blanehard Argynnis Bellona 
NOTES: signed by Blanehard. Entire drawing was copied for Plate 45 in die same layout. The figures represent die eastern North American 
subspecies B. s. myrina (Cramer), S. c. cijbele, and B. b. bellona. 

24 Abbot a. Chlosyne gorgone (Hiibner) Df.Vf.L.P 46 Df.Vf.L.P Abbot Melitaea Ismeria 

b. Euptoieta claudia (Cramer) Df.Vf.L.P 44 Df,Vf,L,P Abbot Argynnis Columbian 

NOTES: the figures of C. gorgone were used for the original description of M. ismeria (Calhoun 2003) (see text). The larva in this drawing is 
not accurately referable to any species, but conceptually resembles that of C. gorgone (Calhoun 2003) (also see drawing 35). 

25 Blanehard a. Euphydryas phaeton (Drury) Dm,Vm 47 Dm.Vm Blanehard Melitaea Phaeton 

b. Phyciodes batesii (Reakirt) Dm 47 Dm Blanehard Melitaea Thaws 

c. Phyciodes tharos (Drury) Dm.Vm 47 Dm.Vm Blanehard Melitaea Tharos 
NOTES: signed by Blanehard. Entire drawing was copied for Plate 47 in die same layout. The published plate identified die male P. batesii 
as a female of P. tharos. The figures are consistent with eastern North American phenotypes. 

26 Abbot Polygonia interrogation's (Fabricius) Dm,Vm,L,P 51 DmVm.L.P Abbot Vanessa C. Aureum 
NOTES: includes figure numbers to be used for B&L Plate 51, but the arrangement was ultimately changed for die final plate. There is also 
a penciled symbol on the drawing that was probably used to instruct the engraver to position die dorsal adult figure at an angle on die plate. 



146 



Journal of the Lepidopterists" Society 



Table 1. Continued 













Figures 


Artist 












B&L 


copied 


credited on 




No. 


Artist 


Species depicted 


Figures 


plate 


for B&L 


B&L plate 


B&L name 



27 Abbot Junonia coenia Hiibner Df.Vf.L.P 

28 Abbot Junonia coenia Hiibner Dm.Vm.L.P 49 Df,Vf,L,P Abbot Vanessa Coenia 
NOTES: Scudder (1888-1889, Pi. 74, fig. 30; PI. 83, fig. 66) copied the larva and pupa. Holland (1898, PI. 4, fig. 66) reproduced Scudder's 
pupa. 

29 Abbot Vanessa virginiensis (Drury) Df.Vf.L.P 48 Df.Vf,L,P Abbot Vanessa Huntera 
NOTES: drawing has a penciled reference to B&L Plate 4S, as well as corresponding figure numbers and legends used on the published 
plate. Also includes Abbot's inscription, "Huntera." 

30 Abbot Vanessa atalanta (L.) Dm,Vf,L,P 
NOTES: includes Abbot's inscription of "Atalanta." 

31 Abbot Nymphalis antiopa (L.) Dm, Vm 

Notes: includes Abbot's inscription of "Antiopa." Kraus ([1964]) figured the entire drawing. 

32 Blanchard a. Roddia vaualbum Denis & Schiff. Df.Vf 50 Df.Vf 

b. Polygonia progne (Cramer) Dm,Vm 50 Dm.Vm 

d. Aglais milbcrti (Godart) Dm.Vm 50 Dm,Vm 

NOTES: signed by Blanchard. Entire drawing was copied for B&L Plate 50 in the same layout. The figures of R. 
the original description of V j. album. 

33 Abbot Limenitis arthemis (Drury) Df,Vf,L,P 53 Df.Vf.L.P 



Blanchard 
Blanchard 
Blanchard 



Abbot 



Vanessa]. Album 
Vanessa Progne 
Vanessa Milberti 



albun 



accompanied 



NOTES: depicts the subspecies L. arthemis astyanax (Fabricius). The text ; 

different genera used. 

34 Blanchard Limenitis arthemis (Drury) Dm.Vm.Df 



Nymphalis Ursula 
(text), Limenitis 
Ursula (plate) 
nd plate were issued separately in B&L, accounting for the 



54 



Dm.Vm.Df 



Blanchard 



Nymphalis Arthemis 
(text), Limenitis 
Arthemis (plate) 
NOTES: signed by Blanchard. Depicts the subspecies L. a. arthemis (Df) and possibly the more western subspecies L. a. rubrofasciata 
(Barnes & McDunnough) (Dm, Vm). The text and plate were issued separately in B&L. accounting for the different genera used. 

35 Abbot Asterocampa clyton (B&L) Dm,Vm,L,P — — — 

NOTES: the larva and pupa in this drawing are incorrect for this species. In his notes (Harvard), Scudder wrote, "butterfly clyton, but the 
larva & chrys. couldn't belong to it." Although the larva is not an accurate depiction of any species, Scudder (1888-1889) copied and 
identified it as P. interrogationis (Plate 74, fig. 27). He copied the pupa as that of Polygonia comma (Harris) (PI. 83. fig. 39). Holland (1898, 
PI. 3, fig. 27; PL 4, fig. 39) reproduced Scudder's figures and identifications. The same incorrect early stages were used for a drawing of this 
species in New Zealand (see drawing 24 for a similarly cryptic larva). The original description of Apatura clyton was accompanied by B&L 
Plate 56, which was copied from an Abbot hostplant drawing with accurate early stages 

36 Abbot Asterocampa ecltis (B&L) Dm,Df,Vm,L,P — — 

NOTES: includes Abbot's inscription, "Portlandia," apparently a misidentification of Enodia portlandia (Fabricius). In his notes (Harvard), 
Scudder also observed that this drawing was "marked portlandia." The original description of Apatura celtis was accompanied by B&L Plate 
57, which was copied from another Abbot drawing. 

37 Dumenil Historis odius (Fabricius) Dm.Vm 52 Dm,Vm Blanchard Aganisthos Orion 
NOTES: includes a penciled reference to B&L Plate 52, as well as figure numbers and legends for the published plate. Although the plate 
credits Blanchard, a handwritten notation by Boisduval on the drawing attributes it to Dumenil ("Dumenil Pinxit."). It is interesting that 
Boisduval used the genus Prepona on diis drawing and referred to Prepona in livraison 22 of B&L, believed published in 1835. However, 
Boisduval is credited with authoring this genus in Boisduval (1836) (see Cowan 1969). The figures represent the continental subspecies H. o. 
Orion (Fabricius), which also occurs in the Lesser Antilles. 

38 Abbot Libytheana carinenta (Cramer) Dm.Vm.L.P — — — 

NOTES: B&L Plate 64 of Libythea motya was copied from an Abbot hostplant drawing with duplicate early stages, but the adults were 
evidently derived from a specimen in Boisduval's collection (see text). Scudder (1888-1889, Pi. 84, fig. 24) copied the pupa, which was 
reproduced by Holland (1898, Pi. 5, fig. 24). See Figs. 29, 30. 

39 Abbot a. Enodia creola (Skinner) Dm — — — 

b. Enodia portlandia (Fabricius) Df.Vf — — — 

NOTES: Abbot incorrectly associated the male of E. creola with the female of E. portlandia. An Abbot hostplant drawing that correctly 
associated the male and female of E. portlandia was copied for B&L Plate 58. 



Volume 58, Number 3 147 



Table 1. Continued 













Figures 


Artist 












B&L 


copied 


credited on 




No. 


Artist 


Species depicted 


Figures 


plate 


for B&L 


B&L plate 


B&L name 



40 Blanchard Satt/rodes appalachia (R. L. Chermock) Dm,Df,Vm,Vf 60 Df,Vf,Dm,Vm Blanchard Satijrus Canthus 
NOTES: signed by Blanchard. Entire drawing was copied for B&L Plate 60 in the same layout. Satyrodes appalachia was not recognized as 
a separate species from Satyrodes eurydice (Johansson) [=Satyrus canthus (L.)] until 1970. The figures represent the southeastern nominate 
subspecies. 

41 Abbot a. Hermeuptychia sosybius (Fabricius) Dm,Df,Vm,L,P 63 Dm,Vm,L,P Blanchard Satijrus Sosybius 

b. Neonympha areolatus (J. E. Smith) Df,Vf,L,P 63 Df,Vf,L,P Blanchard Satyrus Areolatus 

NOTES: Scudder (1888-1889, PI. 83, fig. 11) copied the pupa of N. areolatus, which was reproduced by Holland (1898, PL 4, fig. 10). 

42 Abbot a. Achalarus lyciades (Geyer) Df,Vm,L,P 71 Df,Vm,L,P Abbot Eudamus Lycidas 

b. Epargyreus clarus (Cramer) Dm,Vm,L,P 72 Dm,Vm,L,P Abbot Eudamus Tityrus 

NOTES: includes Abbot's inscriptions, "Lycidas" for A. lyciades and "Tytirus" (a misspelling of the synonym tityrus Fabricius) for E. clarus. 

43 Abbot Urbanus proteus (Linnaeus) Dm,Vm,L,P 69 Dm,Vm,L,P Abbot Eudamus Proteus 
NOTES: includes Abbot's inscription, "Proteus." 

44 Le Conte? Megathymus ijuccae (B&L) larva only — — — — 
NOTES: drawing is likely by J. E. Le Conte (see text). In 1876, entomologist Charles V. Riley published the life history of this species. In 
Scudder's notes (Harvard) there is a sketch of this figure, which Scudder later sent to Riley and asked, "is this Megathymus yuccae'?", to 
which Riley replied, "widiout doubt!" Eudamus yuccae was first "described" (no textual reference) from an Abbot hostplant drawing on 
B&L Plate 70. 

45 Blanchard a. Calpodes ethlius (Stoll) Dm.Vm 75 Dm.Vm Blanchard Eudamus? Olynthus 

b. Polites vibex (Geyer) Df(2),Vf 75 Df(2),Vf Blanchard Hesperia Brettus 

NOTES: signed by Blanchard. Entire drawing was copied in the same layout for B&L Plate 75, which included the original "descriptions" 
(no textual references) of E. olynthus and H. brettus. Boisduval apparently considered the sexes off! vibex to be analogous, as figures 3 & 4 
of dorsal and ventral females were both identified as males on B&L Plate 75. 

46 Abbot Problema bulenta (B&L) Dm,Df,Vm,L,P 67 Dm,Df,Vm,L,P Abbot Hesperia Bulenta 
NOTES: figures copied for B&L Plate 67, the original "description" (no textual reference) of H. bulenta. In his notes for another drawing of 
P. bulenta in New Zealand, Abbot wrote, "Feeds on the Broad grass, Zozani aquatica folding itself up in the leaf changed 25th bred 6di Aug't. 
Frequents Rice fields, ditches, and the sides of ponds in the lower parts of Georgia is not common." The species was not rediscovered until 
1925. Abbot probably collected his specimens near the mouth of the Savannah River where the species still occurs today. 

47 Abbot a. Hylephila phyleus (Drury) Dm,Df,Vm,L,P 78 Dm,Df,Vm,L,P Abbot Hesperia Phyleus 

b. Wallengrenia otho (J. E. Smith) Dm,Df,Vm,L,P 77 Dm,Df,Vm,L,P Abbot Hesperia Otho 

NOTES: Scudder (1888-1889, PI. 77, figs. 19, 34; Pi. 85, figs. 39, 42) copied all the larvae and pupae. The pupae were reproduced by 
Holland (1898, PI. 6, figs. 39, 42). 

48 Blanchard a. Wallengrenia egeremet (Scudder) Dm.Df — — — — 

b. Wallengrenia otho (J. E. Smith) Vf — — — — 

c. Poanes zabulon (B&L) Df,Vf — — — — 

d. Arnblijscirtes aesculapias (Fabricius) Dm.Vm — — — 

NOTES: signed by Blanchard. Drawing predates die description of W. egeremet, which until recendy was generally treated as conspecific 
with W otho. A penciled circle around the Wallengrenia figures and the notation "remplacer" [replace] suggests that Boisduval was going to 
copy them for B&L, but instead used those from Abbot's drawing 47. 

49 Blanchard a. Nastra Iherminier (Latrielle) Df — — 

b. Polites origenes (Fabricius) Df — — — 

c. Polites themistocles (Latrielle) Vf — — — 

d. Ataloped.es campestris (Boisduval) Dm,Vm — — — — 

e. Poanes yehl (Skinner) Dm,Vm — — — — 
NOTES: signed by Blanchard. Females of up to three different species are associated in this drawing (probably Boisduval's 
misidentification). The second dorsal female may represent P. themistocles, but it cannot be identified with certainty. The figures of A. 
campestris represent the subspecies A. c. huron (W. H. Edwards). 

50 Blanchard a. Anatrytone logan (W. H. Edwards) Dm.Df.Vf - - 

b. Polites peckius (W. Kirby) Dm.Vm - - 

NOTES: signed by Blanchard. The figures of A. logon are consistent with the nominate subspecies from eastern North America. 



148 



Journal of the Lepidopterists' Society 



Table 1. Continued 













Figures 


Artist 












B&L 


copied 


credited on 




No. 


Artist 


Species depicted 


Figures 


plate 


for B&L 


B&L plate 


B&L name 



51 Blanchard a. Polites themistocles (Latreille) Dm.Vm 76 Dm.Vm Blanchard Hesperia Cernes 

b. Atrytone arogos (B&L) Dm.Df.Vm 76 Dm.Df.Vm Blanchard Hesperia Arogos 

c. Poanes zabulon (B&L) Dm.Vm 76 Dm.Vm Blanchard Hesperia Zabulon 

NOTES: signed by Blanchard. Entire drawing copied for Plate 76 in the same layout and used for the original "descriptions" (no textual 
references) of H. cernes, H. arogos and H. zabulon. See Fig. 5. The figures of A. arogos represent the nominate subspecies from eastern 
North America. 

52 Blanchard a. Polites baracoa (Lucas) Dm.Vm 

b. Wallengrenia ophites Mabille Dm,Vm 

NOTES: drawing depicts two West Indian species. The P. baracoa figures may represent the Hispaniolan subspecies P. b. loma Evans. 

53 Blanchard a. Euphyes vestris (Boisduval) Dm.DfVf 

b. Lerema accius (J. E. Smith) Dm.DfVf 

c. Oligoria maculata (W. H. Edwards) Dm.Vm 

NOTES: signed by Blanchard. Illegible notations, scrawled in Boisduval's hand, are partially cut off in the right margin. The figures of E. 
vestris are consistent with the eastern North American subspecies E. v. tnetacomet (Harris). 



54 Blanchard a. Panoquina ocola (W. H. Edwards) 

d. Pholisora catullus (Fabricius) 
NOTES: signed by Blanchard. 

55 Abbot a. Erynnis brizo (B&L) 

b. En/nnis juvenalis (Fabricius) 



Dm.Df.Vm 
Dm.Df.Vm 



Dm,Df,Vm,L,P — 

Dm, Df, Vm 65 Dm.Df.Vm Abbot Thanaos Juuvenalis 

NOTES: includes Abbot's inscription, "Juvenalis" (for E. juvenalis). The identities of the sexes of the dorsal figures of E. juvenalis were 
reversed on B&L Plate 65. Thanaos brizo was originally "described" (no textual reference) from B&L Plate 66, which was copied from an 
Abbot host plant drawing with duplicate figures and a hostplant. A penciled circle drawn around the figures of E. brizo suggests Boisduval 
was going to copy them for B&L, but instead used the other Abbot drawing. Scudder (1888-1889, PI. 77, fig. 18; PI. 85, fig. 38) copied the 
larva and pupa of E. brizo, which were reproduced by Holland (1898, PL 6, fig. 38). 



drawings at the University of South Carolina depict 
butterflies. Forty include figures that were 
reproduced on 44 of the 78 plates in Boisduval & Le 
Conte (1829-[1837]), hereafter referred to as B&L. 
The butterfly determinations and information about 
the corresponding published plates are presented in 
Table 1. Nomenclature follows Opler & Warren 
(2002). The butterfly drawings portray at total of 80 
species and most include multiple species (Figs. 3-5, 
10, 36). The 94 moth drawings in this set are still 
under review. 

The drawings at the University of South Carolina 
were rendered on cream-colored wove paper and 
measure approximately 27 cm x 16.5 cm. They are 
mounted on stiff paper backing, matted, and 
contained in six blue half-morocco portfolio cases with 
gilt lettering that incorrectly identify them as the 
original drawings for Smith & Abbot (1797) (Figs. 1, 
2). The portfolio cases were created by rare book firm 
H. P. Kraus of New York, who sold the drawings to the 
University of South Carolina in 1964 (Calhoun 2003). 
The drawings by John Abbot were rendered in a 
horizontal format, with figures of early stages 
positioned above the adults (Figs. 2, 3, 4, 10, 36). 
When J. E. Le Conte commissioned Abbot for these 
drawings in 1813, he requested that hostplants be 



omitted (Rogers-Price 1983). The drawings by 
Blanchard and Dumenil were mostly rendered in a 
vertical format, do not include early stages, and depict 
only one half of dorsal adults . Nearly all of Blanchard s 
drawings have a penciled outline around the figures 
and are signed, "E. Blanchard, pit." (Figs. 5, 5a). 
Blanchards artistic style was highly refined and true to 
life. Although Abbot's figures were rearranged for the 
plates in B&L, virtually all of the published drawings 
by Blanchard and Dumenil were reproduced in their 
original layouts. 

One drawing in this set portrays only the mature 
larva of Megathymus ijuccae (Boisduval & Le Conte) 
with copious annotations in Latin and French by J. E. 
Le Conte (Table 1). It was drawn on a smaller piece of 
paper that was pasted onto a larger sheet. The style of 
this drawing is similar to smaller drawings of moth 
larvae in this set, most of which were probably 
rendered by Le Conte (Calhoun 2003). This is 
supported by the notes of John Abbot, who credited 
Le Conte (as "Mr. Le Compt") for discovering the 
larva of M. yuccae. In Boisduval et al. (1832-1837), 
Boisduval wrote about caterpillar drawings that he had 
received from New York and Savannah, obviously 
referring to Le Conte (from New York) and Abbot 
(who lived for a time in Savannah, Georgia). Boisduval 



Volume 58, Number 3 



149 




FIGS. 1-2. Portfolio cases of original drawings in the Thomas Cooper Library, University of South Carolina. 1, Five of the six cases. 2, Case 
1 opened to show the matted illustrations (Abbot's drawing 2 of E. marcellus is visible), (photos courtesy of Thomas Cooper Library) 



(1836) also noted that Le Conte had executed as many 
caterpillar drawings as Abbot. 

The drawings at the University of South Carolina 
are numbered in pencil and loosely arranged in 
taxonomic order. Most have notations that were 
written by Boisduval and/or Le Conte, including 
names used in B&L. Although I previously suspected 
that the majority of the inscriptions on these drawings 
were by Boisduval (Calhoun 2003), I have since 
confirmed through additional writing samples that Le 
Conte was responsible for many names and other 
notations (Fig. 6). Samuel H. Scudder examined these 
drawings while they were still in Boisduvals possession 
and also observed that, "in some of Abbot's drawings 
which Dr. Boisduval received from Major LeConte is 
a memorandum by the latter" (Scudder 1888-1889). 
"Nobis," or more often the abbreviation "nob.," follows 
many of the species names. This Latin term loosely 
means "of us" or "of me" and was used to claim 
audiorship of new names. Ten of the drawings possess 
penciled references to the corresponding B&L plates 
("planches" in French) (e.g., "Pi. 1") (Table 1, Fig. 10). 
Some individual figures are numbered (Fig. 6) and 
three drawings even include the complete figure 
legends used on the published plates. On several 
drawings, Abbot inscribed the same Latin names 
employed in Smith & Abbot (1797). The source of 
other inscriptions is not readily identifiable. 

Many of the drawings by Abbot that were copied for 
published plates show alterations, particularly to the 
legs and bodies of adults. Beferring to this set of 
drawings, Oberthiir (1920) complained that Abbots 
legs and bodies were more fantasy than reality. In die 
preface to B&L livraison 10, Boisduval addressed 
criticisms from subscribers about the inaccuracy of 



various published figures, particularly relating to 
poorly formed bodies and legs. In a statement under 
the heading "Avis de Lun des Auteurs" [Opinion of 
One of the Authors], Boisduval promised to retouch 
Abbot's drawings and that beginning with livraison 10 
the published figures would no longer exhibit these 
defects (an imprecise translation in Calhoun (2003) 
suggested the engravings were altered). The 
retouched drawings in South Carolina were used for 
plates issued after livraison 10, further supporting this 
connection. The style of the alterations closely 
matches that of Blanchard, who was probably 
instructed by Boisduval to improve and standardize 
the figures for the engravers. 

When S. H. Scudder visited Boisduval, probably in 
1871, he sketched at least 23 of Abbot's larvae and 
pupae in these drawings. Scudder (1888-1889) 
published these facsimiles, some of which were later 
reproduced by Holland (1898) and Klots (1951) (Table 
1). The auction catalog of Sotheby & Co. (1963) 
figured an entire moth drawing from this set (no. 106). 
Kraus ([1964]) and Bogers-Price (1983) figured five 
butterfly drawings (nos. 31 and 3, 16, IS, 21. 
respectively). 

Missing drawings. Published Plates 4-11, 19. and 
20 were copied from missing Abbot drawings that 
were probably rendered in die same format as those in 
South Carolina. Oberthiir (1920) also feared that 
several original drawings from this set were lost. One 
such drawing, depicting Eurema lisa Boisduval & Le 
Conte, was used for B&L Plate 19 and apparently also 
for figures of the early stages of this species in 
Boisduval (1836, Pi. 2). 

There are fifteen plates from Abbot in B&L diat 
included sizable hostplants (nos. 32, 37, 55-59, 61, 62, 



150 



Journal of the Lepidofterists' Society 





..■ 6 rs.,,,..,;.,; 





10 





Figs. 3-12. Original drawings and published plates. B&L = Boisduval & Le Conte (1829-[1837]). 3, John Abbots drawing 3° of S. liparops 
and C. henrid. 4, Abbot's drawing 11° of C. niphon, S. titus, and C. gryneus. 5, Blanchard's drawing 51° of P. themistocles, A. arogos, and P. 
zabulon used for B&L Plate 76. 5a, Blanchard's signature. 5b, Notation crediting Abbot for B&L Plate 64. 6, Ventral S. liparops from Abbot's 
drawing 12° (note "2" above figure); inset is "liparops" in Le Conte's hand. 7, Ventral female of Thecla liparops, B&L Plate 31, fig. 2. 8, B&L 
Plate 16 ofPieris cleomes. 9, Abbot's drawing of A. m. phileta, New Zealand (ref. no. E-272-f-012). 10, Abbot's drawing 7° of A. m. phileta 
(reference to "PI. 16" at lower right) 11, Dorsal male from Abbot's drawing 7° of A. m. phileta. 12, Dorsal male of P. cleomes, B&L Plate 16. (* 
Thomas Cooper Library, University of Soudi Carolina) 



Volume 58, Number 3 



151 



64, 66, 68, 70, 73, 74) (Fig. 24). The original drawings 
for these plates are also missing, but Scudder (1888, 
1888-1889) noted that he obtained from Boisduval 
"three series of manuscript notes entitled 'Notes to the 
Drawings of Insects,' all written in Abbot's own hand, 
and comprising twenty-seven foolscap pages, rather 
closely written, and describing the changes of two 
hundred and one species; of these thirty-eight are 
butterflies." Scudder donated these notes in 1903 to 
the Boston Society of Natural History. In 1946, 
Harvard University obtained them as part of a larger 
acquisition that also included approximately 600 
original Abbot drawings owned by the Society. I 
obtained a copy of these notes, which are now 
deposited in the Houghton Library. The thee sets of 
notes were segregated by Abbot as "a," "b," and "c." 
Written on the cover sheet in Scudder's hand is "Given 
me bv Dr. Boisduval SHS." These notes include 
entries that correspond to species portrayed with 
hostplants in B&L. Abbot must have presented these 
drawings to Le Conte, who then passed them to 
Boisduval along with Abbot's drawings now in South 
Carolina. 

These missing hostplant drawings (depicting both 
insects and hostplants) probably also account for 



abbreviated plants found in other plates in B&L. 
Plates 16, 18, 22, 25-30 included hostplant leaves or 
small sprigs, which served as substrates for figures of 
larvae and pupae (Figs. 8, 15). P. Dumenil was the 
master engraver for the first 30 plates in B&L, 
including the nine with abbreviated hostplants. He 
apparently created two of these (nos. 16, 22) bv 
combining Abbots figures in South Carolina with 
portions of plants from Abbot's missing hostplant 
drawings. Penciled references to these two published 
plates are found on drawings in South Carolina (nos. 7, 
9) and the plants match those in other surviving Abbot 
drawings. For four of the remaining seven plates that 
have no equivalent drawings in South Carolina (nos. 
18, 25, 28, 29), Dumenil must have derived the 
illustrations from the missing hostplant drawings, but 
reproduced only fragments of the plants to remain 
consistent with his other plates. The plant in Plate 30 
(Fig. 15) appears to have been copied from an 
illustration in Smith & Abbot (1797) (see Discussion). 
It was not until Dumenil was replaced by another 
engraver, known simply as Borromee, that Abbots 
hostplant drawings were reproduced in dieir entirety. 
Whereas Dumenil and his association with Boisduval 
are fairly well understood (see Cowan 1969), 



Table 2. Entries from John Abbot's notes (Harvard University) that correspond to hostplant plates in Boisduval & Le Conte (1S29-[1S37]) 
(= B&L). Grammar and spelling are as given in die original notes. Asterisks (°) denote entries that were assigned to plates from limited infor- 
mation. 



B&L 
plate 



Species depicted 



Set and 

entry 

number 



Notation 



16 



18 



22 



25 



Ascia monuste (L.) 



Eureina daira (Godart) 



Zerene cesonia (Stoll) 



Atlides halesus (Cramer) 



b.6 "White Butterfly. Danai Cleome. Feeds on die Cleome pentapliilles. 
changed 17th July, bred 23rd July, nearly half of the female Butterflies 
varies being of a Dingy colour, as figured, They are generally rare, but 
some years at intervals are very plenty in the lower parts of the County. 
I am indebted for die discovery of the caterpillar of this Species to my 
friend Mr. Oemler, who first found it in his garden in Savannah." 

b.8 "Black streaked little yellow Butterfly Feeds on die Cassia 
Araineicrista, Tyed itself up 27th Aug't changed 28th bred 5th Sep'r. 
Both these kinds [diis and Eurcma lisa Boisd. & Le Conte; prior entry 
in notes] is common in all parts ot the County in Autumn, and settles 
so many together at times to suck moist places on the ground, diat I 
have seen 20 in die compass of a hat." 

a.30 "Clouded yellow Butterfly. Feeds on die Plant figured. Tved up lSdi 
April, changed 19di bred 2nd May, continues to breed all die Summer 
and Autumn, Is most common in the pine woods, often settles several 
together to suck the moist places in roads, and other places." 

c.7 "Great Purple hair Streak. Feeds on die Willow Oak, Qucrcus phclhs. 
Tyed itself up 18th Aug't changed 20di bred 6di Sep'r is not common." 



152 



Journal of the Lepidopterists" Society 



Table 2. Continued 



B&L 

plate 



Species depicted 



Set and 

entry 
number Notation 

oS "Red spotted hair streak Butterllv. Feeds on the Flower figured, 
Parsley Haw, pine. Snap beans etc. Tyed itself up 28th April, changed 
20th bred 14di May. Is not very common." 

b.17 "Black hair streak Butterfly. Feeds on the Parsley Haw, and Oaks and 
Hickory. Tyed up 25di April, changed the 28di bred 10th May. The 
Butterfly frequents Chinquepin blossoms and is not uncommon in the 
oak woods." 

c.9° "Little Brown Butterfly. Feeds on the plant figured etc. Tyed up 20th 
June, changed 22nd bred 20th March, is very rare." 

a.37 "Orange Butterfly. Feeds on the Wild Currant Tree, and Alder, the 
Caterpillar is partly covered with a white loose down, Tyed up 12th 
April, changed 14th bred 25th. The Butterfly frequents Swamps, is 
rare." 
a.25, c.2 "Black veined orange Butterfly. Feeds on Willow. Tyed up 30th July, 
changed die 31st bred 7th August, neither the Caterpillar or Butterfly 
is very common, most frequent near Savannah" [a.25]. "Black veined 
Orange Butterfly. Feeds on die Plant figured; and Willow mostly. Tyed 
up 30th July, changed into Chrysalis the 31st bred 7th August. Is not 
very common" [c.2; this is probably the entry for the drawing used by 
B&L]. 

a. 27° "Orange brown Butterfly. Feeds on the plant figured, Tyed up 16th 
May changed 17th bred 2nd June. Is a rare species" ["common name is 
comparable to "Orange coloured Butterfly" for A. clyton drawing in 
New Zealand; die dates are also similar]. 

"Sugar berry Butterfly. Feeds on the Sugar berry (or Hack berry), Tyed 
up 23rd May, changed 24th , bred 12 June. Is a rare Species." 
"Reed Butterfly. Feeds on Reeds, Tyed up 16 June, changed 17th bred 
25th mostly frequents Swamps in different parts of the County, but is 
not a common species." 

"Great Meadow brown Butterfly. Feeds on the grass figured, and odier 
grasses. Tyed up 19th June changed 20th bred 5th July. Frequents the 
pine woods etc. is not common." 

"Ringlet Butterfly. Feeds on the grass figured, and other grasses. Tyed 
up 25[4]di April, changed 24[5]th bred 2nd May. Is plenty in 
Hammocks and the side of Branches, in most parts of the country" [° 
This entry does not refer to Henneuptychia sosybius (Fabricius), as 
entry b.10 is identical to notes for a drawing of this species in New 
Zealand]. 

"Swamp Butterfly. Feeds on the grass figured, and other grasses. Tyed 
up 10th April, changed lldi bred 24th. frequents Swamps and 
Hammocks is not common." 

"Snout Butterfly. Feeds on the Sugar berry, or Hack beny, Tyed up 
28th April changed 29di bred 8th May. Is rare." 
"Lesser Dingy Skipper. Feeds on the Vine figured. Wild Indigo, and 
Oaks, spun up last Oct, changed into Chrysalis in March, bred 21st 
April, Is not so common as die larger kind." 



28 
29 

32 
37 

55 



56 

57 
58 

59 

61 



62 

64 
66 



Strtjmon melinus (Hilbner) 
Satyrium calamus (Hiibner) 

Callophrys irus (Godart) 
Feniseca tarquinius (Fabricius) 

Limenitis archippus (Cramer) 



Asterocampa clyton (Boisduval & Le Conte) 

Asterocampa celtis (Boisduval & Le Conte) 
Enodia portlandia (Fabricius) 

Cercyonis pegala (Fabricius) 

Megisto cymela (Cramer) 



Cyllopsis gemma (Hiibner) 

Libytheana carinenta (Cramer) 
Erynnis brizo (Boisduval & Le Conte) 



a.29 
a.31° 

b.9 

a.32° 



c.6° 

b.ll 

a.33 



Volume 58, Number 3 



153 



TABLE 2. Continued 







Set and 




B&L 




entry 




plate 


Species depicted 


number 


Notation 



68 



Euphyes arpa (Boisduval & Le Conte) 



c.4° 



70 



Megathymus ijuccae (Boisduval & Le Conte) 



a.45 



73 



74 



Autochton cellus (Boisduval & Le Conte) 



Thorybes bathyllus (J. E. Smith) 



c.3 



b.12 



"Georgia Skipper Butterfly. Feeds on the grass figured, and other 
grasses, Spun up 25th April, Frequents the sides of Ponds in the pine 
Woods, is rare" [dates and habitat are consistent with E. arpa; hostplant 
on the B&L plate is giant whitetop, Rhynchospora latifolia (Baldwin) 
W. W. Thomas (Cyperaceae)-a "grass" to Abbot. This skipper normally 
feeds on saw palmetto, Serenoa repens (Bartr.) Small (Palmae), but 
Minno (1994) reared it in the lab on a species of Cyperaceae, 
suggesting Abbot could also have reared it on the sedge. Abbot mav 
have considered E. arpa and Euphyes pilatka (W. H. Edwards), which 
feeds on Cladium jamaicense Crantz (Cyperaceae), to be the same 
species. Nonetheless, the adults and pattern on the head capsule of the 
larva in the published plate are consistent with E. arpa]. 
"Great Georgia Skipper Butterfly. Lives and feeds on die heart and bud 
of the Bear grass or Wild Aloe, closing the top together with a web, and 
in which its changes into Chrysalis, one which changed the 17th May, 
was bred the 20th June. Those that changes in Autumn lives in 
Chrysalis in the Aloe all winter, coming out the last of March & April, 
is rare but most frequent in the lower parts of the Countv. I am 
indebted to Mr. Le Compte for the discovery of die Caterpillar and 
manner of living of this rare and elegant Species. The Chrysalis is 
covered with a land of powder similar to the Underwing Modis." [the 
dorsal female on B&L Plate 70 represents Megathymus cofaqui 
(Strecker)]. 

"Barr'd Skipper Butterfly. Feeds on the Convolvulus figured, spun up 
4th April, bred 25th . Frequents the sides of Swamps, is rare." 
"Brown Skipper. Feeds on the Beggers lice, spun up in die leaves lSth 
Oct'r bred 20th April, is not very comm on." 



Borromee is obscure. Even his full name is unknown. 
He was an Italian artist and engraver active in France 
during the first half of the 19th century and worked as 
a natural history illustrator for the Museum National 
d'Histoire Naturelle in Paris where some of his 
artwork is currently deposited (Meissner 1996). 
Borromee also served as the master engraver for 
Boisduval (18.36). 

I compared the three sets of Abbots notes at 
Harvard with other notes that he prepared for his 
drawings now in London (Scudder 1872, 1888-1889) 
and New Zealand, and have identified entries for 21 of 
the 24 missing hostplant drawings used for plates in 
B&L (Table 2). Two of these drawings were copied for 
plates that accompanied the original descriptions of 
Apatura clyton Boisduval & Le Conte (= Asterocampa 
clyton) (Plate 56) and Apatura celtis Boisduval & Le 
Conte (= Asterocampa celtis) (Plate 57). Four 
drawings were copied for plates that represent the 
"original descriptions" (no accompanying text) of 
Thanaos brizo Boisduval & Le Conte (= Erynnis 



brizo) (Plate 66), Hesperia arpa Boisduval 6c Le Conte 
(= Euphyes aijia) (Plate 68), Eudamus ijuccae 
Boisduval & Le Conte (= Megathymus yuccae) (Plate 
70), and Eudamus cellus Boisduval & Le Conte ( = 
Autochton cellus) (Plate 73). Abbot's accompanying 
notes help to fill the void where no text was provided 
for these published plates. 

The host leaves in Plates 26 and 27 are unaccounted 
for. Ironically, both plates portrav Parhassius m-album 
Boisduval & Le Conte and do not have corresponding 
entries among Abbot's notes at Harvard. The figures 
of P. m-album on drawing 10 in South Carolina are 
numbered, indicating that all the figures on Plate 27 
were copied from this rendering. The leaf on diis 
plate is crude and looks to have been created h\ 
Dumenil merely to improve composition. There is 
also no appropriate entry for Plate 30 that combined 
figures of two species under die name of Thecla 
favonius (J. E. Smidi) (see Discussion). By comparing 
notes from surviving sets of Abbot drawings. I verified 
that the remaining butterfly entries refer to species not 



154 



Journal of the Lepidopterists' Society 



treated in B&L. 

Artists responsible for the published plates. 

The plates in B&L included small printed notations 
that credited the original drawings to Abbot, 
Blanchard, Dumenil, or Le Conte (Fig. 5b). 
Following the name of each artist was printed "pinx." 
or "pinxit," from the Latin meaning "painted by." 
Based on these notations, Bogers-Price (1983) 
attributed 62 plates to Abbot, whereas Gilbert (1998) 
listed 65. dos Passos (1962) seemingly misunderstood 
the notations, believing 62 plates were merely 
"coloured by Abbot." I examined the plates from the 
copy of B&L once owned by dos Passos (Wittenberg 
University, Ohio) and confirmed that 62 plates 
credited Abbot (nos. 1-20, 22, 24-29, 31-34, 36, 37, 39, 
40, 42, 44, 46, 48, 49, 51, 53, 55-59, 61, 62, 64-74, 77, 
78). Twelve plates credited Blanchard (nos. 35, 38, 41, 
45, 47, 50, 52, 54, 60, 63, 75, 76), three credited 
Dumenil (nos. 21, 23, 30), and one credited Le Conte 
(no. 43). For Plates 1-8 and 10-30 Dumenil 
misspelled Abbot as "Abbott." With the exception of 
Plates 36 and 37 where "Obbit" was given, Borromee 
spelled the name correctly (Table 1). 

As demonstrated by the drawings in South Carolina, 
as well as other original and published illustrations by 
Abbot, the wrong artist was credited on seven plates 
(nos. 3, 14, 15, 17, 43, 52, 63) (Table 1). Fifty-seven 
plates can be wholly attributed to Abbot (nos. 1, 2, 4- 

13, 16, 18-20, 22, 24-29, 31-33, 36, 39, 40, 42-44, 46, 
48, 49, 51, 53, 55-59, 61-63, 65-74, 77, 78). Two plates 
from Abbot (nos. 34, 37) also included figures from 
Blanchard. Two plates (nos. 30, 64) were largely 
derived from Abbot, but included adult figures that 
were evidently derived from specimens in Boisduval's 
collection (see Discussion). Ten plates were from 
Blanchard (nos. 35, 38, 41, 45, 47, 50, 54, 60, 75, 76). 
Seven plates were most likely from Dumenil (nos. 3, 

14, 15, 17, 21, 23, 52). 

Discussion 
The origin of figured specimens. Figures for 
nine of the 29 original descriptions in B&L were from 
drawings by Emile Blanchard (Table 1). This finding 
complicates historical notions about the type localities 
of five taxa: Eudamus olynthus Boisduval & Le Conte, 
Hesperia brettus Boisduval & Le Conte, Hesperia 
cernes Boisduval & Le Conte, Hesperia arogos 
Boisduval & Le Conte, and Hesperia zabulon 
Boisduval & Le Conte. No text accompanied Plates 
75 and 76, but based on the belief ttiat the published 
figures were from John Abbot, Bell (1938) and Miller 
& Brown (1981) assigned these taxa the type locality of 
"Georgia." Such assumptions are no longer 



appropriate. 

Abbot usually drew insect specimens that he reared 
and collected himself, but he occasionally illustrated 
specimens obtained from other local naturalists, 
particularly pharmacist Augustus G Oemler of 
Savannah, Georgia, and botanist Stephen Elliott of 
South Carolina. Abbot provided many plant 
specimens for Elliott's herbarium and exchanged a set 
of insect watercolors for specimens that Elliott had 
personally obtained in Georgia and South Carolina 
(Bogers-Price 1983, Gilbert 1998). Specimens 
collected by Abbot were dispersed to many European 
naturalists during the late 18th and early 19th 
centuries, mostly through London jeweler John 
Francillon. Swainson (1840) observed that Abbot's 
insects "were always sent home expanded, even the 
most minute." At least some specimens were labeled 
"Georgia" in Abbot's own hand (Calhoun 2003). Many 
of these specimens found their way into various 
museums, but it is impossible to know if any were used 
as models for his drawings. 

On the other hand, Blanchard and Dumenil 
undoubtedly drew specimens contained in the 
extensive collection of Boisduval, who obtained them 
from multiple sources. Boisduval (1836) wrote that he 
had received from Le Conte an immense quantity of 
species from North America. In turn, some specimens 
that Le Conte gave to Boisduval came from Thaddeus 
W Harris (Scudder 1869) and Abbot. In B&L, 
Boisduval noted that a specimen "nous a ete envoye 
par Abbot" [has been sent to us by Abbot]. Harris also 
had specimens from Abbot, now deposited in the 
Museum of Comparative Zoology, Harvard University. 
Boisduval possessed Abbot specimens, but it may be 
difficult to establish that Blanchard and Dumenil used 
any of them as subjects for their drawings. Six unused 
drawings by Blanchard in South Carolina (nos. 48, 49, 
50, 52, 53, 54) were probably prepared for the planned 
continuation of B&L. To better understand the origin 
of figured specimens in Blanchards drawings, I have 
identified the applicable subspecies in Table 1 where 
possible. 

Four plates attributed to John Abbot in B&L 
portrayed butterflies that he certainly never 
encountered in Virginia or Georgia. Harris (1972) 
wondered how Abbot was able to obtain specimens of 
these exotic species. Unfortunately, only one of the 
original drawings for these plates is included in the set 
in South Carolina. Three plates portray West Indian 
species: Eurytides celadon (Lucas) (as Papilio sinon, 
Pi. 3), Battus devilliersii (Godart) (as Papilio villersii, 
Pi. 14), and Battus polijdamas (L.) (as Papilio 
polijdamas, Pi. 15). The text did not mention Virginia 



Volume 58, Number 3 



155 



or Georgia within the ranges of these species. 
Although B. polydamas is a resident of Florida that 
occasionally strays into Georgia (Harris 1972), and 
Boisduval (1836) dubiously stated that the species was 
very common in Georgia, die plate in B&L actually 
depicted the distinctive subspecies B. p. neodamas 
(Lucas) diat occurs only on the Lesser Antilles islands 
of Guadeloupe and Marie-Galante. These islands 
were under French control during most of the late 
18di and early 19th centuries. The most unusual 
species treated in B&L is the Asian Leptosia nina 
(Fabricius) (as Pieris chlorographa) (Plate 17, figs. 4, 
5). Boisduval hesitantly included this butterfly, as he 
was uncertain diat the two specimens he possessed 
actually came from North America. He soon rectified 
this error, admitting "C'est par erreur, et sur la foi 
d'Hubner, que nous avons figure cette variete dans 
notre Iconographie . . . nous avons maintenant la 
certitude qu'elle vient de File de Java" [it is by 
mistake, and on the faith of Hiibner, that we have 
figured this variety in our Iconography. . .we are now 
certain that it comes from the island of Java] 
(Boisduval 1836). Obviously, Abbot did not collect the 
figured specimens, nor have I found evidence that he 
received such specimens to illustrate from Boisduval, 
Le Conte, or anyone else. Although Abbot obtained 
specimens from New England during his residency in 
America (Remington 1948), it does not appear that he 
ever drew Lepidoptera that originated from anywhere 
outside die region extending from Virginia to Georgia. 
In all probability, these four plates that included 
extralimital species were created entirely by Dumenil, 
who engraved five of the six plates in B&L that 
included Neotropical and Asian species (Pis. 3, 14, 15, 
17, 23). He was also credited as the artist for Plate 23 
of Anteos maenda (Fabricius). Although Borromee 
engraved the sixth plate, Historis odius (Fabricius) (Pi. 
52), it was based on drawing 37 in South Carolina that 
Boisduval attributed to Dumenil (Table 1). Drawing 4 
in South Carolina of E. celadon (for Plate 3) is 
probably also by Dumenil (Table 1). All six plates diat 
included tropical species should tentatively be 
credited to Dumenil. Because only two drawings for 
these plates survive, Dumenil may have engraved the 
remaining three plates directly from Boisduval's 
specimens. The same can be said for Plate 21 that 
included figures for the original description of Colias 
pelidne Boisduval & Le Conte. This plate credited 
Dumenil as the artist and no corresponding original 
figures are included among the drawings in South 
Carolina. Boisduval (1836) noted specimens of this 
species in his personal collection at the time the plate 
was created. Dumenil also provided the original 



drawings and served as the master engraver for many 
of the plates in Dejean & Boisduval (1829-1837) and 
Boisduval et al. (1832-1837). 

The figures for the original description of Pieris 
protodice Boisduval & Le Conte (= Pontia protodice) 
(Plate 17, figs 1-3) were portrayed on the same plate as 
L. nina and credited to Abbot. However, the text 
referred to the occurrence of this species only in New 
York and Connecticut. Boisduval (1836) again placed 
this species further north when he stated that it had 
been found in the vicinity of New York and 
Philadelphia. A drawing in The Natural History 
Museum, London, reveals that Abbot also 
encountered diis species in Georgia, but he 
considered it "very rare" and noted only a single 
capture on 13 May. This drawing depicts the dorsal 
and ventral surfaces of a single female, rather than a 
dorsal male, dorsal female, and ventral female as 
portrayed on the B&L plate. The dorsal figures on the 
plate were also engraved with disproportionately small 
hindwings, an unlikely result if they had been copied 
from an Abbot drawing. Like the figures of L. nina on 
the same plate, Dumenil probably derived those of P. 
protodice from specimens in Boisduval's collection. 

Miller & Brown (1981) were unaware of a supposed 
type of P. protodice, but a male of this species was 
discovered in the NMNH among specimens 
recognized as Boisduval types acquired via the William 
Barnes collection. The printed labels read "EX- 
MUSAEO/Dris. [Doctoris] BOISDUVAL" and 
"Oberthur Collection." The handwritten 

determination label reads "Protodice. B. Sp./Am: 
Sept:." A red-bordered label reads 

"Type/protodice/a/c Hofer." The determination label 
is similar in format to labels by Boisduval, but is not in 
Boisduval's hand. It may have been written by Louis 
M. A. Depuiset, who helped maintain Boisduval's 
insect collection (Clement 1887). The abbreviation 
"B. Sp." probably refers to "Boisduval [as treated in] 
Species Generale" [Boisduval 1836]. The locality 
"Am: Sept:" refers to Amerique Septentiionale 
[northern (North) America]. Boisduval (1836) 
confirmed that his collection contained specimens of P. 
protodice around die time when B&L Plate 17 was 
prepared. The specimen in die NMNH mav further 
support the theory that the published figures were 
derived, not from a drawing by John Abbot, but from 
specimens in Boisduval's collection. Rather than 
"probably Screven Co., Georgia" as proposed by 
Miller & Brown (1981), the type locality of P. 
protodice is hereby amended to New York. }. E. Le 
Conte lived in New York and conceivably collected die 
figured specimens, including the "tvpe" in the 



156 



Journal of the Lepidofterists' Society 



NMNH. 

Boisduval type specimens. Remarks are 
necessary regarding the Boisduval type specimens oi 
North American Lepidoptera. In 1876, three years 
prior to his death, Boisduval bequeathed his 
Lepidoptera collection to Charles M. Oberthiir of 
Rennes, France (Oberthiir 1880). In 1913, American 
lepidopterist William Barnes organized a project for 
the purpose of comparing North American 
Lepidoptera specimens against the types in European 
museums (Barnes & McDunnough 1914, Oberthiir 
1913, 1914). The actual work was conducted by 
Barnes' curator, James H. McDunnough, who visited 
Oberthiir on 13-14 October 1913 to examine the 
Boisduval material. Because Boisduval had designated 
very few type specimens, McDunnough and Oberthiir 
personally selected Boisduval specimens to serve as 
types (Oberthiir 1913, 1914). Oberthiir (1913, 1914, 
1920) figured many of these specimens, chiefly those 
from California described in Boisduval (1852, 1869). 
This selection process was largely based on existing 
determination labels and resulted in some 
misidentified "types" (Brown 1965, see Discussion). 

Upon Oberthiir's death in 1924, his collection was 
sold for the benefit of his heirs. Appointed to organize 
the sale was Carl Hofer (Riley 1927), who may have 
been Oberthiir's curator (Emmel et al. 1998). Nodiing 
further is known of Hofer; even the exhaustive 
historical files of the library of the Deutsches 
Entomologisches Institut, Eberswalde, Germany, lack 
information about his identity (R. Gaedike, pers com.). 
With the exception of the Sphingidae and 
Hesperiidae, William Barnes purchased the Boisduval 
"types" of North American Lepidoptera (Riley 1927, 
Horn et al. 1990). According to information on some 
of the specimen labels (Fig. 14), Barnes received this 
material in 1925. Oberthiir's brother, Rene, purchased 
the specimens of Hesperiidae. In late 1926 and early 
1927, the bulk of the C. Oberthiir collection, about 
750,000 specimens, was secured by the British 
Museum (N. H.) (now The Natural Histoiy Museum, 
London) (Riley 1927, 1964). The specimens of 
Hesperiidae, including North American types, were 
purchased in 1931 by the BMNH from R. Oberthiir 
(Riley 1964). 

In the Entomology Library of The Natural History 
Museum is a loose-leaf typewritten manuscript 
entitled "List of specimens disposed of by C. Hofer 
prior to purchase of remainder by the British Museum 
(Natural Histoiy)." It is stamped "C. HOFER, 36, F8 
DE PARIS, RENNES (FRANCE)," which was 
Oberthiir's address. The list was conceivably prepared 
in 1927 by Norman D. Riley, who was then serving as 



an Assistant Keeper of Entomology in the BMNH. 
Riley (1927) wrote a detailed account of the purchase 
of Oberthiir's collection and noted that facts about 
preceding sales of Oberthiir's specimens were given to 
the BMNH; the typewritten list of specimens likely 
served as a summary of these transactions. 
Unfortunately, entries on the list have faded and 
become difficult to read. One section has a 
handwritten heading of "Dr. Barnes" and inventories 
the specimens that were sent to W Barnes in 1925. 
Apparently based on a similar list from Hofer, the 
specimens sent to Barnes were labeled upon receipt as 
"type a/c [according to] Hofer." Although Emmel et 
al. (1998) believed that Barnes personally penned 
these labels, they were actually prepared by Foster H. 
Benjamin, who curated Barnes' collection from 1922- 
1927. Benjamin signed and dated some of the labels 
on these specimens (Fig. 14). After Barnes' death in 
1930, his collection was purchased the following year 
by the United States Government for $50,000 and 
moved to the National Museum in Washington, D.C. 
(Hewes 1936, Horn et al. 1990). 

Boisduval's specimens from California were 
obtained mostly from a single collector (R J. M. 
Lorquin) and their histoiy is reasonably well 
documented (Emmel et al. 1998), lending credibility 
to their acceptance as holotypes and syntypes. On the 
other hand, the "types" that correspond to taxa 
described in B&L cannot be traced to any particular 
source. Although Boisduval (1836) claimed that he 
possessed specimens of almost all the taxa that he 
described in B&L, he maintained his collection until 
1876 and he may have acquired the selected "type" 
specimens up to 47 years after the original descriptions 
were published. In addition, not all these "types" are 
consistent with the written descriptions, suggesting 
Boisduval based his original characterizations on other 
specimens or John Abbot drawings. Brown (1965) and 
Miller & Brown (1981) considered such specimens to 
be "pseudotypes." Most of the written descriptions in 
B&L were accompanied by Abbot illustrations and it is 
safe to assume that the original drawings were at least 
consulted for all these treatments. The specimens 
selected as "types" for B&L taxa serve as helpful 
vouchers and can convey Boisdual's taxonomic 
concepts, but they cannot automatically be accepted as 
valid holotypes or syntypes. Some of these "types," 
especially of Hesperiidae, may correspond to 
specimens from Boisduval's collection that were drawn 
by Blanchard for plates in B&L. If so, a careful 
comparison against the original drawings in South 
Carolina may establish such specimens as acceptable 
types. 



Volume 58, Number 3 



157 



Drawings by Blanchard and Abbot. Emile 
Blanchard's artwork is meticulous, arguably surpassing 
Abbot in detail and accuracy. It seems inconceivable, 
but he was onlv about 15 years old when his drawings 
in South Carolina were completed. Blanchard was 
extraordinarily gifted and his father was also a natural 
history illustrator. Like contemporary French 
entomologist Pierre Hippolyte Lucas, Blanchard 
began work in the Museum National d'Histoire 
Naturelle in Paris at a young age. Lucas was just 13 
years old when he was hired to apprentice in the 
zoology laboratory while Blanchard was 14 when he 
accepted a temporary position in the entomology 
laboratory (Gaudry 1900, Lesne 1901). Beginning in 
his teens, Blanchard published on many subjects, 
including insects, mammals and fish. He was probably 
introduced to Boisduval when he started working at 
the museum (ca. 1833) and this is consistent with 
Boisduvals use of Blanchard drawings for plates in 
various publications at that time, including Boisduval 
(1833, 1836) and Boisduval et al. (1832-1837). 
Dumenil and Borromee were also the master 
engravers for these works, and Dumenil served as the 
publisher for Blanchard (1840). The first plate issued 
in B&L from an illustration by Blanchard was Plate 35, 
copied from drawing no. 14 in South Carolina and 
published no earlier than 1833 (dos Passos 1962, 
Cowan 1969). Blanchard became one of the most 
celebrated French natural history illustrators of his 
era, but sadly his eyesight deteriorated over the course 
of his lifetime, resulting in total blindness in his later 
years (Gaudry 1900). 

In contrast, Abbot was over 60 years old when Le 
Conte commissioned him for the drawings in South 
Carolina. Out of convenience, Abbot often relied on 
template drawings that he developed earlier in his 
career to produce duplicate renderings of insects, as 
well as birds (Simpson 1993, Gilbert 1998). 
Consequently, Abbots drawings in South Carolina 
share numerous figures with his other original and 
published illustrations. I compared Abbot's 105 
Lepidoptera drawings in South Carolina with his 95 
Lepidoptera illustrations completed between 1816 and 
1818 for English Naturalist William Swainson that are 
now deposited in the Alexander Turnbull Library, 
Wellington, New Zealand. Fully 142 figures of adults 
and early stages are duplicates. At least ten of the 
butterfly species treated in both B&L and Smith & 
Abbot (1797) share identical figures of early stages. 
Baker (1959) noted that plates in B&L resembled 
some of the original Abbot drawings in Emory 
University. In his exhaustive treatment of geometrid 
moths, Packard ( 1876) copied twenty-three figures of 



adults and early stages from "Abbot MS 
[manuscripts]." Many of these figures are identical to 
those on drawings in South Carolina, but Packard 
doubtless copied them from other Abbot watercolors 
once owned by A. G. Oemler and Cambridge botanist 
Asa Gray. At that time, these drawings were deposited 
in the library of the Boston Society of Natural History' 
where Packard served as acting librarian and custodian 
in 1865 (Mallis 1971). They were procured in 1946 by 
Harvard University. 

Hillhouse (1985) aptly described Abbot's artwork as 
"true in color, subtle and full of light, exact in size, and 
with detailed accuracy" Nonetheless, a few of Abbots 
illustrations in South Carolina lack much of the 
painstaking detail of his earlier watercolors. The 
engravers and colorists were often criticized for the 
imprecision of many plates in B&L, which mav have 
resulted in Dumenil s early departure from the project 
(Cowan 1969). It can now be seen that Abbot himself 
was also responsible for some of the inaccuracies. 

Besides M. ismeria (Plate 46), another problematic- 
illustration in B&L was that of Thecla liparops Le 
Conte (=Satyrium liparops) (Plate 31, figs. 1-2). 
Edwards (1872) thought the published figures of T. 
liparops were "a wretched attempt" at copying the 
work of Abbot, adding, "the species has puzzled 
lepidopterists, nothing like that plate having been seen 
in nature." Michener & dos Passos (1942) similarly 
remarked that the ventral figure on the published 
plate "does not agree with any butterfly known to the 
authors." Forbes (1943) assumed that details of 
Abbot's original drawing had been "tampered with" on 
the plate. Gatrelle (2001) also thought die published 
figures of T. liparops were surely inferior to Abbots 
original drawing. Drawing 12 in Soudi Carolina 
demonstrates that Abbot's figures were poorly 
executed and the published plate was a faithful 
reproduction (Figs. 3, 6, 7). Although the colorists 
were sometimes a bit too liberal with their use of blue 
on the plate, the overall design is die same. Abbots 
drawing for T. liparops confounded even Boisduval. 
who argued that it had "la plus grande resemblance 
avec la Favonius de Smith" [the most greatest 
resemblance with the Favonius of Smith]. Boisduval 
so disagreed with Le Conte over the identity of die 
depicted species that he ceded authorship of die name 
to Le Conte. The identity of this taxon remained in 
some doubt for over a century. 

Original moth drawings. Moth drawings in Soudi 
Carolina were almost certainly destined for use in 
preparing the plates for another installment of Histoirc 
Generate. Boisduval planned to continue the project 
with a volume on moths, but it was never realized 



158 



Journal of the Lepidopterists' Society 



(Cowan 1969). French entomologist Achille Guenee 
consulted a set of Abbot drawings lor descriptions and 
plates in his multivolume treatise on moths (Guenee 
1852- 185S), but the whereabouts oi these drawings 
was unknown (Gall & Hawks 2002). In the preface to 
his first volume on noctuid moths, Guenee referred to 
a set of Abbot drawings provided to him by Boisduval. 
Guenee described them as being accompanied by the 
figures of the caterpillar and he erroneously related 
that the renderings were intended for the continuation 
of Smith & Abbot (1797). ObertMr (1920), who was 
personally acquainted with both entomologists, 
revealed that Boisduval had loaned Guenee the Abbot 
drawings now in South Carolina (and probably also the 
missing hostplant drawings). Upon a recent 
examination of these drawings, Lawrence F. Gall 
confirmed that they were likely among those consulted 
by Guenee. A more thorough examination of Guenee 
(1852-1858) is required to determine if any published 
figures were copied from these Abbot drawings. 
Fmile Blanchard also provided some of the original 
drawings for Guenee's published plates. Sixteen 
smaller moth drawings in South Carolina (nos. 58, 66, 
67, 70, 82, 84, 85, 87, 88, 90, 92, 103, 104, 107, 114. 
115) are crude relative to Abbot's work and were 
rendered on darker paper. J. E. Le Conte, a less 
talented illustrator, may have been the artist as 
suggested by a reference to "Leconte" on drawing 90. 
The true identity of Pieris cleomes. This taxon 
was described in B&L on pages 43-45 and figured on 
Plate 16 from a drawing by John Abbot. The written 
description characterized the male and included brief 
remarks about the female and early stages. The plate 
included dorsal and ventral males, dorsal female, larva 
on a host leaf, and pupa (Fig. 8). The text stated, "il 
habite la Georgie et la Virginie, mais il y est assez rare; 
il est plus commun dans la Floride" [it lives in Georgia 



and Virginia, but it is rather rare there; it is more 
common in Florida]. The dorsal adults in the plate 
have more distinct black wing borders than generally 
found in the Florida subspecies Ascia rnonuste phileta 
(Fabricius). Subsequent intei-pretations of cleomes 
varied. The first was Boisduval himself who wasted no 
time in treating cleomes as a variety of Ascia rnonuste 
(L.) (Boisduval 1836). Despite Boisduval's revised 
treatment, Doubleday (1844) listed three specimens 
from Honduras in the British Museum as P. cleomes. 
Bober (1909) called cleomes a form of A. rnonuste 
from "die soudi of North America" and characterized 
it as "somewhat smaller and less blackly marked." 
Talbot (1932) listed it as a subspecies of A. rnonuste 
from the southern United States. Comstock (1943) 
proposed that cleomes "might be properly applied to a 
subspecies with a more northern range than phileta." 
Chermock (1946) also treated cleomes as a subspecies 
of A. rnonuste and suggested that it must have been a 
small population that eventually became extinct. Klots 
(1951) observed that cleomes "resembled Hi. rnonuste 
rather than phileta," and likewise believed it was 
"possibly a now extinct or diluted subspecies of the 
coastal plain from Georgia onwards." dos Passos 
(1964) synonymized cleomes under nominate A. 
rnonuste. Harris (1972) distinguished cleomes as 
having a "slightly wider and continuous black border 
on the forewings" and thought that Abbot may have 
figured a heavily marked individual of A. m. phileta. 
Howe (1975) referred to "unresolved problems related 
to cleomes" and treated it as a possibly extinct 
subspecies that did not possess a dark form of the 
female as in A. m. phileta. Miller & Brown (1981) and 
Ferris (1989) listed cleomes as a subspecies of A. 
rnonuste, but Ferris thought it might apply to a "pale 
migratory form of A. m. phileta." Most recently, 
Gatrelle (2000) also considered this butterfly to 




FIGS. 13-14. Unacceptable "type" specimen oi Pieris cleomes. 13, Dorsal (left) and ventral of male A. m. orseis from Boisduval's collection. 
14, Specimen labels. 



Volume 58, Number 3 



159 



represent the "never before collected" subspecies A. 
m. cleomes. 

Abbot's drawing 7 of A. monuste in South Carolina 
includes a penciled reference to Plate 16 of B&L (Fig. 
10). Boisduval wrote the name "Pieris orseis God." on 
the drawing, ob\iouslv in his attempt to compare the 
figures with what is now recognized as Ascia monuste 
orseis (Godart). He also discussed orseis in the texts of 
B&L and Boisduval (1836). Sets of Abbot drawings in 
The Alexander Tumbull Library (Wellington, New 
Zealand), The Houghton Library (Harvard 
University), and The Natural History Museum, 
London, contain exact duplicates of a larger rendering 
of A. monuste that portrays the same adults and early 
stages as those in drawing 7 in South Carolina and 
B&L Plate 16 (Figs. 8-10). However, the larger 
drawings also portray a dark female (Fig. 9). This dark 
form is characteristic of A. m. phileta and contradicts 
die notion of Howe (1975) that cleomes lacks dark 
females. The published plate also included a plant leaf 
that matches a portion of the hostplant in the larger 
drawings (Figs. 8, 9). A comparison of Abbot's notes 
demonstrates that Boisduval possessed yet another 
duplicate of Abbot's larger rendering of A. monuste 
(Table 2). Abbot's notes for each drawing are as 
follows: 

1. For John Francillon, ca. 1805-1810 (The Natural History 
Museum, London; see Gilbert 1998, PI. 29): "The White 
Butterfly. Papilio Danaii candidi. The caterpillar feeds on the 
plant figured. It tyed itself up 16th July and changed in the 
chrysalis 17th and bred 23rd. near half the female Butterflies 
varies being a clingy black as figured. They continue in plenty 
about Savannah all diis last summer but I have rarely seen any for 
the last twelve years. I am indebted for this discovery of the 
caterpillars to my friend Mr. Oemler who first found it in his 
garden in Savannah." [although this volume of drawings is dated 
1804, Abbot did not meet Oemler until 1805]. 

2. For Augustus G. Oemler, ca. 1810 (Houghton Library, Harvard 
University; see Rogers-Price 1983, catalog fig. 10): "Pap. Danai 
Cleome. Feeds on the Cleome Pentaphilles. Tyed up 16th July. 
Changed 17th bred 2.3rd nearly half of the female Butterflies 
varies being a dingy black as figured. Some years they are in 
plenty about Savannah, but in others very rare if to be met with 
at all. I am indebted for the discovery of this Caterpillar to my 
friend Mr. Oemler who first met with it in his garden in 
Savannah." 

3. For J. E. Le Conte, ca. 1815-1820 (given to Boisduval) 
(Houghton Library, Harvard University; notes only): "White 
Butterfly. Danai Cleome. Feeds on the Cleome pentaphilles. 
Changed 17th July, bred 23rd July, nearly half of the female 
Butterflies varies being of a Dingy colour, as figured. They are 
generally rare, but some years at intervals are very plenty in the 
lower parts of the Country, I am indebted for the discovery of die 



caterpillar of the Species to my (riend Mr. Oemler, who first 

found it in his garden in Savannah." 

4. For William Swainson, ca. 1817 (Alexander Tumbull Library, 

Wellington, New Zealand; Fig. 9): "Papilio Danai Cleome. Feeds 

on the Cleome Pentaphilles. Tyed up 16th July, changed 17th, 

bred 23rd. many of the female Butterflies varies being of a ding}' 

black as figured. This Butterfly is some Summers very plenty in 

Savannah but is rare in the Inland parts." 

The Latin names that Abbot used for this species 

were based on the Linnaean classification system, 

where Papilio was the genus, Danai (or Danaii) was a 

group that included the Pieridae, Candidi was a 

subdivision of white butterflies, and Cleome was the 

name coined by Abbot based on the hostplant. The 

text for P. cleomes in B&L stated, "Cette chenille \it 

dans les jardins, sur le cleome pentaphylla" [this 

caterpillar lives in the gardens, on the cleome 

pentaphylla] and was obviously excerpted from 

Abbot's notes for the hostplant drawing that Boisduval 

possessed (3 above; Table 2). Abbot correcdy 

identified the hostplant in his drawings as Cleome 

gynandra L. (= C. pentaphylla) (Capjjaraceae). Le 

Conte wrote "Cleome" on drawing 7 in Soudi 

Carolina, undoubtedly in reference to the name or 

hostplant given in Abbot's notes. The adult figures of 

P. cleomes in B&L were evidently copied from drawing 

7 (Fig. 10), while the host leaf was taken from die 

missing hostplant drawing that was analogous to 

surviving copies (Fig. 9). 

Abbot s observations mirror the modern occurrence 
of A. in. phileta in coastal Georgia, which Harris 
(1972) defined as "fairly common at times, especially 
near or on the coastal islands, it is very sporadic in 
occurrence. Further inland it becomes infrequent to 
very rare." The text of B&L even referred to the more 
common occurrence of the butterfly in Florida where 
A. m. phileta is most abundant. This subsj^ecies is 
migratory and adults sometimes reach more northern 
coastal areas in great numbers, such as in June 1881 
when thousands were recorded in Bluffton, Soutii 
Carolina (Williams 1930). Bluffton, Beaufort County, 
South Carolina is located 40 km (25 mi) north of 
Savannah, Georgia where Abbot observed A. monuste. 
Adults of A. in. phileta are extremely variable and 
individuals from peninsular Florida routinely present 
the same dorsal and ventral characteristics as die 
figures in Abbot's drawings. Abbot's illustrations and 
observations, as well as the comments in B&L, are 
referable to A. in. phileta. Pieris cleomes is therefore 
synonymous with A. in. phileta and the engraver, P. 
Dumenil, simply exaggerated some of die pattern 
elements (Figs. 11, 12). This finding is more plausible 
than the fanciful theory that P. cleomes represented an 



160 



Journal of the Lepidopterists" Society 



extinct subspecies. Miller & Brown (1981) gave the 
type locality of P. cleomes as "probably Screven Co., 
Georgia." In view of Abbot s notes, I hereby amend 
the type locality to Savannah, Chatham Count)', 
Georgia. As the identity of the intended butterfly is 
now apparent, and P. cleomes is not involved in any 
complex zoological problem, there is no exceptional 
need to designate a neotype to objectively define the 
taxon. 

Finally, it must be mentioned that there is a male A. 
rnonuste from Boisduval's collection in the NMNH 
that was sent to W. Barnes as the type of P. cleomes 
(Fig. 13). It was listed as "Pieris rnonuste cleomes Bdv. 
& Lee." on the typewritten list of specimens that were 
purchased from the Oberthiir collection by W. Barnes 
(list in The Natural Histoiy Museum, London). The 
original determination label, not written by Boisduval 
(Depuiset?), reads "Monuste L. H. B./Orseis. God./v. 
Cleomes. B. Sp./ America" (Fig. 14). The three letters 
"L. H. B." probably refer to the name rnonuste as 
published by Linnaeus, Hiibner, and Boisduval. 
"Orseis" and "Cleomes" were written in smaller 
handwriting directly below "Monuste," obviously as 
synonyms following the treatment in Boisduval (1836). 
The line "v. Cleomes. B. Sp." likely means "variete 
Cleomes [as treated by] Boisduval [in] Species 
Generale." Also present are three conjoined red- 
bordered labels, written by F H. Benjamin, that read, 
"This is the type of cleomes a/c Hofer, but surely not 
true type as it violates the O.D. and figs, of Bdl.-Lec. 
(1925-J. H. Benj.)" (Fig. 14). As" indicated by 



Benjamin, this specimen is contrary to the original 
description of P. cleomes. In the text of B&L, 
Boisduval compared P. cleomes to monuste and orseis, 
noting that the ventral hindwings of cleomes are less 
brown, the dorsal forewings lack elliptical white spots 
at the apex, and the dorsal hindwings of the male lack 
a series of black marginal spots. The "type" specimen 
in the NMNH (Fig. 13) boldly exhibits all these 
features and is consistent with the South American 
subspecies A. m. orseis. Again, this demonstrates that 
such Boisduval "type" specimens for B&L taxa were 
arbitrarily selected on the basis of existing 
determination labels that included applicable names. 

The true identity of Thecla favonius in 
Boisduval & Le Conte (1829-[1837]). Plate 30 in 
B&L was identified as Thecla favonius from die "les 
parties meridionales des Etats-Unis" [the southern 
parts of the United States]. P. Dumenil was credited 
as the artist, but in the text Boisduval attributed the 
figures to Abbot. The plate depicted dorsal and 
ventral males, dorsal female, larva on a host leaf, and 
pupa (Fig. 15). In an "Observation" following the 
treatment of T. favonius in B&L, Boisduval wrote that 
he doubted his favonius was the same species 
described and illustrated as Papilio favonius in Smith 
& Abbot (1797) (= Satyrium favonius) (Fig. 16). He 
suspected that his favonius was synonymous with 
Papilio melinus Hiibner (= Strymon melinus), whereas 
the butterfly described in B&L as Thecla liparops was 
akin to the favonius of Smith & Abbot (1797). To 
further complicate matters, Boisduval also figured S. 



^W 



% 



+ 




Cg 



15 



*r ^ 




16 







20 \ 




21 




22 



Figs. 15-22. Thecla favonius, Plate 30 in Boisduval &Le Conte (1829-[1837]) (B&L) and Papilio favonius, Plate 14 in Smith & Abbot 
(1797) (S&A). 15, T. favonius, B&L. 16, P. favonius, S&A. 17, Dorsal male S. melinus, B&L. 18, Ventral male S. melinus, B&L. 19, Dorsal 
female, B&L. 20, Dorsal female, S&A. 21, Larva and pupa, B&L. 22, Larva and pupa, S&A. 



Volume 58, Number 3 



161 



melinus on Plate 28 and described it as Thecla hyperici 
Boisduval & Le Conte. The association of these 
various figures and names created nomenelatural 
chaos that persisted for decades. Morris (1862) 
complained, "There is an almost inextricable confusion 
in the determination of these species." Sixty years 
later, Oberthiir (1920) bemoaned that BoisduvaPs 
treatment resulted in "un manque facheux de clarte" 
[an annoying lack of clarity]. Harris (1841) was the 
first to associate the favonius of B&L with S. melinus 
when he noted that it was the same as the species he 
described as Thecla hamuli, now considered a 
subspecies of S. melinus. In his copy of B&L, Cyril F. 
dos Passos wrote on Plate 30, "= melinus Hbn." 
However, diis synonymy is only partially correct. 

Oberthiir (1920) accurately determined that the 
dorsal male in B&L Plate 30 was S. melinus (Figs. 17, 
18), but observed that the female was a different, 
unidentified species. A comparison of this female 
figure with other illustrations by Abbot reveals that it is 
equivalent to the female of S. favonius on Plate 14 of 
Smith & Abbot (1797), but was inexplicably colored to 
portray a darker butterfly (Figs. 19, 20). The larva, 
pupa, and host leaf also correspond to the plate of S. 
favonius in Smith & Abbot (1797) (Figs. 21, 22). 
Therefore, the favonius of B&L is synonymous with 
both S. melinus and S. favonius. Abbot (in Smith & 
Abbot 1797) called the figured hostplant "fork leaved 
blackjack." Although J. E. Smith (in Smith & Abbot 
1797) identified it as Quercus rubra L. (Fagaceae), it 
most closely resembles Quercus laevis Walter 
(Fagaceae), for which "black jack" is a colloquial name 
in the region where Abbot resided (M. A. Garland, B. 
P. Wunderlin pers. com.). Nineteenth century Florida 
botanist Alvan W. Chapman suspected that Abbot's 
references to "black jack" represented Quercus 
catesbaei Michx. (Fagaceae) (Scudder 1888-1889), 
now considered a synonym of Q. laevis. Abbot's 
original drawings for Smith & Abbot (1797) are 
currently deposited in the John Work Garrett Library 
of the Johns Hopkins University. The Linnean Society 
of London preserves his accompanying notes. 

The story of B&L Plate 30 does not end there. 
Boisduval remarked in B&L that the larva in the 
Abbot drawing used for Plate 30 was the same as the 
one figured in Smith & Abbot (1797), but the adults 
were consistent with T. hyperici (= S. melinus). This 
infers that Abbot mistakenly inserted adult figures of 
S. melinus into one of his hostplant drawing of S. 
favonius. This assertion seems unlikely, especially if 
we are also to believe that Abbot altered the coloration 
of his female S. favonius to more closely resemble S. 
melinus. Furthermore, the figures of S. melinus in 



Plate 30 are dissimilar to Abbot's other representations 
of this species. There are also no entries in Abbot's 
notes at Harvard that correspond to a third drawing of 
S. melinus. As credited on Plate 30, Dumenil was 
probably responsible for these figures and he based 
them on a specimen in Boisduval's collection. Abbot's 
notes at Harvard also lack an appropriate entry for a 
drawing of S. favonius, suggesting Dumenil copied the 
female and early stages from Plate 14 of Smith & 
Abbot (1797). Perhaps Boisduval desired to reconcile 
favonius and instructed Dumenil to include both 
inteipretations of this taxon. Plate 30 is 

unquestionably the most peculiar plate in B&L. 

The true identity of Melitaea ismeria. 
According to Cowan (1969), the most accurate 
publication date for M. ismeria in B&L is 1835, not 
1833 as reiterated by other authors. This includes 
Calhoun (2003), in which I provided evidence that M. 
ismeria is synonymous with C. gorgone, rather than C. 
nycteis as proposed by Gatrelle (1998). The original 
description of M. ismeria in B&L is an accurate 
portrayal of the figures in Abbot's drawing 24 in South 
Carolina. Based on a faults' translation of the 
description of the ventral wings, Gatrelle (2003) 
argued that the written account was likelv derived 
from a specimen of C. nycteis. Following is the 
description of the ventral wings as it appeared on 
pages 168-169 of B&L with a translation that 
corresponds to Abbot's original drawing (Fig. 29) (see 
Calhoun (2003) for a color reproduction): "Le dessous 
des superieures differe du dessus en ce que, avant le 
bord posterieur, il y une bande blanche maculaire, 
precedee de trois ou quatre taches de sa couleur. Le 
dessous des ailes inferieures est fauve, avec des taches 
blanches vers la base, puis une bande mediane 
irreguliere, transverse, et enfin des lunules marginales 
de la meme couleur; celles-ci sont separees de la 
bande transverse par une serie de points noiratres 
correspondant a ceux du dessus. La frange de toutes 
les ailes est noiratre entrecoupee de blanc" [The 
underside of the forewing differs from die upperside 
in that, before the posterior edge, is a white macular 
band (a), preceded by three or four spots of its color 
(b). The underside of the hindwing is fawn, with white 
spots towards the base (c), dien an irregular median 
band, transverse (d), and lastly marginal lunules of die 
same color (e); tiiese are separated from die transverse 
band (d) by a series of blackish points (f) 
corresponding to diose of the upperside. The fringe of 
all the win°;s is blackish intersected by white (?)]. 
Gatrelle (2003) misapplied the fawn ("tawny") color to 
the median band and lunules of die hindwing. In so 
doing, he confused diese characters with other pattern 



162 



Journal of the Lefidofterists' Society 




Figs. 23-33. Chlosyne specimens and Abbot figures. 23, Dorsal C. nycteis female, 16.vi.1996, Jackson Co., Florida (leg. J. V. Calhoun) 
[=C. ismeria ismeria of Gatrelle (1998)]. 24, Dorsal C. gorgone female, 28.iv.2003, Hancock, Burke Co., Georgia (leg. J. V. Calhoun) [=C. g. 
gorgone of Gatrelle (1998)]. 25, Dorsal C. g. carlota, 7.V.1972, Jasper Co., Georgia (FSCA). 26, Abbot's dorsal female," London. 27, Abbot's 
dorsal female, New Zealand (ref. no. E-272-f-017). 28, Abbot's dorsal female, South Carolina 00 . 29, Abbot's original ventral figure for M. 
ismeria (characters correspond to text). 30, Ventral C. gorgone, 28.iv.1995, Orangeburg Co., South Carolina (ex. ovum, FSCA) [=C. g. 
gorgone of Gatrelle (1998)]. 31, Abbot's ventral figure", London. 32, Abbot's ventral figure. New Zealand. 33, Abbot's ventral figure. South 
Carolina" °. (°© The Natural History Museum, London; ° "Thomas Cooper Library, University of South Carolina) 



elements. Furthermore, the original description 
clearly defined the lunules as being separated from the 
median band by blackish fioints, not "punctuated" by 
points as indicated by Gatrelle's translation. 

Drawing 24 in South Carolina is one of at least four 
drawings by John Abbot that depict analogous figures 
of C. gorgone, though his attention to detail varied. 
Abbot referred to Burke County, Georgia in his notes 
for three of these drawings. Gatrelle (1998) 
characterized pojxilations of C. gorgone currently 
found in extreme eastern Burke County and adjacent 
portions of South Carolina as a univoltine (April-early 
May) subspecies, C. g. gorgone. The dorsal surface of 
a female C. gorgone that I collected in April 2003 in 
eastern Burke County is reminiscent of Abbot's female 
figures (Figs. 24, 26-28), which are unlike C. nycteis 
from the region (Fig. 23). The ventral surfaces of 
some individuals of the single-brooded phenotype of 
C. gorgone are also consistent with Abbot's ventral 
figures (Figs. 30-33). However, Abbot's drawings 
more closely resemble specimens tentatively- 
recognized as the widespread subspecies C. g. carlota 
(Reakirt) (Fig. 25). The ventral surface of a pre-1840 
Georgia specimen of C. gorgone in The Natural 
Histoiy Museum, probably collected by Abbot and 
identified by Gatrelle (2003) as C. g. carlota, is 
extremely similar to Abbot's ventral illustrations 
(Calhoun 2003, figs. 23, 24). Populations attributed to 
C. g. carlota tend to be slightly paler and are 
multivoltine in Georgia, with adults flying from April 
to September (Harris 1972). A record from Houston 



County, located within the same physiographic section 
of Georgia as Burke County, is dated 3 June (Harris 
1972). This is compatible with the adult emergence 
date of 26 May given in Abbot's notes (though Abbot's 
rearing conditions could have altered development). 
This can also account for the blooming Helianthus 
divaricatus (L.) (Asteraceae) hostplant in three of 
Abbot's duplicate drawings; this plant flowers after the 
flight period of the single-brooded populations of C. 
gorgone. It is notable that the dorsal painting of a 
male C. g. carlota by Howe (1975, PL 40, fig. 1) is 
remarkably consistent with the male figure by Abbot 
(Calhoun 2003, figs. 3, 22). The " proximity of 
occurrence and relationship of the perceived C. 
gorgone jihenotypes in Georgia are unknown. Abbot 
traveled over large portions of southeastern Georgia in 
search of specimens and his reference to Burke 
County does not exclude the possibility that he 
encountered C. gorgone in other areas. Gatrelle 
(2003) believed that the adult butterflies in Abbot's 
drawings are unlike any taxon in die eastern United 
States and projjosed that they represent "composites" 
of C. gorgone and C. nycteis. I see no evidence of this. 
The right half of Abbots dorsal figure of C. gorgone 
in South Carolina is more refined, thus it was used to 
create the dorsal engraving of M. ismeria in B&L 
(Calhoun 2003, figs. 8,9). However, the left half (Fig. 
28) is more faithful to Abbots other versions of the 
same figure. A close examination of the adult figures 
in this drawing revealed no subsequent alterations to 
wing shape or design. Abbot simply rendered these 



Volume 58, Number 3 



163 



figures with less devotion to his template, contributing 
to 170 years of confusion. 

Doubleday (1847) named Melitaea nycteis in 
connection with an accurate engraving of the female 
holotvpe (in The Natural History Museum, London), 
but failed to provide a written description. He did not 
consider M. nycteis to be the same as M. ismeria, 
which he associated with the species now recognized 
as C. gorgone (Calhoun 2003). Boisduvals own 
concept of C. nycteis apparently also differed from 
that of M. ismeria. Boisduval (1869) listed Melitaea 
nycteis and characterized it as approaching the 
appearance of M. ismeria, but he did not suggest they 
were conspecific. Boisduval published this treatment 
34 years after he described M. ismeria, and 22 years 
after Doubleday had named M. nycteis. Although 
Boisduval (1869) referred to nycteis from California 
where it is not known to occur, he was likely familiar 
with this widespread North American species by that 
time. There was little potential for misidentification, 
as Boisduval (1852, 1869) considered the most similar 
California species to represent different taxa (see 
Emmel et al. 1998). Boisduval was presumably 
confused about the source of some of his specimens, 
as he also listed several other species not validly 
recorded from California (J. F. Emmel pers. com.). 
The specimens listed by Boisduval as C. nycteis have 
not yet been located. If correctly identified, they were 
likely from the eastern United States where the 
majority of Boisduvals American specimens 
originated. During the preparation of Doubleday et 
al. (1846-1852), in which M. nycteis was first named 
and figured, Boisduval provided the authors access to 
his personal collection and even loaned them 
specimens for reproduction on their color plates 
(Oberthur 1880). Like many entomologists of his day, 
Doubleday took specimens to Boisduval for 
evaluation. Doubleday mentioned one such trip in 
1841 when he wrote, "should I go merely for a short 
visit to France I mean to take a host of things for 
Boisduvals inspection" (Scudder 1869). Doubleday 
freely loaned and gave away specimens to other 
researchers, sometimes to the detriment of the 
collections in his care (Salmon 2000). There was 
ample opportunity for Doubleday to discuss the 
identity of M. ismeria and M. nycteis and compare 
specimens with Boisduval. 

Additional new evidence indicates that Boisduval 
ultimately possessed at least one specimen that he 
identified as ismeria. The typewritten list in The 
Natural History Museum, itemizing the Boisduval 
"types" that were sent to W. Barnes in 1925, has a 
ticked entry for "Phyciodes ismeria" The use of the 



genus Phyciodes reflects prevailing usage at the time 
the list was created (ca. 1927), as Boisduvals 
determination labels typically included only the 
species name. The specimen has not been found in 
the NMNH and its fate remains a mystery, yet there 
are two valuable clues to its identity. J. H. 
McDunnough, who ostensibly selected this specimen 
in 1913 to serve as the type of M. ismeria, and F. H. 
Benjamin, who would have accessioned this "type" 
into the Barnes collection in 1925, both subsequently' 
coauthored checklists of Lepidoptera with Barnes in 
which they considered ismeria to be synonymous with 
gorgone (Barnes & McDunnough 1917, Barnes 6c 
Benjamin 1926). Prior to moving the massive Barnes 
collection from Decatur, Illinois to Washington, D. C, 
workers spent two weeks "ramming home" the 
300,000 insect pins to prevent the attached specimens 
from jarring loose in transit (Hewes 1936). The 
missing "type" of M. ismeria was perhaps a casualty of 
this process. 

The true identity of Libythea motya in 
Boisduval & Le Conte (1829-[1837]). Abbot is also 
credited with rendering the original drawing for Plate 
64 in B&L of Libythea motya (Fig. 34). Many audiors, 
such as Morris (1862), Seitz (1916), Riley (1975), 
Miller & Brown (1981), and Smith et al. (1994) 
incorrectly attributed the original description of motya 
to this plate in B&L, but Hiibner (1819-[1S27]) was 
actually the first to figure this Cuban species, as 
Hecaerge motya (Plate [137], figs. 1-2). No textual 
reference accompanied the plate of Libythea motya in 
B&L. Because of the perceived connection of the 
original drawing to Abbot, the figured specimens were 
thought to have come from Georgia. Harris (1972) 
speculated that Abbot may have captured a rare 
Cuban stray or acquired it from a source outside 
Georgia. Miller & Brown (1981) observed that the 
illustration seems to depict the Cuban butterfly, but 
suspected it actually represented a "genetic 
throwback" of the eastern North American subspecies, 
Libytheana carinenta bachmanii (Kirtland). 

The plate of Libythea motya in B&L was ob\iouslv 
based on an Abbot hostplant drawing of L. c. 
bachmanii that was a duplicate of another Abbot 
illustration now in New Zealand (Fig. 35). Abbots 
manuscript notes for the drawing in New Zealand are 
identical to his notes at Harvard Universitv that likelv 
accompanied the drawing used for the published plate 
(Table 2). Boisduval gave Scudder the notes now at 
Harvard and Scudder's handwritten name "bachmanii" 
is found next to Abbots entry for this species. For his 
treatment of bachmanii, Scudder (1888-1889) 
obtained the information "Georgia 'rare' (Abbot)" 



164 



Journal of the Lepidopterists' Society 




34 



35 








39 








44 







Figs. 34-45. Libytheana c. bachmanii and L. motya, 34, B&L Plate 64 identified as Libijthea niotya. 35, John Abbot's drawing of L. c. 
bachmanii, New Zealand (ref. no. E-272-f-018). 36, Drawing 38° of L. c. bachmanii. 37, Larva and pupa, B&L Plate 64. 38, Larva and 
pupa from Abbot's drawing. New Zealand. 39, Larva and pupa from Abbot's drawing 38*. 40, Dorsal figure of L. c. bachmanii. New Zealand. 
41, Dorsal figure of L. motya, B&L Plate 64. 42, Dorsal (left) and ventral of male L. motya specimen from Boisduval's collection. 43, 
Ventral figure of L. motya, B&L Plate 64 (image reversed). 44, Ventral figure of L. c. bachmanii, New Zealand (image reversed). 45, Three 
labels from Boisduval's specimen of L. motya. (° Thomas Cooper Library, University of South Carolina) 

from these notes. Drawing 38 in South Carolina also species, Hi/pataus terena (Godart) the occurrence of 

portrays identical figures of L. c. bachmanii, but which in the United States is unknown; the caterpillar 

without the sjjrig of the hostplant, Celtis tenuifolia and chrysalis, however, are from Abbot s drawings, and 



Nutt. (Celtidaceae) (Fig. 36). The early stages and 
hostplant on B&L Plate 64 are identical to Abbot's 
drawings of L. c. bachmanii (Figs. 37-39), but the 
adult figures are quite different (Figs. 40, 41, 43, 44). 
The wings are broader and the palpi are more 
elongated. The forewing apical spots are inconsistent 
with his other drawings and the hindwings are 
rounded, not squared and deeply scalloped as in L. c. 
bachmanii. The dark brown coloring on the dorsal 
hindwings is much less extensive and the ground color 
of the ventral hindwings is dark, speckled with black- 
spots. These deviations from Abbot's figures are too 
great to simply dismiss as a poor engraving. 

An overlooked discovery by S. H. Scudder provides 
extraordinary insight into this mystery. Scudder (1888- 
1889) wrote, "I have examined in Boisduval's 
collection the butterfly figured in Boisduval and Le 
Conte's work on North American butterflies, under 
the name of Libijthea motya, and it is the West Indian 



represent our common species [L. c. bachmanii]." 
Scudder most likely saw this specimen during his trip 
to Paris when he also examined the Abbot drawings 
now in South Carolina and obtained Abbots notes at 
Harvard. Although he identified the specimen as the 
Hispaniolan species Libytheana terena, Scudder 
(1875) also called the published figures "a Cuban 
species," thus he considered L. motija to be 
synonymous with L. terena. Contemporaries of 
Scudder, such as Gundlach (1881), also placed L. 
motya within the synonymy of L. terena. The 
relationship of these taxa is still uncertain, but minor 
genitalic differences and a lack of intermediates 
suggest they are separate species (Kawahara 2001, 
Kawahara pers com.). Scudder must have used 
drawing 38 in South Carolina (Fig. 36) to confirm that 
the larva and pupa in B&L Plate 64 (Fig. 34) were 
from Abbot. Drawing 38 is listed in Scudder's 
personal notes (Harvard University) and he also 



Volume 58, Number 3 



165 



published a copy of the figured pupa (Scudder 1888- 
1889) (Table 1). Probably following Scudder s 
observations, Kirby (1896) also noted that only the 
early stages of B&L Plate 64 (figs. 3, 4) represented L. 
c. bachnwnii. 

To confirm Scudder's claim, the National Museum 
of Natural History (USNM, Washington, D C.) and 
The Natural History Museum, London, were searched 
for specimens of L. motya and L. terena from 
Boisduval's collection. Nothing was found in the 
NMNH, but a single male L. motya was discovered in 
drawer no. 5445 of the Oberthiir collection in London 
(Fig. 42). It has two printed labels that read, "EX- 
MUSAEO/Dris. BOISDUVAL" and "Ex Oberthiir 
coll./Brit. Mus. 1927 — 3." It also has two handwritten 
labels that read, "Motya h. Bd./am. Sept." and 
"Hecaerge Motya/Hubn. (Samml. exot./schmett. II 
Vol. pl.34./fig.3,4) = ne ressemble pas / beaucoup a la 
fig. de Hubn./est intermediaire entre la / fig. 1 & la fig. 
3" [(Sammlung exotischer schmetterling Vol. II, Plate 
34, figs. 3, 4) = does not much resemble the figure of 
Hiibner, is intermediate between the fig. 1 and the fig. 
3]. The determination label (Fig. 45) was written by 
Boisduval and is consistent with his other known labels 
(Horn et al. 1990). The abbreviations 'h." and "Bd." 
probably refer to the name motya as published by 
Hiibner and Boisduval. Again, his abbreviation "am. 
Sept," refers to Amerique Septentrionale (northern 
[North] America). The other handwritten label is 
from C. M. Oberthiir (Horn et al. 1990) and refers to 
the figures of Hiibner (1819-[1827]), who illustrated 
two species under the name of Hecaerge motya: L. 
motya (figs 1-2, as the male) and L. terena (figs. 3-4, as 
the female). This is probably the same specimen that 
Scudder identified as the model for B&L Plate 64 
(Figs. 41-43). Boisduval evidently ordered the 
reproduction of all the elements of Abbot s hostplant 
drawing of L. c. bachmanii, but substituted the adults 
with figures taken from his own specimen. Libythea 
motya of B&L is therefore synonymous with both 
Libytheana motya (adults) and L. c. bachmanii (early 
stages). Scudder (1888-1889) noted that, "bachmanii 
was also in Boisduval's collection, separated from the 
other [motya], but without name." Libythea 
bachmanii was not described until 1851 and previous 
authors considered all American specimens to 
represent L. motya (following B&L, not Hiibner). 
Boisduval obviously chose to illustrate the butterfly 
that most closely resembled the male figures in 
Hiibner (1819-[1827]). There is no surviving original 
drawing of Boisduval's L. motya, suggesting Borromee 
engraved the figures directly from this specimen. This 
additional evidence disassociates John Abbot from 



these figures of Libythea motya and offers yet another 
glimpse into the complicated production of the 
legendary iconography by Boisduval and Le Conte. 

Acknowledgements 

I would like to thank Patrick G. Scott (Thomas Cooper Li- 
brary, University of South Carolina) for providing access to the 
original drawings and for allowing me to reproduce them. I 
am grateful for his hospitality during my visit to the library. 
Mark Maier digitized the drawings and supplied high-resolu- 
tion scans when required. My research on these drawings 
would not have been possible without their kindness and gen- 
erosity. Suzanne Smailes (Thomas Library. Wittenberg Uni- 
veristy Springfield. Ohio) patiently supplied countless scans of 
published plates. Marian Minson (Alexander Turnbull Library, 
Wellington, New Zealand) provided scans and photocopies of 
original Abbot material in her care and granted permission for 
their reproduction. Fred E. Lohrer (Archbold Biological Sta- 
tion, Lake Placid, Florida), Thomas R. Caswell (Architecture 
and Fine Arts Library, University of Florida, Gainesville), and 
especially Beverly L. Pope (Division of Plant Industrv Library, 
Gainesville, Florida) provided literature witiiout delay. Susan 
Halpert (Houghton Library, Harvard Universitv) helped me to 
obtain copies of John Abbot's unpublished notes. Phillip R. 
Ackery and Kim Goodger (The Natural Historv Museum, 
London) searched for specimens and photographed Boisdu- 
val's specimen of L. motya. Robert K. Robbins (National Mu- 
seum of Natural History Smithsonian Institution, Washington, 
D. C.) kindly provided information, as well as digital photos 
and scans of Boisduval "type" specimens. John B. Heppner 
(Florida State Collection of Arthropods, Division of Plant In- 
dustry, Florida Dept. of Agriculture & Consumer Services, 
Gainesville) allowed access to specimens in his care. Margaret 
Humphrey (Macleay Museum, University of Sidnev) spent 
much valuable time searching for Abbot specimens. Lucv Hall 
and Richard Kielb (Entomology Library, The Natural History 
Museum, London) helped with manuscripts and Abbot draw- 
ings. Mark A. Garland (Division of Plant Industry. Gainesville, 
Florida) and Richard P. Wunderlin (Universitv of Soudr 
Florida) furnished identifications of the plants in Abbot's origi- 
nal and published drawings. Akito Y. Kawahara (Universitv- of 
Maryland) offered valuable insight into his research on the 
Libytheinae. Robert S. Cox (American Philosophical Societv) 
provided digital scans of J. E. Le Conte letters for handwriting 
comparisons. Lawrence F. Gall identified the moth depicted 
on drawing 13 in South Carolina. John F. Emmel bestowed 
advice on California butterflies. Reinhard Gaedike 
(Deutsches Entomologisches Institut, Ebersvvalde, Gernianv 1 
searched for historical information. Lawrence F. Gall. Ger- 
ardo Lamas, and David M. Wright provided thorough and 
helpful reviews of the manuscript. Finally, I express my sin- 
cere thanks to Larry Gall and Dave Wright for many lively and 
informative discussions about Abbot and otiier earlv lepi- 
dopteiists. I am indebted to all. 

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168 Journal of the Lepidopterists' Society 



1 1 Nov. 2003). Received for publication 5 September 2003; revised and accepted 

Williams, C. B. 1930. The migration of butterflies. Oliver and 14 April 2004 

Bovd. London. 473 pp. 



TECHNICAL COMMENT 



Journal of the Lepidopterists' Society 
58(3), 2004, 169-172 

A CRITICAL RESPONSE TO THE PAPER "TOUGH AFRICAN MODELS AND WEAK MIMICS: NEW 
HORIZONS IN THE EVOLUTION OF BAD TASTE" BY P. DEVRIES PUBLISHED IN THIS JOURNAL, 

VOL. 57(3), 2003 



P. DeVries has published two papers in the last two 
years about the existence of a strong association be- 
tween "bad taste" of butterflies and toughness of wings 
(DeVries 2002, 2003). One of these papers, "Tough 
African models and weak mimics: new horizons in the 
evolution of bad taste," was published in this Journal 
(57: 235-238). Here, I present a critical review of both 
of DeVries's papers and an opposing point of view. 

DeVries postulates that the evolution of "bad taste" 
(distastefulness) is, in some way, directly connected 
with the development of tough wings, and that "a 
toughened wing integument may be a general trait as- 
sociated with the evolution of distastefulness in butter- 
flies." He argues that toughness of wings appears to be 
an essential component of butterfly resistance to bird 
attacks. He claims that he presented experimental 
proof of his concept of "a wing toughness spectrum 
that has evolved in parallel with the palatability spec- 
trum" and that "toughness of the wings makes butter- 
flies resistant to handling by predators." I fully dis- 
agree with these concepts. I consider them the result 
of conclusions made on the basis of an experimental 
design that does not mimic natural conditions. 

Under the conditions of the experimental design 
used by Dr. DeVries, a dead butterfly is firmly "fixed in 
the grip of a clothing peg with all four wings closed in 
a natural resting position" leaving free only part of the 
wings. A clip assembly (the artificial metallic beak) is 
attached to the hind wings distal margin in such a way 
that the jaw grips the wings of the dead butterfly be- 
tween veins Cul and 2A. Weight is applied on the arti- 
ficial beak until there is a tear in the wings and the 
metallic beak, with the applied weight and the part of 
the torn wing remaining into its grip, falls into a col- 
lecting receptacle. This weight determines the wing 
tear weight (DeVries 2002, 2003). This weight was 
found to be in the range of many hundreds of times 
that of the butterfly tested - 40.0 g for the unpalatable 
Amaurus niavius (a weight that surpasses that of most 
insectivorous birds), L5g for Acraea insignis, and 7.5 g 
for the palatable Bicyclus sufitza and Junonia terea. 

Under natural conditions the butterfly is not firmly 
fixed as it is under the conditions of the experimental 
design used by DeVries. Usually, when caught by a 
bird, a butterfly hangs freely, with only one wing fixed 
by the grip of the beak. The body of the live butterfly 



and the remaining three wings remain free. There is 
practically no weight applied; the weight of the freely 
hanging butterfly is negligible. Thus, the force respon- 
sible for the tear of the wing under natural conditions 
is the strength applied by the violent struggling of the 
freely hanging butterfly to escape from the grip of the 
beak. Obviously, the stronger the butterfly, the higher 
is the chance the caught wing will sustain a tear and 
the butterfly will fly away with only relatively small 
damage to the wing. If the wing breaks under the 
weight of the insect, a bird could never catch success- 
fully and consume a butterfly. Thus, under natural 
conditions, the "wing tear weight" (wtw) is the force 
applied by the struggling butterfly to free itself from 
the grip of the beak. It is a very dynamic, pulling, tear- 
ing force applied under different conditions than diose 
in the experimental design used. It is not a gradual in- 
crease of added weight on the firmly fixed wings of a 
dead butterfly. 

If this force could be measured in grams and ap- 
proximated that of the weight applied under the con- 
ditions of the experiment causing a tear in the wing, 
the experimental design used by DeVries could reflect 
natural conditions. DeVries (2002, 2003) claims that 
"by estimating the force necessary to tear wings" his 
reports "corroborate the hypothesis that wing tough- 
ness may be a corollary of unpalatabilitv in butterflies." 
However, he does not estimate die force applied on 
the beak of the bird by the struggling butterfly leading 
to a tear in the wing at the point where the beak holds 
the wing. Instead he considers that it is the weight ap- 
plied on the wing that leads to a tear. 

In general, palatable butterflies characteristically 
have a short, stout fatty body, relatively shorter wings, 
wide thorax and a last erratic flight. In contrast, but- 
terflies considered unpalatable are characterized bv 
Ions; slender bodies, elongated wings, narrow thoraxes, 
fluttering wing beats, and a slow flight in a straight and 
regular path (Marshall 1909; Chai 19S6, 19SS; Chai & 
Srygley 1990; Srygley & Chai 1990; Pinheiro 1996). 
The flight pattern of palatable butterflies is highly cor- 
related with thoracic muscle mass (Chai & Srvglev 
1990, Srygley 1994). In fact, most of their wide tho- 
racic cage (85-95% of wet thoracic mass) is filled with 
massive flight muscles for quick take off, acceleration 
and increased flight speed (Hocking 1985; Ellington 



170 



Journal of the Lepidopterists' Society 



1991). Evidently, butterflies considered palatable pos- 
sess a high struggling ability, more strength and thus a 
better chance to slip out of the beak or escape its grip 
— leaving the bird with only a small piece of wing in 
the beak. In contrast, the markedly elongated slender 
thorax of butterflies considered unpalatable is associ- 
ated with weaker flight muscles (less muscle mass), 
which explains their characteristic flight pattern. No 
doubt, they are less capable of opposing the strong 
grip of the beak. Evidently, a palatable butterfly, hav- 
ing a low wtw (weaker toughness of the wing), is better 
protected than a distasteful one from being eaten by a 
bird by escaping only with a small defect in its wing. 
Thus, the considered distasteful butterflies, contrary to 
DeVries's thesis, are less capable of escaping from the 
grip of the beak, i.e., more vulnerable to predation by 
birds, despite their higher wtw. 

Two questions arise: Why should unpalatable but- 
terflies, despite their supposed strong chemical de- 
fense and warning aposematic coloration, evolve wings 
with a high wtw — a physical attribute that makes them 
more vulnerable to predator attack than palatable but- 
terflies which, instead of a chemical defense and warn- 
ing (aposematic) color patterns for evading a predator, 
rely on their cryptic color patterns and a fast erratic 
flight? Why should palatable butterflies with their 
characteristic fast erratic flight be attacked by birds 
and comprise their usual diet but unpalatable butter- 
flies, with their characteristically fluttering wing beats, 
slow flight in a straight and regular path and wings 
with high wtw, be avoided by predatory birds? It is a 
paradox that prey that is easy to catch and with a high 
wtw is avoided and prey that is most difficult to catch 
and possesses wings with low wtw is preferred by birds 
and forms part of their regular diet. 

I argue that a bird does not reject a butterfly on the 
basis of aposematic color pattern and a supposed 
chemical defense, but rather on die basis of a charac- 
teristic morphological and behavioral pattern, which 
provides the bird with a signal whether the prey is ac- 
tually profitable or unprofitable as a food source (see 
Kassarov 2003b, c). Only the flight muscles, the repro- 
ductive organs, the digestive tract and the abdominal 
fat have a nutritional value; the remaining chitinous in- 
tegument, including the wings, is not metabolized. In 
contrast to the narrow thorax and long slender body of 
unpalatable butterflies, palatable butterflies character- 
istically have a wide thorax filled with powerful flight 
muscles and a stout, fatty abdomen. 

It is well known that butterflies considered unpalat- 
able have a tough, very resilient body with a rubbery 
consistence. Wiklund and Jarvi (1982) suggested that, 
because many aposematic species are tough and diffi- 



cult to kill (Cott 1940; Edmunds 1974), body tough- 
ness (they do not mention the wings) would reduce 
the risk of a lethal attack and allow them to escape. 
Birds are very seldom, if at all, able to attack the but- 
terfly's body directly. The relatively small body is well 
hidden between the large wings and thus protected by 
them from a direct attack. This fact is especially true 
for aerial hawkers, the main bird predators of butter- 
flies, who catch their prey on the wing. To reach the 
body, the bird has to lose energy first to catch the but- 
terfly and then, as most bird species do, dismember 
the butterfly (another energy and time-consuming 
process) before finally swallowing it. Whether the 
body is tough or not tough does not change the fate of 
the butterfly; a dismembered butterfly is a dead but- 
terfly. If toughness of the integument protects an in- 
sect from being eaten by birds, Coleoptera with their 
"armored" integument should be the best-protected 
insects. In fact, these insects belong to the regular diet 
of birds regardless of whether they are hawkers catch- 
ing their prey on the wing or terrestrial gleaners. 

There are no published data concerning a causal re- 
lationship between toughness of the integument and 
chemical compounds that may render the insect dis- 
tasteful. Such a relationship could exist if based on a 
chemical reaction; for example, polymerization of the 
chemical compound responsible for a chemical de- 
fense that leads to hardening of the chitinous integu- 
ment. It seems highly improbable, however, that 
chemical compounds that supposedly render a butter- 
fly distasteful could cause the integument to become 
tough and resilient simultaneously (see Kassarov, 
2003a). 

Thus, how could toughness (high wing tear weight) 
of the wing be a "corollary of unpalatability" as De- 
Vries (2002, 2003) postulates? It is rather a corollary of 
palatability. If there is "an evolutionary correlate be- 
tween toughness of wings and unpalatability," it is logi- 
cal to expect that there should be an evolutionary 
corollary between weakness of wings and palatability. 
Neither is correct. Chemical defense (distastefulness) 
of the butterfly and toughness of wings are two attrib- 
utes that evidently do not act in concert but against 
each other. The weaker the wing (the lower the wtw), 
die better the chance the butterfly will escape and vice 
versa - the tougher the wing, the lower the chance that 
die butterfly will escape. The only way the butterfly 
can escape is by the wing breaking at the point where 
the beak holds it. Thus, low wtw facilitates escape. If 
the wing does not break, the bird will subdue the but- 
terfly, i.e., the butterfly will be a dead butterfly. If taste 
is die factor responsible for the rejection of a distaste- 
ful butterfly by a bird predator, why should nature ere- 



Volume 58, Number 3 



171 



ate conditions for the parallel evolution of a physical 
attribute (toughness of wing) acting against the sup- 
posed chemical defense? The bird's ability to taste a 
butterfly via beak mark tasting was discussed in detail 
elsewhere (Kassarov 1999). It was shown that an in- 
sectivorous bird is not able to taste a butterfly via beak 
mark tasting. 

There are many more flaws in DeVrises's experi- 
mental design. Using only a single size artificial metal- 
lic beak (10mm x 3.68mm) makes a reliable compari- 
son of wing tear weights in butterflies with different 
sizes hardlv possible. The smaller the wing the larger 
will be die torn area of the wing in proportion to its 
size and the lower will be the wtw: the larger the wing, 
the smaller will be the torn area and the higher the 
wtw. The smaller the part of wing gripped by the arti- 
ficial beak, compared to the remaining free part of the 
wing, the lower will be the wing tear weight. The 
closer die artificial beak is placed to the periphery of 
the distal margin of the wing, the weaker will be the 
measured toughness of the wing (the wtw). For com- 
parable results, an equal part of the artificial bill 
should grip the wings of the different butterflies tested 
(for example, 10.0 mm inward from the outer margin 
of the wing), and, what is more important, an equal 
part of the wing of the butterflies with a different size 
tested should be out of the grip of the clothing peg 
(only one size clothing peg was used). The greater die 
part of die wings of the firmly held butterfly (with the 
wings closed in a natural resting position) secured in 
the jaws of a wooden clothing peg, the higher will be 
the wtw. For an assessment of the toughness of the 
wings of different species belonging to different gen- 
era, the artificial beak used in the experiment should 
grip an equal portion of wing. Whether the artificial 
beak is placed in the space between two veins or in a 
space including one or more veins affects markedly the 
value of the wtw. The "vein tear weight" can be ex- 
pected to be markedly higher than the wing tear 
weight measured with the beak placed in the space be- 
tween two veins. The smaller the wing, the smaller is 
the space between two veins. Using the same size arti- 
ficial beak and clothing peg leads inevitably to mis- 
leading results. DeVries did not use same sized winged 
butterflies. Thus, the position of the artificial beak on 
the wing (angle of attachment, amount of wing 
gripped, etc.) is most important for receiving compa- 
rable results. The presentation of the experimental de- 
sign in the methods section of DeVries's (2002, 2003) 
papers is very vague, inviting many questions in regard 
to its reliability 

DeVries (2002, 2003) reports no significant relation- 
ship between wing length and wtw among species. 



This finding is misleading. It does not reflect the con- 
ditions observed in nature. As mentioned above, under 
his experimental design, the artificial beak is anchored 
in the space between Cul and 2A (hind wings) of the 
firmly fixed four wings in the jaws of a wooden clothing 
peg. However, under natural conditions, i.e., the but- 
terfly hanging free (not fixed), held only at the point of 
the grip of the beak, the length of the wing will play a 
significant role. The strength of the wing will depend 
on where it is held by the beak. The closer to the apex 
(away from the base) the weaker the wing. The force 
applied on the wing by the struggling butterfly in- 
creases and the weight of the butterfly also starts to 
play a role in the process of tearing. I have in my col- 
lection of several thousand Heliconius (a genus with 
markedly elongated elegant wings) a great number of 
specimens with wing damage considered to be the re- 
sult of a bird attack. Only in a few of them is the dam- 
age located in the space between Cul and 2A. 

Under die conditions oi the experiment the 
strength with which the artificial metal beak holds die 
wing of each tested dead butterfly remains constant. 
The initial reaction of the bird to the violent effort of 
the prey to escape from the grip of the beak is disre- 
garded. If the insect manages to escape, it is usually 
immediately after being caught — a very dynamic 
event. 

Obviously, if unpalatable butterflies have a high wtw 
in contrast to the low wtw of the palatable butterflies 
(DeVries 2002), the supposedly unpalatable models 
should also have a higher wtw in contrast to that of die 
palatable mimics. In DeVries (2003), an aposematic 
model (Amaurus albimaculata) was found to have sig- 
nificantly tougher wings than its putative Batesian 
mimic (Pseudacreae lucretia); die mimic was found to 
have significantly tougher wings than its non-mimic 
relative, a palatable species belonging to a different 
genus (Cymothoe herminia). Note diat the experimen- 
tal design used to measure the wing tear weights is die 
same in both papers. No doubt, the results of the ex- 
periments performed in both papers will be the same, 
and the conclusions also. The onlv difference between 
die two papers is that onh/ one species of unpalatable 
butterfly considered the model was tested against onh/ 
one species considered a putative mimic (a palatable 
butterfly) and one non-mimic palatable butterfly 
(2003), instead of die two palatable and three unpalat- 
able species, again belonging to different genera, but 
not considered models and mimics (2002). 

If mimics have higher wtw than non-mimics, all 
mimetic butterflies in the genus should have higher 
wtw than the non-mimics in die same genus. Mean 
wtw differed significantly among different individuals 



172 



Journal of the Lepidopterists' Society 



of the species tested. Figure 1 of DeVries (2002) 
shows that the highest wtw of P lucretia (N = 23) was 
far above the lowest wing tear weight of A. albimacu- 
lata. The same was found for the wing tear weight of 
C. herminia (N = 14) compared to that of P. lucretia. 
Does this marked amplitude between the highest and 
die lowest wtw in different individuals belonging to 
the same species indicate differences in toughness of 
their wings? Different distastefulness? If there is a 
corollary between wtw and palatability, there should 
be a significant difference in palatability and flight pat- 
tern among individual species (mimetic and non- 
mimetic) belonging to the same genus. I do not know 
a butterfly genus comprising species morphologically 
different or with different flight patterns. Why hould 
mimetic species have a higher wtw than non-mimetic 
species belonging to the same or different genera? Is 
there a corollary between wtw and the ability of a sp- 
cies to mimic a model? Is a certain level of wtw nec- 
essary to enable a species to mimic a model? 

DeVries states that his method provides a means for 
asking whether model butterflies are tougher than 
mimics, and if non-mimic butterflies are the weakest 
ones. He also states that "by exploring the parallel be- 
tween the palatability spectrum and wing toughness 
we may potentially open new horizons in the evolution 
of bad taste." Obviously I fully disagree! I consider 
the results obtained by DeVries (2002, 2003) an exper- 
imental artifact. The conclusions drawn are valid only 
for the conditions of die experiment. They cannot, and 
should not, be extrapolated to the different conditions 
existing during an attack of a bird on a butterfly in na- 
ture. 

Literature Cited 

Chai, P. 1986. Field observation and feeding experiments on the re- 
sponse of Rufous Tailed Jacamars (Galbula nificunda) to free- 
flying butterflies in a tropical rainforest. 

. 1988. Wing coloration of free-flying Neotropical butterflies as 

a signal learned by a specialized avian predator. Biotropica 20: 
20-30. 

. and R. B. Srygley. 1990. Predation and the flight, morphology, 

and temperature of Neotropical rain-forest butterflies. Am. 



Nat. 135: 748-765. 

COTT, II. B. 1940. Adaptive coloration in animals. Methuen, Lon- 
don. 

DeVriws, P. J. 2002. Differential wing toughness in distasteful and 
palatable butterflies: direct evidence supports unpalatable 
theory. Biotropica 34: 176-181 

. 2003. Tough African models and weak mimics: new horizons 

in the evolution of bad taste. J. Lepidopterists' Soc. 57: 235-238 

EDMUNDS, M. 1974. Defence in Animals: A survey of antipredator 
defences. Longman, Burnt Mill. 

Ellington, C. P. 1991. Limitation on animal flight performance. J. 
Exp. Biol. 160: 71-91. 

Hocking, B. 1958. Insect flight. Sci. Am. 199: 92-98. 

Kassarov, L. 1999. Are birds able to taste and reject butterflies 
based on "beak mark tasting"? A different point of view. Be- 
haviour 136: 965-981. 

. 2003a. Notes on the evolution of unpalatability in butterflies 

by means of individual selection. J. Res. Lepid. 37: 71-73. 

. 2003b. Are birds the primary selective force leading to evolu- 
tion of mimicry and aposematism in butterflies? An opposing 
point of view. Behaviour 140: 433-451. 

. 2003c. Is aposematism a valid concept in predator-prey rela- 
tionship between birds and butterflies? A different point of 
view. Tropical Lepid. 2003, in press. 

Marshall. G. A. K. 1909. Birds as a factor in the production of 
mimetic resemblance among butterflies. Trans. Rov. Entomol. 
Soc. Lond. 1909:329-383. 

Pinheiro, C. E. G. 1996 Palatability and escaping ability in 
Neotropical butterflies: Tests with wild kingbirds (Tyrannus 
melancholicus, Tyrannidae). Biol. J. Linn. Soc. Lond. 59: 351- 
365. 

Srygley, R. B. & P. Chai. 1990. Flight morphology of Neotropical 
butterflies: palatability and distribution of mass to the thorax 
and abdomen. Oecologia 84: 491-499. 

SRYGLEY, R. B. 1994. Locomotor mimicry in butterflies? The associ- 
ation of position of centers of mass among groups of mimetic 
unprofitable prey. Phil. Trans. Roy. Soc. Lond. (B), 343: 145- 
155. 

WlKLUND, C. & T JARVI. 1982. Survival of distasteful insects after 
being seized by naive birds. A reprisal of the theory of apose- 
matic coloration evolving through individual selection. Evolu- 
tion 36: 998-1002. 



LUKA KASSAROV, Research Associate, Florida State Collection of 
Arthropods. DPI, FDACP, P.O. Box 147100, Gainesville, Florida, 
32614 -7100. USA. Corresponding address: 130 Spruce Street 28B, 
Philadelphia, PA 19106: E-mail: lukakassaroo@cs.com 

Received for publication 17 January 2004; revised and accepted 16 
May 2004 



GENERAL NOTES 



Journal of the Lepidopterists' Society 
58(3), 2004, 173-177 

DEFENSIVE FLOCCULENT EMISSIONS IN A TIGER MOTH, HOMOEOCERA STICTOSOMA 

(ARCTIIDAE:ARCTIINAE) 



Additional kev words: antipredator. Euchromiini. Panama, subabdominal pouch, moth, chemical defense. 



Tiger moths (Arctiidae) exliibit a wide range of an- 
tipredator adaptations, including ultrasound reception 
and production, reflex immobilization, wasp mimicry, 
and chemical secretions in the form of foams and liq- 
uids (see Beebe & Kenedy 1957, Blest 1964, Roth- 
schild et al. 1979, Fullard 1990, Conner 1999, Weller 
et al. 1999). While performing field studies in Panama 
over tire course of several years, we observed what ap- 
pears to be a novel mode of defense for the Lepi- 
doptera- the expulsion of abdominal 'flocculent' mate- 
rial in a neotropical tiger moth, Homoeocera 
stictosoma Druce (Arctiidae, Arctiinae, Euchromiini) 
(Jacobson & Weller 2002). In this note we summarize 
our observations on the behavior associated with floc- 
culent release, and describe some structural and 
chemical characteristics of the flocculent fibers, and 
the subabdominal pouches where they are produced 
and stored. 

Certain species of the arctiid tribe Euchromiini pos- 
sess conspicuous pouches on the ventral anterior ab- 
domen. These pouches have been described only in 
males to date, and are variously referred to as 'ventral 
valves' or 'subabdominal pouches' (see Weller et al. 
2000). Depending on the species, there may be one or 
two pouches, which contain either non-deciduous hair 
pencils or deciduous scales (Weller et al. 2000). The 
latter, when discharged from the pouch, appear as 
fluffy white or yellowish cotton, called 'flocculent' by 
Blest (1964). Despite the widespread occurrence of 



subabdominal flocculent throughout the Euchromiini 
(R. Simmons, pers. com.), little is known of its func- 
tion. Conner et al. (2000) provided experimental evi- 
dence for its role in courtship in one species, Cosmo- 
soma myrodora Dyar, whereby males increase their 
probability of mating by enveloping the female with 
flocculent. Females covered with this material, in 
turn, are thought to be chemically protected against 
spiders, due to the high pyrrolizidine alkaloid content 
of the flocculent filaments. Flocculent has also been 
documented to play a role in courtship in two other 
species, Syntomeida melanthus, which actively re- 
leases flocculent (Sanderford 1992), and S. ipomoeae, 
which flashes its flocculent briefly, but does not release 
it during courtship (Johnson 2002). In C. myodora 
and other flocculent-bearing species examined, there 
is no direct evidence that flocculent functions as a 
mechanism of defense for the possessor, because its 
production has not been observed to be evoked by 
handling or otiier mechanical disturbances of the indi- 
vidual (Blest 1964, Conner et al. 2000, R. Simmons, 
pers. com,, Yack, pers. obs.). Homoeocera stictosoma 
(Fig. 1) is exceptional in this respect. Blest (1964) 
commented that this species emits flocculent upon 
'light restraint of an individual'. Our observations, out- 
lined below, corroborate Blest's earlier hvpodiesis. 

Moths were collected at ultraviolet and mercury va- 
por lights in neotropical lowland rainforest on Barro 
Colorado Island by A. Aiello and R. Silberglied in 




Fig. 1. A. A male Homoeocera stictosoma, collected on Barro Colorado Island, Panama. B. A male H. sticto- 
soma discharging flocculent while being restrained. An arrow points to the subabdominal pouch, where flocculent 
is rapidly scooped out by the thoracic legs. 



174 



Journal of the Lepidopterists' Society 



March 1980, and by J. Yack in June and October 1998 
and September 1999. Moths were collected from 
sheets and placed in plastic vials where they remained 
for up to 2 days until being tested. Behavioural obser- 
vations were made on a total of 9 (3 in 1980; 3 in 1998; 
3 in 1999) individuals at the study site, and subsequent 
anatomical investigations of the subabdominal 
pouches were carried out on 3 specimens fixed in alco- 
holic Bouin's (Pantin 1946) or C&C fixative (Chau- 
thani & Callahan 1966), and one dried specimen 
cleared in warm 10% KOH for 10 minutes. Floccu- 
lent material was collected from two live specimens, 
glued to aluminum stubs and dried overnight. The 
specimens were coated with gold-palladium lor 80 sec- 
onds using a Pelco SC-4 Plasma Coater, and examined 
with an Amray 1810 Scanning Electron Microscope. 
For chemical analysis, the flocculent was extracted 
with methanol (2 ml) overnight at room temperature 
and subsequently analyzed by positive-ion electrospray 
ionization mass spectroscopy, using a Micromass 
(Manchester, U.K.) Quattro I mass spectrometer. 
Moths were identified as male H. stlctosoma according 
to their distinctive wing and body markings using Ru- 
bio & Pesantez (1997) and Draudt (1915), and through 
comparison with specimens at the United States Na- 
tional Museum with the assistance of R. Simmons. 

Independent behavioral observations were made by 
one of the authors [AA] in 1980, and another author 
[JEY] during 1998 and 1999. In 1980, observations on 
H. stlctosoma were made while doing palatabilitv ex- 
periments with moths on small orb weaver spiders. 
During these trials, various tiger moth species were 
tossed from their holding vials into spider webs, and 
observations were made on the spider's response. To 
the author's surprise, when a H. stlctosoma struck the 
web, it immediately used its legs to pull out abdominal 
flocculent, and thereby escaped the spider's web, leav- 
ing it covered in flocculent. Moths were recaptured 
and the trials were repeated until the moths ran out of 
flocculent material. Similar observations were made 
by J.E.Y. in 1999, whereby a male tossed into a spider 
web immediately released flocculent and escaped, 
leaving the web covered with flocculent fibers. In 
1998 and 1999, all 6 moths tested discharged floccu- 
lent upon being restrained. In no cases was flocculent 
released while moths were collected or stored in plas- 
tic vials, but immediately upon being restrained, all in- 
dividuals instantaneously responded by releasing 
clouds of flocculent that surrounded the moth, adher- 
ing to the forceps and/or fingers being used to hold die 
moth (Fig. 1). Flocculent emission discontinued when 
the moth was placed back in the holding container, but 
could be provoked repeatedly upon further restraint 



until the subabdominal pouches were empty. 

In H. stlctosoma, the flocculent fibers are tightly 
packed inside two subabdominal pouches formed by 
abdominal sternites I, II, and III (Fig. 2). The largest 
of the two, the anterior pouch, is covered ventrally by 
an enlarged sternal plate (SII) that partially overlays 
the anterior edge of SIII. The opening to the anterior 
pouch is directed posteriorly. The outermost edge of 
SII is lined with clear, rounded, non-deciduous scales 
that may be interpreted as scent scales (cf. Weller et al. 
2000). The opening to the smaller posterior pouch oc- 
curs between SIII and SIV, with shallow pockets ex- 
tending both anteriorly and posteriorly. The posterior 
edge of SIII is also lined with transparent, rectangular 
scales that are somewhat smaller than those associated 
with SII. Corresponding to the pattern found in other 
euchromiines, female H. stlctosoma lack pouches (R. 
Simmons, pers. com.). In a detailed examination of 
the subabdominal pouches in the euchromiine-ctenu- 
chine clade, Weller et al. (2000) described two main 
types: a single pouch, and a double pouch that corre- 
sponds to that of H. stlctosoma. Species that possess a 
constricted abdomen (i.e. mimicry of the hy- 
menopteran petiole - wasp mimics) consistently have 
either a single pouch or no pouch at all, suggesting that 
there is a trade off between wasp mimicry and the de- 
velopment of subabdominal pouches. Homoeocera 
stlctosoma, accordingly, lacks a restricted abdomen. 

Although the flocculent fibers are highly com- 
pressed within the pouches, when dispersed, they ap- 
pear as lightweight 'fluff that floats around the moth. . 
Upon examination with the scanning electron micro- 
scope, the flocculent material of H. stlctosoma is fila- 
mentous, composed of flattened, sculptured scales 
(Fig. 3). Within the Euchromiini, the morphology of 
flocculent varies considerably, from ribbon like scales 
to those that resemble twisted chains or even solid 
cvlinders (Sanderford 1992, Boada 1997, Conner etal. 
2000). Those described here for H. stlctosoma appear 
to be constructed for lightness and strength, and are 
described as being thin and ribbon like with a network 
pattern, and covered with a thin translucent film. 
They most closely resemble the flocculent of Gym- 
nelia salvinl Butler, Sarosa sp., Mijrmecopsls crabonis 
Druce, and Pseudophex polistes Hubn. (Boada 1997). 
In H. stictosoma, the flocculent also bears a rather 
strong phenolic odour when released. Mass spectra 
obtained showed strong ions of m/z 134 and 222, char- 
acteristic for a number of pyrrolizidine alkaloids (PAs), 
therefore suggesting the presence of PAs in the floccu- 
lent. The small amounts of flocculent available for 
analysis however, did not yield enough material for 
further mass spectroscopic or NMR spectrometric in- 



Volume 5S, Number 3 



175 



A 




Fig. 2. Schematic representations, based on camera lucida drawings, of the ventral (A) and left lateral (B) views 
of the abdomen of a male H. stictosoma. Flocculent material is densely packed within two subabdominal pouches, 
the openings of which are depicted by irregularly spaced short lines. Rectangular-shaped scales, interpreted as 
scent scales (SS) line the edges of the openings to the subabdominal pouches. The larger anterior pouch is defined 
by sternite II and is directed anterior!}', while the posterior pouch is directed anteriorly and posteriorly Scale bar: 
1 mm. 



vestigations to corroborate this hypothesis. 

Defensive secretions, including regurgitation, defe- 
cation, autohemoraging, and the expulsion of foams 
and sprays are commonly used among insects (Whit- 
man et al. 1990), but to our knowledge, reflexive dis- 
charge of 'scales' has not been documented. Our ob- 
servations of H. stictosoma indicate that flocculent is 
employed as a defense against spiders and vertebrate 
predators for the following reasons. 1. Flocculent is 
consistently discharged upon restraint of an individual. 
2. The flocculent scales adhere to the 'attacker' and 
have a distinct phenolic odour, suggesting a chemical 
defense. 3. Moths discharge the flocculent when col- 
liding with a spider web, and this is followed by their 
escape, suggesting that the flocculent coats the web so 
that the moth does not stick to it. The hypothesis that 
flocculent functions as an 'instant release' from spider 
webs and/or an 'anti-consumption' device should be 
validated experimentally by performing feeding trials 
with various vertebrate and invertebrate predators, 
and a detailed analysis of the flocculent chemistry. 
Also, the role that flocculent plays in the courtship be- 
haviour of H. stictosoma, and in the courtship and/or 
defensive behavior of the other 19 listed Homoeocera 
species (Druce 1881-1900; Draudt 1915) should be ex- 
plored. 

Despite the widespread occurrence of subabdomi- 



nal pouches in the Euchromiini, little is known of the 
taxonomic distribution or functional significance of 
these interesting structures and their constituent floc- 
culent material. To date, the use of flocculent in die 
context of courtship has been implicated in onlv 3 
species- C. myrodora (Conner et al. 2000), Syn- 
tomeida ipomoeae (Johnson 2002) and S. melanthus 
(Sanderford 1992). In C. myrodora, the flocculent 
may also function indirectly for defense, by protecting 
the 'adorned' female from spiders (Conner et al. 
2000). Homoeocera stictosoma is the only species re- 
ported to date that unequivocally discharges flocculent 
upon being restrained. Two other species, C. tcuthras 
and C. myrodora, have been suggested to use "fluff 
from the ventral valve" as part of a defensive beha\ior 
(Adams 1990), but no other details were prorided, and 
subsequent attempts to induce flocculent production 
in C. myrodora have proved unsuccessful (Conner et 
al. 2000). It is not possible at this time to know how 
these various traits evolved, due to our lack of under- 
standing of the functional distribution of flocculent 
material within the Euchromiini, and the phvlogenetic 
relationships among Euchromiini species. It is inter- 
esting to note that some Arctiidae extrude non-decid- 
uous, brightly colored cervical, genital or abdominal 
hair tufts upon being restrained, and it is believed diat 
the use of these structures in a defensive context de- 



176 



Journal of the Lepidopterists' Society 




Fig. 3. Filamentous 'scales' from flocculent material 
in a male H. stictosoma. A. Rounded tip of a filament. 
B. Middle portion of a filament. C. The bases of 
three filaments where they have broken off of the ab- 
dominal pouch. Scale bars: 10 pm. 

rives secondarily from a mating function (Blest, 1964). 
Similarly, one can envision that the defensive emission 
of flocculent, as we have proposed for H. stictosoma, 
evolved secondarily from a courtship function. 

We gratefully acknowledge Dr. R. Simmons at the 
USDA/ARS/PSI/Systematic Entomology Laboratory, 
Smithsonian Institution, Washington D.C. for assis- 
tance with specimen identification and for her valuable 
comments on the manuscript. This research was sup- 
ported by funding to T.E.Y. by National Science and 
Research Council of Canada, and the Smithsonian 
Tropical Research Institute. 



Literature Cited 

Adams, J.K. 1990. The defenses of adult tiger moths ( Lepidoptera: 
Arctiidae): Phylogenetic and ecological factors influencing the 
array of defenses in individual species. Ph.D. Dissertation. 
University of Kansas, Lawrence, Kansas. 

Beebe.W. & R. Kenedy. 1957. Habits, palatability, and mimicry in 
thirteen ctenuchid moth species from Trinidad, B.W.I.. Zoo- 
logica (New York) 42:147-158. 

Boada, R. 1997. Courtship and defense of the scarlet-bodied wasp 
moth Cosmosoma myrodora Dyar (Lepidoptera: Arctiidae) 
widi notes on related Euchromiines. M.Ss. Thesis. Wake For- 
est University, Winston-Salem, North Carolina. 

BLEST, A.D. 1964. Protective display and sound production in 
some new world arctiid and ctenuchid moths. Zoologica 
49:161-181. 

Chauthani, A.R. & P.S. Callahan. 1966. A dissection technique 
for studying internal anatomy of different stadia of Noctuidae. 
Ann. Ent. Soc. Amer. 59: 1017-1018. 

Conner, W.E. 1999. 'Un chant d'appel amoureux': Acoustic com- 
munication in moths. J. Exp. Biol. 202:1711-1723. 

Conner, W.E., R. Boada, F.C. Schroeder, A. Gonzalez, J. Mein- 
WALD & T. Eisner. 2000. Chemical defense: Bestowal of a 
nuptial alkaloidal garment by a male moth on its mate. Proc. 
Nat. Acad. Sci. 97:14406-14411. 

Draudt, M. 1915. Family Syntomidae. pp. 33-230. In Macrolepi- 
doptera of the World, Division II: The Macrolepidopteraofthe 
American Region. A. Seitz (Ed.) Volume 6. The American 
Bombyces and Sphinges. 

DRUCE, H. 1881-1900. In Biologia Centrali-Americana or Contribu- 
tions to the Knowledge of the Fauna and Flora of Mexico and 
Central America. Godman, F.D. and Salvin, O. (Eds.) (1879- 
1908). P.P. Calvert. London. Insecta. Lepidoptera-Heterocera. 
Vol. I (1881-1900), xxxi + 490 pp.; Vol. II (1891-1900) 622 pp.; 
Vol. Ill, 101 plates. 

Fullard, J.H. 1990. The sensory ecology of moths and bats: Global 
lessons in staying alive. In: Insect defenses. Adapative mechan- 
isms and strategies of prey and predators. Evans, D.L., and 
Schmidt, J.O. (Eds.) State University of New York Press, Albany 

NY. 

Jacobson, N.L. & S.J. Weller. 2002. A cladistic study of the tiger 
moth family Arctiidae (Noctuoidea) based on larval and adult 
morphology. Thomas Say Publications. Entomological Society 
of America, Lanham, MD. 

Johnson, R.M. 2002. Mimicry and courtship in the yellow-banded 
wasp moth Syntomeida ipomoeae Harris (Lepidoptera: Arcti- 
idae). MS Thesis. Wake Forest University. Winston-Salem, NC. 

PANTIN, C.F.A. 1946. Notes on the microscopical technique for zo- 
ologists. Cambridge: Cambridge University Press. 

Rothschild, M., R.T. Aplin, P.A. Cockrum, J.A. Edgar, P. Fair- 
weather & R. LEES 1979. Pyrrolizidine alkaloids in arctiid 
moths (Lep.) with a discussion on host plant relationships and 



Volume 58, Number 3 



177 



the role of these secondary plant : 
Biol. J. Linn. Soc. 12:305-326. 



lbstan 



the Arctikla 



Schmidt, J.O. (Eds.) State University of New York Press. Albany 
NY. 



RUBIO, F. P. & I.M. PesaNTEZ 1997. Mariposas del Ecuador. Vol. 1. 
Generos. Pontificia Universidad Catocila del Ecuador, Quito. 

Sanderford, M.V. 1992. Acoustic communication of tire polka-dot 
wasp moth, Syntomeida epilais Walker (Lepidoptera, Arctiidae, 
Ctenuchinae). Ph.D. dissertation, Wake-Forest University, 
Winston-Salem, NC. 

Weller, S.J., N.L. Jacobson & W.E. Conner 1999. The evolution 
of chemical defenses and mating systems in tiger moths (Lepi- 
doptera: Arctiidae). Biol. J. Linn. Soc. 68:557-578. 

Weller, S.J., R.B. Simmons, R. Boada & W.E. Conner. 2000. 
Abdominal modifications occurring in wasp mimics of the 
Ctenuchine-Euchromiine Clade (Lepidoptera: Arctiidae). 
Ann. Entomol. Soc. Am. 93:920-928. 

Whitman, D.W., M.S. Blum & D.W Alsop. 1990. Allomones: 
Chemicals for defense. In: Insect defenses. Adapative mechan- 
isms and strategies of prey and predators. Evans, D.L., and 



Jayne E. Yack, Tiffany A. Timbers (Department of 
Biology, Carleton University, Ottawa, Ontario, 
Canada, K1S 5B6; email: jyack@ccs. carleton. ca), 
William E. Conner {Department of Biology, Wake 
Forest University, Winston-Salem, North Carolina, 
USA, 27109; email: conner@wfu.edu), Annette 
AlELLO (Smithsonian Tropical Research Institute, Box 
2072, Balboa, Ancon, Republic of Panama; 
email:AIELLOA@tivoli.si.edu) and Frank C. 
Schroeder (Department of Chemistry and Chemical 
Biology, Cornell University, Ithaca, NY, USA; 
email:is31@come]l.edu) 

Received for publication 12 January 2004; revised 
and accepted 4 June 2004. 



178 



Journal of the Lepidopterists' Society 



Journal of the Lepidopterists' Society 
58(3), 2004, 178-182 

CATOCYCLOTIS AEMULIUS ADELINA (RIODINIDAE) REVISITED: IT AIN'T NECESSARILY SO 

Additional key words: genitalia, aUometric changes, Neotropies, Brazil. Costa Rica. 



Given their popularity 'with collectors and entomol- 
ogists, butterflies are considered one of the taxonomi- 
cally best-known groups of all insects. Nevertheless, 
revisiting apparently well known taxa with a modern 
eye may yield surprises. Consider the case here. The 
metalmark butterfly currently known as Catocyclotis 
aemulhis adelina (Butler, 1872) was originally de- 
scribed in the genus Lemonias Hubner, 1907, and con- 
sidered by Butler (1872) and Godman & Salvin (1879) 
to be a Central American species closely allied to 
South American L. aemulhis (Fabricius, 1793). Sub- 
sequently in the influential Genera Insectorum, 
Stichel (1911) downgraded adelina to a subspecies and 
placed both taxa together under his genus Catocyclotis 
Stichel, 1911, and maintained this arrangement m die 
Lepidopterorum Catalogus (Stichel 1930-1931). 
Thus, for over 90 years adelina has been regarded as a 
subspecies of the uncommon, yet broadly distributed 
species aemulius. Nevertheless, to paraphrase the fa- 
mous song from George Gershwin's folk opera Porgy 
& Bess, "... the things that you're liable, to read in the 
Bible, it ain't necessarily so". 

During a routine comparative study of male and fe- 
male genitalia we found conspicuous differences be- 
tween C. aemulius adelina and C. aemulius aemulius 
that correspond with noticeable differences in wing 
color pattern. These observations suggested that diese 
taxa represent two distinct species, as originallv pro- 
posed by Butler. The purpose of this study is to re- 
assess the status of taxa currently placed in the genus 
Catocyclotis, provide a diagnosis and illustrations of 
male and female genitalia of aemulius and adelina, and 
point to directions for future research. 

Catocyclotis aemulius (Fabricius, 1793) 

(Figs. 1-3) 

Type species of the genus Catocyclotis by original 
designation (Stichel 1911). 

Description: Male (Fig. 1) - Dorsal side, FW brown with a 
single-peaked, pale yellow mark distally on anal edge; brown areas of 
both FW and HW conspicuously marbled by thin light lines; HW 
extensively pale yellow (but less so than adelina), matching dorsal 
coloration of abdomen, and bearing marginal spots in cells Rs, Ml, 
M2, Cul and Cu2 (not always a full complement of spots is present); 
abdominal tergites 1-2 brown, 3-8 yellow. Ventral side, HW white 



and bearing marginal spots as on dorsal side. Female (Fig. 1) - wing 
pattern and color as in male; female abdomen dorsallv brown with 
scattered pale scales that vary in density. 

Genitalia: Male (Fig. 2) - seventh sternite with thin, elongated 
rami; aedeagus long with 11 spine-shaped cornuti; uncus with elon- 
gated marginal spines (longer than adelina); gnathos tip spatulate in 
ventral view; valva with two defined processes, dorsal process 
smoothly arched before tip in ventral view (humped in adelina), 
ventral process slightly projected, edge of ventral process with abun- 
dant, thickened setae (less abundant and thinner in adelina); sclero- 
tized transtilla broader than in adelina; saccus tip narrow in ventral 
view (narrower than adelina). Female (Fig. 3) - ostium bursa with 
a sclerotized point that reaches the edge of abdominal segment 8; 
antrum (defined here as the portion of the ductus bursa posterior to 
the ductus seminalis) elongated, sclerotized ventrally, wrinkled near 
ostium bursa, and with an anterior enlargement bearing internal 
clusters of spines; corpus bursa rounded (even in mated females) 
with symmetrically positioned signa. 

Distribution: Brazil, Ecuador? 

Material examined: Milwaukee Public Museum (MPM) - 
BRAZIL: 1 male, [Santa Catarina] Joinville, 8 June 1955; 1 male, 
S[anta] Catarina 13 May 1933; 1 male, Rio de Janeiro, Gavea, 21 
May 1956; 2 males, Rio [de Janeiro] 20 May 1934; 2 males, 1 dis- 
sected, Rio [de Janeiro] 15 May 1941; 1 male, Rio [de Janeiro] 15 
May 1941; 1 male, Rio [de Janeiro] 17 June 1931; 1 male, Estado do 
Rio [de Janeiro], Guapy 29 May 1940; 1 male, [Rio de Janeiro] 
Colony Guapy 13 May 1964; 1 male, dissected, [Rio de Janeiro] 
Petropolis 4 February 1962; 1 male, dissected, Barreira 18 October 
1955; 1 male, no data, acquired from P. Gagarin; 1 female, Estado 
do Rio [de Janeiro] Guapy 13 May 1940; 1 female, Estado do Rio 
[de Janeiro] Guapy 13 May 1940; 2 females, 1 dissected, [Rio de 
Janeiro] Mundo Novo 15 May 1940; 1 female, Rio [de Janeiro] 
Paineira[s] 22 May 1932; 1 female, [Rio de Janeiro] Petropolis 14 
August 1963; 1 female, [Rio de Janeiro] Petropolis 23 October 1965; 
1 female, [Rio de Janeiro] Petropolis 14 November 1963; 1 female, 
Rio [de Janeiro] 15 may 1941; 1 female Gavea, Rio [de Janeiro] 15 
July 1935. American Museum of Natural History (AMNH) - 
BRAZIL: 1 male, Rio de Janeiro 4 February [19]66; 1 male, Brazil, 
Rio [de Janeiro] 18 August [19]11; NO DATA: 1 female. 

Biology: Early stage biology unknown. This 
species is sexually monomorphic, and it inhabits forest 
areas between sea level (Xerem and Rio de Janeiro, 
both in Rio de Janeiro state, K. Brown pers. com.) and 
900 m (label data above). 

Remarks: Stichel (1911) and Seitz (1916) stated 
that aemulius occurred in 'south Brazil' (i.e., Rio de 



Volume 58, Number 3 



179 



Janeiro and surrounding areas) while adelina was dis- 
tributed from Costa Rica to Ecuador. Although we did 
not examine material from Ecuador, the male speci- 
men from Napo illustrated in D'Abrera (1994; note 
missing abdomen) shows a single-peaked yellow mark 
distally on the FW anal edge, brown areas of both FW 
and HW conspicuously marbled bv thin light lines, and 
H\V marginal dots - all traits of aemulius. Nonethe- 
less, in the D'Abrera (1994) illustration the HW color 
is orange as in adelina. This raises the questions of 
whether this represents true geographical variation, if 
the color is a printing artifact, or if the specimen be- 
longs to a different species. Although some of our 
Brazilian material is old and potentially faded, the 
AMNH collection includes a male specimen collected 
by K. S. Brown in Rio de Janeiro in 1966 that is virtu- 
ally identical in color to a male collected in the same 
locality in 1911. Therefore, based on D'Abrera (1994) 
we tentatively expand the previous notion of aemulius 
distribution to include Ecuador, but caution that the 
constancy of HW color should be verified with more 
Ecuadorian material. 

Catocijclotis adelina (Butler, 1872), revised status 

(Figs. 1-3) 

Description: Male (Fig. 1; color Illustration in DeVries 19S7, pi. 
18) - Dorsal side, FW brown with a double-peaked dark orange 
mark distally on anal edge; brown areas of FW and HW faintly mar- 
bled by thin light lines (conspicuously marbled in aemulius), HW 
not marbled in some specimens; HW extensively orange (more so 
than aemulius), matching dorsal coloration of abdomen, and lacking 
marginal spots; abdominal tergite 1 brown, 2-8 orange. Ventral side, 
HW white and normally lacking marginal spots, but in one specimen 
from Colombia small spots were present in cells M2, Cul and Cu2. 
Female (Fig. 1) - wing pattern similar to male, but HW orange 
color replaced by pale yellow (see DeVries 1987, pi. 18) or nearly 
white in worn individuals; female abdomen dorsally brown with scat- 
tered pale yellow scales that vary in density. 

Genitalia: Male (Fig. 2) - seventh stemite with short, broad 
rami; aedeagus short and lacking comuti; uncus with elongated mar- 
ginal spines (shorter than aemulius); gnathos tip narrow in ventral 
view; valva with two defined processes, dorsal process humped be- 
fore tip in ventral view (smoothly arched in aemulius), edge of ven- 
tral process with thin setae (abundant, thickened in aemulius); scle- 
rotized transtilla narrower than in aemulius; saccus tip in ventral 
view broader than adelina. Female (Fig. 3) - ostium bursa widi a 
sclerotized point that does not reach the edge of abdominal segment 
8; antrum (defined here as the portion of the ductus bursa posterior 
to the ductus seminalis) short, sclerotized ventrally, and broadened 
near ostium bursa; corpus bursa elongated with asymmetrically po- 
sitioned signa. 

Distribution: Costa Rica, Panama, Colombia. 



Material examined: DeVries Collection - COSTA RICA: 1 
male, dissected, Puntarenas, Las Alturas 24 May 1991; 1 male. 
Puntarenas, Las Alturas 25 May 1991; 2 males, Puntarenas, Las Al- 
turas 25 August 1991; 1 male, Moravia de Chirripo 16 April [19]83: 
1 female, dissected, [Puntarenas, Las Alturas] site IOC 22 Decem- 
ber [19]84. American Museum of Natural History (New York) - 
COSTA RICA: 1 male, Cairo 27 August [19]31; COLOMBIA: 1 
male, S.A., Felipe Ovalle. Q, no date; 1 male, dissected, Amazonas. 
Rio Cocoma 27 August 1946. 




aemulius female 




adelina female 



Fig. 1. Male and female habitus of Catoa/clotis aemulius and 
adelina, dorsal view on die left, ventral on the right. Arrows point to 
diagnostic characters mentioned in the text. Locality data from top 
to bottom: aemulius male, Brazil (acquired from P. Gagarin; 
aemulius female, Brazil, Rio de Janeiro, Guapv; adelina male, Costa 
Rica, Cairo; adelina female, Costa Rica, Puntarenas, Las .Alturas. 



180 



Journal of the Lepidoptekists' Society 




adelina 






aemulius 




adelina 



aemulius 



adelina 



Fig. 2. Male genitalia of Catocyclotis aemulius and adelina: in lateral view, genitalic capsule and detail of aedeagus; in ventral view, details of 
the uncus, gnathos, valvae, saccus, and seventli sternite. Valva on the left shows shape and distribution of setae, dense stippling was applied to 
valva on the right to show areas that are more heavily sclerotized. Locality data: C. aemulius, Brazil, Barreira; C. adelina, Costa Rica, Puntare- 
nas, Las Alturas. 



Volume 58, Number 3 



181 



aemulius 






adelina 



FlG. 3. Female genitalia of Catocijclot is aemulius and adelina: in lateral view, positioning of antrum+ductus bursa, ductus seminalis, and cor- 
pus bursa inside female abdomen, a detail of the ductus bursa of C. aemulius shows internal spines, setae show extent of sclerotization, ab- 
domens were dissected open and their outline does not represent actual abdominal width; in ventral view, ostium bursa and seventii stemite. 
papillae anales are represented schematically. Locality data: C. aemulius, Brazil, Rio de Janeiro, Mundo Novo; C. adelina, Costa Rica, Puntare- 
nas, Las Alturas. 



Biology: This species is reported to occur in forest 
areas between 800 and 1600 m (DeVries 1987). The 
sexes are dimorphic. The caterpillar of C. adelina and 
its natural history bear a strong similarity to Nymphid- 
ium (e.g., cachrus, hematostictum), and will be de- 
scribed in detail elsewhere (K. Nishida in prep.). The 
observation that adelina produces a clicking sound 
while in flight (D'Abrera 1994) is of particular interest, 
and should be verified. 

Remarks: The Costa Rican specimen studied by 
Penz & DeVries (1999) corresponds to adelina. 

Discussion. Although they have similar wing pat- 
terns, differences in wing markings were useful for 



separating aemulius and adelina. The shape of the 
FW anal marking (single- or double-peaked), together 
with the extent of yellow or orange in the H\Y plus ab- 
domen, and presence/absence of HW marginal spots, 
allow species determinations without dissection. 
These external color pattern characters can be easilv 
used to sort specimens in collections. 

The two Catoci/clotis species studied here showed 
dramatic differences in male and female genitalia (Fig. 
2 and 3). For example, the aedeagus of aemulius is 
much longer than that of adelina (Fig. 2), and corre- 
sponds to the longer female antrum+ductus bursa in 
aemulius (Fig. 3) - such correspondence has been ob- 



182 



Journal of the Lepidopterists' Society 



served among many other butterfly species (CMP un- 
published). Although aemulius and adelina are closely 
related, most parts of their genitalia differ, particularly 
with respect to allometric proportions. These observa- 
tions suggest to us that when Stichel (1911) considered 
adelina a subspecies of aemulius, he did so without 
comparing their genitalia. 

Two lines of inquiry suggest that Catocyclotis may 
include more than two species. Originally Stichel 
(1911) placed elpinice Godman, 1903 in Catocyclotis, 
but subsequently transferred it to his sentiformes sec- 
tion of what is now considered Adelotypa (Stichel 
1930-31). Of interest is that the specimen of Adelo- 
typa elpinice illustrated by D'Abrera (1994) bears 
strong phenotypic similarities to C. aemulius and 
adelina. While not conclusive, such observations indi- 
cate that elpinice may, upon closer study, be trans- 
ferred back to Stichel's (1911) home for it in Catocy- 
clotis. Secondly, Hall & Harvey (2002) suggested that 
Catocyclotis may include species currently classified in 
Mycastor Callaghan, 1983, but they did not allude to 
characters or taxa in support of their idea. Based on a 
cursory comparison ol C. aemulius and adelina with 
descriptions and illustrations of male genitalia of My- 
castor by Callaghan (1983) we offer the following ob- 
servations: (a) Catocyclotis and Mycastor have a scle- 
rotized transtilla between the valvae - a trait also 
present in Nymphidium (Penz & DeVries 1999 and in 
prep.); (b) M. leucarpis and Catocyclotis share a spiny 
uncus; (c) M. leucarpis and scurrilis seem to have a 
valva similar to that of Catocyclotis; (d) M. scurrilis has 
a saccus similar to Catocyclotis, and the seventh stem- 
ite resembles that of adelina. These characters lend 
support to the idea that Mycastor includes taxa better 
placed in Catocyclotis . However, it is evident that the 
total number of species embraced by Catocyclotis can 
only be verified through a comprehensive phyloge- 
netic analysis that includes many Nymphidiini genera 
and species. 



We thank Eric Quinter (American Museum of Natural History) 
for the loan of specimens under his care, K. Rozema for ideas on ri- 
odinid systematics, and anonymous reviewer for suggestions that im- 
proved the manuscript. Special thanks to Kenji Nishida for sharing 
his unpublished information on the biology of C. adelina. This study 
was supported in part by NSF-DEB 03-16505. This paper is dedi- 
cated to Adelina "Moreninha" Penz, George Gershwin, and Sportin' 
Life. 

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Butler, A.G. 1872. Descriptions of new butterflies from Costa 
Rica. Cistula Entomologica 1:72-90. 

Callaghan, C.J. 1983. A new genus of riodinid butterflies. Bull. Al- 
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D'ABRERA. B. 1994. Butterflies of the Neotropical region. Part VI, 
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Devries, P. J. 1997. Butterflies of Costa Rica and their natural his- 
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Godman, F.D. & O. Salvin. 1879-1901. Insecta, Lepidoptera, 
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Hall, J.P.W. & D.J. Harvey. 2002. Basal subtribes of the Nymphidi- 
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Hemming, F. 1967. The generic names of the butterflies and their 
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Hist.) Ent. Suppl. 9: 1-509. 

Penz CM. &: P.J. Devries. 1999. Higher level phylogeny of the 
tribe Lemoniini (Lepidoptera, Riodinidae): a preliminary as- 
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32. 

Seitz, A. E. 1916-20. Erycinidae. pp. 617-738. Macrolepidoptera of 
the World, vol. 5. Alfred Keman, Stuttgart. 

STICHEL, H. 1911. Riodinidae. Genera Insectorum P. Whytsman. 

Stichel, H. 1930-1931, Lepidopterorum Catalogus.Pars 40, 41, 44. 
W. Junk, Berlin. 

CARLA M. PENZ AND PHIL DEVRIES, Department of In- 
vertebrate Zoology and Center for Biodiversity Studies, Milwaukee 
Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 
53233, USA. Current address: Department of Biological Sciences, 
University of New Orleans, 2000 Lakeshore Drive, New Orleans, 
Louisiana 7014S, USA 



BOOK REVIEWS 



Journal of the Lepidopterists' Society 
58(3), 2004, 183-184 

The Geometrid Moths of Europe. Volume 4. 
Larentiinae II (Perizomini & Eupitheciini) by 
Vladimir Mironov. Apollo Books, Stenstrup, Denmark. 
463 pages, including 16 color plates. Hardback. 2003. 
ISBN 87-88757-40-4. Price: DKK 720,00. 

This is die second volume to appear in a series of six 
on the geometrid moths of Europe, a series that 
promises to provide a definitive, copiously illustrated 
foundation for future studies on the family. This vol- 
ume is of particular importance as it includes the most 
diverse but taxonomically most challenging genus in 
the fauna, Eupithecia Curtis. This genus contributes 
128 species (out of 1300 worldwide) to the European 
fauna. These make up about 85% of those treated in 
the volume, there being five more Eupitheciini and 18 
Perizomini. 

The author is eminently suited to produce such a 
work, having worked extensively on the Palaearctic 
fauna of the group, and provides an essentially eastern 
perspective, vital when so many of the generic centers 
of richness of the Palaearctic occur outside Europe. 
He divides Eupithecia into 35 species groups that are 
defined on genitalia characters. 

All the species are illustrated 1.5 times natural size 
in 16 color plates. Several specimens of each species 
are illustrated, particularly where variability is encoun- 
tered, and the text also includes half-tones that indi- 
cate key pattern characteristics distinguishing closely 
related species. The male and female genitalia are also 
illustrated by line drawings of very high clarity and 
quality. They have a distinctive style, but a colleague 
who also works extensively on the group testifies to 
dieir accuracy. In many instances the aedeagus vesica 
is shown everted. These illustrations are numbered 
with the species number in the main text, which makes 
it easy for the reader to track a given species through 
them, the only exceptions being the half-tones and ve- 
nation diagrams in the body of the text. The distribu- 
tion of each species is illustrated by a map within or 
close to the relevant text. This consists of spot localities 
within a more generally shaded range, though the 
spots also indicate outlying records. 

The text itself is comprehensive, with: synonymy; 
description of facies and genitalia; details of distribu- 
tion; phenology and biology, including host-plant 
records; records of parasitoids; habitat; particular diag- 
nostic features relative to similar species; remarks on 
taxonomic matters, particularly 25 newly established 
synonymies. 

There is a summary checklist that also notes 53 



species from areas adjacent to Europe that potentially 
could occur there, though these are not illustrated. 
The reference list is extensive. There is an index to sci- 
entific names occurring in the main text, though this 
does not include synonyms, only taxa recognised at 
specific or subspecific level, so readers confused as to 
the fate of taxa newly established as synonyms will 
need to refer to the Abstract on p. 8. This Abstract also 
provides a summary of species newly recorded in par- 
ticular European countries in the main text. The ex- 
tensive list of Acknowledgements is testimony to the 
thoroughness and industry on which the volume is 
based and to the wealth of information resources dis- 
tilled into it. 

One topic where I was hoping for enlightenment 
but was somewhat disappointed was the treatment of 
the higher classification. The Larentiinae are the one 
geometrid group where the greatest species richness 
and tribal diversity occurs in temperate latitudes. It is 
from these latitudes, therefore, that clear definition of 
higher taxa is likely to be established. A treatment in 
depth of a major component of the larentiine fauna at 
a continental level might be expected to offer some in- 
sight in this area. 

Xue & Scoble (2002), investigating the larentiine 
tribe Asthenini at a global level, found difficulty in dis- 
tinguishing this tribe from the Eupitheciini, a problem 
that had previously confronted this reviewer (Hol- 
loway, 1997: 120). This book notes in the introduction 
that the Perizomini are also closely related to the Eup- 
itheciini, and a genus is transferred to the Perizomini 
as it shows 'all the characteristic morphological fea- 
tures of the tribe 1 . In the 'Remarks' section of die 
tribal account, the Perizomini are distinguished from 
all other geometrids primarily by the presence of 
labides in the male genitalia. The corresponding ac- 
count of the Eupitheciini lacks 'Remarks' but notes in 
the description of the male genitalia that there are 
labides! Labides are also seen in the Asthenini (Hol- 
loway, 1997; Xue & Scoble, 2002: 79-80). The female 
in Perizomini is distinguished by a heavily sclerotised 
band arising from spurs from the anterior apophvses 
and encircling the antrum; this is not present as a com- 
plete ring in the Eupitheciini, though the spurs are 
present. Another feature that appears to distinguish 
the two tribes is presence in the Eupitheciini of modi- 
fication to the eighth stemite of the male. 

This prompts me to air a more general complaint 
about sections headed 'Diagnosis' in taxonomic publi- 
cations. I hope to find highlighted in such sections die 
diagnostic features that will enable me to distinguish 
the taxon concerned unambiguously from all odiers. 
either individually or in combination. However, I fre- 



184 



Journal of the Lepidopterists' Society 



quently find myself wading through a lengthy general 
description of features that I suspect are also widely 
distributed outside the taxon in question, yet are un- 
qualified by diagnostic remarks. In this particular 
book, the real diagnosis is usually found in the 'Re- 
marks' and 'Similar species' sections and in the text fig- 
ures; these are of sufficient quality to be truly diagnos- 
tic, the features indicated with 'Peterson' pointers, 
with the excellent illustrations of genitalia in support. 
The book is well designed and printed in clear type, 
and the color values of the plates appear accurate. I 
can fully recommend it, and expect it to be indispens- 
able in maximising accuracy in future identifications of 
European Eupitheciini. 

Literature Cited 

Holloway, J.D. (1997) The moths of Borneo: family Geometridae, 
subfamilies Sterrhinae and Larentiinae. Malayan Nature Jour- 
nal 51: 1-242. 

Xue, D. & SCOBLE, M.J. (2002) A review of the genera associated 
with the tribe Asthenini (Lepidoptera: Geometridae: Larenti- 
inae). Bull. nat. Hist. Mus. Lond. (Ent.). 71: 77-133. 

JEREMY D. HOLLOWAY, Department of Entomology, The 
Natural Histonj Museum, Cromwell Road, London SW7 5BD, U.K. 

Journal of the Lepidopterists' Society 
58(3). 2004, 184-185 

Las Mariposas de Machu Picchu. Guia ilustrada 
de las Mariposas del Santuario Historico Machu 
Picchu, Cuzco, Peru, by Gerardo Lamas. Published 
by the Fondo National para Areas Naturales Protegi- 
das por el Estado (PROFONANPE), Lima, Peru, [iv] 
+ 221 pages, 1 map, 569 color images (34 color plates). 
Softcover, glossy paper, 21.0 x 29.7 cm, 2003. ISBN 
9972-778- 10-X. Available for US $25.00 (not includ- 
ing postage), from PROFONANPE, Fondo National 
para Areas Protegidas por el Estado; Prolongation 
Arenales 722. Lima-18, Peru. Phone: (511) 212-1010; 
Fax: (511) 212-1957; e-mail: prf@profonanpe.org.pe; 
www.profonanpe.org.pe. 

Claiming over 3800 species of butterflies, Peru 
arguably hosts the greatest diversity of butterfly 
species of any nation on Earth (Lamas 2000). While 
this new book treats only a small fraction of Peru's 
species, it treats a fascinating cross-section of Peru's 
butterfly diversity, of mid- and high-elevation Andean 
species. 

Entirely in Spanish, the twelve-page introduction 
includes a brief historical review of natural history 
studies at Machu Picchu, including details of all 
previous lepidopterological expeditions to the area. A 
brief description of habitats found at Machu Picchu 
accompanies notes on butterfly morphology, diversity 
and biogeography in Cuzco and surrounding 



Departments. The introduction concludes with 
information on the book's color illustrations, various 
sources of related information available on the 
internet, and acknowledgments. 

Following the introduction, 69 pages are dedicated 
to detailed accounts for 194 genera and 377 butterfly 
species recorded at Machu Picchu. All taxon names 
include authorship and date of description, and 
taxonomy generally follows Lamas (in press). For each 
included genus, a brief summary of its total diversity 
and distribution is provided, and in a few cases, 
references to detailed revisions are included. For each 
species, notes on identification, distribution, habitats 
and foodplants at Machu Picchu are provided, along 
with citations to other publications where taxa have 
been cited as occurring at Machu Picchu. 

The first 6 color plates include 54 images of live 
butterflies and larvae (of two papilionids), taken in the 
wild by David H. Ahrenholz. The following 28 color 
plates include 488 images of pinned specimens, 
representing all 377 species currently recorded from 
Machu Picchu. For most species, only one image is 
provided, though a few species that display strong 
sexual dimorphism are represented by multiple 
images. Images are not life size, but each includes a 1 
cm scale bar to give an idea of true size. 

Following the color plates is a brief two-page 
bibliography and four appendices. The first appendix 
is dedicated to descriptions of new taxa. A total of 13 
new subspecies are described, in the Pieridae (2), 
Riodinidae (1), Satyrinae (3), Biblidinae (1), and 
Ithomiinae (6). The second appendix includes a list of 
all species recorded at Machu Picchu, indicating the 
distribution of each at 6 elevational intervals, from 
1500 m. to over 4000 m. elevation. The third appendix 
includes a list of 1373 butterfly taxa currently recorded 
from the Department of Cuzco, and highlights 278 
species that are expected to eventually be found at 
Machu Picchu. The fourth appendix provides 
specimen label data for each of the 488 illustrated 
pinned specimens. 

This is perhaps the first book to treat and illustrate 
the entire known butterfly fauna of any single South 
American locality. Due to its large size, it is not really 
a field guide, but nevertheless, this book belongs on 
the bookshelves of anyone interested in Neotropical 
butterflies, whether or not they have intentions of 
traveling to Machu Picchu or elsewhere in Peru. 
Some of die illustrated taxa are not illustrated 
elsewhere, and the excellent quality of the illustrations 
make the book visually quite appealing. Considering 
its low price, this book is a bargain; the 108 color 
images of correctly determined skippers, alone, make 



Volume 58, Number 3 



185 



the book well worth the cost. All information in the 
book is accurate, the taxonomy and nomenclature is 
up-to-date (though about mid-2002), and I was unable 
to find any errors. In conclusion, I strongly 
recommend this book. 

Literature Cited 

Lamas, G. 2000. Estado actual del conocimiento de la sistematica 
de los Iepidopteros, con especial referenda a la Region 
Neotropical, pp. 253-260, 3 tabs. In: Martin-Piera, F., J. J. Mor- 
rone & A. Melic (Eds.). Hacia un Proyecto CYTED para el In- 
ventario y Estimation de la Diversidad Entomologica en 
Iberoamerica: PrIBES 2000. Zaragoza, Sociedad Entomologica 
Aragonesa. 

Lamas, G. (Ed.). In press. Checklist: Part 4A. Papilionoidea - Hes- 
perioidea in: Heppner, J. B„ (Ed.). Atlas of Neotropical Lepi- 
doptera. Scientific Publishers, Gainesville, Florida. 

ANDREW D. WARREN, Department of Zoology, Oregon State 
University, Coixallis, OR 97331 USA. 

Journal of the Lepkfopterists' Society 
58(3), 2004, 1S5-1S6 

The Butterflies of Zambia, by Alan Heath, 
Michael A. Newport & David Hancock. 2002. Pub- 
lished by the African Butterfly Research Institute, 
Nairobi, Kenya, and The Lepidopterists' Society of 
Africa, xvii + 137 pages, 3 maps, CD-ROM (PC for- 
mat) with 2287 color images. Softcover, 20.8 x 29.1 
cm, ISBN 0-620-29211-3. Available from The Lepi- 
dopterists' Society of Africa for US $50.00, price in- 
cludes airmail shipping from South Africa. Send book 
order with shipping address, along with evidence of 
completed electronic fund transfer (EFT), to Dave L. 
McDermott, Public Relations Officer, dave@copy- 
wise.co.za. Direct EFT of US $50.00 to: NEDBANK, 
Johannesburg, South Africa, Florida Branch; Branch 
number 190-541; Account number 1095-032617, indi- 
cate "for Zambia book" as reference. 

How does a team of authors working with a limited 
budget go about publishing a tremendous volume of 
information on 839 species of tropical African 
butterflies, including multiple color illustrations of 
virtually every species? The authors of this book found 
a solution, using a CD-ROM for all illustrations. The 
book consists of a short introductory text, 137 pages of 
annotated checklist, a bibliography and index. The 
CD contains two folders. The "Zambia images" folder 
includes 2287 color .jpg images of museum specimens 
of Zambian butterflies, arranged alphabetically. The 
"Zambia labels" folder contains a .pdf file giving label 
data for each specimen. 

The 17 introductory pages (pp. i-xvii) provide 
general information on recent books treating the 
butterfly fauna of other southern African nations, the 



underlying philosophy of the authors towards the 
presentation of the book, and a brief explanation of the 
classification and nomenclature used in the book. 
Several pages discuss Zambia's topography, climate, 13 
major different vegetation types, and a brief political 
history of the nation. The introduction also includes 
information on butterfly conservation, the history of 
butterfly collecting in Zambia, extensive details on the 
annotated checklist over its 26 year development, a list 
of die collections consulted, a list of abbreviations used 
in the checklist that indicate other published sources 
with illustrations of treated species, and 
acknowledgments. A brief summary of taxonomic 
changes made in the annotated checklist is provided, 
however, discussion of these is provided in the 
checklist itself. Revised status is proposed for 12 taxa, 
and one new species of Charaxes Ochsenheimer is 
described by S. F. Henning. The introductory pages 
conclude with a three-page gazetteer of Zambian 
localities, and three maps of Zambia. 

The 122-page annotated checklist fills the majoritv 
of the book, and treats all species recorded from 
Zambia in detail. A brief summary is given for each 
family subfamily and genus that occurs in Zambia, 
including the authorship for each taxon, and in many 
cases, brief taxonomic discussions. Below each 
generic summary, Zambian species and subspecies are 
listed, again, with authorship indicated for everv taxon. 
Below each listed species or subspecies are 
abbreviations indicating other works in which that 
taxon has been illustrated, along with a diagnosis 
listing important characters for identification. For 
each taxon, a summary of distributional and 
phenological data is provided, known foodplants are 
listed, and for some taxa, a lengthy discussion of 
variation or tavonomic issues is included. Two pages 
are dedicated to a discussion of unsubstantiated 
records. An extensive bibliography of over 100 
sources, and 12 pages of index are given at the end of 
the checklist. 

The CD-ROM is in a jacket attached to die inside of 
the back cover, and can be read only by a PC, not by 
Macintosh machines. For many species, images of 
dorsal and ventral surfaces of one specimen are 
provided, especially when those species display little 
or no sexual or seasonal variation. For many species, 
dorsal and ventral surfaces of male and female 
specimens are illustrated. For some species, seasonal 
or geographic variation is illustrated in detail, with up 
to a dozen images. Images are high quality and show 
well-prepared specimens. 

The butterfly images and list of specimen data can 
be printed, and images can be copied and arranged 



186 



Journal of the Lepidopterists' Society 



into plates by the user if so desired, for side-by-side 
use with the text, away from a computer. I did this, 
and was able to fit 56 images on each printed page, for 
a total of 43 color plates. I made a short legend for 
each plate, placed a printout with the data for the 
images (as provided on the CD) at the end of my 
plates, and had them bound together at the corner 
copy store. 

While it did require some time to examine and 
arrange all the images on the CD, it was a useful crash- 
course in Zambian butterfly diversity and 
identification. I quickly learned that there are an 
amazing number of confusingly similar Zambian 
species of Acraea Fabricius (74 species), Bicyclus 
Kirby (21 species), Neptis Fabricius (20 species), 
Charaxes (50 species), lolaus Hiibner (32 species), 
Deudorix Hewitson (21 species), Anthene Doubleday 
(23 species) and Lepidochnjsops Hedicke (28 species). 
I found few errors in labeling of the images, and a few 
instances of what appear to be "duplicate" images on 
the CD; for example the dorsal and ventral images of 
"male" Xanthodisca vibius (Hewitson), Catopsilia 
florella (Fabricius) and Neocoenijra cooksoni Druce 
are actually duplicates of the female images of those 
species. However, considering the total of 2287 
images on the CD, there are remarkably few mistakes, 
and overall, the library of images on the CD is 
extraordinarily useful. 

This book contains a tremendous wealth of 
information on the identification, distribution, biology, 
taxonomy and nomenclature of Zambian butterflies in 
particular, and of central and southern African 
butterflies in general. Anyone interested in these 
subjects will want to own this volume, as will anyone 
interested in the natural history of southern Africa. 

ANDREW D. WARREN, Department of Zoology, Oregon State 
University, Corvallis, OR 97331 USA 



Date of Issue (Vol. 58, No. 3): 27 October 2004 



EDITORIAL STAFF OF THE JOURNAL 

Michael E. Toliveh, Editor 
Department of Biology 
Eureka College 
Eureka, Illinois 61530 USA 
miketol@eureka.edu 
Brian Scholtens, Assistant Editor Phil DeVries, Book Review Editor 

Biology Department Department of Biological Sciences 

College of Charleston, 66 College Street, University' of New Orleans 

Charleston, South Carolina 29424-0011, USA New Orleans, Louisiana 70148 USA 

scholtensb@cofc,edu pdevries@UNO.edu 

Associate Editors: 

Gerardo Lamas (Peru), Kenelm W. Philip (USA), Rorert K. Robbins (USA), Felix Sperling (Canada), 

David L. Wacner (USA), Christer Wiklund (Sweden), Carla Penz (USA), Andrew Warren (USA), 

NOTICE TO CONTRIBUTORS 

Contributions to the Journal may deal widi any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes. Techni- 
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Sheppard, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp. 
1961a. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10:165-216. 

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CONTENTS 

Papilionoidea of the Evergreen Tropical Forests of Mexico Jose L. Salinas-Gutierrez, 

Armando Luis-Martinez and Jorge Llorente-Bousquets 125 

Histoire Generale et Iconographie des Lepwopteres et des Chenilles de l'Amerique 
Septentrionale by Boisduval & Le Conte (1829-[1837]): Original Drawings used for the 
Engraved Plates and the True Identities of Four Figured Taxa John V. Calhoun 143 

Technical Comment 

a critical response to the paper "tough african models and weak mimics: new horizons 

IN THE EVOLUTION OF BAD TASTE" BY P. DeVrIES PUBLISHED IN THIS JOURNAL, VOL. 57(3), 2003 

Luka Kassarov 169 

General Notes 

Defensive Flocculent Emissions in a Tiger Moth, Homoeocera Stictosoma 
(Arctiidae:Arctiinae) Jayne E. Jack, Tiffany A. Timbers, William E. Conner, Annette 
Aiello and Frank C. Schroeder 173 

Catocyclotis Aemulius Adelina (Riodinidae) Revisited: it Ain't Necessarily so Carla 
Penz and Phil DeVries 178 

Book Reviews 

The Geometrid Moths of Europe. Volume 4. Larentiinae II (Perizomini & Eupitheciini) 
Jeremy D. Holloivay 183 

Las Mariposas de Machu Picchu. Guia ilustrada de las Mariposas del Santuari HistOrico 
Machu Picchu, Cuzco, Peru Andrew D. Warren 184 

The Butterflies of Zambia Andrew D. Warren 185 




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