Location via proxy:   [ UP ]  
[Report a bug]   [Manage cookies]                

Orugas Ecuador

Download as pdf or txt
Download as pdf or txt
You are on page 1of 15

Ecology, Natural History, and Larval Descriptions of Arctiinae

(Lepidoptera: Noctuoidea: Erebidae) from a Cloud Forest in the


Eastern Andes of Ecuador
Author(s): Suzanne B. Rab Green, Grant L. Gentry, Harold F. Greeney, and Lee A.
Dyer
Source: Annals of the Entomological Society of America, 104(6):1135-1148. 2011.
Published By: Entomological Society of America
DOI: http://dx.doi.org/10.1603/AN10165
URL: http://www.bioone.org/doi/full/10.1603/AN10165

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the


biological, ecological, and environmental sciences. BioOne provides a sustainable online
platform for over 170 journals and books published by nonprofit societies, associations,
museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates
your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial
use. Commercial inquiries or rights and permissions requests should be directed to the
individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers,
academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.
SPECIAL FEATURE

Ecology, Natural History, and Larval Descriptions of Arctiinae


(Lepidoptera: Noctuoidea: Erebidae) From a Cloud Forest in the
Eastern Andes of Ecuador
SUZANNE B. RAB GREEN,1 GRANT L. GENTRY,2 HAROLD F. GREENEY,3 AND LEE A. DYER4

Ann. Entomol. Soc. Am. 104(6): 1135Ð1148 (2011); DOI: http://dx.doi.org/10.1603/AN10165


ABSTRACT Adults and larvae in the subfamily Arctiinae (family Erebidae), along with their host
plants and the parasitic wasps and ßies that attack them, are important components of most terrestrial
food webs, but basic taxonomic and life-history knowledge for the arctiines of the Neotropics is still
poorly known. This is true for most groups of Lepidoptera in hyperdiverse countries such as Ecuador.
To examine host afÞliations and natural enemies in diverse ecosystems, we collected 6,243 arctiine
caterpillars (representing 821 larval morphospecies) at elevations ranging from 400 to 3,500 m as part
of a larger ecological and taxonomic survey of trophic associations centered at the Yanayacu Biological
Station in Napo province, eastern Ecuador. Here, we provide a brief review of the systematics of
Neotropical Arctiinae, provide basic information on host afÞliations and parasitism rates of arctiines
reared from Napo province in Ecuador and describe the immature stages of 16 species encountered
in the region.

KEY WORDS Arctiinae, Ecuador, larval morphology, parasitoid, trophic interactions

Tropical butterßies are quite well studied, but there al. 2011) have published a number of studies examin-
are major gaps in the taxonomy of most moth groups ing diversity of tropical Arctiinae, particularly in the
and limited knowledge about ecological interactions Andes, and it is clear from these studies that natural
between Lepidoptera and associated plants and ani- history of tropical Arctiinae is lacking. Here, we pres-
mals (summarized by Dyer et al. 2010). Tropical in- ent natural history data and larval descriptions for a
ventories of multitrophic interactions, focusing on representative sample of arctiine larvae that were col-
common taxa of lepidopteran larvae to uncover pat- lected and reared on the eastern slope of the Ecua-
terns of plantÐ herbivoreÐ enemy interactions, have dorian Andes.
contributed to important advances in ecology and Natural History and Interaction Ecology of Arctii-
evolutionary biology (Janzen et al. 2005, Novotny et al. nae. The Arctiinae or “tiger moths” (superfamily Noc-
2006). Here, we report results from one such inven- tuoidea), containing roughly 12,000 species, comprise
tory, with a focus on a particularly diverse and eco- 6% of lepidopteran species diversity worldwide (Wat-
logically interesting taxon, the Arctiinae (Lepidop- son and Goodger 1986, Goodger and Watson 1995)
tera: Noctuoidea: Erebidae). and are an important component of most tropical
What has historically been called the family “Arc-
communities. Adult sizes vary from mostly medium-
tiidae” is now recognized as a subfamily within the
sized (3Ð 6 cm in length) to fairly large (⬎6 cm), and
family Erebidae, based on a phylogenetic framework
they possess a diverse spectrum of colors and shapes,
for assigning the content and arrangement of taxa
within the Noctuoidea (Lafontaine and Schmidt 2010, from brightly colored to cryptic, sometime forming
Zahiri et al. 2010). The Arctiinae is a diverse taxon, and mimicry complexes with other taxa (Conner 2009).
the ecology and evolutionary biology of this subfamily The Neotropical region hosts a higher diversity of
is relatively well studied in temperate ecosystems (re- Arctiinae than any other geographic area, with a rough
viewed by chapters in Conner 2009) but less so in the estimate of 6,000 described species (Watson and
tropics. Brehm and colleagues (Brehm 2007, Beck et Goodger 1986, Scoble 1992). There are ⬇12,000 de-
scribed species worldwide, grouped into 750 genera
1 Corresponding author: Division of Invertebrate Zoology, Amer-
(Common 1990). The number of undescribed arctiine
ican Museum of Natural History, New York, NY 10024 (e-mail:
species in all zoogeographical regions is estimated to
rabgreen@amnh.org). be 10,000 species (Heppner 1991, Kristensen et al.
2 Department of Biology and Environmental Sciences, Samford
2007). It is likely that many of these undescribed
University, Birmingham, AL 35229. species occur in the Andes, a region with particularly
3 Yanayacu Biological Station and Center for Creative Studies,

Napo, Ecuador, c/o Foch 721 y Amazonas, Quito, Ecuador. high lepidopteran species richness and a large pro-
4 Department of Biology, University of Nevada, Reno, NV 89117. portion of undescribed species (Brehm et al. 2005).

0013-8746/11/1135Ð1148$04.00/0 䉷 2011 Entomological Society of America


1136 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Arctiines are common throughout the Neotropics, early works form the basis for modern species-level
occurring at elevations ranging from sea level up to identiÞcations and taxonomy.
⬇3,500 m (Piñas et al. 2000, Hilt and Fiedler 2005). In Early arctiine taxonomists relied heavily on super-
southern Ecuador, a clear diversity peak occurs at Þcial characteristics of adults such as wing venation
1,600 m, but arctiines are probably most diverse at and color patterns, whereas important identiÞcation
lower altitudes (Fiedler et al. 2008). Arctiine commu- tools, such as genitalia, were not used in systematics
nities in the eastern Andes of southern Ecuador are until the 1940s. Arctiinae are often sexually dimorphic,
highly diverse in late successional to young secondary polymorphic, or geographically variable; thus, wing
forest; openness of habitat and elevation are the most pattern characters are problematic for classiÞcation.
important predictors of community diversity (Hilt and Species-level investigations of arctiine systematics in-
Fiedler 2005). It is not clear whether such habitat clude the works of Travassos (1943Ð1964), Rego Bar-
preference is correlated with adult preference for ros (1956 Ð1971), Forbes (1917Ð1957), Watson (1967-
oviposition sites or with speciesÕ host speciÞcity 1995), and deToulgoët (1967Ð2000). DeToulgoët
(Brehm and Fiedler 2001), but these and related ques- published 76 papers in which he described nearly 200
tions can only be answered through natural-history Neotropical species and provided revisionary treat-
observations and studies of larval ecology. ments of groups in need of reclassiÞcation. Forbes
Habitus in arctiine caterpillars varies tremendously (1939, 1941, 1960) had a marked impact on the higher
(Wagner 2005, Wagner 2009). In most cases they are classiÞcation and hypotheses of generic afÞnities, fol-
densely setose, contrastingly and brightly colored, and lowed by other authors, including Franclemont
often have elongate planta with heteroideous cro- (1983), Ferguson (1985), and Watson and Goodger
chets. All segments except the head contain verrucae (1986). Taxonomic revisions and phylogenetic studies
(or “warts”) on which secondary setae are arranged in of species groups include Forbes (1941), Field (1975),
tufts at the primary setal locations. This gives them Watson (1975, 1977, 1980), Dietz and Duckworth
their “furry” appearance and the well-known common (1976), Dietz (1994), DaCosta and Weller (2005),
name “woolly bears.” Aposematic coloration is used to DaCosta (2007), Zaspel (2004), Zaspel and Weller
advertise larval toxicity in arctiine caterpillars. Other (2006), and Schmidt (2007). A major goal of our study
defenses include the sequestration of chemicals, is to present larval descriptions with potentially useful
characters that have been neglected in taxonomic
group feeding, unusual postures, thrashing, stridula-
studies of Neotropical Arctiineae.
tion, odors, secreted toxins on the integument or setae,
defecation, and host switching or other spatial or tem-
poral alterations in feeding behavior (Rothschild et al. Methods
1970, Gentry and Dyer 2002, Stireman and Singer 2003,
Collecting, Rearing, and Curation. We collected
Wagner 2009). Adults of many species participate in
caterpillars from primary forests, partially reforested
mimicry complexes with toxic beetles (Simmons pastures, and roadsides in Napo province of eastern
2009), other Lepidoptera (Aiello and Brown 1988), Ecuador, in the vicinity of the Yanayacu Biological
and stinging Hymenoptera (Simmons and Weller Station and Center for Creative Studies ([YBS] 00⬚
2002). Studies of such larval defenses and adult mim- 35.9⬘ S, 77⬚ 53.4⬘ W; altitude, 2,100 m), 5 km west of
icry complexes as well as investigations on interaction Cosanga. Altitudes and geographical coordinates for
diversity all beneÞt from improved documentation of sampling sites were measured with hand held GPS
natural history. The larval descriptions, host plant units. In addition, we collected caterpillars and adults
data, and data on parasitism rates presented here pro- along an altitudinal transect from 400 m (Tena) to
vide a foundation for further ecological studies with 3,500 m (Papallacta), searching for caterpillars in the
these species. understory on all accessible shrubs, forbs, and trees.
Taxonomy of Arctiinae. Arctiines have long at- We searched opportunistically, as well as systemati-
tracted the attention of collectors and systematists; cally within 10-m-diameter plots. All caterpillars were
yet, despite extensive representation in collections, reared on leaves of the plant from which they were
the classiÞcation of the group remains controversial collected (see species descriptions for complete host
and great uncertainty exists (Weller et al. 2009). Tax- plant information), until they either died or an adult
onomic work on Neotropical arctiines during the late moth or parasitoid eclosed. Levels of parasitism were
nineteenth and early twentieth centuries was carried calculated as the percentage of parasitoids reared from
out by many investigators, including: Paul Dognin the total number of results (emerged adults and para-
(1889 Ð1924), Arthur G. Butler (1867Ð1882), Francis sitoids). Host plant speciÞcity was quantiÞed using the
Walker (1854 Ð1869), George F. Hampson (1898 Ð average number of plant species on which an individ-
1920), Herbert Druce (1883Ð1911), Harrison G. Dyar ual arctiine larval morphospecies was collected. Mod-
(1892Ð1925), Lionel Walter Rothschild (1909 Ð1937), iÞcations of WhittakerÕs ␤ diversity also were calcu-
Adalbert Seitz (1919 Ð1925), and William Schaus lated for all arctiine morphospecies as a proxy for
(1889 Ð1941) (Walker 1865, Schaus 1901, Druce 1910, measuring the degree of specialization (Dyer et al.
Rothschild 1910, Dognin 1913, Hampson 1914). Pub- 2007) across host plant species.
lications by some of these authors provided useful Freshly eclosed adult specimens were freezer-killed
detailed color illustrations; however, most of their and either individually wrapped in glassine envelopes
species descriptions were brief. Nevertheless, these or Þeld pinned. They were subsequently spread and
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1137

genital dissections were carried out for selected indi- Table 1. Identified species of Arctiinae reared from immature
viduals, according to standard protocols (Winter stages to adult in the eastern Andes of Ecuador
2000). Specimens were sorted to morphospecies, with
Author Ma Fa Ua
males and females being grouped separately, and iden-
tiÞed using publications and reference collections. Arctiini: Phaegopterina
Idalus veneta Dognin 11 12 3
The following collections and their type material were Symphlebia palmeri Rothschild
consulted for identiÞcations: American Museum of Amaxia sp. 1
Natural History, New York (AMNH); National Mu- Neonerita haemasticta Dognin 3
seum of Natural History, Washington, DC (USNM); Melese innocua Dognin 2 4
Melese nr.amastris Druce 3 5 1
Natural History Museum, London, United Kingdom Melese peruviana Rothschild 1
(BMNH); and the Muséum National dÕHistoire Na- Bertholdia griseopalpis Rawlins 1 1
turelle, Paris, France (MNHN). Literature sources for Bertholdia nr. Simplex Rawlins 1
all generic and speciÞc names were checked and are Bertholdia ockendeni Rothschild 1 1
Bertholdia partita Rawlins 6 6 5
cited here, along with type localities from the original Bertholdia sp. 1
descriptions. Pachydota affinis Rothschild 2 1
Specimens from this study have been deposited in Pachydota nr. albiceps Walker 21
collections at the Museo Ecuatoriano de Ciencias Na- Pelochyta gandolfii Schaus 13 14 9
Pelochyta nr. albipars Hampson 1 1
turales (MECN), the AMNH, the USNM, and in a Pelochyta nr. cinerea Walker 1
synoptic collection at the University of Nevada, Reno, Elysius hades Druce 3 8 2
NV. A synoptic collection, which will eventually con- Elysius melanoplaga Hampson 2 2
tain at least one specimen representing each form, also Elysius nr. ochrota Hampson 2
Elysius ochrota Hampson 9 12 8
is located at the MECN. Adult specimens were pho- Elysius sp. 1
tographed at the AMNH by using a Microptics-USA Ischnocampa nr.obscurata Hampson 1 2
photomicrographic apparatus, equipped with InÞnity Ischnocampa nr. lugubris Schaus 5 7
K2 optics, a Nikon D1X digital camera, and a Nikon Ischnocampa sordidor Rothschild 3 7
Ischnocampa sp. 1 2
60-mm AF Micro Nikkor. The scale bar shown with Opharus consimilis Hampson 6 10 4
adult photographs is in millimeters. All larval descrip- Opharus nr. flavicostata Dognin 4 1
tions apply to last instars and were based on Þeld notes Opharus procroides Walker 3 1
and larval images from our rearing project. All images Opharus subflavus Toulgot 1
Bernathonomus sp. 1 1
were produced by a variety of digital cameras in the Phaegoptera Rothschild 19 17 9
Þeld. The terminology used to describe the larval decrepidoides
stages was consistent with that used by Stehr (1993). Phaegoptera nr. discisema Hampson 1 1
Praeamastus albipuncta Hampson 1 1
Praeamastus minerva Dognin 10 15 7
Results and Discussion Amastus coccinator Schaus 8 4
Amastus mirificus Toulgot 1 2
Larval Natural History and Host Plant Affiliations. Amastus nr. hyalina Dognin 11 6 1
From January 2002 to June 2010, we collected 6,243 Amastus persimilis Hampson 3 8 2
Amastus reticulatus Toulgot 1
arctiine caterpillars (representing 821 larval mor- Amastus thermidora Dognin 1 2
phospecies) and successfully reared 2,313 to an adult Turuptiana obliqua Walker 1 1
moth or parasitoid (Table 1). The majority of these Halysidota atra rindgei Watson 9 2 4
collections were from cloud forest sites at elevations Halysidota humosa Dognin 1
Halysidota intensa Rothschild 1
ranging from 1,800 to 2,400 m. Based on the rearings, Lophocampa atriceps Hampson 5 1 1
mean developmental time from early third instar to Lophocampa nr. distincta Rothschild 24 25 11
pupation was 36.1 ⫾ 0.9 d, and the pupal stage lasted Lophocampa petulans Dognin 2
36.4 ⫾ 0.8 d until eclosion. Lophocampa sp. A 2 3
Lophocampa sp. B 1
Diet breadth of arctiines is broad compared with Lophocampa sp. C 4 2
most other Neotropical Lepidoptera (Dyer et al. Leucanopsis Schaus 6 7
2007). In our samples, caterpillars were collected and apicepunctata
reared on plants in 2.4 ⫾ 0.8 families, 2.3 ⫾ 0.7 genera, Leucanopsis guascana Schaus 2
Leucanopsis luridioides Rothschild 1
and 2.6 ⫾ 0.9 species. The WhittakerÕs ␤ measure of Leucanopsis notodontina Rothschild 1 1
specialization (described in Dyer et al. 2007), varies Leucanopsis nr. stipulata Rothschild 1 2
from 0 to 1, with values ⬍0.9 indicating a very general Leucanopsis nr. Rothschild 4 5
diet) was 0.84 ⫾ 0.0004, which is more generalized subterranea
Leucanopsis nr. tabernilla Schaus 1
than any other lepidopteran group at the study area. Leucanopsis oruba Schaus 1 2
Arctiine caterpillars were collected on plants from 76 Leucanopsis siegruna Schaus 6
families, and the families that yielded the most indi- Arctiini: Arctiina
vidual larvae were Poaceae (mostly Chusquea scan- Hypercompe nemophila Herr.-Schaeffer 1 1
Hypercompe nr. nigriloba Hulstaert 1
dens Kunth), Asteraceae, Fabaceae, Rosaceae, and Hypercompe obscura Schaus 3 2
Melastomataceae (Table 2). The arctiine host plants in Hypercompe robusta Dognin 1
our study (Table 2) are often common in disturbed Chlorhoda metamelaena Dognin 1
habitats, such as on the borders of roads, trails, and
creek beds. Although this seems to be consistent with
1138 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Table 1. Continued Table 2. Plant families on which arctiine caterpillars from the
eastern Andes in Ecuador were collected and reared
Author Ma Fa Ua
Plant family N Plant family N
Arctiini: Ctenuchina
Dasysphinx mucescens Felder 1 1 Poaceae 1885 Polygalaceae 7
Dasysphinx sp. A 1 Asteraceae 906 Myrtaceae 6
Dasysphinx sp. B 1 Fabaceae 387 Smilacaceae 6
Gymnelia bricenoi Rothschild 1 1 Rosaceae 366 Amaranthaceae 5
Chrostosoma dolens Walker 1 Melastomataceae 338 Malphigiaceae 5
Cosmosoma flavicornis Druce 1 Betulaceae 223 Mirsinaceae 5
Cosmosoma thiacia Hampson 1 Urticaceae 221 Monimiaceae 5
Holophaea sp. 1 Dryopteridaceae 199 Onagraceae 5
Holophaea endoleuca Dognin 1 1 4 Unknown 190 Sapindaceae 5
Holophaea lycone Druce 2 Euphorbiaceae 122 Bromeliaceae 4
Rhynchopyga bicolor Dognin 1 Piperaceae 118 Malvaceae 4
Saurita melanifera Kaye 1 1 Ericaceae 116 Meliaceae 4
Saurita mosca Dognin 86 79 15 Gesneriaceae 99 Mimosaceae 4
Phaeo cephalena Druce 2 2 1 Lauraceae 98 Thymeliaceae 4
Argyroeides augiades Druce 1 Gunneraceae 91 Begoniaceae 3
Hyalomis thyria Druce 3 2 Solanaceae 80 Brunelliaceae 3
Diptilon chrysocraspis Hampson 8 5 Heliconiaceae 69 Convolvulaceae 3
Antichloris nr. bricenoi Rothschild 13 17 7 Moraceae 68 Flacourtiaceae 3
Antichloris nr. caca Huebner 1 Rubiaceae 68 Verbenaceae 3
Napata pseudolelex Rothschild 2 Juglandaceae 50 Bignoniaceae 2
Cyanopepla alonzo Butler 1 1 Araceae 46 Cunoniaceae 2
Cyanopepla bertha Druce 5 9 Cyclanthaceae 42 Hydrangeaceae 2
Cyanopepla buckleyi Druce 5 9 1 Pteridophyta 35 Lamiaceae 2
Cyanopepla hurama Butler 3 1 Cannaceae 34 Leguminosae 2
Desmotricha imitata Druce 9 15 4 Clusiaceae 32 Oxalidaceae 2
Mevania basalis Walker 27 21 3 Myricaceae 31 Rutaceae 2
Mevania larissa Druce 2 4 2 Campanulaceae 29 Sabiaceae 2
Hyparphara junctura Walker 18 28 4 Actinidiaceae 27 Amaryllidaceae 1
Delphyre tetilla Dognin 1 Boraginaceae 27 Commelinaceae 1
Eucereon casca Dognin 2 Cecropiaceae 19 Coriariaceae 1
Eucereon davidi Dognin 1 Hippocrataceae 13 Costaceae 1
Eucereon minerva Dognin 2 Arecaceae 12 Cucurbitaceae 1
Eucereon ockendeni Druce 1 Zingiberaceae 12 Lecythidaceae 1
Eucereon zizana Dognin 1 Dennstaedtiaceae 11 Magnoliaceae 1
Correbia nr. rufescens Rothschild 1 Bombacaceae 10 Musaceae 1
Correbia sp. 1 Aquifoliaceae 9 Orchidaceae 1
Hyaleucerea morosa Schaus 1 Araliaceae 8 Tropaeolaceae 1
Hyaleucerea nr. lemoulti Schaus 1 Myristicaceae 8 Zamiaceae 1
Arctiini: Pericopina Passißoraceae 7
Hypocrita plagifera Felder 1
Crocomela erectistria Warren 2 1 a
Number of individual caterpillars collected from plants in the
Antiotricha furonia Druce 2 family; some caterpillars were collected and reared on unknown
Antiotricha integra Walker 7 families.
Dysschema nr. Walker 9 5
dissimulata
Dysschema nr. mosera Druce 2 sented by three hymenopteran families, Braconidae
Dysschema nr. bivittata Dognin 1
Dysschema nr. talboti Dognin 4 1 9
(N ⫽ 197), Eulophidae (N ⫽ 9), and Ichneumoni-
Dysschema palmeri Druce 9 4 dae (N ⫽ 119), and one dipteran family, Tachinidae
Dysschema sp. 3 (N ⫽ 312). The overall level of parasitism on arc-
Lithosiini: Lithosiina tiines was 29.6%, which is intermediate compared
Nodozana cocciniceps Dognin 1
Metalobosia holophaea Dognin 2
with other commonly collected externally feeding
Marsypophora erycinoides Felder 1 immatures, such as Pyralidae (37.3%), Noctuidae
Marsypophora sp. 1 (36.9%), Saturniidae (35.3%), Hesperiidae (28.9%),
Ardonea sp. 1 Nymphalidae (24.9%), Notodontidae (22.9%), and
Unknown genus 1
Geometridae (15%).
M, number of male specimens reared; F, number of female spec- Parasitism on the arctiines in this region was not
imens reared; U, sex undetermined. remarkably high compared with parasitism rates in
temperate areas with more predictable weather pat-
terns (Stireman et al. 2005). The relative impor-
sampling results reported by Hilt and Fiedler (2005), tance of different parasitoid taxa was similar to that
with openness or secondary growth predicting high found at other sites and in other caterpillar groups.
incidence of arctiines, we did not set out to test this Tachinids were the dominant parasitoid taxon, fol-
hypothesis. Our data simply provide a sample of host lowed by braconids and ichneumonids (Stireman et
plants used at the study sites (Table 2), along with a al. 2005). Interestingly, there were no recorded
report of host plant afÞliations. cases of attack by myrmithid nematodes, which are
Parasitism. In total, 685 parasitoids were reared a common parasitoid in tropical lowlands (Gentry
from arctiine caterpillars, with most being repre- and Dyer 2002).
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1139

Fig. 1. Larvae and associated adults of arctiines reared from the study site in the eastern Andes of Ecuador. V, ventral
view; D, dorsal view. Top to bottom: Idalus veneta, Symphlebia palmeri, Neonerita haemasticta, and Melese peruviana. (Online
Þgure in color.)

Larval Descriptions. The caterpillars discussed here Idalus veneta Dognin, 1901
are species that we were able to positively identify; (Fig. 1)
some are the most commonly collected species,
Tribe. Arctiini.
whereas others were rarely collected but represent Subtribe. Phaegopterina.
important groups within the subfamily. The host Type Locality. Colombia.
plants listed with each caterpillar are the plants on Identification. Compared with 么 Type No. 30465
which they were discovered and successfully reared. U.S.N.M.; Idalus venata Dognin type. Type 么 Idalus
Rearing times for different species varied but could veneta Dognin by A. Watson, 1966. Genitalia slide
have been affected by the rearing conditions, includ- AW19.
ing differences in temperature, humidity, absence of Dissections. Specimens dissected by S.R.G.:
natural enemies, and nutritional value of cut leaves. Reared 么 # 37061 located at AMNH, genitalia slide
The descriptions here are of Þfth (ultimate) instar SRG #110.
caterpillars; earlier instars for many species in this Collected (YBS) 么 located at AMNH, genitalia slide
region look entirely different from the later instars, SRG #109.
with some changing characteristics across instars, such Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
as integument color; density, length, and color of sec- W; altitude, 2,200 m.
ondary setae; and number of verrucae bearing setae. Description. Head mostly shining black but with
The species (Table 1, Figs. 1Ð 4) are arranged in the epicranial suture prominent and pale pink and with
same taxonomic order as they appear in the checklist posterior portions of head showing a rosy cast; body
of Watson and Goodger (1986): 1) Phaegopterina, 2) ground color deep, velvety black, intersegmental
Arctiina, and 3) Pericopina. Ctenuchina are in se- areas slightly rosy; A2ÐA8 each bearing a distinct,
quential order as listed by Seitz (1925). angled, red hatch mark ventrolaterally on anterior
1140 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Fig. 2. Larvae and associated adults of arctiines reared from the study site in the eastern Andes of Ecuador. V, ventral
view, D, dorsal view. Top to bottom: Praeamastus albipuncta, Amastus thermidora, Amastus nr. hyalina, and Halysidota atra
rindgei. (Online Þgure in color.)

portion of segment, markings on A2 and A8 brighter Reared 么 # 42631 located at AMNH, genitalia slide
than others; body with moderately dense, mid- SRG #178.
length to long, stiff, slightly plumose (appearing Reared 么 # 42648 deposited at AMNH, genitalia slide
serrate along shaft), black setae; thoracic and ter- SRG #179.
minal abdominal segments with a few similar, but Reared 么 # 39560 located at AMNH, genitalia slide
longer, setae; verrucae match ground color on all SRG #174.
segments; thoracic legs shining black; prolegs trans- Reared 么 # 42649 located at AMNH, genitalia slide
parent, dark red. SRG #177.
Natural History. The host plant for the one indi- Reared 么 # 42624 located at AMNH, genitalia slide
vidual that has been reared was an unknown species SRG #175.
of Bombacaceae. Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
W; altitude, 2,200 m.
Description. Larva robust; head uniformly shining
Symphlebia palmeri Rothschild, 1910 black; body ground color velvety dark purple dor-
(Fig. 1) sally, velvety black laterally; dorsum bearing indis-
tinct white lines crossing the body near interseg-
Tribe. Arctiini. mental areas, spiracles small, bright white; body
Subtribe. Phaegopterina. with scattered mid-length, soft, white, slightly plu-
Type Locality. San Antonio, western Colombia. mose setae, sparsest on thorax and posterior seg-
Identification. Compared with type collection at ments, which include similar but longer setae mixed
BMNH, collections at AMNH, MNHN, and USNM. in; dorsum with sparse, long, soft, slightly plumose
Dissections. Specimens dissected by S.R.G.: gray setae, several longer setae posteriorly and an-
Reared 乆 # 10756 located at AMNH, genitalia slide teriorly; segments T2ÐA8 bearing two to four dense
SRG #105. tufts of short, soft, deep orange setae creating a
Reared 么 # 42649 located at AMNH, genitalia slide nearly uniform covering across dorsum when larvae
SRG #177. are at rest, but are more obviously broken into tufts
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1141

Fig. 3. Larvae and associated adults of arctiines reared from the study site in the eastern Andes of Ecuador. V, ventral
view; D, dorsal view. Top to bottom: Lophocampa atriceps, Hypercompe obscura, Crocomela erectistria, and Dysschema palmeri.
(Online Þgure in color.)

during movement; thoracic legs black; prolegs dark Description. Final instar slender; head black with
purple with pale pads. uniformly sized yellow speckling, denser dorsally;
Natural History. The larvae of this species feed and body ground color velvety black above spiracular line
rest in groups. Known host plants include Gunnera sp. and dirty white below, thorax with a broad mid-dorsal
(Gunneraceae), Miconia sp., and Tibouchina lepidota orange-brown stripe, irregular yellow spiracular stripe
Aublet (Melastomataceae). along length of body, spiracles black, T3ÐA9 with a
thinner yellow stripe, the superior supraspiracular
margin broken into irregular spots; dorsum with
sparse, long, soft, slightly plumose gray setae, several
Neonerita haemasticta Dognin, 1906
longer setae posteriorly and anteriorly bear white,
(Fig. 1)
plumose tips, laterally and ventrally with sparse, stiff,
Tribe. Arctiini. short and mid-length, nonplumose white setae, A2ÐA8
Subtribe. Phaegopterina. each with four dorsal tufts of short, dense, strongly
Type Locality. Carabaya, southeastern Peru. plumose white setae, darker near bases, A1 bearing
Identification. Compared with 么 Type No. 34414 only one pair of tufts, on either side of midline and
U.S.N.M., slide AW287. twice as long all other tufts but similar in length to the
Dissections. Specimen dissected by S.R.G.: posterior pair of tufts on A7, dorsal verrucae dull
Collected (YBS) 么 located at AMNH, genitalia slide yellow with small black spots at base of setae, verrucae
SRG #72. below spiracular line matching surrounding cuticle
Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘ color; thoracic legs orange-brown; prolegs pale, dull
W; altitude, 2,200 m. orange with pale purple pads.
1142 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Fig. 4. Larvae and associated adults of arctiines reared from the study site in the eastern Andes of Ecuador. V, ventral
view; D, dorsal view. Top to bottom: Dysschema nr. dissimulata, Saurita mosca, Phaeo cephalena, and Desmotricha imitata.
(Online Þgure in color.)

Natural History. The larvae of this species are sol- projected anteriorly, these orange at bases, black dis-
itary feeders and have been reared on Columnea ericae tally, T2 also bearing a pair of similar dorsal tufts which
Mansfeld (Gesneriaceae), Chusquea scandens Kunth are slightly denser, longer, and becoming plumose
(Poaceae), Rubus sp. (Rosaceae), and Miconia sp. and apically; verrucae dull red-orange with small darker
T. lepiota (Melastomataceae). spots at setal bases; thoracic legs dark gray; prolegs
pale, dull red-orange with pale pads.
Natural History. The larvae of this species are sol-
Melese peruviana Rothschild, 1909
itary feeders and have been reared on Erato polym-
(Fig. 1)
nioides de Candolle (Asteracae) and Miconia sp.
Tribe. Arctiinae. (Melastomataceae).
Subtribe. Phaegopterina.
Type Locality. Carabaya, southeastern Peru.
Praeamastus albipuncta Hampson, 1901
Identification. Compared with type collection at
(Fig. 2)
BMNH, collections at AMNH, MNHN, and USNM.
Dissections. No previously published dissections. Tribe. Arctiini.
Description. Final instar slender; head uniformly Subtribe. Phaegopterina.
orange; body ground color beige, T3ÐA8 with a dark Type Locality. Tovar, Venezuela.
gray or black lateral wash, darkest close to interseg- Identification. Compared with type collection at
mental areas; very faint dark beige markings forming BMNH, collections at AMNH, MNHN, and USNM.
an indistinct mid-dorsal stripe, dorsum of T1 with dull Dissections. No previously published dissections.
orange wash; body sparsely covered with stiff, mid- Description. Larva slender; head dull brown with
length setae, dull orange near bases and darker at tips, very Þne reticulated pattern; body ground color olive
slightly longer on posterior and anterior segments, T2 green, thorax bearing a pale yellow, saddle-like patch
with sparse, supraspiracular tufts of long, soft hairs, covering posterior portion of T2 and anterior portion
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1143

of T3, patch extending to edge of dorsum on T2 but tus thermidora Dognin by A. Watson, 1967. Genitalia
down near spiracles on T3, large, elongate, triangular slide AW544.
or crescent shaped patches of same color cover most Dissections. Specimens dissected by S.R.G.:
of lateral portions of A3ÐA7, patch on A3 extending Reared 乆 #36190 located at AMNH, genitalia slide
anterodorsally onto A2 to form a thin stripe across SRG #137.
dorsum, patch on A7 extending below spiracle as a thin Collected (YBS) 么 located at AMNH, genitalia slide
line onto A8, spiracles small, pale yellow outlined with SRG #104.
light brown; body bare except for a few short, stiff Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
red-brown setae arising from supraspiracular verrucae W; altitude, 2,200 m.
from A1ÐA9, thorax and A8 ÐA10 with slightly denser Description. Larva robust; head uniformly reddish
setae of same type, interspersed with a few black, soft, brown with paler areas epicranially and around ecdy-
very long, plumose-tipped setae, these setae denser, sial suture; body ground dull gray-brown with indis-
projecting forward from dorsum of T2; T1Ð7 with tinct cream-colored and black markings dorsally and
dense mid-length tufts of reddish brown, soft setae; latero-ventrally, most prominent on thorax and ter-
one laterally and one subventral but close enough to minal abdominal segments, dorsum of A10 black, spir-
appear as one large tuft on each segment; dorsum of acles bright crimson but mostly obscured by setae;
A2ÐA7 with similar but shorter tufts, two on either side body with moderately dense, short, stiff setae, reddish
of midline in tightly packed groups so as to appear as brown at bases, dark centrally on shaft, and with red-
a single dorsal tuft, A2 with only one pair of similar brown tips, dorsal verrucae bearing additional dense
tufts, A1 with four tufts but these consisting of long tufts of shorter, soft, red-brown setae giving dorsum a
hairs merging into a dense, mid-dorsal spike-like tuft, fuzzy appearance, whereas rest of body appears spiny,
A8 with a similar arrangement of tufts of equal length thorax with a few long, soft, red-brown setae; thoracic
but angled backwards in a tail-like fashion, T3 bearing legs black; prolegs matching body ground color with
four very long, prominent, densely plumose-tipped, darker pads.
black setae, these arising in pairs from dorsal verrucae Natural History. The larvae of this species are sol-
and generally held erect; thoracic legs brown; prolegs itary feeders and have been reared on E. polymnioides
same color as body with paler pads. (Asteracae) and Miconia sp. (Melastomataceae).
Natural History. We have reared larvae of this spe-
cies on a large number of host plants, including Sau-
Amastus nr. hyalina Dognin, 1889
rauia bullosa Wawra (Actinidaceae), Anthurium sp.
(Fig. 2)
(Araceae), Baccaris sp. (Asteraceae), Brunnelia sp.
(Brunneliaceae), Erythrina edulis Triana ex M. Mi- Tribe. Arctiini.
cheli (Fabaceae), T. lepidota (Melastomataceae), Subtribe. Phaegopterina.
Rubus sp. (Rosaceae), Smilax sp. (Smilacaceae), Type Locality. Loja, Ecuador.
Boehmeria pavoni Weddell (Urticaceae), and Cythar- Identification. Compared with 么 Type No.30425
exylum sp. (Verbenaceae). We have reared tachinids U.S.N.M.; Amastus hyalina Dognin type. Type 么 Pha-
and meteorine braconids from this species. The ta- egoptera hyalina Dognin by A. Watson, 1967. Genitalia
chinid develops internally in the larva and emerges slide AW605. Our dissected specimen differed signif-
from the host pupa to pupate away from the host; up icantly from the type.
to 17 braconids develop internally and pupate away Dissections. Specimens dissected by S.R.G.:
from the host larva. At least Þve other caterpillars with Reared 么 # 33717 located at AMNH, genitalia slide
the same morphology and setal arrangements, but SRG #167.
different (sometimes quite different) coloration, have Reared 么 # 37871 located at AMNH, genitalia slide
been collected from the same host plants as P. albi- SRG #170.
puncta. The adults are likewise similar and generally Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
identiÞed as P. albipuncta. It is still unknown whether W; altitude, 2,200 m.
these are different species or if P. albipuncta has vari- Description. Larva robust; head uniformly red-
ably colored larvae. The adults are similar in appear- brown; body ground color beige with thick, indistinct
ance to species of the genera Praeamastus and Amas- darker brown spiracular strip and dark velvety black
tus. Their ambulatory behavior and the way they hold mid-dorsal stripe on thorax and posteriorly from A7,
their wingsÑfolded, overlapping, on their backsÑ segments A1ÐA7 with rounded black patches extend-
gives them the silhouette of a very large vespid wasp. ing ventrally to edge of subdorsal area, T1 and ab-
dominal spiracles bright golden-orange; body with
pale beige, moderately dense, mid-length, soft setae
arising from beige verrucae, densest below spiracles
Amastus thermidora Dognin, 1913
and on dorsal thorax, thorax and terminal abdominal
(Fig. 2)
segments additionally with a few sparse, long, soft,
Tribe. Arctiini. pale setae, A1ÐA7 each with two (subdorsal) dense
Subtribe. Phaegopterina. tufts of short, soft, red-brown setae, greatly reduced
Type Locality. Loja, Ecuador. on A1, A8 with moderately dense, mid-dorsal tuft of
Identification. Compared with 乆 Type No. 30404 short, soft, light brown setae; thoracic legs red-brown;
U.S.N.M.; Amastus thermidora Dognin. Type 乆 Amas- prolegs pale with dark gray pads.
1144 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Natural History. The larvae of this species are sol- Reared 么 # 26209 located at AMNH, genitalia slide
itary feeders. The most common host plant at our study SRG #138.
site is C. scandens (Poaceae), but additional host plants Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
include Erythrina smithiana Krukoff (Fabaceae), Piper W; altitude, 2,200 m.
cf. schuppii Gentry (Piperaceae), Vismia tomentosa Description. Larva elongate, tapering sharply be-
Ruiz & Pavon (Clusiaceae), Diplazium costale variety hind head, giving it a slightly hump-backed appear-
robustum Sodiro (Dryopteridaceae), and Rubus sp. (Ro- ance; head uniformly shiny black; body ground color
saceae). black, all body segments with sparse, Þne, bright white
ßecking, these forming an indistinct, thin, mid-dorsal
line from T2ÐA6; body with sparse, long to very long,
Halysidota atra rindgei Watson, 1980
soft, Þnely plumose white setae, dorsum of T3ÐA1 and
(Fig. 2)
A7ÐA8 bearing clusters of erect, densely packed tufts
Tribe. Arctiini. of mid-length, soft black setae, posterior tufts shorter
Subtribe. Phaegopterina. than anterior tufts; thoracic legs black; prolegs black
Type Locality. Rṍo Tabaconas, 6,000 feet, northern with red-brown pads.
Peru. Natural History. The larvae of this species feed in
Identification. Compared with 么 Type No. 34414 groups until the third instar and move to separate
U.S.N.M., slide AW287. leaves or plants as solitary feeders in later instars. The
Dissections. Specimen dissected by S.R.G.: most common host plant at our study sites is Boehmeria
Reared 么 # 2089 located at AMNH, genitalia slide spp. (Urticaceae, including B. bullata Kunth, B. cau-
SRG #198. data Swartz, B. pavoni Weddell, and B. ulmifolia Wed-
Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘ dell), but additional host plants include Phenax rugosus
W; altitude, 2,200 m. Weddell and Myriocarpa sp. (Urticaceae); C. scandens
Description. Larva robust, tapering slightly poste- (Poaceae); Erythrina smithiana Krukoff (Fabaceae);
riorly; head uniformly shiny dark brown; body ground Diplazium costale variety robustum Sodiro (Dryopteri-
color dark brown to black, T2ÐT3 with small orange daceae); Psammisia sodiroi Hoerold (Ericaceae), T. lepi-
markings dorsally; body with dense, soft, brown, short dota, Miconia sp., and Monochaetum lineatum D. Don
setae intermixed with sparse mid-length setae espe- (Melastomataceae); Geonoma orbignyana Martius (Are-
cially laterally; many black setae forming darker areas caceae); Niphidium sp. (Pteridophyta); Guettarda aff.
dorsally on body and on most of thorax, T2 and T3 with ochreata Schechtendal (Rubiaceae); Mikania sp. (Aster-
pairs of dorsal and subdorsal hair pencils of soft, very aceae); Centropogon sp. (Campanulaceae); Capanea
long, pale orange setae, those on T2 oriented forwards affinis Fritsch (Gesneriaceae); Acalypha sp. (Euphorbi-
and those on T3 swept back, these hair pencils move- aceae); and Piper schuppii Gentry (Piperaceae).
able and forming a star-like halo that hides head; A8
with a pair of similar hair pencils oriented posteriorly,
verrucae matching ground color of surrounding cuti-
Hypercompe obscura Schaus, 1901
cle; legs black, prolegs matching body color with pale
(Fig. 3)
pads.
Natural History. We have reared this species from Tribe. Arctiini.
eggs on Nectandra species (Lauraceae). Early and Subtribe. Arctiina.
middle instars have a bright pink ground color and Type Locality. Peru.
sparser secondary setae, and look completely different Identification. Compared with 么 Lectotype
from the Þnal instar caterpillar. Additional host plants No.11122 U.S.N.M.; Turuptiana obscura Schaus type.
include Castilla sp. (Moraceae), E. edulis Triana ex M. Lectotype 么 Turuptiana obscura Schaus by A. Watson,
Micheli and Desmodium sp. (Fabaceae), Juglans sp. 1967. Genitalia slide AW490.
(Juglandaceae), Rubus sp. (Rosaceae); Wercklea ferox Dissections. Specimens dissected by S.R.G.:
Hooker (Malvaceae), C. scandens (Poaceae), and Reared 么 # 35661 located at AMNH, genitalia slide
Piper baezanum de Candolle and Piper sp. (Piper- SRG #190.
aceae). The larvae are parasitized by Distatrix spp. Collected (YBS) 么 located at AMNH, genitalia slide
(Braconidae: Microgastrinae) and unknown species SRG #143.
of tachinid ßies; both the braconids and tachinids Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
develop internally and pupate outside of the host. W; altitude, 2,200 m.
Description. Larva robust; head uniformly shiny
dark brown to black; body ground color dark brown
Lophocampa atriceps Hampson, 1901
with a pale gray wash giving it a frosted appearance,
(Fig. 3)
T1 and abdominal spiracles bone-colored; body with
Tribe. Arctiini. reddish brown, moderately dense, mid-length, stiff
Subtribe. Phaegopterina. setae arising from shiny gray verrucae, thorax, sides,
Type Locality. Colombia. and posterior segments additionally with a few sparse,
Identification. Compared with type collection at long, soft, pale setae emanating from subdorsal and
BMNH, collections at AMNH, MNHN, and USNM. lateral verrucae; thoracic legs black; prolegs matching
Dissections. Specimen dissected by S.R.G.: body color with gray pads.
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1145

Natural History. The larvae of this species are sol- subspiracular verrucae from T3 to A8 bright crimson,
itary feeders. Host plants include Barnadesia parvi- dorsal verrucae on A8 dark red, all others black; tho-
flora Spruce ex Bentham & Hooker and Adenostemma racic legs, prolegs, and pads black.
harlingii King & Robinson (Asteraceae) and Browallia Natural History. The larvae of this species are sol-
speciosa Hooker and Solanum sp. (Solanaceae). itary feeders. The most common host plants are Oreo-
panax spp. (Araliaceae) and C. ericae (Gesneriaceae).
Additional hosts include C. scandens (Poaceae); Allo-
Crocomela erectistria Warren, 1904
plectus tetragonoides Mansfeld (Gesneriaceae); Schef-
(Fig. 3)
flera dielssi Hams (Araliaceae); and Dendrophorbium
Tribe. Arctiini. lloense Jeffrey and Critoniopsis occidentalis Cuatreca-
Subtribe. Pericopina. sas (Asteraceae).
Type Locality. Baños, Ecuador.
Identification. Compared with collections at
AMNH, USNM, and BMNH. Dysschema nr. dissimulata Walker, 1865
Dissections. Specimen dissected by S.R.G.: (Fig. 4)
Reared 么 # B841 located at AMNH, genitalia slide
Tribe. Arctiini.
SRG #133.
Subtribe. Pericopina.
Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
Type Locality. Bogotá, Colombia.
W; altitude, 2,200 m.
Identification. Compared with type collection at
Description. Larva slender; head uniformly shiny
BMNH, collections at AMNH and USNM.
black; body ground color black, paler in intersegment
Dissections. No previously published dissections.
regions, T2, A2ÐA7 bearing thick, bright yellow trans-
Description. Larva slender; head uniformly shiny
verse stripes anterior to intersegmental areas, A2 and
black; body ground color dark, velvety purple, dark
A7 also bearing similar transverse stripes posterior to
purple at intersegmental areas, body bearing dull
intersegmental areas; body with sparse, short to mid-
white, broken dorsal, subdorsal, and spiracular lines on
length, fairly stiff, black setae, anterior and posterior
T2 to A8, T1 and A8 with more extensive dull yellowish
segments also bearing a few very long, soft, slightly
white patterning, especially laterally, on T1 forming a
plumose setae, some with white tips, verrucae shining
distinct patch behind head, A10 also with dorsal pale
metallic blue-black; thoracic legs black; prolegs shiny
markings, T3 and A4 with light orange stripes extend-
blue-black, pads dark gray.
ing to spiracular area, spiracles dull orange; body with
Natural History. The larvae of this species feed in
sparse, short to mid-length, fairly stiff, black setae,
groups. Only one group has been collected at our
each segment also bearing a few longer, soft white
study site, feeding on an unknown plant in the Bor-
setae, white setae longest and most concentrated on
aginaceae.
dorsum of A8, dorsal setae on A3ÐA6 orange near
bases, most verrucae shiny metallic blue-black, excep-
Dysschema palmeri Druce, 1910 tions being those verrucae on T3, A4, and A8 and those
(Fig. 3) below spiracles on A3ÐA6, these exceptions bright
orange; thoracic legs black; prolegs shiny blue-black,
Tribe. Arctiini.
pads dark gray.
Subtribe. Pericopina.
Natural History. The larvae of this species are sol-
Type Locality. Male syntype: San Antonio, 5,800
itary feeders. The most common host plants are in
feet, western Colombia; female syntype: Puzuzo, east-
the Asteraceae: Critoniopsis occidentalis, Barnadesia
ern Peru.
parviflora, Clibadium glabrescens Blake, Mikania sp.,
Identification. Compared with type collection at
Adenostemma harlingii King & Robinson, and Erato
BMNH, collections at AMNH and USNM.
polymnioides. Other hosts include C. scandens
Dissections. Specimens dissected by S.R.G.:
(Poaceae) and unknown plant species in the So-
Reared 么 # 30445 located at AMNH, genitalia slide
lanaceae. Larvae are parasitized by tachinid ßies,
SRG #135.
which develop internally and pupate away from the
Locality. Ecuador, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘ W;
larva.
altitude, 2,200 m.
Description. Larva slender; head uniformly shiny
black; body ground color velvety black, dark purple at
Saurita mosca Dognin, 1897
intersegmental areas, body bearing dorsal, spiracular,
(Fig. 4)
and subspiracular broken lines of bright white dashes
from T3 to A8, T3 with a wide, bright crimson stripe, Tribe. Arctiini.
broken mid-dorsally and extending to spiracular area, Subtribe. Ctenuchina.
A8 with crimson spot around spiracular verrucae ex- Type Locality. Loja, Ecuador.
tending slightly dorsally to subdorsal area; spiracles Identification. Compared with type collections at
small and black; body with sparse, mid-length, fairly BMNH and USNM, collection at AMNH.
stiff, black setae, each segment also bearing a few long, Dissections. Specimens dissected by S.R.G.:
slightly plumose, soft white setae and sparse, short, Reared 么 # 36704 located at AMNH, genitalia slide
stiff white, nonplumose setae laterally, spiracular and SRG #130.
1146 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Reared 么 # 12414 located at AMNH, genitalia slide Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘
SRG #131. W; altitude, 2,200 m.
Reared 么 # 36687 located at AMNH, genitalia slide Description. Larva robust, tapering sharply at tho-
SRG #132. rax; head uniformly yellow; body ground color yellow
Locality. Ecuador, Napo, YBS, 00⬚ 35.9⬘ S, 77⬚ 53.4⬘ except for A1 and A7, which are dull brownish yellow,
W; altitude, 2,200 m. A2ÐA6 paler than other yellow segments, nearly white
Description. Larva slender; head shiny black later- dorsally; spiracles small and dark brown; thorax nearly
ally but pale whitish around ecdysial suture, dividing bare except for a few short, soft, pale, white setae;
black into two hemispheres; body ground color dark A8 ÐA10 with only slightly denser setae but these mid-
gray with indistinct white highlights, giving it a moldy length to long, soft and white except for a few dark
appearance, anal plate white, dorsal verrucae on T1 ones interspersed, some of the dark setae are slightly
with a faint pink cast; body with sparse, soft, black, plumose, A1ÐA7 with moderately dense, soft, white,
mid-length, curved and slightly plumose setae, ante- mid-length to long setae, sparser and shorter laterally,
rior and posterior segments with a few additional sim- A1 with subdorsal tufts of densely packed, soft, plu-
ilar, but longer, setae, T2 with six long dorsal setae mose-tipped, mid-length setae, tufts becoming black
ending in dense plumose tufts, two tufts white-tipped, dorsally; lateral setae white, A7 bearing similar dense
the rest black, A8 with a dorsal pair of white-tipped tufts of setae but slightly shorter near supraspiracular
setae, A1 and A7 with dense lateral tufts of short dull line; verrucae dark brown; thoracic legs, prolegs, and
orange setae, tufts so densely packed as to appear as pads same color as body.
solid projections; each tuft consisting of two densely Natural History. The larvae of P. cephalena are sol-
packed paddle-shaped hair pencils, one above the itary feeders. This species has been reared twice at our
other; verrucae matching ground color of surrounding study site, both times on Mikania micrantha Kunth
cuticle; thoracic legs and prolegs matching body color. (Asteraceae).
Natural History. The larvae of this species are sol-
itary feeders. The most common host plant is C. scan-
dens (Poaceae), but this species feeds on a large
number of plants, including Sarcorhachis sydowii Desmotricha imitata Druce, 1883
Trelease, Piper baezanum, Piper augustum Rudge, (Fig. 4)
and Piper cf. schuppii (Piperaceae); Miriocarpa
Tribe. Arctiini.
sp. and Boehmeria bullata (Urticaceae); Casearia
Subtribe. Ctenuchina.
sp. (Flacourtiaceae); Geonoma orbignyana (Are-
Type Locality. Chiguinda, western Ecuador.
caceae); Diplazium costale (Dryopteridaceae);
Identification. Compared with type collections at
Erato polymnioides (Asteraceae); Heliconia sp. (Heli-
BMNH and USNM, collections at AMNH.
coniaceae); Palicourea thyrsiflora Ruiz & Pavon (Ru-
Dissections. No previously published dissections.
biaceae); Nectandra cissiflora Nees (Lauraceae); Al-
loplectus tetragonoides (Gesneriaceae); Anthurium sp. Description. Larva somewhat robust, ßat across
(Araceae); and an unknown species in the Magnoli- dorsum and roughly square in cross-section; head uni-
aceae. The larvae are heavily parasitized by braconids, formly bright orange, slightly paler around epicranial
ichneumonids, and tachinids, all of which develop suture; body ground color bright white dorsally and
internally. Most of these parasitoids pupate away from dull yellow-white ventrally, strongly marked with
the host, except for some ichneumonids, which pupate black producing an indistinct checker-board dorsally
inside it. One tachinid species attacks the larva and and thick black stripes laterally, broken only by white
emerges as a prepupa from the host pupa. All the areas around base of supraspiracular verrucae; dorsum
others are larval-larval parasitoids. Parasitoid taxa that of thorax mostly white and unpatterned; body verru-
have been identiÞed include Carcelia sp. (Tachinidae: cae either black or white, matching patterning on
Exoristinae), Meteorus sp. (Braconidae: Meteorinae), surrounding cuticle; body covered with long to very
and an unknown species of Campopleginae (Ichneu- long, soft, slightly plumose setae, shorter setae entirely
monidae). white, longest setae gray apically; thorax and terminal
abdominal segments bearing several very long setae,
these gray with white tips; thoracic legs, prolegs, and
pads clear white to yellow-white.
Phaeo cephalena Druce, 1883 Natural History. Larvae are solitary. The primary
(Fig. 4) host plant is C. scandens (Poaceae), but D. imitata also
Tribe. Arctiini. has been reared on Baccharis latifolia Ruiz & Pavon
Subtribe. Ctenuchina. (Asteraceae). This species is a good wasp mimic when
Type Locality. Sarayacu, Ecuador; Antioquia, Co- in ßight. UnidentiÞed parasitoids in the families Bra-
lombia. conidae, Ichneumonidae, and Tachinidae attack the
Identification. Compared with type collections at larvae. All parasitoids develop internally, the ichneu-
BMNH and USNM, collections at AMNH. monids also pupate internally, whereas the braconids
Dissections. Specimen dissected by S.R.G.: and tachinids pupate outside of the larva. The bra-
Reared 么 # 35151 located at AMNH, genitalia slide conids emerge to pupate in groups of up to 30 prepupal
SRG #127. larvae.
November 2011 RAB GREEN ET AL.: DIVERSITY OF CLOUD FOREST ARCTIINAE 1147

Acknowledgments Fiedler, K., G. Brehm, N. Hilt, D. Süssenbach, and C. L.


Häuser. 2008. Variation of diversity patterns across
For assistance in the Þeld, we thank Rafael Granizo, Marco moth families along a tropical altitudinal gradient, pp.
Gualinga, Wilmer Simbaña, Lucia Salazar, and dozens of 167Ð180. In E. Beck, J. Bendix, I. Kottke, F. Makeschin,
Earthwatch volunteers. We thank the PBNHS, Tim Metz, Jay and R. Mosandl (eds.), Gradients in a tropical mountain
Peltz, and the Humboldt Crew for logistical support. For ecosystem of Ecuador (Springer Ecological Studies 198).
extensive comments and guidance on earlier versions of this Springer, Berlin, Germany.
manuscript, we thank Jim Miller. This study was funded by Field, W. D. 1975. Ctenuchid moths of Ceramidia Butler,
National Science Foundation grants DEB-0346729 and DEB- Ceramidiodes Hampson, and the caca species group of
1020509 and grants from Earthwatch Institute; Lepidoptera Antichloris Hübner. Sm. C. Zool. 198: 1Ð 45.
Research Foundation, Inc.; and the University of Nevada. Forbes, W.T.M. 1939. The Lepidoptera of Barro Colorado
This is publication 171 of the Yanayacu Natural History Re- Island, Panama. B. Mus. Comp. Zool. Harv. 85: 234 Ð321.
search Group. Forbes, W.T.M. 1941. The position of Uthetheisa galapagen-
sis (Lepidoptera, Arctiidae). J. NY Entomol. S. 49: 101Ð
110.
References Cited Forbes, W.T.M. 1960. The Lepidoptera of New York and
neighboring states. IV. Agaristidae through Nymphalidae,
Aiello, A., and K. S. Brown. 1988. Mimicry by illusion in a including butterßies. Memoir 371. Cornell University Ag-
sexually dimorphic, day-ßying moth, Dysschema jansonis ricultural Experiment Station, Ithaca, NY.
(Lepidoptera: Arctiidae: Pericopinae). J. Res. Lepidopt. Franclemont, J.G.F. 1983. Arctiidae, pp. 114 Ð119. In R. W.
26: 173Ð176. Hodges, T. Dominick, D. R. Davis, D. C. Ferguson, J. G.
Beck, J., G. Brehm, and K. Fiedler. 2011. Links between the Franclemont, E. G. Munroe, and J. A. Powell (eds.),
environment, abundance and diversity of Andean moths. Checklist of the Lepidoptera of America North of Mexico,
Biotropica 43: 208 Ð217. including Greenland. E. W. Classey Ltd. and Wedge En-
Brehm, G. 2007. Contrasting patterns of vertical stratiÞca- tomological Research Foundation, London, United King-
tion in two moth families in a Costa Rican lowland rain dom.
forest. Basic Appl. Ecol. 8: 44 Ð54. Gentry, G. L., and L. A. Dyer. 2002. On the conditional
Brehm, G., and K. Fiedler. 2001. Host plant relationships of nature of Neotropical caterpillar defenses against their
geometrid moths in an extremely species rich mountain natural enemies. Ecology 83: 3108 Ð3119.
rainforest in South Ecuador. Deutsche Gesellschaft für Goodger, D. T., and A. Watson. 1995. The Afrotropical tiger-
allgemeine und angewandte Entomologie, Düsseldorf, moths: an illustrated catalogue with generic diagnoses
Germany. and species distribution of the Afrotropical Arctiinae
Brehm, G., L. M. Pitkin, N. Hilt, and K. Fiedler. 2005. Mon- (Lepidoptera: Arctiidae). Apollo Books, Stenstrup, Den-
tane Andean rain forests are a global diversity hotspot of mark.
geometrid moths. J. Biogeogr. 32: 1621Ð1627. Hampson, G. F. 1914. Catalogue of the Lepidoptera Pha-
Common, I.F.B. 1990. Moths of Australia. Melbourne Uni- laenae in the British Museum. Supplement 1: Catalogue
versity Press, Melbourne, Australia. of the Amatidae and Arctiadae (Nolinae & Lithosianae)
Conner, W. E. 2009. Tiger moths and woolly bears: behav- in the collection of the British Museum. British Museum
ior, ecology, and evolution of the Arctiidae. Oxford Uni- (Natural History), London, United Kingdom.
versity Press, New York. Heppner, J. B. 1991. Faunal regions and the diversity of
DaCosta, M. A., and S. J. Weller. 2005. Phylogeny and clas- Lepidoptera. Trop. Lepidopt. 2 (Suppl. 1): 1Ð 85.
siÞcation of Callimorphini (Lepidoptera: Arctiidae: Arc- Hilt, N., and K. Fiedler. 2005. Diversity and composition of
tiinae). Zootaxa 1025: 1Ð94. Arctiidae moth ensembles along a successional gradient
DaCosta, M. A. 2007. Phylogenetic studies of Utetheisa Hüb- in the Ecuadorian Andes. Divers. Distrib. 11: 387Ð398.
ner, the rattle box moth, and other arctiines (Lepidop- Janzen, D. H., M. Hajibabaei, J. M. Burns, W. Hallwachs, E.
tera: Noctuoidea: Arctiidae). Ph.D. dissertation Univer- Remigio, and P.D.N. Hebert. 2005. Wedding biodiver-
sity of Minnesota, St. Paul. sity inventory of a large and complex Lepidoptera fauna
Dietz, IV, R. E. 1994. Systematics and biology of the genus with DNA barcoding. Philos. Trans. R. Soc. B 360: 1835Ð
Macrocneme Hübner (Lepidoptera : Ctenuchidae). Univ. 1845.
Calif. Publ. Entomol. 113: 1Ð121. Lafontaine, J. D., and C. Schmidt. 2010. Annotated check
Dietz, IV, R. E., and W. D. Duckworth. 1976. A review of list of the Noctuoidea (Insecta, Lepidoptera) of North
the genus Horama Hübner and reestablishment of the America north of Mexico. Zookeys 40: 1Ð239.
genus Poiliopastea Hampson (Lepidoptera: Ctenuchi- Novotny, V., P. Drozd, S. E. Miller, M. Kulfan, M. Janda, Y.
dae). Sm. C. Zool. 215: 1Ð53. Basset, and G. D. Weiblen. 2006. Why are there so many
Dognin, P. 1913. Hétérocères nouveaux de lÕAmérique du species of herbivorous insects in tropical rainforests? Sci-
Sud. Mém. Soc. Entomol. Belg. 22: 1Ð54. ence 313: 1115Ð1118.
Dyer, L. A., M. S. Singer, J. T. Lill, J. O. Stireman, III, G. L. Piñas, F. S., S. B. Rab-Green, G. Onore, and I. Manzano.
Gentry, R. J. Marquis, R. E. Ricklefs, H. F. Greeney, D. L. 2000. Mariposas del Ecuador. vol. 20. Familia Arctiidae.
Wagner, H. C. Morais, et al. 2007. Host speciÞcity of Subfamilias: Arctiinae y Pericopinae. PontiÞcia Univer-
Lepidoptera in tropical and temperate forests. Nature sidad Católica del Ecuador, Quito, Ecuador.
448: 696 Ð 699. Rothschild, L. W. 1910. Catalogue of the Arctianae in the
Dyer, L. A., T. R. Walla, H. F. Greeney, J. O. Stireman, III, Tring Museum, with notes and descriptions of new spe-
and R. F. Hazen. 2010. Diversity of interactions: A met- cies. Novitates Zool. 17: 1Ð 85.
ric for studies of biodiversity. Biotropica 42: 181Ð189. Rothschild, M., T. Reichstein, J. von Euw, R. Aplin, and
Ferguson, D. C. 1985. Contributions toward reclassiÞcation R.R.M. Harman. 1970. Toxic Lepidoptera. Toxicon 8:
of the world genera. The tribe Arctiini, part 1: Introduc- 293Ð299.
tion and revision of the Neoarctia-Grammia group (Lep- Schaus, W. 1901. Descriptions of some new species of Het-
idoptera: Arctiinae). Entomography 3: 181Ð275. erocera. Ann. Mag. Nat. Hist. 7: 265Ð270.
1148 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 104, no. 6

Schmidt, B. C. 2007. Systematics of Grammia tiger moths British Museum (Natural History), London, United King-
(Lepidoptera: Noctuidae). Ph.D. dissertation, University dom.
of Alberta, Edmonton, AB, Canada. Watson, A. 1975. A reclassiÞcation of the Arctiidae and Cte-
Scoble, M. J. 1992. The Lepidoptera: form, function and nuchidae formerly placed in the thyretid genus Automolis
diversity. Oxford University Press, Oxford, United King- Hübner. Bull. Br. Mus. (Nat. Hist.) (Entomol.) Suppl. 25:
dom. 1Ð104.
Seitz, A. 1925. Syntomidae: general topics, vol. 6, pp. 33Ð320. Watson, A. 1977. A review of the suffusa species-group of
In A. Seitz (ed.), The Macrolepidoptera of the world. Hypercompe Hübner, with a description of a new species
Alfred Kernen, Stuttgart, Germany. affecting Gossypium hirsutum L. (Cotton). Rev. Fac.
Simmons, R. B. 2009. Adaptive coloration and mimicry, pp. Agron. Univ. Cent. Venez. 9: 137Ð147.
115Ð126. In W. E. Conner (ed.), Tiger moths and woolly Watson, A. 1980. A revision of the Halysidota tesselaris spe-
bears: behavior, ecology, and evolution of the Arctiidae. cies-group. Bull. Br. Mus. Nat. Hist. Entomol. 40: 1Ð 65.
Oxford University Press, New York. Watson, A., and D. T. Goodger. 1986. Catalogue of the Neo-
Simmons, R. B., and S. J. Weller. 2002. What kind of signals tropical Tiger moths. Occas. Pap. Syst. Entomol. Br. Mus.
do mimetic tiger moths send? A phylogenetic test in tiger Nat. Hist. 1: 1Ð71.
moths (Lepidoptera: Arctiidae: Euchromiini). Proc. R. Weller, S., M. DeCosta, R. Simmons, K. Dittmar, and M.
Soc. Lond. Biol. Lett. 269: 983Ð990. Whiting. 2009. Evolution and taxonomic confusion in
Stehr, F. W. 1993. Immature insects. Kendall/Hunt, Arctiidae, pp. 11Ð30. In W. Conner (ed.), Tiger moths and
Dubuque, IA. woolly bears: behavior, ecology, and evolution of the
Stireman, III, J. O., L. A. Dyer, D. H. Janzen, M. S. Singer, Arctiidae. Oxford University Press, New York.
J. T. Lill, R. J. Marquis, R. E. Ricklefs, G. L. Gentry, W. Winter, W. D. 2000. Basic techniques for observing and
Hallwachs, P. D. Coley, et al. 2005. Climatic unpredict- studying moths and butterßies. Lepidopt. Soc. Mem. 5:
ability and parasitism of caterpillars: Implications of 1Ð 444.
global warming. Proc. Natl. Acad. Sci. U.S.A. 102: 17384 Ð Zahiri, R., I. J. Kitching, J. D. Lafontaine, M. Mutanen, L.
17387. Kaila, J. D. Holloway, and N. Wahlberg. 2010. A new
Stireman, III, J. O., and M. S. Singer. 2003. Determinants of molecular phylogeny offers hope for a stable family level
parasitoid-host associations: Insights from a natural ta- classiÞcation of the Noctuoidea (Lepidoptera). Zool. Scr.
chinid-lepidopteran community. Ecology 84: 296 Ð310. 40: 158 Ð173.
Wagner, D. 2005. Caterpillars of eastern North America: a Zaspel, J. M. 2004. Phylogenetic and faunistic studies of the
guide to identiÞcation and natural history. Princeton Uni- tiger moth genera Virbia Walker and Holomelina Her-
versity Press, Princeton, NJ. rich-Schäffer (Lepidoptera: Arctiidae). M.S. thesis, Uni-
Wagner, D. L. 2009. The immature stages: structure, func- versity of Minnesota, Minneapolis.
tion, behavior, and ecology, pp. 31Ð53. In W. E. Conner Zaspel, J. M., and S. J. Weller. 2006. Review of the generic
(ed.), Tiger moths and woolly bears: behavior, ecology, limits of the tiger moth genera Virbia and Holomelina
and evolution of the Arctiidae. Oxford University Press, (Arctiidae: Arctiinae: Arctiini). Zootaxa 1159: 1Ð 68.
New York.
Walker, F. 1865. List of the Specimens of lepidopterous
insects in the collection of the British Museum part 31. Received 24 October 2010; accepted 20 May 2011.

You might also like