A Survey of The Butterfly Fauna of Jatun Sacha, Ecuador (Lepidoptera: Hesperioidea and Papilionoidea)
A Survey of The Butterfly Fauna of Jatun Sacha, Ecuador (Lepidoptera: Hesperioidea and Papilionoidea)
A Survey of The Butterfly Fauna of Jatun Sacha, Ecuador (Lepidoptera: Hesperioidea and Papilionoidea)
Debra L. Murray
Abstract. The first extensive butterfly survey of the upper Río Napo ba-
sin in eastern Ecuador was conducted from 1990 to 1993. A total of 811
species was recorded at Jatun Sacha Biological Reserve. Based on species
richness comparisons with a similar site in southern Peru and extrapola-
tions from ithomiine diversity, Jatun Sacha is estimated to have approxi-
mately 1300 species of butterflies. Species richness is compared with two
other Amazonian sites (Pakitza, Peru, 1300 species and Cacaulandia, Bra-
zil, 843 species). Species and generic compositions are more similar be-
tween Pakitza and Jatun Sacha than Cacaulandia. This similarity may be
due to environmental factors. A greater percentage of Nymphalidae and
a lower percentage of Hesperiidae and Lycaenidae occur at the two some-
what disturbed sites (Jatun Sacha and Cacaulandia) than the less disturbed
site (Pakitza). Of the 228 species common to all three sites, more nympha-
lid butterfly species were found than expected based on observed spe-
cies in each family.
INTRODUCTION
The Amazon basin covers an area approximately 6 million square kilo-
meters and houses the world’s greatest diversity of plant and animal life
(Erwin 1988, Dinerstein et al. 1995). Insects are the most diverse taxon in
the neotropics, yet they have been poorly studied in this vast area (National
Academy 1992, Lamas 1989 and ref. therein, Raven 1988, Reid & Miller
1989). Even for taxonomically well known insect groups, such as the but-
terflies, there exist large gaps in our understanding of tropical species rich-
ness and factors influencing diversity (DeVries 1994, Ackery 1986). One
major hindrance is the lack of basic information available on natural his-
tory and species distributions for most Amazonian butterflies (Ackery 1986,
DeVries 1994, DeVries et al. 1997). Inventories from specific localities can
be useful in investigating changes in species compositions across landscapes,
but most of the current faunal information on Amazonian butterfly com-
munities are from Peru (Lamas 1985, 1989, Robbins et al. 1996) and areas
in Brazil (Brown 1984, 1991, Emmel & Austin 1990, Mielke 1994). There
are few published surveys of butterfly faunas in eastern Ecuador and Co-
STUDY SITES
Jatun Sacha Biological Station is located 30 km east of the base of the
Andes (01° 04’S; 77° 36’W) and lies between the confluence of the Napo
and Arahuno rivers, its natural boundaries. Elevation varies from 400m to
450m. The uplands, typified by steep, low hills and narrow ridges with small
streams in the valleys, comprise the majority of the land. There is also a
small tract (100 hectares) in the Rio Napo floodplain with alluvial soils and
seasonal flooding. The Holdridge system would classify the lowland forests
of this area as Tropical Wet Forest (Cañadas 1983). Rainfall data, recorded
since 1986, averages 3700mm annually, with no definite dry season. How-
ever, April through July are generally the wettest months and December
through February the driest months. Major floods of streams and rivers
occurs throughout the year but are more common during the wetter
months. Soil fertility is relatively rich for tropical wet forests, especially in
phosphorous and calcium, when compared to other lowland forest sites
(Clinebell et al. 1995). Storms are infrequent in the area but often cause
multiple treefalls, leaving the forest in various stages of succession (D. Neill
& W. Palacios, unpublished).
The land-use patterns in the vicinity of Jatun Sacha have undergone rapid
changes in the last decade. Before the early 1980’s the area was sparsely
populated by native Quichuans and accessed only by rivers. A road built in
1986 bisected the reserve at its northern end along the Río Napo and greatly
increased access to the area. The influx of small scale farmers and portable
sawmills resulted in deforestation in areas accessible by the road. Currently,
tracts of land owned by farmers adjacent to the road typically have 40 to 70
44 J. Res. Lepid.
percent of the land cleared. Tracts in the interior are more pristine, from
50 to 100 percent primary forest. Jatun Sacha continues to expand its re-
serve and purchases lands in a piecemeal fashion as funds and land become
available. Thus the reserve is a patchwork of habitats. Its central core is
mostly primary forest (70%), and its edges are a mosaic of primary forest,
secondary forest, scrub, and pasture land (D. Neill & W. Palacios, unpub-
lished).
A brief description is presented below of the two comparative sites, Pakitza
and Cacaulandia. More complete descriptions are available from Erwin
(1991) for Pakitza and Emmel and Austin (1990) for Cacaulandia. Pakitza
is a biological station in the Reserved Zone of Parque Nacional Manu. It is
located in Madre de Dios drainage basin in Peru along the foothills of the
eastern Andes in a similar geographical zone as Jatun Sacha. The butterfly
survey for Pakitza was comprehensive and yielded 1300 species (Robbins et
al. 1996). The survey from Cacaulandia was conducted on a private ranch
in Rondonia, Brazil. Located in the rolling hills and flat plains of the Ama-
zon basin, it has both intact forest and disturbed areas. A total of 843 spe-
cies of butterflies was recorded by Emmel and Austin (1990), although con-
tinued surveys have increased this total number to approximately 1500 spe-
cies (Austin & Emmel 1996, cited as “unpublished data”). The area of
Cacaulandia is ecologically less similar to Jatun Sacha than Pakitza, but faces
similar pressures from development.
ignated with question marks. A synoptic collection has been deposited in the Museo
de Ecuatoriana Nacional in Quito, Ecuador.
For comparative work among the three sites, the percent of species occurring in
each family was tabulated, and a test for homogeneity across the families was calcu-
lated using a 2x2 contingency table. To compare similarity in species assemblages
between the three sites, coefficient of community indices (Pielou 1974) were cal-
culated in pairwise comparisons between Jatun Sacha and Pakitza, Jatun Sacha and
Cacaulandia, and Pakitza and Cacaulandia. Only those identified to species (spe-
cies similarities) or genus (generic similarities) were used in calculations. Lycaenidae
was not used in due to poor taxonomic resolution at the genus level and lack of
identifications in the Cacaulandia survey (59 of the 87 species were unidentified).
Using these adjusted species numbers, percentages were again calculated for fam-
ily compositions, which were used in contrasting the expected and observed spe-
cies common to all three sites.
RESULTS
A total of 811 species were recorded at the reserve by the end of 1993
(Appendix 1). The taxonomic composition of the butterfly fauna is as
follows: Hesperiidae, 198 spp. (25%), Papilionidae, 26 spp. (3%),
Pieridae, 27 spp. (3%), Nymphalidae, 307 spp. (38%), Riodinidae, 194
spp. (24%), and Lycaenidae, 59 spp. (7%). Within Nymphalidae, 56 spe-
cies of Ithomiinae are those reported by Beccaloni (1995), who con-
ducted a thorough study of this group. Temporal variations in richness
and abundance were generally noted for the butterfly families, although
quantitative data was collected only for the fruit-feeding nymphalids. The
fruit-feeders were more common during the wetter months (DeVries et al.
1997), and many specimens collected during this period were fresh, indi-
cating a recent emergence. During this same time period, other families
were observed to be much less abundant, although certain species could
be common (Eurybia dardus, Urbanus simplicius, “Thecla” tephraeus gr).
Hesperiidae, Riodinidae, and to some extent, Lycaenidae, were more abun-
dant as the rainfall decreased in August and September. Differences were
noted in the abundance of families and individual species from year to year,
46 J. Res. Lepid.
Jatun Sacha-Pakitza 49 81
Jatun Sacha-Cacaulandia 45 75
Pakitza-Cacaulandia 38 6
DISCUSSION
The survey conducted at Jatun Sacha was aimed at developing a baseline
understanding of the butterfly community of the area. A large portion of
the fauna undoubtably remains unsampled. This conclusion is supported
by the fact that unrecorded species were collected up to the end of the sur-
vey time. In addition, preliminary identifications for certain groups have
probably underestimated the number of butterfly species actually collected.
Because field collection was not standardized, estimations of the total spe-
cies richness at Jatun Sacha can not be generated through rigorous statisti-
35:42–60, 1996 (2000) 47
versity in certain groups, such as Nymphalidae (Brown 1982; but see also
DeVries et al. 1997). Butterfly species common to open, disturbed areas are
rare or absent at Pakitza (Robbins et al. 1996), but are quite common at
Jatun Sacha along the road bisecting the reserve. The low species richness
of Hesperiidae and Lycaenidae recorded at Jatun Sacha and Cacaulandia
could also reflect disturbance, especially at Jatun Sacha. A lepidopterist who
has been collecting in the Upper Napo area since 1978 has noted a great
decrease in the species and abundance of the Hesperiidae over the last
decade as developmental pressures increased (S. Nicolay, pers. comm.).
From the comparisons of the overlapping species, nymphalid species were
most common and found at greater numbers than expected. This suggests
broader distributions of nymphalids than other butterfly families. This may
be due to the wide dispersing capabilities of many nymphalids, which have
been correlated with greater distributions (Hanski et al. 1993). It could also
reflect broader hostplant ranges for nymphalids or more specialized, and
hence, localized host use by other butterfly families. With our limited knowl-
edge of host use even in well studied areas such as Costa Rica (DeVries 1987,
1996; DeVries et al. 1994), examining these broader biogeographical pat-
terns must await further investigations (but see Ackery 1988).
Human influence outside of Jatun Sacha most likely has impacted the but-
terfly fauna. Species inventories conducted while the area contains a high
percentage of pristine forest could be compared with future inventories in
a potentially much more disturbed landscape. Because degradation of the
upper Napo basin will continue, there is a critical need for more research.
For too many species, little is known beyond their site records. A great deal
remains to be discovered to complete our understanding of the butterfly
fauna, not only in documentation of the species diversity, but also their
ecology, evolution, and population dynamics.
Acknowledgements. I thank the following for contributing specimens to the list: Dave
Arenholz, George Beccaloni, Marion Murray, Andrew Neild, Stan Nicolay, Carla
Penz, Karina Soria, Alejandro Suárez, Gabriel Tapuy, and the students of the Save
the Rainforest. The survey list would not have been possible without the time and
effort to identify specimens by Stan Nicolay (Hesperiidae), George Beccaloni
(Ithomiinae), Lee Miller (Satyrinae), Don Harvey, Dave Arenholz, and Phil DeVries
(Riodinidae), and Bob Robbins (Lycaenidae). This manuscript was improved by
changes suggested by George Beccaloni (Natural History Museum), Chris Carlton
(Louisiana State University), Phil DeVries (University of Oregon), Sam Messier
(University of Colorado), David Neill (Missouri Botanical Garden), Dorothy Prowell
(Louisiana State University), and Bob Robbins (National Museum of Natural His-
tory). This work was possible through the support of Fundacíon Jatun Sacha and
the United States Peace Corps.
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APPENDIX 1
Pyrrhopyginae: 4
Elbella theseus Bell, 1933
Passova passova Evans, 1951
Pyrrhopyge proculus cintra Evans, 1951
Pyrrhopyge aziza lexos Evans, 1951
52 J. Res. Lepid.