Perrycollins, pdf006
Perrycollins, pdf006
BUTTERFLIES OF PEDERNALES,
MICHOACAN, MEXICO,
WITH NOTES ON SEASONALITY
AND FAUNISTIC AFFINITIES
(LEPIDOPTERA: PAPILIONOIDEA AND
HESPERIOIDEA)
MANUEL A. BALCAZAR L.
Florida State Collection of Arthropods,
Division of Plant Industry, FDACS, P. O. Box 147100, Gainesville, Florida 32614, USA
ABSTRACT.- A list of 205 species of butterflies and skippers is presented for Pedernales, Michoacan, in western Mexico. The fauna belongs to five
families: Nymphalidae with 31% of the total number of species, Hesperiidae with 28%, Lycaenidae 24%, Pieridae 11% and Papilionidae 7%. A low
species richness and a high species richness seasonally, which seem to be related to the dry and humid seasons, respectively, was found. The fauna
in the area has a clear affinity to that of the Pacific Slope and especially to other localities of the Balsas Basin.
RESL'MEN.- El presente trabajo es el primer listado lepidopterofaunistico para una localidad en el estado de Michoacan basado en recolecciones
sistematicas. Un total de 205 especies son registradas por primera vez para Pedernales, Michoacan. Aunque la lista no es completa, se considera que
continen alrededor de un 80-85% del total de especies presente en el area. Del total de especies, un 31% pertenece a la familia Nymphalidae, 28%
a Hesperiidae, 24% a Lycaenidae, 11% a Pieridae y 7% a Papilionidae, composicion que concuerda con datos reportados para regiones neotropicales.
Durante 1986, se encontraron dos epocas bien claras en cuanto a su riqueza especifica con base en la presencia de imagos. una baja que va de enero
a junio, y otra alta de Julio a diciembre. Al comparar el numero total de especies presentes por mes contra el diagrama ombrotermico, se encontro
relacion entre la epoca humeda y alta riqueza especifica y entre la epoca seca y baja riqueza. Si bien, esta correspondencia no es exacta, ya que los
dos perfodos de riqueza se encuentran desplazados ligeramente a la derecha de sus correspondientes epocas de humedad. Los meses de mayor riqueza
son octubre y septiembre, al final de la epoca humeda y el de menor riqueza es abril, justo despues del mes mas seco. Un analisis de afinidades con
otras faunas utilizando el mdice de Simpson y metodos feneticos dio como resultado dos fenogramas, uno para cada superfamilia, el de Papilionoidea
con un mdice de correlacion cofenetica muy pobre (r = 0.65) y el de Hesperioidea pobre (r = 0.77). Se encontraron dos conglomerados de localidades
para la Papilionoidea, uno formado por localidades de la vertiente del Golfo y el Soconusco, Chiapas, y otro unicamente de localidades de la vertiente
del Pacifico. El Valle de Mexico no quedo incluido en ninglin conglomerado. En el caso de Hesperioidea, solo se observa el conglomerado de las
localidades del Pacifico. En ambos fenogramas, Pedernales aparece mas cercano a Rancho Viejo-Tepoztlan, Morelos que a ninguna otra localidad.
KEY WORDS: Anacardiaceae, Burseraceae, Caribbean, distribution. Ebenaceae, fauna, Gramineae, Hesperiidae, Leguminosae, Lycaenidae,
Mesoamerica, Moraceae, Neotropical, Nymphalidae. Papilionidae, Pieridae, Rubiaceae, Rutaceae, Sapindaceae, Sapotaceae. taxonomy, zoogeography.
Regional lists of butterflies are of great importance, because 1976), for which only one paper has been published (de la Maza,
they provide basic information to taxonomists, zoogeographers 1975. 1976). Another aim of this study was to conduct a compar-
and ecologists showing when and where to look for certain taxa. ative analysis of the butterfly fauna of Pedernales with other
Unfortunately, this sort of work is not often accomplished Mexican localities. Finally, it is intended that this paper serve as
(Clench, 1979). There are just a few Lepidoptera papers dealing a baseline for studying future changes of the faunistic composi-
with the fauna of western Mexico, and of these, only two deal tion in the area.
with localities within Michoacan State. The first papers were by
Brown (1943; 1944a. b. c; 1945), based upon material collected STUDY AREA
during the second and third expeditions of Hoogstraal in 1939 and The area studied in this paper is a narrow strip located in
1940, where 60 species of Papilionoidea were reported. The central Michoacan, in the south end of "Municipio de Tacambaro"
second paper, by Mather (1967), reports 31 species of Hesperioi- (Fig. 1), 19°07'30" - 19°10'00" north latitude and 101°26'57" -
dea. The object of this study was to examine butterfly species 101°28'25" west longitude. Due to its location on the south slope
richness in a Michoacan locality resulting from the first study of the Volcanic Transversal Axis, the elevation varies from
based upon systematic collections. In addition to the importance 1100m in the north to 850m in the south. From a floristic stand-
of having a regional list, the study area itself is important because point, it is located in the Caribbean Region, within the Balsas
it is located in the Balsas River Basin, a very important district Depression province (Rzedowski, 1978) and faunistically in the
from a zoogeographic standpoint (Hoffmann, 1940; Halffter, Nayarit-Guerrero province, according to Alvarez and Lachica
94 BALCAZAR: Michoacan Butterflies TROPICAL LEPIDOPTERA
Temperature Precipitation
170
-•-Temperature
+ Precipitation
ESNumber of species
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Month
Fig. 2. Climate of Pedernales and total number of species found per month.
(CETENAL, 1970. Station 16-010).
METHODS
Monthly visits were performed from February 1986 to January
1987, in addition to previous visits in 1984 and 1985, that in total
amounted to 42 collecting days. Specimens were collected with
Fig. 1. Localization of study area (modified from Correa [1979:41])
aerial nets, and by traps with rotten fruit (as described by Rydon,
1964). Part of the material was conserved in glassine envelopes
(1974). The weather is warm subhumid. with the coldest month in tin-coated boxes, and the rest was mounted with entomological
having an average temperature over 18°C. Summer rains are pins following Beutelspacher's (1983b) directions.
predominant, with winter rains comprising less than 5% of the Identifications were by comparison with material of the
annual total, resulting in a quotient precipitation/temperature Coleccion Entomologica del Institute de Biologfa (UNAM,
between 43.2 and 55.3. The annual fluctuation of average Mexico) and with the works of Beutelspacher (1984). Comstock
temperatures is 5 to 7°C and there is a Ganges temperature march (1961), Godman and Salvin (1878-1901), Higgins (1981), Jenkins
(CETENAL, 1970). A dry period from November to May and a (1983), McAlpine (1971), L. Miller (1974) and Seitz (1924).
wet period from June to October are observed in the ombrotermic Genitalic preparations were made following Hardwick (1950) and
diagram (Fig. 2). Seven soil types are present in the area, being Borror et al. (1981).
predominant cromic vertisol, ranker, ocric andosol and litosol, the Due to the lack of a comprehensive systematic arrangement for
textural classes are fine and medium. Despite the destruction of the Neotropical butterflies, several sources were used: Ackery
the original vegetation in the great majority of the zone, two (1984), Ackery and Vane-Wright (1984), Beutelspacher (1984),
kinds of vegetation types can be distinguished (sensu Rzedowski, DeVries et al. (1985), Hodges et al. (1983), Hoffmann (1940,
1978): subdeciduous tropical forest and deciduous tropical forest 1941), Kristensen (1975), J. Miller (1987), L. Miller (1968), and
(Fig. 3). The former is restricted to the humid ravines, and some Scott (1985).
of the typical trees are Enterolobium cyclocarpum (Jackg.) For the comparative faunal study, Simpson's similarity index
"parota", Pithecellobium dulce Roxb. "pinzan", Inga spuria was used; and for the cluster analysis, the unweighted pair-group
"jinicuil" (Legummosa.e),Masticodendron capiri "capiri" (Sapota- method using arithmetic averages (UPGMA) was employed. It
ceae), Thouindium decandrum (H. et B.) "charapo" (Sapindaceae) was intended that for the faunal comparison all the available
and Ficus spp. "sirandas" (Moraceae). The deciduous tropical published papers up to 1990 were to be used [Soconusco, Chiapas
forest occupies the rest of the area except for the agricultural (Hoffmann, 1933); Los Tuxtlas, Veracruz (Ross, 1967); Las
lands; the most striking trees of this vegetation belong to the Minas, Veracruz (Beutelspacher B., 1975), Rancho Viejo-
genera Acacia (Leguminosae) and Bursera ("copales") (Bursera- Tepoztlan, Morelos (de la Maza, 1975, 1976); El Valle de
ceae). In the flat lands is grown "sugar cane" (Saccharum Mexico; Chamela, Jalisco; El Chorreadero, Chiapas (Beutel-
officinarum) (Gramineae), and in small plots fruit trees such as spacher, 1980, 1981, 1983a) and Teocelo, Veracruz (Llorente el
"mango" (Mangifera sp., Anacardiaceae), "banana" (Musa sp., a/., 1986). More recent faunistic papers (de la Maza, 1988; de la
Musaceae). "black zapote" (Diospyros ebanaster, Ebenaceae), Maza et al., 1989; Luis et al., 1991; Vargas et al., 1991) will be
"coffee" (Coffea sp., Rubiaceae) and several citrus trees (Citrus included in a more comprehensive phenetic study of faunal
spp., Rutaceae). relationships (Balcazar, in prep.).
Vol. 4 No. 2 1993 BALCAZAR: Michoacan Butterflies 95
The results are based on more than 1850 collected and checked
specimens. The months when adult specimens of the species were 200
collected, and their number, are pointed out in the list.
Clench (1979) stated that a regional list, like the one presented 150 -
here, will never be complete, but as the number of collecting
man-hours in the study area increases, the closer to completeness
100 -
the list becomes. Unfortunately, there is no accurate record of this
parameter, and the total theoretical amount of species could not
50 -
be calculated with Clench's formula (Clench. 1979). However, in
Fig. 4 there is a clear decrease of the curve slope for the last
collecting months, so one can suppose it is close to the asymp- Feb Jun Aug Sep Oct Nov Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan
tote. Based on this, it is considered that the list includes about 80-
Field trips
85% of the total number of species present in the area. 1984 1985 1986 1987
The family with the greatest number of species in the area was Fig. 4. Cumulative number of species collected in the study area.
Nymphalidae. with 31% of the total, followed by Hesperiidae
with 28%, Lycaenidae 24%, Pieridae 11%, and Papilionidae 7%. reported number of species can be considered representative of
These data agree with those of Heppner (1987) for these families their respective areas. Out of the localities of the Gulf Slope
in the Neotropical region, with the exception that Hesperiidae and [which is richer in species numbers. (Hoffmann, 1940)], Peder-
Lycaenidae exchange positions. The numbers of species reported nales is the third richest locality, surpassed only by "Rancho
for other localities in faunistic studies like the present are shown Viejo-Tepoztlan", Morelos, and "El Soconusco", Chiapas, which
in Table 1. Although these studies are very heterogeneous in represent larger areas and include more vegetation associations.
techniques, duration and collecting effort, as well as the study It is interesting to show that in Pedernales, an area of approxi-
area size, elevation and number of vegetation associations, the mately 12km2, a Papilionoidea number is found equivalent to 30%
•1
Total taxa:
Shared taxa:
HESPERIOIDEA
Total taxa:
Shared taxa:
Fed. R.V. Cha. Cho. V.M. Min. Soc. Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov D
Fed. Month
R.V. 39 • Papilionidae SSPieridae SNymphalidae CDLycaenidae •Hesperiidae
Cha. 23 27 ...
Cho. 17 20 13 ___
V.M. 15 26 8 5 Fig. 6. Areas diagram showing the behavior of families species richness along
Min. 13 17 9 5 12 year in Pedernales.
Soc. 11 13 9 7 7 8
a high richness season from July to December (Fig. 2). There
of the total known for the US and Canada (Miller and Brown. a relationship between the wet season with high species richne
1981), and that almost all the species can be found in the "Arroyo and between the dry season and low richness, but this relatior
Frio" ravine, which is less than 500m long. not exact, since both richness periods lag behind precipitat
It is very difficult to get random non-selective samples for periods. The months with higher richness are October £
butterflies that do not favor one species over another and bias the September, at the end of the humid period. The month with lo\
measure of relative abundance (Shapiro, 1975). This becomes
clear from the number of collected and observed specimens, a followed in general by Papilionoidea. For all the families the h
commonly reported phenomenon in the tropics (Owen, 1971; richness periods start just after the beginning of the wet seasor
Lamas, 1981). There is a great number of species represented by July, and reach their higher richness peaks at the end (Pieridae
only one specimen, while just for a few species many specimens
were collected. Out of 250 species, 67 were represented in the
Fig. 7. SATYRINAE: 1. "Euptychia" polyphemus cyclops (Butler): ITHOP
sample by just one specimen and 78% of the species are repre-
NAE: 2. Greta morgane (Geyer); NYMPHALINAE: 3. Chlosyne riobalse
sented by 10 or fewer specimens. The most abundant species Bauer: 4. Pyrrhogyra nearea hypsenoer (Godman & Salvin): 5. Hamadi
were Eurema daira (Godart) (Pieridae), with 139 specimens, amphinome mexicana (Lucas); 6. Prepona demophoort pallantias Fruhstoi
followed by Chlosyne lacinia (Geyer) (Nymphalidae), with 69 THECLINAE: 7. Pseudoelycaena damo (Druce): 8. Panthiades battus ji
(Fig. 6). (Reakirt): 9. P. meton (Cramer); 10. Oenomaus ortygnus lauta (Draudt);
"Thecla" mycon Godman & Salvin; 12. Ocaria ocricia (Hewitson); 13. "Thet
tephraeus (Geyer): 14. "Thecla" syedra Hewitson; 15. Ipidecla miadora D
SEASONALITY RIODININAE: 16. Rhetus arcius beutelspacherii Llorente; 17. Calept.
Two clear cut seasons were found in Pedernales, based on perditalis (Barnes & McDunnough): 18. Baeotis hisbon zonata Felder;
species [adult] richness: a low season from January to June, and Melanis pixe (Boisduval).
98 BALCAZAR: Michoacan Butterflies TROPICAL LEPIDOPTERji
35
31
Fig. 8. RIODININAE: 20. Emesis mandana (Cramer): 21. Lasaia sessilis Schaus; 22. Theope pedias isia Godman & Salvin: PYRRHOPYGINAE: 23. Pyrrhopyg
chalybea chalybea Scudder: PYRGINAE: 24. Astraptes fulgerator (Walsch): 25. Typhedanus ampyx (Godman & Salvin); 26. Nascus phocus (Cramer): 27. Pholisoi
mejicanus (Reakirt); HETEROPTERIKAE: 28.Dallafaulafaula (Godman); 29. D. bubobon Dyar: HESPERIINAE: 30. Mellana helva (Moschler); 31. Vidiusperigem
(Godman); 32. Nyctelius nyctelius nyctelius (Latreille); 33. Vettius fantasos (Stoll); PYRGINAE: 34. Autochton neis (Geyer): HESPERIINAE: 35. Atrytonopsis det
(Edwards).
September. Nymphalidae and Lycaenidae in October), with only tially governed in the tropics by an alternation between rainy an
one exception at the beginning of the dry season (Papilionidae in dry seasons (Emmel and Leek, 1969; Owen, 1971; Austin, 197S
November). After the two first months of the dry season, the four Wolda, 1988), as opposed to temperate regions where shar
families show a clear decrease in species richness. January is the differences in temperature and photoperiod characterize th
month with lowest richness for both Pieridae and Papilionidae, seasonality (Shapiro, 1975). The findings cannot be considere
while for Lycaenidae the richness continues decreasing until conclusive due to the short length of the study, since accordin
April. For Nymphalidae there are two low points in January and to Wolda (1988) this is only possible if the pattern is repeats
April. Hesperioidea (Hesperiidae) show the lowest richness in year by year.
April and May, while September is the month with highest Apparently about 55% of the species were bi- or multivoltin
richness (Fig. 7) for that family. in 1986, which is a common phenomenon in the tropics (Woldi
These findings agree with the general idea that the life cycle of 1988). It was not possible to establish if the species collected <|
the butterflies, and consequently their species richness, is essen- observed on just a few opportunities were resident or migrant; bi
Vol. 4 No. 2 1993 BALCAZAR: Michoacan Butterflies 99
Hesperiidae which represent 7. 11, 31, 24 and 28% respectively Borror, D. J., D. M. Delong, and C. A. Triplehorn
of the total number of species. These data agree with what has 1981. An introduction to the study of insects. 5th ed. Philadelpljj
been reported for the Neotropical region. A low species richness Saunders Coll. Publ. 827pp.
season from January to June, and a high richness season from Brown, F. M.
July to December, were found, which seem to be related to the 1943. Notes on Mexican butterflies I. Papilionidae. /. New Yo
Ent. Soc. ,51:161-178.
humid and dry seasons, respectively. A few species are illustrat-
1944a. Notes on Mexican butterflies II. Pieridae. /. New York A
ed in Fig. 7-8. Soc., 52:99-119.
After the faunal similitude cluster analysis (Fig. 9), just one 1944b. Notes on Mexican butterflies III. Danaidae. /. New Yo
phenogram was found for each superfamily faunal lists, that for Ent. Soc. , 52:237-246.
Papilionoidea with a very poor co-phenetic correlation index (r = 1944c. Notes on Mexican butterflies IV. /. New York Ent. 5<J
0.65) and poor for Hesperioidea (r = 0.77). Two clusters were 52:343-360.
found for Papilionoidea, the first one constituted by localities of 1945. Notes on Mexican butterflies V. /. New York Ent. Soc,
the Gulf slope, and El Soconusco; and the second just by 53:31-46.
localities of the Pacific Slope; Valle de Mexico does not cluster Clench, H.
with any of the groups. Hesperioidea localities phenogram only 1968. Butterflies from Coahuila, Mexico. /. Lepid. Soc. (
shows one cluster for Pacific localities. In both cases, Pedernales
is closer to Rancho Viejo-Tepoztlan than to any other locality. 1979.
Angeles), 22:227-231.
How to make regional lists of butterflies: some thoughts.
Lepid. Soc. (Los Angeles), 33:215-231.
1
Comision de Estudios del Territorio Nacional (CETENAL).
ACKNOWLEDGMENTS 1970. Carta de Climas, Mexico 14Q-V. Mexico City: Secreta
de la Presidencia. Scale 1:50,000.
I wish to thank Dr. C. Beutelspacher, Institito de Biologia, Comstock, W. P.
Universidad Nacional Autonoma de Mexico (IBUNAM, Mexico 1961. Butterflies of the American tropics: The genus Anaea.
City), for his kind advice and suggestions during the achievement York: Amer. Mus. Nat. Hist. 214pp.
of this study. I am grateful to J. B. Heppner, Florida State Correa, P. G.
Collection of Arthropods (FSCA, Gainesville, FL), and T. C. 1979. Atlas geogrdfico del Estado de Michoacan. Mexico Ci
Emmel, Dept. of Zoology, University of Florida (Gainesville, FL), EDDISA. 92pp.
for critically reading the manuscript. DeVries, P. J., I. J. Kitching, and R. I. Vane-Wright
1985. The systematic position of Antirrhea and Caerois. wil
comments on the higher classification of the N\i
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Vol. 4 No. 2 1993 BALCAZAR: Michoacan Butterflies 101