Lepidoptera: Geometridae: Sterrhinae: Cyllopodini)

REVISION OF THE GENUS CYLLOPODA (LEPIDOPTERA: GEOMETRIDAE: STERRHINAE: CYLLOPODINI)

DELANO ST. AUBYN LEWIS

A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2006

Delano St. Aubyn Lewis

"He that is at peace with his God is at peace with himself and the world."

Delano S. Lewis

ACKNOWLEDGMENTS

This study could not be accomplished without the expert guidance and support

from my graduate committee. I thank Dr. Thomas Emmel for his untiring efforts at

making my sojourn at the University of Florida a smooth one. The expert advice and

hands-on training received from Dr. Charles Covell Jr. were without equal. Dr. Andrei

Sourakov’s many practical suggestions also greatly enriched my experience in

conducting this revision.

I would also like to express my sincere gratitude to the staff of the McGuire Center

for Lepidoptera and Biodiversity, Florida Museum of Natural History, for their helpful

considerations, advice and discussions, and Mr. Alan Chin-Lee for his assistance with

photography. I would be negligent not to mention that my interactions with personnel at

the National Museum of Natural History-Smithsonian Institution, American Museum of

Natural History, Naturhistoriska Riksmuseet, Zoologische Staatsammlungen, The Natural

History Museum and El Instituto National de Biodiversidad made an otherwise

impossible task much easier. Dr. Patricia Gentili-Poole was particularly helpful, making

my visit to the Smithsonian Institution most profitable. I also would like to thank my wife

for her patience and understanding and my family for their continued support in all my educational endeavors.

TABLE OF CONTENTS

ACKNOWLEDGMENTS ...... iv

LIST OF FIGURES ...... vi

ABSTRACT...... vii

CHAPTER

1 INTRODUCTION ...... 1

The Family Geometridae ...... 1 The Subfamily Sterrhinae...... 4 The Tribe Cyllopodini ...... 5 The Genus Cyllopoda ...... 6 Scope of the Project...... 7

2 REVISION OF THE GENUS CYLLOPODA...... 9

Introduction to the Genus ...... 9 Materials and Methods ...... 9 Method of Identification...... 11 Historical Background...... 11 Cyllopoda Dalman...... 12 Key to the Species of Cyllopoda ...... 14 The Species of Cyllopoda...... 16

3 DISCUSSION...... 61

Synonymic Checklist...... 64

REFERENCES ...... 67

BIOGRAPHICAL SKETCH ...... 72

LIST OF FIGURES

Figure page

2-1 Wing venation in Cyllopoda...... 52

2-2.1 Cyllopoda adults...... 53

2-2.2 Cyllopoda adults (continued)...... 54

2-3.1 The male genitalia and aedeagus of Cyllopoda ...... 55

2-3.2 The male genitalia and aedeagus of Cyllopoda (continued)...... 56

2-3.3 The male genitalia and aedeagus of Cyllopoda (continued)...... 57

2-4 Female genitalia of C. claudicula...... 58

2-5 Female genitalia of C. gibbifrons ...... 59

2-6 Female genitalia of C. jatropharia jatropharia...... 60

2-7 Female genitalia of C. osiris osiris ...... 60

3-1 Holotype of Atyria albifrons...... 66

Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science

REVISION OF THE GENUS CYLLOPODA (LEPIDOPTERA: GEOMETRIDAE: STERRHINAE: CYLLOPODINI)

Delano St. Aubyn Lewis

December 2006

Chair: Thomas Emmel PhD Major Department: Entomology and Nematology

This revision is the first part of a comprehensive revision of the tribe Cyllopodini.

Taxonomic relationships, evolutionary history, mimicry, and biogeographic influences on convergence are future studies that hinge on this work. Some of the synonymy that exists in the genus Cyllopoda is resolved and a contribution to a better understanding of the relationships within this genus is accomplished. Morphological taxonomic techniques were used, leading to: four new synonymies, Cyllopoda versicolor, Cyllopoda claudicula catabathmus, Cyllopoda ovata and Cyllopoda protmeta eurychoma; the revision and elevation to species level of Cyllopoda osiris; the use of new combinations Cyllopoda osiris osiris and Cyllopoda osiris protmeta; the designation of a neotype for Cyllopoda osiris; the designation of lectotypes for Cyllopoda angusta, Cyllopoda claudicula,

Cyllopoda claudicula catabathmus, Cyllopoda jatropharia puta, and Cyllopoda postica;

and the designation of paralectotypes for Cyllopoda angusta, Cyllopoda claudicula,

Cyllopoda jatropharia puta, and Cyllopoda postica.

vii CHAPTER 1 INTRODUCTION

The Family Geometridae

The family Geometridae is one of the three largest families of the order

Lepidoptera with a global representation of about 21,000 described species (Scoble et al.

1995; Minet & Scoble, 1999). This family belongs in the super-family Geometroidea along with Drepanidae and Uraniidae, and is subdivided into six sub-families. The moths of this family are broad-winged, butterfly-like in appearance and relatively easy to separate from other moths. When the adults of some species rest, the wings are held outspread (not folded roof-like over the body). The wings in many other species may be held vertically with the dorsal surface of the wings touching. Adults can be most effectively recognized and defined by the presence of paired tympanal organs at the base of the abdomen, which are very distinctive (Cook and Scoble, 1992; Kennel and Eggers,

1993; Minet, 1983). They lie in deep ventrolateral walled cavities called cavi tympani

(Cook and Scoble 1992) but may be reduced or lost in brachypterous females (Minet &

Scoble, 1999).

Geometridae eggs in many species are laid horizontally, with longitudinal axis lying parallel, or almost parallel, to the substrate. However, those of some other species

(some Sterrhinae) are laid in a barrel-shaped fashion with the longitudinal axis perpendicular to the substrate. These two types of egg orientation may occur in the same genus. Egg shape may be ovoid or elliptical when viewed from the lateral side, and circular or oval when viewed from above (Minet & Scoble, 1999), and they may or may

1 2

not be sculptured. They are laid singly or in batches wrapped around twigs, usually glued

to plant substrate, inserted in crevices of bark, or simply dropped from the female

positioned over or near the host plant (Minet & Scoble, 1999). No single character or

combination of characters is known that can be used to separate eggs of Geometridae

from eggs of other families in Lepidoptera.

While it is easy to tell the adults apart from other closely related species, the best character used to differentiate the family is found in the larvae. Geometrid larvae are distinctive from those of other families because of their looping mode of locomotion

(Noctuidae has some semi-loopers). They move this way because they have only two pairs of abdominal pro-legs. In fact, the derivative of the mane Geometridae comes from the looping habit of the larvae, which in the original Greek means “earth measuring”. The larvae are typically cylindrical with slender bodies, the length of which is about 10 times the width (Minet & Scoble, 1999). At times, they may be stout and flattened. The cuticle is smooth or granular and there are often stripes running the length of the body. Humps, filaments or other processes may be present. A full complement of primary setae is present but secondary setae are not usually very pronounced. In a few genera, there is an extra setum above the spiracle, and some have it below. The spiracles are elliptical with the T1 and T8 spiracles larger than the rest. The mature larvae range in size from 15 to 60 mm. The head in the first instar is rounded with this shape changing in later instars to a more rectangular or bifid shape. The mouthparts are hypognathous, and the spinneret is usually shorter than the prementum. The front-clypeus is as wide as long or wider than long with the margins straight and extending no more than two-thirds the epicranial notch. Ecdysial lines are very close to front-clypeal margins, meeting just above the

3 frontoclypeal apex. Six stemmata (simple larval eyes) are present. On the thorax, the T1 shield is indistinct. D1 and D2 are present on all segments. SD2 is caudad and very close to SD1. XD1, XD2, and SD1 are usually in a fairly vertical line. SD1 is sometimes brighter but with a smaller diameter than SD2. L group is bisetose on T1, and trisetose on

T2 and T3. The SV group is bisetose on T1, and unisetose on T2 and T3. The pro-legs are typically found only on abdominal segments A6 and A10, the others having been lost.

Some species may have a full complement of pro-legs with some reduction in size. If there is uncertainty about the familial identification of the larva, a closer examination of the abdomen can be done. Due to the mode of movement employed by geometrid larvae, all are ‘exposed’ feeders (as opposed to borers and miners). Some live in folded leaves or use debris to cover themselves. They feed primarily on deciduous trees and shrubs, but some feed on ferns, coniferous foliage and coniferous seeds. They often exhibit twig and stipule mimicking behavior where the body is held out at an angle from the twig. Many geometrids are common, but few are economically important. They are not a major pest group, but some are pests of fruit trees and a few can be extremely damaging to forest trees. Their major ecological impact can be seen in the tropics or subtropics where most species exist. Given appropriate conditions, certain species can reach pest status because of their defoliation capabilities. They, however, have great environmental significance as primary consumers and as food for parasitoids and vertebrate predators; also, because they have a tendency towards habitat specificity, they are good indicators of environmental change (Minet & Scoble, 1999).

The pupae are typically lepidopteran - the obtect type, lacking exposed maxillary palpi and mandibles. They are exposed, with no cocoon, or they are concealed in detritus.

At times, they may be suspended from the tail (cremaster) with no median silk girdle, or not suspended but attached at the tail and secured by a median girdle of silk. Some are subterranean while others pupate on the surface of the ground. Typically, pupation occurs in a flimsy cocoon in debris or in the soil from which they do not protrude.

The adults are small to large moths that are primarily nocturnal, but some are crepuscular and others diurnal. Most have slender bodies of medium length, but short and long bodied individuals do exist. The abdomen is often plain, but can be brightly patterned. The antennae range from short to medium in length (0.4-0.6 times length of forewing), and in males may be bipectinate, dentate, simple, simple-ciliate, or fascicle- ciliate (fasciculate). Antennae are usually simple in the females. They have a fully developed un-scaled proboscis, no ocelli, and hairy eyes. The maxillary palps usually have one segment, but sometimes two; labial palps are ascending. Chaetosemata are present. The wingspan ranges from 16 to 69 mm (10-19 times width of thorax). The forewings are usually broad (about 2 times as long as wide), apically blunt or pointed, and the hindwings range from being narrower than the forewings to about the same breadth, and are broadly rounded or broadly angular. The venation of the forewings and hindwings differ (Hausmann, 2001), but the forewing ground color and lines often continue through the hindwings. Discal spots are often present.

The Subfamily Sterrhinae

The Sterrhinae are often referred to as ‘waves’ because of the narrow, uneven lines on their wings (Sihvonen and Kaila, 2004). There are about 2,800 described species in this group and its species diversity decreases the farther one goes from the equator. They are described as mostly slender bodied, concolorous, pink or pale fawn, with many white or yellow. Fine lines extend across the forewing and continue across the hindwing. It is

unclear if the fusing of the Sc+R1 and Rs veins near the wing base, before their sharp divergence, can be used to effectively diagnose the subfamily (Minet & Scoble, 1999).

The discal spot is frequently present in both pairs of wings and they usually have a reduced uncus or gnathos (Minet & Scoble, 1999). Some Sterrhinae lack discal spots in both wings and may have a bright atypical yellow and black pattern as seen in the diurnal

Cyllopodini (Covell, 1983); however, most Sterrhinae are nocturnal and conform to the general pattern of having discal spots.

This is a poorly defined subfamily, with literature on their taxonomic classification into tribes being scattered and very limited in scope (Sihvonen and Kaila, 2004). The concentration on regional faunas has led to the absence of a global integrated tribal structure for this subfamily (Hong-Xiang et al 2004). Many family group names have been introduced without descriptions because historically, most of the data concerning their phylogeny was deduced from checklists and other similar works (Sihvonen and

Kaila, 2004).

The most widely accepted treatment of this subfamily arranges seven tribes into two lineages. This classification is based predominantly on the anatomy of the male and female abdomen. Larval and pupal traits also support this treatment (Sihvonen, 2004).

Genetic analysis of this group is also limited in scope and the keys that exist for the tribes of Sterrhinae do not resolve the relationships between the tribes (Sihvonen and Kaila,

2004). The latest treatment of the Subfamily Sterrhinae is Sihvonen and Kaila (2004), which includes a detailed look at the history of its tribal classification.

The Tribe Cyllopodini

The Cyllopodini are a group of mimetic day-flying moths with a distinctive bright yellow and black coloration of the fore and hindwings. Restricted to Central and South

America, they are often found mixed with unrelated but very similar looking Dioptinae

moths in collections (Prout, 1938) and can also be seen in nature flying with similar

looking species. This behavior suggests that this coloration may be aposematic and there

seems to be considerable convergence on one particular patterning, or variations thereof, in this tribe, suggesting the conferral of some protective function. Prout in Seitz (1938) recognized that there is considerable divergence among the genitalia of some individuals in this group which will likely impact future taxonomic studies.

This tribe consists at present of the genera Cyllopoda Dalman, Atyria Hübner,

Atyriodes Warren, Formiana Druce, Paratyria Warren, Smicropus Warren, and Xanthyris

Felder & Felder. The genera Myrice Walker and Oncopus Herrich-Schaffer are placed at the end of the series of Cyllopodini by Prout (1934), but their association with the tribe is uncertain. Since the description of this tribe by Prout (1938), no revisionary studies have been made of this group. Scoble (1999) suggested that much revisionary taxonomy is needed at the generic and species levels for the family Geometridae, arguing that such work would probably be the most valuable contribution that can be made to geometrid systematics at this time.

The Genus Cyllopoda

The genus Cyllopoda Dalman, 1823, is the type genus of the tribe Cyllopodini. The adults differ from the other members of the tribe and subfamily in that they possess the unique combination of the following characters: moderate to long palpi; bipectinate male and simple female antennae; the male hindtibia is either spurless or with vestigial spurs and may be more or less modified, and usually with hair a pencil; the female hindtibia has only two apical spurs; the forewing has the 2nd subcosta arising from the discal cell

and the hindwing has the 2nd subcosta stalked; there are two areoles on the forewing, the

second one often being small (Prout in Seitz, 1931), sometimes absent. The type species

for this genus is Cyllopoda claudicula Dalman, 1823, from Brazil with the holotype

deposited in Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRS).

Scope of the Project

This study is the first part of what I hope will be a comprehensive revision of the

tribe Cyllopodini, with future work focusing on phylogenetic studies, mimicry, chemical

ecology, and gaining knowledge of their life histories. Any study in which modern

revisionary techniques are employed, such as investigation of genitalic structures or

phylogenetic analyses, will be instrumental in resolving some of the synonymy that exists

in the family (Scoble, 1999), and will contribute to a global integration of the tribal

structure for this subfamily (Hong-Xiang et al 2005). This current revision of the genus

Cyllopoda will be carried out by means of morphological taxonomic techniques only.

Genetic techniques are an important supplement to morphological techniques and I intend

to pursue genetic analysis at a later date and on a much wider scope including closely

related tribes.

Porter (1986) argues that adult characteristics present numerous problems for phylogenetic reconstruction because “superficial” characters show much parallelism, plesiomorphy, and convergence. He also states that the use of the adult genitalia may pose problems as they are often very similar within genera and very different between tribes, so that there is a loss of phylogenetic significance. He used many larval characters in his study of the tribe Nemoriini. However, it is very difficult to obtain immature stages from a representative cross section of the Cyllopodini since it is neotropical. There is almost no life history information in the literature because of the difficulty in securing

8 living individuals for rearing from the neotropics and almost no rearing has been done there. Attempts will be made to study immature stages if living material can be obtained.

Mimicry complexes of butterflies and some species of diurnal moths have been investigated (Sheppard et al. 1985; Brown, 1988; Beccaloni, 1997; Vane-Wright et al.,

1999); however, no work has been published on mimicry complexes involving this tribe of geometrids. It has been established that the yellow and black coloration pattern in nature functions as aposematic or warning coloration (Rowe and Skelhorn, 2005). This aposematic coloration seen in the adults in this tribe may be a consequence of them being distasteful and involved in Müllerian mimicry, a result of them being Batesian mimics of similarly patterned species, or a combination of both types of mimicry. Investigation of the mimicry patterns and behavior of the adults and larvae of Cyllopodini is one of the future projects that would naturally follow the present study and which no doubt would contribute significantly to geometrid systematics and ecology as well as our knowledge of the nature of mimicry.

CHAPTER 2 REVISION OF THE GENUS CYLLOPODA

Introduction to the Genus

The genus Cyllopoda is the type genus of the tribe Cyllopodini which has six other

included genera (Atyria, Atyriodes, Formiana, Paratyria, Smicropus, and Xanthyris), and

a total of 93 currently recognized species and subspecies. Two other genera, Myrice and

Oncopus, are of uncertain association with the tribe and include seven species and one

subspecies. The genus Cyllopoda currently includes 15 species and five subspecies. The

majority of species now recognized in Atyria were formerly placed in Cyllopoda because

of their close resemblance.

Members of this genus seem to be involved in an extensive mimicry complex with

other members of the tribe Cyllopodini as well as other Geometridae, their color patterns

being aposematic. A few species from the families Notodontidae (Miller 1991),

Hesperiidae, Nymphalidae, Arctiidae, Riodinidae, Noctuidae, and Pyralidae, as well as

other Lepidoptera, may also be involved in this complex. The mimicry complex involved

in the Cyllopodini has not yet been investigated and no life history information on this

genus or the tribe has been published. Members of this genus can be found mainly in

South America from Colombia to Bolivia in the south and east to Suriname and Brazil.

They can also be found in Central America and Trinidad and Tobago.

Materials and Methods

Most of the material studied was borrowed from the following institutions:

National Museum of Natural History, Smithsonian Institution, Washington DC, USA

9 10

(USNM); American Museum of Natural History, New York, NY, USA (AMNH);

Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRS); Zoologische

Staatsammlungen, Munich, Germany (ZSBS); The Natural History Museum, London,

UK (BMNH); Instituto National de Biodiversidad, San Jose, Costa Rica (INBio);

Pontificia Universidad Catolica del Ecuador, Quito, Ecuador (PUCE); and Dr. Jack

Schuster, Guatemala. The remaining material is the property of the McGuire Center for

Lepidoptera and Biodiversity, Florida Museum of Natural History, Gainesville, FL, USA.

Identification of the material was accomplished by comparison with photographs of the type species in the BMNH, the NRM, and the Museum of Evolution, Uppsala,

Sweden, and with photographs and examination of type specimens in the USNM.

To study the genitalia, preparations were made by first exposing the abdomen to

10% Potassium Hydroxide (KOH). After maceration, the abdomens were denuded of

scales and stained with chlorazol black. The genitalia were then separated from the

abdominal pelt and both were temporarily stored in alcohol or glycerol to allow

examination from various perspectives. They were afterwards mounted on microslides in

Euparol. To study the wing venation, wings were denuded of scales and examined under

a stereoscopic microscope equipped with a camera lucida. The terminology for male and

female genitalia follows Covell (1970) and Klots (1970), and the terminology for wing

venation follows Hausmann (2001). Measurements of forewing and genitalia follow

those of Covell (1970). Images of the genitalia were captured using a Micro-optics system with a Canon EOS 1D Mark II camera and processed in Adobe Photoshop

Elements®. Drawings of wing venation were made by means of a stereoscopic microscope equipped with a camera lucida. Illustrations of the genitalia were done by

11 placing Vellum-Accent paper (Translucent Clear, 20lbs) over black and white prints of the photographs to get the outline. The illustrations were then completed by looking at the actual specimens to better understand their three dimensional structure.

Specimen data are recorded as they appear on labels. The information on each label is enclosed with double quotes (“), lines on each label are separated by a forward slash

(/). Information on separate labels is separated by a comma (,) before the double quotes.

Additional information is enclosed in square brackets ([]), and information for different specimens are separated by a semicolon (;).

Method of Identification

The identification of the specimen was done by comparing photographs of the type specimen at the Natural History Museum (BMNH), Naturhistoriska Riksmuseet

(NHRS), or the National Museum of Natural History (USNM) to each specimen.

Additionally, types at the USNM were viewed in person. Original descriptions were also used, along with plate 17 from Prout (1938). In addition to type material, 51 specimens were identified as Cyllopoda and studied for this revision.

Historical Background

The genus Cyllopoda was described by Dalman (1823) with the type species

Bombyx claudicula (Dalman) 1823. The description of the genus was, however, preceded by the description of Phalaena jatropharia by Linnaeus in 1758 and Phalaena osiris by

Cramer in 1777. These are now included in Cyllopoda, having been moved to that genus upon its creation. In the same year that the description of the genus was published (1823),

Hübner described what he called Atyria jatrophae, which was later recognized as an emendation of Cyllopoda jatropharia (Linnaeus). Walker (1854) described a new genus,

Flavinia, now recognized as a synonym of Cyllopoda, and a species Chrysauge postica,

now recognized as Cyllopoda postica (Walker). In 1885, Druce described Flavinia

roxana, now recognized as Cyllopoda roxana (Druce). From this point on, the genus

name Cyllopoda was exclusively used, signifying its acceptance by the scientific

community. The only other synonym described in this time period was Cyllopoda

latimargo Warren (1897), a synonym of Cyllopoda osiris (Cramer). The remaining

species were all described in Cyllopoda from 1897 to 1938 by Warren, Dognin, Prout,

and Strand. Prout (1938) was the only comprehensive treatment of this group to date.

Cyllopoda Dalman

Cyllopoda Dalman, 1823: 102. Walker, 1856: 1778. Warren, 1895: 84, 85. Prout, 1910:

239, 240. Sick, 1937: 400-412. Prout, 1934: 131. Scoble, 1999: 214. Type

claudicula Dalman by original designation.

Flavinia Walker, 1854: 369. Druce, 1885: 150. Type postica Walker by original

designation.

Diagnosis: Male antenna bipectinate, female simple. Male hindleg reduced, with

or without hair pencils, lacking spurs, or with modified spurs in the form of two apical

lobes on the tibia; female hind tibia with two apical spurs. Two areoles in the front wing.

The forewing has the 2nd subcosta arising from the discal cell and the hindwing has the

2nd subcosta stalked.

Description: Frons sometimes protuberant, mostly black, sometimes white or

yellow. Interantennal ridge and antennal shaft black with or without sprinkling of yellow

or white scales at the base of antenna. Male antenna bipectinate, female simple; collar

black with or without sprinkling of yellow and white scales; palpus porrect, smooth.

Thorax above black; pectus white, yellow or black; tegula, yellow, black, or with yellow

at the base only, with long black hair scales at the tips. Legs white, tan or black with or

without suffusion; male hindleg reduced with or without hair pencil on the tibia, tarsi

may be reduced or modified, spurs absent or modified as round apical projections on the

tibia; female hindleg with two apical spurs.

Venation typically Sterrhinae with one or two areoles (Figure 2-1). Wings from

above almost always with a prominent bar on the forewing, and almost always without a

black bar on the hindwing, black border may or may not surround the entire wing.

Forewing with a rounded tip, Pattern almost always repeated below (Figure 2-2)

Male genitalia (Figure 2-3): At least twice as long as it is wide; Tegumen slightly to moderately sclerotized; socii as short oval lobes or as long, petiolate lobes with short to long hairs, in some the socii are not immediately obvious; gnathos moderately to heavily sclerotized, usually arising as two arms from the lateral edges of the teguman but may also arise as a cone as in the nominate species; uncus slightly to moderately sclerotized, lateral sides curved or straight, sometimes with an indentation at the base, the tip may be cleft to produce a bifid appearance, may be expanded greatly laterally or it may be simple, sometimes spoon-shaped, the lateral edges often with long hairs; valva simple, lightly setose, the last third usually thinner than the first converging into sharp point; tip of valva may be slightly or greatly curved dorsally; dorsal margin of the valva may possess an area with teeth midway along its length; vinculum moderately to heavily sclerotized, surrounding the base of the valva; saccus membranous giving rise to the annulus which may be membranous, slightly sclerotized or moderately sclerotized; top of the annulus may be separated into two halves; juxta slightly to moderately sclerotized projecting cephalad of the vinculum; aedeagus slightly to greatly curved, often appearing u-shaped, occasionally almost sigmoid, portions slightly to heavily sclerotized; tip

14 pointlike, or almost same diameter as the anterior portion; the ductus ejaculatorius entering at the posterior end, close to the posterior end or almost one thirds the distance from the posterior end. Cornuti present or absent.

Female genitalia (Figures 2-4 to 2-7): Long, with simple papilla anales containing short to elongated hairs; apophyses posteriores and apophyses anteriores slightly thickened, the apophyses posteriores two to four times as long as the apophyses anteriores; genital plate circular or v-shaped just caudad to the ostium bursae or leading straight into the ostuim bursae; ductus bursae membranous to heavily sclerotized; ductus seminalis meets the ostium bursae about midpoint, a slightly sclerotized sac may be located where the ductus seminalis joins the ostium bursae; corpus bursae sac-like, with a paired signum; signum as rows of short hooks or a row of large spines extending almost along the entire length.

Variation: Some individuals may have the basal areole much smaller in comparison to the proximal. Others may only have one areole. The frons in some are more protuberant than others.

Immature Stages: Unknown.

Distribution: This genus is Neotropical. Its members are predominantly in South

America, with a few species in Central America and Trinidad and Tobago.

Key to the Species of Cyllopoda

1. Wings yellow and black . . . . . (2)

Wings orange and black . . . . . roxana

2. Wings black with a large yellow spots on the forewing and two smaller spots on

the hindwing ...... angusta

Wings predominantly yellow with black wing borders . (3)

3. Forewing and hindwing above with a prominent bar . . (4)

Forewing with prominent bar from above, absent in hindwing (5)

4. Bar in hindwing repeated below . . . . claudicula

Bar in hindwing not repeated below . . . . radiata

5. Black border on forewing on all margins . . . (6)

Black border on forewing not along inner margin . . (8)

6. Hindwing with black border on all margins . . . (7)

Hindwing with black border on outer margin only . . (9)

7. Forewing with a bright black line from wing base to almost middle of bar, along

Cu ...... nigrivena

Black line along Cu pale . . . . . expansifascia

8. Abdomen with yellow lateral lines of narrowing posteriorly postica

Abdomen without yellow lateral lines . . . (10)

9. Basal patch of yellow scales in forewing half wing length . bipuncta

Basal yellow patch in forewing less than half wing length . (11)

10. Pectinations of male antenna long, socii short and oval . (12)

Pectinations on antenna short, socii long, petiolate / corpus bursae with one row

of large spines on each side . . . . . gibbifrons

11. Basal patch of yellow scales in forewing triangular . . latiflava

Basal patch of yellow scales in forewing one third wing length and almost oval in

shape ...... breviplaga

12. Hindwing borders broad, at times almost one third the wing . (13)

Hindwing borders never broad, less than one quarter the wing (14)

13. Forewing with thick black bar . . . . osiris osiris

Forewing without bar ...... osiris protmeta

14. Black wing borders less than one quarter wing length jatropharia jatropharia

Black wing borders one quarter wing length . . jatropharia puta

The Species of Cyllopoda

Cyllopoda claudicula (Dalman)

(Figures 2-2.1.E and 2-2.1.F)

Bombyx (Cyllopoda) claudicula Dalman 1823: 102. Walker, 1856: 1778. Kirby, 1892:

403. Prout, 1916: 173; 1934: 132. Sick, 1937: 400- 412.

Callimorpha dichroa Perty, 1833: 161.

Chrysauge dichroa: Walker, 1854: 371. Kirby, 1892.

Cyllopoda claudicula catabathmus Prout 1938: 120. Scoble, 1999: 214. syn. nov.

Cyllopoda claudicula claudicula: Prout, 1938: 120. Scoble, 1999: 214.

Cyllopoda dichroa: Prout, 1916: 174.

Flavinia dichroa: Strand, 1920: 136.

Diagnosis : This species can be differentiated from the others in this genus by the black bar present on the hindwing that is repeated on the underside. The hindleg of the male is much more reduced than the others and lacks a hair pencil.

Male (Figure 2-2.1.E): Frons black with yellow and white scales near labial palpus; interantennal ridge and antennal shafts black; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus black, almost same length as frons.

Thorax with yellow lateral lines; pectus white to yellow; tegula yellow, edged with black scales, with long black hair scales at the tips; legs white with tan suffusion on anterior

surfaces of forelegs, base yellow; hind tibia compressed, short, lacking spurs and hair

pencil. Abdomen black dorsally, and yellow ventrally, with median black bar.

Forewing with two areoles, apex rounded. Wings above yellow with broad black

border all around and a prominent black bar in both forewing and hindwing. Bar in

forewing broad, from midpoint costal margin to anal angle; hindwing with bar narrow

from base of wing to broad at anal angle along M. Pattern on the wings repeated on the

underside. Length of forewing 1.7 to 1.8 cm.

Male genitalia (Figure 2-3.1.A): About 3 mm long and 1.5 mm wide. Tegumen

moderately sclerotized; socii length about one-half the length between them, short, oval,

and moderately sclerotized with short hairs; gnathos heavily sclerotized, tip drawn out

into a sharp point, base moderately sclerotized arising from the caudal margin about

midpoint of the tegumen; uncus moderately sclerotized, sides straight, moderately broad,

the tip slightly bifid, with two moderately setose lateral lobes; valva simple, long,

relatively smooth, the last third thinner than the first two thirds, drawn out into lightly setose tips; vinculum moderately sclerotized, v-shaped, surrounding the base of the valva; saccus membranous giving rise to a membranous annulus close to the base of the valva; juxta moderately sclerotized projecting cephalad of the vinculum; aedeagus about 3 mm long, u-shaped in posterior third; moderately sclerotized, the tip pointed and more sclerotized than the next two thirds; the ductus ejaculatorius entering about one-third the distance from posterior end.

Female (Figure 2-2.1.F): Similar to male with simple antenna, lacking lateral yellow lines on the thorax, hind tibia with two apical spurs. The black wing border and

18 bar on the wing are broader and more diffused than in the male. Length of forewing 1.8 to 1.9 cm.

Female genitalia (Figure 2-4): About 4.8 mm long and 1 mm wide. Papilla anales simple and elongated with short hairs; apophyses posteriores and apophyses anteriores slightly thickened, the apophyses posteriores about twice as long as apophyses anteriores; genital plate circular, well defined, just caudad to the ostium bursae; ostium bursae as a circular opening on the eighth abdominal segment; ductus bursae slightly sclerotized leading to the appendix bursae which is leads into the corpus bursae and is hardly distinguishable; ductus seminalis meets the ostium bursae about midpoint; corpus bursae sac-like, with paired signum; signum as rows of short hooks extending almost along the entire lenght, ending about 1 mm from the end of the corpus bursae.

Variation: Some individuals may be slightly more melanized than the typical.

They may have no yellow markings on the tegula or the frons and the legs may be darker.

Slight sexual dimorphism obvious.

Distribution: Blumenau, Santa Caterina, and Rio Janeiro in Brazil; Chile.

Types: I have examined a photograph of a syntype for C. claudicula catabathmus at the BMNH label data: "49. 29./Blumenau/Sta, Cath./Brazil/18.V.'29/F. Schade",

"Cyllopoda/claudicula/catabathmus/♂ type Prout", one of four specimens, three males and one female. I designate the specimen with the above label data as the lectotype for C. claudicula catabathmus. The type series for C. claudicula consists of two syntypes, one male and one female, in NHRS. The original description identifies a male and a female, but does not specify which the holotype is. I designate the male specimen with label data

“Type Dalmani/An. ent. P. 102” as the lectotype for this species. The second specimen, a

female, is not labeled and I designate it as the paralectotype. A third specimen, a female,

with label data “Brazil”, can also be found there. This specimen is most likely not a part

of the type series.

Material examined: "Tijuco", "Cyllopoda/claudicula/Dalm.",

"Rothschild/Bequest/B.M. 1939-1.", "Geometridae/genitalia slide/No. 18465 ♂"

(BMNH); "1939/Chile" [male] (AMNH); "1939/ Brazil", "No. 7739/Collection/Hy.

Edwards" [male] (AMNH); "Callimorpha/dichroa Perty/(x) [male] (NHRS); "Rio

Janeiro", "Flavinia/dichroa/Perty" [male] (NHRS); "Rio Janeiro",

"Collection/WmSchaus" [male] (USNM); "Petropolis, Brazil ", "Collection/WMSchaus"

[female] (USNM); "Rio Janeiro","Collection/ WmSchaus" [ female] (USNM);

"Ty","EdwTOwen/Collection" [ female] [handwriting difficult to read](USNM); "St.

Catherina","Flavina dichroa","Cyllopoda claudicula Dalm", "Rothschild/Bequest/B.M.

1939-1.", "C. claudicula catabathmus" [male] (BMNH).

Method of determination: Examination of photographs of the syntype.

Cyllopoda angusta Warren

(Figure 2-2.1.A)

Cyllopoda angusta Warren 1897: 419. Prout, 1934: 132; 1938: 120. Scoble, 1999: 214.

Diagnosis: Individuals can easily be differentiated from others in the genus in that it has predominantly black wings with one small area of yellow scales on the forewing and two even smaller areas of yellow scales on the hindwing.

Male (Figure 2-2.1.A): Frons, interantennal ridge, and antennal shaft black; male antenna bipectinate; lower parts of the face yellow; tegula black; legs whitish. Abdomen above black, underside whitish.

Wings above predominantly black; forewing with a large, transverse, oblong yellow spot near the tip of the wing but not touching borders; hindwing with two yellow spots, the larger oval spot at the lower end of the cell, the smaller, a short streak almost touching the inner margin close to wing base. Length of forewing 1.8 cm.

Male genitalia: No specimen was secured on loan; description of the adult body made by comparing the photograph of the syntype with the original description.

Female: Unknown.

Distribution: Known only from type locality of Reyes, Bolivia.

Types: Two syntypes, one male and one female are in the BMNH. I have examined the photograph of the male with label data: "REYES,/7.8.95/Stuart.",

"Cyllopoda/angusta/Type ♂ Warr". The original description does not state which of the two individuals the holotype is. I designate the specimen with the above label data as the lectotype of the species and the female as a paralectotype.

Method of determination: Examination of photographs of the male syntype.

Cyllopoda angustistiga Warren

(Figure 2-2.1.B)

Cyllopoda angustistriga Warren 1904: 14. Prout, 1934: 133; 1938: 121. Scoble, 1999:

214.

Diagnosis: Individuals can be separated from other members of the genus by the pattern on the forewing. Similar to C. latiflava in general appearance but differ in having the first third of the forewing almost completely black, with a small patch of yellow scales on the inner margin from the wing base almost to the anal angle.

Male (Figure 2-2.1.B): The type is labeled as a female; however, the antenna

appear bipectinate. I treat it here as a male. Wing pattern similar to Cyllopoda latiflava but differs in that the forewing has the black bar extended into a larger patch, covering

almost the entire basal section of the wing except for a small yellow patch on the inner

margin from the wing base to almost where the anal vein (A) meets the wing margin.

Length of forewing 1.6 cm.

Male genitalia: No specimen was secured on loan; description of the adult body was made by comparing the photograph of the type with the original description.

Female: Unknown.

Distribution: Chiri-Mayo and the River Slucuri in South East Peru; 1000 feet.

Types: I have examined the photograph of the holotype in the BMNH with label data: "Chiri-Mayo/S. E. Peru, 1000'./VII. 01. Dry/(Ockenden)",

"Cyllopoda/angustistriga/Type ♀ Warr." This was designated the holotype by original description. However, I believe this specimen to be a male based on the structure of the antenna. The other specimen examined is a male from the River Slucuri in South East

Peru.

Method of determination: Examination of photographs of the holotype.

Cyllopoda bipuncta Warren

(Figure 2-2.1.C)

Cyllopoda bipuncta Warren, 1906: 409. Prout, 1934: 133; 1938: 120. Scoble 1999: 214.

Diagnosis: Cyllopoda bipuncta has a wing pattern similar to C. breviplaga, C. latiflava, C. jatropharia and C. osiris. It can be differentiated from C. jatropharia and C. osiris by shorter pectinations of its antenna and narrower black borders on the wings. It

also appears to be smaller than the typical specimen of these two species, based on the

materials studied. It can be separated from C. breviplaga and C. latiflava by the length

and width of the basal yellow cell created by the black wing borders and the black bar in

the forewing. This cell is longer (half the wing length) in C. bipuncta than in C. breviplaga (one third the wing length) and narrower in C. bipuncta than in C. latiflava where it is more triangular.

Male (Figure 2-2.1.C): Frons yellow; interantennal ridge and antennal shaft black; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus whitish, shorter than length of frons. Pectus yellow; tegula yellow, edged with black scales, with long black hair scales at the tips; legs white with tan suffusion on anterior surfaces of forelegs, yellow bases; Abdomen black dorsally, white ventrally with lateral yellow lines of scales narrowing posteriorly.

Wing above most closely resembles Cyllopoda breviplaga; differs in that the bar in the forewing is closer to the tip, making the yellow cell produced by it and the border on the inner margin longer than in C. breviplaga. The black border on the hindwing also thicker than in C. breviplaga. Pattern repeated below. Length of forewing 1.6 cm

(wingspan: 3.5 cm).

Male genitalia (Figure 2-3.1.B): About 3 mm long and 1.2 mm wide. Tegumen moderately sclerotized; socii very long, petiolate, lightly sclerotized, about the same length as the distance between them, with moderately long hairs; gnathos moderately sclerotized, arising from the tegumen very close to the socii as two projections that meet medially, forming a pointed tip with two short, small terminal spines; uncus arising just above the socii, moderately sclerotized, sides straight, narrow, the tip slightly bilobed;

23 lateral lobes with short hairs; valva lightly setose, curved at the tips, moderately sclerotized, the dorsal edges abruptly constricting to form the tip about two thirds the distance from the base; vinculum moderately sclerotized, surrounding the base of the valva; saccus membranous giving rise to a moderately to heavily sclerotized annulus that extends to about half the length of the tegumen, splitt where the aedeagus exits; juxta narrow, lightly to moderately sclerotized projecting cephalad of the vinculum; aedeagus about 2.5 mm long, slightly curved, lightly sclerotized, the ductus ejaculatorius entering almost at the posterior end; one cornuti seen.

Female: Unknown.

Distribution: Peru.

Types: The holotype at the USNMH was examined and photographs obtained.

The label data for the holotype is: “Peru", "Type/No. 9163/ U.S.N.M.", "not B.M.”,

"Cyllopoda/bipuncta/Type ♀ [should be ♂?]", "Collection/ WmSchaus."Holotype designated by original description.

Method of determination: Examination of specimen and photographs of the holotype.

Cyllopoda breviplaga Dognin

(Figure 2-2.1.D)

Cyllopoda breviplaga Dognin 1906: 108. Prout, 1934: 133. Sick, 1937: 400 - 412. Prout,

1938: 121

Cyllopoda breviplaga breviplaga: Scoble, 1999: 214.

Cyllopoda breviplaga versicolor: Prout, 1934: 133; 1938: 121. Scoble, 1999: 214.

Cyllopoda versicolor Dognin 1908: 17. syn. nov.

Diagnosis: Similar to C. bipuncta, C. latiflava, C. jatropharia and C. osiris in

wing pattern. It can be differentiated from C. jatropharia and C. osiris based on shorter

pectinations of the antenna and narrower black borders on the wing. It appears smaller

than the typical individuals of these two species, based on the material studied. It can be

separated from C. bipuncta and C. latiflava by the length and size of the basal yellow cell

created by the wing borders and the black bar in the forewing. This cell is shorter in C.

breviplaga (one third the wing length and almost oval) than in C. bipuncta (half the wing length), and much smaller in C. breviplaga than in C. latiflava where it is more triangular.

Male (Figure 2-2.1.D): Frons protuberant, black with white scales around lower edges of eyes; interantennal ridge and shaft black; antenna bipectinate; collar black; palpus black, shorter than length of frons; pectus white to yellow; tegula black throughout with long black hair scales; legs white with tan suffusion on anterior surfaces and yellow at the bases; hind tibia with reduced tarsi, hair pencil present on posterior surface of rod- shaped tibia; no spurs. Abdomen black dorsally, white ventrally with lateral yellow lines of scales narrowing posteriorly.

Forewing with two areoles, apex rounded. Wings above yellow, with broad black border all around in forewing and only at outer margin in hindwing; prominent broad black bar in forewing only, from midpoint costal margin to anal angle; hindwing without bar. Pattern repeated below. Length of forewing 1.8 cm.

Male genitalia (Figure 2-3.1.C): About 3 mm long and 1 mm wide. Tegumen moderately sclerotized; socii short, oval, and moderately sclerotized with long hairs; gnathos moderately to heavily sclerotized, arising from the tegumen close to the socii as

two projections that meet medially into a fan-like tip; tip of the gnathos moderately

sclerotized, base heavily sclerotized; uncus arising just above the socii, moderately

sclerotized, slightly indented laterally, narrow, with hairs at the top of the lateral margins;

uncus with a slight cleft at the top; valva lightly setose, slightly curved at the tip, which makes up last one third of the valva, the remaining two thirds thicker than the tip; vinculum moderately sclerotized, surrounding the base of the valva; saccus membranous giving rise to a moderately sclerotized annulus that extends up to one third the length of the genitalia; juxta moderately sclerotized projecting slightly cephalad of the vinculum, slightly broad; aedeagus about 3.5 mm long, heavily curved, u-shaped, the anterior end with the tip moderately sclerotized, the ductus ejaculatorius entering at about the posterior one quarter.

Female: Unknown.

Variation: Some males may be paler than others.

Distribution: Tarapota and Charape, Peru; Ecuador.

Types: I have examined, and secured photographs of the holotype at the USNM, with label data: "Tarapota/Peru/Aty fis. 03", "Type No./33039/U.S.N.M.", "Dognin/

Collection", "Varite de/quicha/le Crois/and 03/No", "not decided/yet/Warren X.N05",

"Cyllopoda/(breviplaga)/sp. nov./Warren mass 03", "Cyllopoda/ breviplaga/type ♂ Dn.",

"perhaps/described/now ?." This was designated as holotype by original description. I have examined the genitalia and type for C. versicolor, and compared its original description (Dognin, 1908) to notes by Prout (1938). The label data for this specimen is:

“Cyllopoda/versicolor/type ♂ [illegible]”, “Numbala/Equatur /1885/abbi gaujon”,

“Cyllopoda/sp nov. ♂/Warren 10.01”, ”Type No./33040/USNM”, “Dognin/Collection”,

”Cyllopoda/(versicolor)/sp. nov.” This specimen differs from C. breviplaga in color only,

the yellow on the wings being much paler, almost white. I agree with Prout (1938) and

treat this specimen as an albinistic form of C. breviplaga.

Material examined: "Charape: N. Peru/4,000 ft. Sept. Oct. 1912./A. & E. Pratt.",

"breviplaga Dogn.", "Det. By L.B.Prout", "Joicey/Bequest./Brit.Mus./1934-120.", "C. b.

breviplaga" [male] (BMNH).

Method of determination: Examination of the holotype.

Cyllopoda expansifascia Prout

(Figure 2-2.1.G)

Cyllopoda expansifascia Prout 1917: 391-392; 1934: 133; 1938: 321. Scoble, 1999: 214.

Diagnosis: Similar to C. nigrivena, C. claudicula and C. radiata in having the

black border on hindwing all around. Differs from C. claudicula and C. radiata in that it

lacks a black bar in the hindwing and possesses a thin line of black scales from base of

wing to prominent bar along Cu. This line is broader in C. nigrivena.

Male (Figure 2-2.1.G): Frons black; interantennal ridge and shaft black; antenna

bipectinate; collar black; palpus black, shorter than length of frons; pectus white to tan;

tegula black throughout with long black hair scales; legs white with tan suffusion on

anterior surfaces, yellow bases; male hind tibia club-shaped with reduced tarsi, hair

pencil present on posterior surface, spurs reduced in the form of two round apical

projections. Abdomen black dorsally with laterally sprinkling of yellow scales, white

ventrally with lateral sprinkling of yellow scales.

Forewing with two areoles, apex rounded. Wings above yellow, with broad black

border all around and a prominent black bar in forewing only; thin line of black scales

from base of wing to prominent bar along Cu; bar in forewing broad, from midpoint

costal margin to anal angle; hindwing with no bar; a sprinkling of black scales at base.

Pattern repeated below. Length of forewing 1.7 to 2.0 cm.

Male genitalia: Similar to Cyllopoda roxana (Figure 2-3.1.D). About 3 mm long and 1.5 mm wide. Tegumen moderately sclerotized; socii very short, oval, and moderately sclerotized with short hairs; gnathos heavily sclerotized, arising from the tegumen close to the socii as two projections that meet medially, forming a tubular tip; uncus arising just above the socii, moderately sclerotized, slightly tapered towards the tip, narrow, with hairs at the top of the lateral margins; valva lightly setose, slightly curved at the tip, area with two teeth present on the dorsal margin about half way from the tip which is heavily sclerotized, the remaining two thirds moderately sclerotized; vinculum moderately to heavily sclerotized, surrounding base of valva; saccus membranous giving rise to a moderately sclerotized annulus that extends up to midpoint; annulus narrower than in C. roxana; juxta moderately sclerotized projecting cephalad of the vinculum, broad; aedeagus about 3 mm long, heavily curved, almost sigmoid, the anterior and posterior moderately sclerotized, the center heavily sclerotized and tubular, the ductus ejaculatorius entering about one third from the posterior end.

Female: Unknown.

Distribution: Charaplaya, Bolivia; Eastern Slopes of the Andes, Charape, San

Remon and Rio Colorado, Peru. 1300 meters.

Types: I have examined a photograph of the holotype at the BMNH with label data: "Charaplaya,/Bol. 1300 m./VI. 01. (Simons)/650 W. 160 S.",

"Cyllopoda/expansifascia/♂ Prout/type." Holotype designated by original description.

The author also mentions two specimens from San Remon, Peru, one from Charape,

North Peru, and specimens from Rio Colorado (unknown amount). I have only examined the male specimen from Charape with label data "E. Slopes of Andes./Charape, N.

Peru./June. PRATT. 1912.", "Det. By L.B.Prout", "Joicey/Bequest./Brit.Mus./1934-120.",

"C. expansifascia" in the BMNH. These other specimens are designated as paratypes by original description.

Method of determination: Examination of photographs of the holotype; examination of a male paratype.

Cyllopoda gibbifrons Prout

(Figure 2-2.1.H and 2-2.1.I)

Cyllopoda gibbifrons Prout, 1917: 391. Prout, 1934: 133; 1938: 121. Scoble, 1999: 215.

Diagnosis: Small, with wing pattern similar to C. jatropharia and C. osiris.

Differs in having very short pectinations on the male antenna, much smaller body, and thinner black wing borders.

Male (Figure 2-2.1.H): Frons protuberant, tan to black with white scales around eyes; interantennal ridge and antennal shaft black with sprinkling of white scales at base of antennal shafts; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus black with tan suffusion on ventral surface, shorter than length of frons; pectus white; tegula black throughout with long black hair scales; legs white with tan suffusion on anterior surfaces, base yellow; compressed, short hindleg, with reduced tarsi, hair pencil present on posterior surface of tibia, reduced spurs in form of two round apical projections on tibia. Abdomen black dorsally, with lateral sprinkling of yellow scales, white ventrally, with lateral sprinkling of yellow scales.

Forewing with two areoles; apex rounded. Wings above yellow, with broad black

border around edges of forewing, except the inner margin and a prominent black bar in

forewing only; sprinkling of black scales from base to prominent bar along Cu; bar in

forewing broad, from midpoint costal margin to anal angle; hindwing with no bar; black

border present on the outer margin only. Forewing Pattern repeated below but hindwing

with yellow space border where pattern on underside stops and pattern from above shows

through. Length of forewing 1.5 to 1.6 cm.

Male genitalia (Figure 2-3.2.E): About 3 mm long and 1.5 mm wide. Tegumen

moderately sclerotized; socii long, petiolate, and moderately sclerotized with long hairs;

gnathos with base heavily sclerotized, the remaining portions moderately to heavily

sclerotized, arising close to the socii as two projections that meet medially into a

moderately sclerotized tip; uncus arising just above the socii, moderately sclerotized,

sides straight, narrow, with hairs at the top of the lateral margins; valva lightly setose,

slightly curved at the tip, which makes up last one third of the valva, the remaining two

thirds thicker than the tip; vinculum moderately sclerotized, surrounding the base of the

valva; saccus membranous giving rise to a moderately sclerotized annulus that extends up

to one third the length of the tegumen; juxta narrow, moderately sclerotized projecting slightly cephalad of the vinculum; aedeagus about 2 mm long, heavily curved, u-shaped, the posterior end moderately sclerotized, the ductus ejaculatorius entering almost at the posterior end.

Female (Figure 2-2.1.I): Similar to male with simple antenna and hind tibia with

two apical spurs. Black wing borders slightly broader. Length of forewing 1.6 cm.

Female genitalia (Figure 2-5): About 6.2 mm long and 2 mm wide. Papilla anales

simple and elongated, with short hairs; apophyses posteriores and apophyses anteriores

slightly thickened, the apophyses posteriores about four times as long as apophyses

anteriores; genital plate well developed and slightly sclerotized, just caudad to the ostium

bursae; ostium bursae leads from the genital plate, at the end of the eighth abdominal segment; ductus bursae slightly sclerotized leading to the appendix bursae which enters the corpus bursae and is hardly distinguishable; ductus seminalis enters about midpoint between the ostium bursae and the appendix bursae, where it enters a slightly sclerotized extended area; corpus bursae sac-like, with paired signum; signum as a row of long spines extending almost most of its length ending about 1 mm from the end of the corpus bursae.

Variation: Females may have a yellow pectus. The abdomen may also be black

dorsally with the ventral surfaces yellow and white at the tip. The black border and bar in

the wings are also slightly broader than in males.

Distribution: Suapure, Maipures and Orinoco, in Venezuela; and Tobago.

Types: Holotype by original description in the BMNH with locality data:

"Suapure,/Venez./1.3.99./(Klages)", "Cyllopoda/gibbifrons/♂ Prout/ type." Prout (1917)

also mentions seven male paratypes with the same locality data and one male paratype

from Maipures, Orinoco, Venezuela. I have examined a male from the BMNH with data:

"Maipures,/Orinoco,/Dec. 98./(Cherrie)", "Cyllopoda/postica/Wlk", "Rothschild/

Bequest/ B.M 1939-1.", "C. gibbifrons". I believe this specimen to be one of the

paratypes mentioned by Prout.

Material examined: "2/25/99", "U.S.N.M./Acc39806" [female] USNM).

Method of determination: Examination of a photograph of the holotype and examination of one male paratype.

Cyllopoda jatropharia (Linnaeus)

This specie is the first to be named in this genus, and in the tribe to which it belongs. Formerly, there were three subspecies, but based on slight differences of the wing pattern and locality, two subspecies are here recognized. The third subspecies, being recognized as having significant differences based on comparisons of wing patterns and genitalia, has been revised and raised to the species level.

Cyllopoda jatropharia jatropharia (Linnaeus)

(Figure 2-2.1.J and 2-2.1.K)

Phalaena jatropharia Linnaeus, 1758: 523. Clerck, 1768 (2). Fabricius, 1775: 629; 1781:

250; 1787: 193; 1794: 154. Göze, 1781: 214. Gm, 1790: 2469.

Atyria jatropharia: Ch, 1882: 166. Kirby, 1892: 403.

Atyria jatrophae (Hübner), 1823: 31 (Emendation of jatropharia)

Atyriodes jatropharia: Dognin, 1900: 214.

Cyllopoda jatropharia: Prout, 1908: 78. Kay & Lamont, 1927: 110. Prout, 1934: 132.

Sick, 1937: 400-412. Prout, 1938: 120.

Cyllopoda jathropharia jathrpharia: Scoble, 1999: 215.

Diagnosis: Pattern on wings similar to C. gibbifrons and C. osiris. Much larger than C. gibbifrons and with much longer pectinations on the male antenna. Can be separated from C. osiris by the more slender black wing borders, especially in the hindwing, and more slender forewing, especially in the males.

Male (Figure 2-2.1.J): Frons slightly protuberant, black with patch of white scales, white scales around lower edges of eyes; interantennal ridge and antennal shaft black with sprinkling of white scales at base of antenna; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus black, about same length as frons;

pectus white to yellow; tegula yellow, with long black hair scales at the tips; legs tan,

yellow bases; hindleg with highly reduced tarsi; hair pencil on posterior surface of a club-

shaped tibia, without spurs. Abdomen black dorsally with lateral yellow stripes, yellow

ventrally.

Forewing with two areoles, apex rounded. Wings above yellow, with broad black

border around edges of wings except the inner margin, and a prominent black bar in

forewing only; bar in forewing broad, from midpoint of costal margin to anal angle;

hindwing with no bar; black border present on the outer margin only. Pattern repeated

below; hindwing underside with black patch at the base. Length of forewing 1.8 to 1.9

cm.

Male genitalia (Figure 2-3.2.F): About 2.5 mm long and 1 mm wide. Tegumen

moderately sclerotized; socii short, oval, and moderately sclerotized with long hairs,

about as long as half the distance between them; gnathos heavily sclerotized, arising from

the tegumen near base as two slender projections that meet medially, with five very short

apical spines; uncus moderately sclerotized, round, moderately broad, with very long

hairs at the top of the lateral margins; uncus arising just above the socii, constricted

laterally; valva simple, slightly curved at the tip, relatively smooth, the last third slightly

thinner than the first two thirds, lightly setose; vinculum moderately sclerotized,

surrounding the base of the valva; saccus membranous giving rise to a moderately

sclerotized annulus very near base of the valva; juxta moderately sclerotized projecting

cephalad of the vinculum; pedunculi moderately sclerotized projecting cephalad of the

vinculum; aedeagus about 1.2 mm long, very slightly curved, moderately sclerotized, the

anterior end about the same width as the posterior, the ductus ejaculatorius entering at the

posterior end.

Female (Figure 2-2.1.K): Similar to male; slightly more robust, with simple

antenna and two apical spurs on the hindleg. Lower edges of eyes with yellow scales;

some females have a yellow pectus. Forewing may white scales at the tip. Length of

forewing 1.8 to 2.1 cm.

Female genitalia (Figure 2-6): About 8.5 mm long and 2 mm wide. Papilla anales

simple and elongated with short hairs; apophyses posteriores about three times as long as

apophyses anteriores; genital plate slightly sclerotized, just caudad to the ostium bursae;

ostium bursae as a circular opening on the eighth abdominal segment; ductus bursae

slightly sclerotized leading to the appendix bursae; corpus bursae as a blind ending sac;

ductus seminalis leads from about midpoint between the ostium bursae and the appendix bursae; corpus bursae sac-like, with paired signum; signum as rows of short hooks on the side of the corpus bursae, extending through mid-region of corpus bursae.

Variation: Sexual dimorphism, where females are larger than males and have more rounded wings. Female forewing may also have a few white scales at the tip.

Distribution: Turrialba, Heredia, Cartago and La Florida in Costa Rica; Northern

Peru; Venezuela; Bolivia; Guayana; Trinidad and Tobago. From 500 to 2250 feet.

Types: The holotype is located in Evolutionsmuseet, Uppsala Universitet,

Sweden (UZUI). This specimen does not appear yellow and black. This may have

34 resulted from the loss of pigments over the last two and one half centuries. It also bears some resemblance in patterning to Celerena sp. and bears a label written by Aurivillius as

Celerene perithea. The original label, “Jatropharia”, was written by Thunberg. The label data for this specimen is: “Jatropharia/Mus. Gust. Ad.”,”Celerene pe-/rithea

(Cram.)/Auriv.1881.” This specimen is part of a royal donation by King Gustaf IV Adolf.

Material examined: : "Rentema Falls,/Upper Maranon;/N. Peru 1000ft/A. & E.

Pratt.", "jatropharia L./det. L.B.P." [male] (BMNH); "Feb./'07", "Peralta/ 2000ftCR",

"Collection/Wm Schaus" [male] (USNM); "Torrialba. C.R./6.24.71" [male]

(MGCL);"COSTA RICA: Prov./Heredia, 3kms.SW/ Puerto Viejo 75m/Finca La

Selva/Oct. 17, 1973 Opler" [ male] (AMNH); "July/'07", "La Florida/ CR

500ft","Collection/WmSchaus" [female] (USNM); "II-8-77/TURRIALBA,

CARTAGO,/COSTA RICA", "Borrowed from/Oregon St-Univ./Collection

7/2001/CVCovell Jr" [female] (INBio); "Costa Rica : Cartago Prov./ V,14,1985 Opler"

[female] (AMNH); "Turrialba/CR. 6-27-71" (MGCL); "Rio Songo/ Bolivia/ 750 m/ Coll.

Fassl" [female] (NHRS).

Method of determination: Examination of a photograph of the holotype.

Cyllopoda jatropharia puta Strand

(Figure 2-2.1.L)

Cyllopoda puta Strand, 1920: 137; 1927: 24.

Cyllopoda jathropharia puta Prout, 1934: 132; 1938: 120. Scoble, 1999: 215.

Diagnosis: As in the nominate subspecies, but with slightly broader black borders, especially on the hindwing.

Male (Figure 2-2.1.L): Similar to the nominate subspecies; differing in that it may have yellow scales as well as white on the sides of the collar, the palpus may also have white suffusion on the ventral surfaces, and the tegula has the yellow at the base only.

Also, the black wing borders are slightly wider. Length of forewing 1.6 to 1.9 cm.

Male genitalia: Almost the same as the nominate subspecies. Differ in having the gnathos less sclerotized.

Female: Unknown.

Distribution: Trinidad and Tobago; Surinam; Rio Branco in Brazil.

Types: The author cited three males from Trinidad and designated “the largest specimen” as the type. I secured on loan a specimen with locality data: "Trinidad/F.

Birch","Cyllopoda/jathropharia/f. puta/♂ Strand","L B. Prout Coll./ B.M.1939-643","C. jathropharia puta" from the BMNH which I believe to be a paratype. Since a holotype was not designated, I designate this specimen as lectotype and the other two as paralectotypes.

Material examined: "St Anns/ Trinidad/ Antoin Peirre/1912", "Dognin/

Collection" [male] (USNM); "Coll. Of by/Albert S. Pinkus", "Lady Chancellors Rd./St.

Anns-Trinidad/Mar 8 1933" [male] (AMNH); "Surinam", "Barnet/Lyon." [male]

(NHRS). "Rio/Branco", "Amazon/Roman", "[third label illegible]" [male] (NHRS).

Method of determination: Examination of a male paratype.

Cyllopoda latiflava Warren

(Figure 2-2.2.M)

Cyllopoda latiflava Warren, 1905: 312. Prout, 1934: 133; 1938: 121. Scoble,1999: 215

Diagnosis: Similar to C. bipuncta, C. breviplaga, C. jatropharia and C. osiris in wing pattern. It can be differentiated from C. jatropharia and C. osiris by its shorter antennal pectinations and narrower black borders on the wings. It is also smaller than the typical individuals of these two species. It can be separated from C. bipuncta and C. breviplaga by the length and size of the basal yellow cell created by the wing borders and the black bar in the forewing. This cell is more triangular in C. latiflava than in C. breviplaga where it is one third the wing length and almost oval, and C. bipuncta where it extends to half the wing and is more slender.

Male (Figure 2-2.2.M): Frons protuberant, black with white scales around lower edges of eyes; interantennal ridge and antennal shaft black with sprinkling of white scales at base of antenna; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus black with tan suffusion on ventral surface, shorter than length of frons; pectus white to yellow; tegula yellow, edged with black scales, with long black hair scales at the tips; legs white with tan suffusion on anterior surfaces, yellow bases; hindleg with reduced tarsi, hair pencil present on posterior surface of rod-shaped tibia, no spurs. Abdomen black dorsally, white ventrally with yellow lateral lines of scales broader at the base of abdomen than at its posterior.

Forewing with two areoles; apex rounded. Wings above yellow with broad black border all around in forewing, and only at outer margin in hindwing; prominent broad black bar in forewing only, from midpoint costal margin to anal angle; hindwing without bar. Pattern repeated below. Length of forewing 1.7 to 1.9 cm.

Male genitalia (Figure 2-3.2.G): About 3.5 mm long and 1.5 mm wide. Tegumen moderately sclerotized; socii very short, oval, and moderately sclerotized with short

hairs; gnathos heavily sclerotized, arising from the tegumen near socii as two projections

that meet medially, forming a fan-like tip; uncus arising just above socii, moderately

sclerotized, slightly constricted at base, slender, with hairs at the top of the lateral

margins, somewhat spoon-like; valva lightly setose, greatly curved at the heavily

sclerotized tips, the remaining two thirds moderately sclerotized; vinculum moderately

sclerotized, surrounding the base of the valva; saccus membranous giving rise to a

moderately sclerotized annulus that extends up to one thirds the tegumen from the base of

the valva; juxta moderately sclerotized projecting cephalad of the vinculum, broad;

aedeagus about 3 mm long, slightly curved, moderately sclerotized, the ductus

ejaculatorius entering at almost at the posterior end.

Female: Unknown.

Variation: Some males may have the basal areole in the forewing smaller than

the distal one.

Distribution: Colombia; Catarina, Brazil.

Types: Warren described this species, based on one male specimen from

Colombia. I have examined a photograph of this specimen located in the BMNH with

locality data: "Colombia", "Cyllopoda/ latiflava/ type ♂ Warr.” This specimen is the holotype by monotypy.

Material examined: "FELDER/ COLLN.", "Cyll./latiflava/♂ Warr.",

"Rothschild/ Bequest /B.M.1939-1.", "C. latiflava" (BMNH); "Muzo./Colombia/1.20",

"Dognin/ Collection" [male] (USNM); "Muzo/Colombia", "Dognin/Collection" [male]

(USNM); "Sta.Catharina/ coll.Wernicke ", "Staatssamml./Muenchen",

"Micropos/simplex C. Feld./♂" [male] (ZSBS); "Anapoima/ amdinamanca/7.VI.1975/

Colombi" [male] (ZSBS).

Method of determination: Examination of a photograph of the holotype.

Cyllopoda nigrivena Prout

(Figure 2-2.2.N)

Cyllopoda nigrivena Prout, 1917: 392; 1934: 133; 1938: 121. Scoble, 1999: 215.

Diagnosis: Similar to C. expansifascia, C. claudicula and C. radiata in having the

black border on hindwing all around. Differs from C. claudicula and C. radiata in that it

lacks a black bar in the hindwing and possesses a thick line of black scales from base of

wing to prominent bar along Cu. This line is thinner in C. expansifascia.

Male: Unknown.

Female (Figure 2-2.2.N): Frons slightly protuberant, black, with white scales

around lower edges of eyes; interantennal ridge and antennal shaft black with a few white

scales at antenna base; antenna simple; collar black with lateral sprinkling of yellow on

both sides; palpus black. Pectus yellow; tegula black with yellow scales on the inner

edges, tipped with long hair scales; legs tan with yellow base. Abdomen black dorsally with yellow median stripe, ventrally black, yellow median stripes separating dorsal from ventral.

Forewing with two areoles, apex rounded. Wings above yellow with broad black border all around and a prominent black bar in forewing only; line of black scales from base of wing to prominent bar along Cu; bar in forewing broad, from midpoint of costal margin to anal angle; hindwing with no bar. Pattern repeated below. Length of forewing

1.8 to 2.0 cm.

Female genitalia: About 9.5 mm long and 2.5 mm wide; similar to C. gibbifrons

(Figure 2-5); differs in that the genital plate, ostium bursae, and ductus bursae much more heavily sclerotized, and the signa in the corpus bursae fewer.

Distribution: Novo Friburgo, Rio de Janeiro, Brazil.

Types: Holotype by monotypy in the BMNH has the following labels: "Novo

/Friburgo.", "Cyllopoda/nigrivena/♀ Prout/type."

Material examined: Female specimen with label: "Zoolog./Staatsslg. [on left of label] /BRASILIEN/Rio de Janeiro/ 17.II.51/leg. H. Ebert" (ZSBS).

Method of determination: Examination of a photograph of the holotype.

Cyllopoda osiris (Cramer), stat. rev.

This species has a wide range. Formerly, it was treated as a subspecies of C. jatropharia by Prout (1908) after it was originally described as a species by Cramer.

Examination of the genitalia has led this author to return it to species status. Two subspecies are easily separated by wing pattern and genitalia. New synonyms have also been established.

Cyllopoda osiris osiris (Cramer), stat. rev.

(Figure 2-2.2.O and 2-2.2.P)

Phalaena osiris Cramer, 1777: 28. Stoll, 1782: 26. Guer, 1818: 25.

Atyria osiris: Heinrich-Schafer, 1856: 20. Möschler, 1877: 659.

Callimorpha osiris: Verloren, 1837: 53.

Chrysauge osiris: Walker, 1854: 370.

Cyllopoda jatropharia var. osiris: Prout, 1908: 78; 1910: 230. Sick, 1937: 412. Prout,

1934: 132.

Cyllopoda jatropharia osiris: Prout, 1938: 120. Scoble, 1999 : 215.

Cyllopoda latimargo Warren, 1897: 420 [Synonym of osiris]

Cyllopoda ovata Warren, 1907: 198. Prout, 1938: 120. Scoble, 1999: 215 syn. nov.

Cyllopoda jatropharia var. ovata : Prout, 1934: 133.

Cyllopoda protmeta eurychoma Prout, 1938: 120. Scoble, 1999: 215. syn. nov.

Cyllopoda protmeta eurychoma ab. osiriodes Prout: 1938: 120

Diagnosis: Wing pattern similar to C. gibbifrons and C. jatropharia. Much larger than C. gibbifrons and with much longer pectinations on male antenna. Can be separated from C. jatropharia by the broader black wing borders, especially in the hindwing, and more robust forewing, especially in males.

Males (Figure 2-2.2.O): Frons slightly protuberant, black with small patch of white scales, yellow scales around lower edges of eyes; interantennal ridge and antennal shaft black with a sprinkling of yellow scales at the base of the antenna; antenna bipectinate; collar black with lateral sprinkling of yellow on both sides; palpus black, about same length as frons, white suffusion on the ventral surface; pectus yellow; tegula black with yellow at base, with long black hair scales at the tips; legs tan, with yellow bases; hindlegs with reduced tarsi, hair pencil present on posterior surface of club-shaped tibia, no spurs. Abdomen black dorsally with lateral yellow stripes, yellow ventrally.

Forewing with two areoles, apex of forewing rounded with yellow and/or white scales at the tip. Forewings above yellow with broad black border around edges of wing except the inner margin and a prominent black bar; bar in forewing broad, from midpoint costal margin to anal angle; hindwing with no bar; very broad black border present on the

41 outer margin only, almost half of wing. Pattern repeated below; hindwing underside with black patch at the base. Length of forewing 1.9 to 2.1 cm.

Male genitalia (Figure 2-3.2.H): About 2 mm long and 1 mm wide. Tegumen moderately sclerotized; socii very short, oval, and moderately sclerotized with long hairs; gnathos heavily sclerotized, arising from the tegumen close to the midpoint as two slender projections that meet medially, with three very short apical spines; uncus moderately sclerotized, sides straight, moderately broad, uniform towards a square tip with long hairs; valva simple, slightly curved at the tip, relatively smooth, the last third thinner than the first two thirds, lightly setose; vinculum moderately sclerotized, surrounding the base of the valva; saccus membranous giving rise to a moderately sclerotized annulus close to the base of the valva; juxta moderately sclerotized projecting cephalad of the vinculum; pedunculi moderately sclerotized and broad; aedeagus about

1.2 mm long, slightly curved, moderately sclerotized, the anterior end about the same width as the posterior, the ductus ejaculatorius entering at the posterior end; one spinose cornutus.

Female (Figure 2-2.2.P): Similar to male with simple antenna and two apical spurs on the hindlegs. Length of forewing 1.9 to 2.3 cm.

Female genitalia (Figure 2-7): About 7 mm long and 2 mm wide. Papilla anales simple and elongated with short hairs; apophyses posteriores and apophyses anteriores slightly thickened, the apophyses posteriores about three times as long as apophyses anteriores; genital plate slightly sclerotized, just caudad to the ostium bursae; ostium bursae as a circular opening; ductus bursae slightly sclerotized leading to the appendix bursae; corpus bursae as a blind ending sac; ductus seminalis leads from about midpoint

42 between the ostium bursae and the appendix bursae, heavily sclerotized, expanded area where ductus seminalis enters ductus bursae; corpus bursae sac-like, with paired signa; signum as row of short hooks, extending through mid-region of corpus bursae.

Variation: May be without yellow scales at the base of antenna or suffusion on the palpus, and the base of the legs may be yellow or black. Legs may be tan or white with or without suffusion. Females slightly larger than males. In some individuals, the bar on the forewing appears to be fading, the middle of the bar becoming thin, sometimes broken.

Distribution. – Bogueron, Ecuador; Surinam; “British Colombia [sic]”; Rio

Essequebo, Lethem and Rockstone, Guyana; La Union, Huacamayo and Iquitos, Peru;

Dabadie, Trinidad. From 2000 to 3100 feet.

Types: I have examined photographs of the male holotype of C. ovata with label data: "R. Huacamayo,/Carabaya, dry s.,/3100 ft., June 04./(G. Ockenden).",

"Cyllopoda/ovata/ Type ♂ Warr."; C. protmeta erychoma female holotype with label data: "Type", "Amazones/Iquitos/M. de Mathon.", "Cyllopoda/prometa/eurychoma ♂

Prout/type"; C. protmeta eurychoma ab. osiriodes female holotype with label data:

"Iquitos,/U. Amazon,/May 1932./(G. Klug.)", "Cyllopoda/ protmeta/ab. Osiriodes/♂ type

Prout"; C. latimargo female holotype with locality data: "Rio Demerara/ British Guiana",

"Cyllopoda/latimargo/Type ♀ Warr" all from the BMNH. I have also examined a photograph of the allotype of C. protmeta eurychoma with label data: "Allo-/ type",

"Iquitos,/U. Amazon,/August 1932./(G. Klug.)", "Cyllopoda/ protmeta/ eurychoma/ ♀

Prout/ allotype" from the same collection.

Cramer (1777) did not specify a holotype for C. osiris, claiming that it was from

Surinam and located in Mr. B. Vriends cabinet. It should be noted that Surinam in the

18th Century could mean land now a part of French Guiana and Guyana, as ship captains often did not specify an exact locality. The only reference to the sex of the specimen is that has thread-like antennae; this would make it female. A drawing of the holotype

(Cramer 1777) was provided by the author but, the collection of Mr. B. Vriends was sold to van Lennep in 1791 after Vriends death that same year. Parts of the van Lennep collection, then a part of the van Eyndhoven collection, had been taken to the Zoological

Museum in Tring by C. Felder who acquired a portion in 1861. Walter Rothschild purchased Felder’s collection and incorporated it into his own. This collection, also housed in the Museum in Tring, was bequeathed to the BMNH in 1937. A part of the van

Eyndhoven collection was also bought by Verloren van Themaat. The collection of

Verloren van Themat was donated to the ‘Ned. Heidemaatschappij’ in Arnhem, now the

Koninklijke Nederlandsche Heidemaatschappij, in 1939 and was housed in a villa in

Zeist near Utrecht. This villa, and the collection, was unfortunately destroyed (Chainey

2005). A check with museums in Europe revealed that the whereabouts of the holotype for C. osiris is unknown and I suppose that it has been lost. I designate the specimen with label data: "Rockstone,/Essequebo.", "Cyllopoda/latimargo/Wlk./4. 420",

"Collection/WmSchaus", "osiris/Cr./115 E" [female] (USNM) as neotype.

Material examined: "Rockstone,/Essequebo.", "Cyllopoda/latimargo/Wlk./4.

420", "Collection/WmSchaus", "osiris/Cr./115 E" [female] (USNM);

"Lethem/Guyana/m-m. Vallex/ 8/8/81", "ATYRIA ALBIFRONS", "J. Bowe/Collection -

2003" [female] (MGCL) ; "Iquitos/ Ecuador/23.III.1987", "Dr. Luke Kassarov/donation

to FSCA/collection" [male] (MGCL); "La Union,/R. Huacamayo,/Carabaya, 2000 ft.,/wet

s., Jan. 1905/(G. Ockenden).", "Cyllopoda/jatropharia/Linn.",

"Rothschild/Bequest/B.M.1939-1.", "C. ovata" [male] (BMNH); "18.5.62 leg.J.S./Bogueron Abd, Peru", "Cyllopoda/ovata Warr;/Det. C. Covell83" [male]

(MGCL); "9/2/99", "U.S.N.M/Acc39806" (USNM); "Collection of by/Albert S. Pinkus",

"Dabadie-Trinidad/April 10 1933" [female] (AMNH).

Method of determination: Examination of photographs of the holotypes of all synonyms.

Cyllopoda osiris protmeta (Prout), stat. nov.

(Figure 2-2.2.Q)

Cyllopoda protmeta Prout, 1938: 120.

Cyllopoda protmeta protmeta: Scoble, 1999: 215.

Diagnosis: Differs from the nominate subspecies in lacking the black bar in the

forewing.

Male (Figure 2-2.2.Q): External characters as in the nominate subspecies,

differing from it in the absence of a bar on the forewing. The location of the bar in the

nominate subspecies can be seen as a thickening of the black border on the costal edge of

the forewing in this subspecies. Length of forewing 1.9 cm.

Male genitalia: Same as nominate subspecies, only slightly more heavily

sclerotized.

Female: Same as nominate subspecies differing in the absence of the black bar on

the forewing. Length of forewing 2.1 cm.

Female genitalia: No dissections were examined.

Distribution: Pebas, Amazonas in Peru; Ecuador.

Types: Holotype by original designation in the BMNH with label data: "Pebas/

Amazones/M. de Mathan/finX.bre & 1.erTr1880", "Cyllopoda/protmeta/ ♀ Prout/type."

This specimen was designated the holotype by original description.

Material examined: "Ecuador. Buckley", "intensa/wlk", "Cyllopoda/protmeta/♂

Prout", "Rothschild/Bequest/ B.M.1939-1.", "C. p. protmeta" (BMNH);"Sarayaco/

Ecuador", "Collection/ WmSchaus" [male] (USNM).

Method of determination: Examination of a photograph of the holotype.

Cyllopoda postica (Walker)

(Figure 2-2.2.R)

Chrysauge postica Walker, 1854: 371.

Flavinia postica: Dognin, 1891: 39. Kirby, 1892: 404.

Atyria postica: Prout, 1916: 174.

Cyllopoda postica: Prout, 1934: 133; 1938: 120. Scoble, 1999: 215.

Diagnosis: Similar to C. gibbifrons and C. jatropharia in pattern on wings.

Differs in having the black wing borders wider than C. gibbifrons and the basal yellow

patch on the forewing larger than C. jatropharia. Also, lateral yellow abdominal lines

narrowing posteriorly separates it from these two species. These lines are not present in

the other similarly pattered species. The forewing also sometimes has only one areole.

Male (Figure 2-2.2.R): Frons protuberant, black with white scales around lower edges of eyes; interantennal ridge and antennal shaft black; antenna bipectinate; collar black; palpus white, shorter than length of frons; pectus white to yellow; tegula yellow, edged with black scales, with long black hair scales at the tips; legs white with tan

suffusion on anterior surfaces, yellow bases; hindleg with reduced tarsi, hair pencil

present on posterior surface of rod-shaped tibia, no spurs. Abdomen black dorsally, white ventrally with yellow lateral narrowing posteriorly.

Forewing of three out of four specimen examined with one areole; apex rounded.

Wings above yellow, with broad black border around edges of wing except the inner margin, and a prominent black bar in forewing only; sprinkling of black scales from base of wing to prominent bar along Cu; bar in forewing broad, from midpoint of costal margin to anal angle; hindwing with no bar; black border present on the outer margin only. Pattern repeated below. Length of forewing 1.6 to 2.0 cm.

Male genitalia (Figure 2-3.3.I): About 5.5 mm long and 1.6 mm wide. Tegumen moderately sclerotized; socii very long, petiolate, lightly sclerotized, about the same length as the distance between them, with moderately long hairs; gnathos moderately sclerotized, arising from the tegumen very close to the socii as two projections that meet medially, forming a fan-like tip, with two projections leave it dorsally; uncus bilobed,

arising just above the socii, moderately sclerotized, constricted at base, narrow, the tip

drawn out into lateral lobes protruding from the uncus; lateral lobes with long hairs; valva

lightly setose, greatly curved at the tips, moderately sclerotized, the base slender,

narrowing towards the tips; vinculum moderately sclerotized, surrounding the base of the

valva; saccus membranous giving rise to a moderately to heavily sclerotized annulus that

extends to about midway the length of the tegumen, split where the aedeagus exits; juxta

broad, lightly to moderately sclerotized projecting cephalad of the vinculum; aedeagus

about 3.5 mm long, slightly curved, moderately to heavily sclerotized, the ductus

ejaculatorius entering at the posterior end, the tip heavily sclerotized.

Female: Similar to male with simple antenna and two apical spurs on the

hindlegs. Length of forewing 1.9 cm.

Female genitalia: Similar to C. osiris but with signa broader, covering a larger area and with a more sclerotized genital plate, and ostium bursae.

Variation: Some individuals have more white on the frons, with some white at the base of the antenna. The collar may also be a mixture of white, yellow, and black scales. The tegula may also lack yellow scales, being entirely black. The abdomen may also be white ventrally turning into black posteriorly.

Distribution: Peru; San Jose de Cúcuta, Colombia, on the border with Venezuela.

Types: The author did not designate a holotype for this species, stating only that it was from Venezuela and that there were five specimens, a - e. I suspect an error in the type locality, as Cúcuta lies on the Colombian border close to Venezuela. I have examined a photograph of a male syntype at the BMNH with label data: "Vene-/zuela",

"[underside of label] 47/g", "GE POSTICA." I designate this specimen the lectotype. The other specimens are designated as paralectotypes.

Material examined: "Cucuta,/Venezuela.", "Cyllopoda/postica/Wlk.",

"Rothschild/Bequest /B.M.1939-1.", "C. postica" [male] (BMNH); 2 specimens "Peru",

"Collection/ WmSchaus" [male] (USNM); “VENEZUELA, 1100m./Rancho

Grande/Estado Aragua/June 22, 1984/C.V. Covell Jr.”, ”gibbifrons/Prt.?/postica/wlk.?”,

”CV Covell coll./MGCL Acc./2004-12” [female] (MGCL).

Method of determination: Examination of a photograph of a syntype.

Cyllopoda radiata Warren

(Figure 2-2.2.S)

Cyllopoda radiata Warren, 1906: 410. Prout, 1938: 120. Scoble, 1999: 215.

Diagnosis: Similar to C. claudicula; differing in that the black bar in the hindwing

is not repeated on the underside and slightly longer pectinations on the male antenna; the hindleg is compressed, with the first tarsal segment large and the others reduced; tibia

bears a hair pencil.

Male (Figure 2-2.2.S): Frons protuberant, black with patch of white scales, white

scales also around back lower edges of eyes; interantennal ridge and antennal shaft black

with sprinkling of yellow scales at base of antenna; antenna bipectinate; collar black with

lateral sprinkling of white on both sides; palpus black with white suffusion on ventral

surface, shorter than length of frons. Thorax black above with light sprinkling of yellow

scales; pectus white to yellow; tegula black throughout with long black hair scales; legs

white with tan suffusion on anterior surfaces, yellow bases; hindleg highly compressed

and thick with first tarsal segment large, others very reduced, hair pencil on femur and

tibia, no spurs. Abdomen black dorsally with yellow lateral stripes divided by a tin black

line, white ventrally.

Forewing with two areoles, apex rounded. Wings above yellow with broad black

border all around and a prominent black bar in both wings; bar on forewing broad, from

midpoint of costal margin to anal angle; yellow streak on forewing in black margin

between Sc and R just before bar; hindwing with bar narrow from base of wing to broad

at anal angle along M. Forewing Pattern repeated below but hindwing with yellow space

border where pattern on underside stops and pattern from above shows through.

Hindwing bar only faintly expressed below. Length of forewing 1.9 cm.

Male genitalia (Figure 2-3.3.J): About 4 mm long and 1.5 mm wide. Tegumen

lightly sclerotized; socii very short, oval, and lightly sclerotized with long hairs; gnathos

heavily sclerotized, arising from the tegumen close to the socii as two projections that

meet medially, forming a fan-like tip; uncus bilobed, arising just above the socii, lightly

sclerotized, slightly constricted laterally close to the base, narrow, the tip drawn out into

lateral lobes; lateral lobes with long hairs; valva lightly setose, slightly curved at the tips,

lightly sclerotized, the tips thinner than the remaining two thirds; vinculum moderately

sclerotized, surrounding the base of the valva; saccus membranous giving rise to a

membranous annulus close to the base of the valva; juxta narrow, moderately sclerotized

barely projecting cephalad of the vinculum; aedeagus about 2 mm long, slightly curved,

moderately sclerotized, the ductus ejaculatorius entering at almost at the posterior end

with one spinose cornutus.

Female: Unknown.

Distribution: Brazil.

Type: I have examined this holotype at the USNM (catalogue Number 9164).

This is the holotype by original designation.

Material examined: "Rio", "Det. By/L. B. Prout", "Joicey/Bequest./Brit.Mus./

1934-120.", "C. radiata" [male] (USNM).

Method of determination: Examination of the holotype.

Cyllopoda roxana (Druce)

(Figure 2-2.2.T)

Flavinia roxana Druce, 1885: 529. Kirby, 1892: 404.

Diagnosis: Similar to C. latiflava in wing pattern, but colored orange and black instead of yellow and black.

Male (Figure 2-2.2.T): Frons protuberant, black with white scales around lower edges of eyes; interantennal ridge and antennal shaft black; antenna bipectinate; collar black; palpus black with white suffusion on ventral surface, shorter than length of frons; pectus white to tan; tegula black with long black hair scales; legs white; tan suffusion on anterior surfaces, white bases; hindleg compressed, short with reduced tarsi, hair pencil present on posterior surface of tibia, spurs in form of two round projections at the base of tibia. Abdomen black dorsally with lateral orange stripes; white ventrally.

Forewing with two areoles, apex rounded. Wings above orange with black border all around and a prominent black bar in forewing only; bar in forewing from midpoint costal margin to anal angle; hindwing without bar, with thin black border on the inner margin. Pattern repeated below. Length of forewing 1.8 to 1.9 cm.

Male genitalia (Figure 2-3.1.D): Similar to C. expansifascia. About 3 mm long and 1.5 mm wide. Tegumen moderately sclerotized; socii very short, oval, and moderately sclerotized with short hairs; gnathos heavily sclerotized, arising from the tegumen close to the socii as two projections that meet medially, forming a tubular tip; uncus arising just above the socii, moderately sclerotized, sides straight, narrow, with hairs at the top of the lateral margins; valva lightly setose, slightly curved at the tip with two teeth present on the dorsal margin about half way from the heavily sclerotized tip, the remaining two thirds moderately sclerotized; vinculum moderately to heavily sclerotized, surrounding the base of the valva; saccus membranous giving rise to a moderately sclerotized annulus that extends up to midpoint the tegumen; juxta moderately sclerotized

51 projecting cephalad of the vinculum, broad; aedeagus about 3 mm long, heavily curved, almost sigmoid, the anterior and posterior moderately sclerotized, the center heavily sclerotized and tubular, the ductus ejaculatorius entering about one third from the posterior end. Tip not as rounded as C. expansifascia. No cornuti seen.

Female: Unknown.

Distribution: Cosnipata and Quillabamba, Cuzo in Peru.

Types: The author did not clearly designate a holotype, only stating the habitat as

Valley of Cosnipata, East Peru, but the holotype is assumed by monotypy. Label data for the holotype: "Cosnipata/Valley/H. Whitely", "Flavinia/roxana/type Druce.", "Holo- type".

Material examined: Two male specimens from:”PERU,

Cuzo:/Quillabamba/13.III.47/J. C. Pallister" (AMNH).

Method of determination: Examination of a photograph of the holotype.

Figure 2-1. Wing venation in Cyllopoda. Scale bar: 1 cm. Illustration by D. S. Lewis. Wing venation terminology: costa (C); subcosta (Sc); radius (R); radial sector (Rs); media (M); cubitus (Cu); anal (A).

Figure 2-2.1. Cyllopoda adults: A) C. angusta (Lectotype BMNH) male, photographed by C. V. Covell Jn; B) C. angustistriga (Holotype BMNH) female, photographed by C. V. Covell Jn; C) C. bipuncta (Holotype USNM) male, photographed by P. Gentili-Poole; D) C. breviplaga male; E) C. claudicula male; F) C. claudicula female; G) C. expansifascia male; H) C. gibbifrons male; I) C. gibbifrons female; J) C. jatropharia jatropharia male; K) C. jatropharia jatropharia female; L) C. jatropharia puta male. Scale bar in C: 1 cm. Scale bar in all others graduated at 1 mm above image. D to L photographed by A. Chin-Lee and D. S. Lewis.

Figure 2-2.2. Cyllopoda adults: M) C. latiflava male; N) C. nigrivena female; O) C. osiris osiris male; P) C. osiris osiris female; Q) C. osiris protmeta male; R) C. postica male; S) C. radiata male; T) C. roxana male. Scale bar graduated at 1 mm above image. Photographs by A. Chin-Lee and D. S. Lewis.

Figure 2-3.1. The male genitalia and aedeagus of Cyllopoda. A) C. claudicula; B) C. bipuncta; C) C. breviplaga; D) C. roxana. Scale bar: 1 mm. Illustrations by D. S. Lewis.

Figure 2-3.2. The male genitalia and aedeagus of Cyllopoda: E) C. gibbifrons; F) C. jatropharia jatropharia; G) C. latiflava; H) C. osiris osiris. Scale bar: 1 mm. Illustrations by D. S. Lewis.

Figure 2-3.3. The male genitalia and aedeagus of Cyllopoda: I) C. postica; J) C. radiata. Scale bar: 1 mm. Illustrations by D. S. Lewis.

Figure 2-4. Female genitalia of C. claudicula. Scale bar 1 mm. Illustration by D. S. Lewis.

Figure 2-5. Female genitalia of C. gibbifrons. Scale bar 1 mm. Illustration by D. S. Lewis.

Figure 2-6. Female genitalia of C. Figure 2-7. Female genitalia of C. jatropharia jatropharia osiris osiris. Scale bar: 1 Scale bar: 1 mm. mm. Illustration by D. S. Illustration by D. S. Lewis. Lewis.

CHAPTER 3 DISCUSSION

This revision attempts to resolve some questions of synonymy that exist in the

genus Cyllopoda and sets the framework for a much more comprehensive revision, that

of the tribe, which will no doubt contribute to an integration of the tribal structure for the

subfamily Sterrhinae. The major difficulty faced in doing this revision was the lack of

acquisition of specimens to study. To that end, two major entomological holdings in the

United States were visited and loans secured from them. These specimens, however, as

well as others borrowed from institutions in England, Germany, Sweden, Costa Rica,

Ecuador and a private collection in Guatemala, proved limited in number.

The examination of the morphology of these specimens revealed that there are

certain characters that vary greatly in this genus and could not be relied upon as

diagnostic of any one species. One such character was the presence of one or two areoles

and the size of the basal areole in comparison to the distal where there were two. It was

seen that in any one species, individuals varied greatly in the characteristics of the areole

in the forewing; however, the majority of individuals possess two areoles, which is used

as a characteristic feature of this genus. Another character that seemed fairly variable was

the color of the scales on the ventral side of the palpus, the pectus, and the legs. There

seemed to be much inter- and intra-specific variation. Color of scales is not a reliable trait

to be used in this genus, and perhaps tribe, as these individuals seem to be involved in mimicry. To this end, little weight was placed on small variations of scale color. Until there is a better understanding of the nature of the aposematic coloration present in this

61 62

genus and tribe, small variations in color patterns should not be used as a means of

separating these species. The original descriptions of the species in this genus also proved

greatly inadequate in separating these species. Larger scale variations in color patterns

like the presence or absence of borders on wings did prove useful in separating species as

well as the structure of the male hind tibia, the presence or absence of vestigial spurs in

the males, the presence or absence of hair pencils, and the structure of the male and

female genitalia.

It was decided that the subspecies C. claudicula catabathmus was only a variation

of the nominate subspecies. The structure of the genitalia is identical to the nominate

subspecies and the structure of the aedeagus fell within the range observed for the nominate subspecies. They also both occurred within the same range, the only major difference being that C. claudicula catabathmus was more melanic than C. claudicula claudicula. This subspecies was synomized with the nominate subspecies C. claudicula claudicula (Figures 6 & 7).

No specimens of the species C. angusta (Figure 2) were secured for investigation.

Only the type photograph of this species was available. The re-description of this species was done using this photograph and information from the original description. This species is readily distinguished from the others in the genus, as it is predominantly black.

C. angustistriga (Figure 3) proved to be elusive also, as no specimens could be secured on loan. A re-description was constructed using the original description and the photograph of the type specimen. The original label on the type specimen indicates that

Warren (1904) thought this specimen to be a female; however, the structure of the antenna indicates that if this species is indeed a Cyllopoda, it should be a male as it

possesses bipectinate antennae. The sex of the specimen is unsubstantiated as

observations of the wing coupling mechanism could not be determined.

Prout (1938), in the only comprehensive treatment of the genus Cyllopoda and

tribe Cyllopodini to date, found that C. breviplaga versicolor differed from the nominate subspecies in that it was paler. He found “no other outstanding difference” and suggested that it was merely an albinistic form of the nominate subspecies (Figure 5). Investigations of the genitalia and other body characteristics revealed no significant difference and I have therefore concluded it is not a valid subspecies.

The species C. expansifascia (Figure 8) proved troubling. The genitalia of the

males are identical to the genitalia of C. roxana (Figure 21), as is the structure of the

hindleg. Both possess similar black markings on the wings, only one is yellow and the

other orange (Figures 8 & 21). There is a possibility that they could be the same species,

as their ranges are quite close to each other. Because I did not have females of either

species to compare, and information on their natural history is not available, I am

unwilling at this time to synonimyze them, and will maintain them as separate species

until further information proves otherwise. The wing patterns of C. expansifascia also

closely resemble that of C. nigrivena (Figure 15), but that is as far as the comparison

goes. Only a female of C. nigrivena was observed; thus comparisons of the genitalia

could not be made. The genitalia of C. nigrivena female, however, closely resemble, but

are not exactly like those of C. gibbifrons (Figure 33), which creates a problem in

understanding the relationships between these species. This problem requires the

examination of many more specimens and perhaps information on their natural history to

solve. Until then, these species are being kept separate.

On the investigation of the patterns of coloration in the C. jatropharia species group (Figures 11, 12 & 13), it was observed that the species formerly recognized as C. jatropharia osiris was significantly larger and had broader margins than the nominate subspecies, (Figures 11 and 12), and the other subspecies C. jatropharia puta (Figure 13).

When the genitalia of the males and females were compared, C. jatropharia osiris males and females (Figures 27, 29, 34, & 35) proved significantly different from the rest while they were identical to that of C. ovata. C. osiris (Figure 17) is revised to species level and

C. ovata and C. protmeta eurychoma treated as synonyms. Based on the structure of the male genitalia and the absence of the bar in the forewings of C. protmeta protmeta

(Figure 18), this is now recognized as a subspecies of C. osiris.

In the examination of the photographs of the types in the tribe Cyllopodini, one individual that has been described as Atyria albifrons Prout (Figure 36) appears to have bipectinate antenna. The genus Atyria has historically been very closely associated to

Cyllopoda, some of its members having first been described as such. A distinguishing character that sets them apart is the shape of the antenna, Atyria having fasciculate antennae. Additional investigations are needed to determine A. albifrons’ rightful placement.

The problems which still exist in this group will only be solved when more specimens are available, the type specimens are all dissected and examined, and their natural history information worked out.

Synonymic Checklist

CYLLOPODA Dalman, 1823; Type species: Bombyx claudicula Dalman, 1823.

FLAVINIA Walker, 1854

angusta Warren,1897

angustistiga Warren, 1904

bipuncta Warren, 1906

breviplaga Dognin, 1906

versicolor Dognin, 1908

claudicula (Dalman), 1823

catabathmus Prout, 1938

expansifascia Prout, 1917

gibbifrons Prout, 1917

jatropharia jatropharia (Linnaeus), 1758

. jatrophae (Hübner), 1823 (Emendation of jatropharia)

jatropharia puta Strand, 1920

latiflava Warren, 1905

nigrivena Prout, 1917

osiris osiris (Cramer), 1777

latimargo Warren, 1897

ovata Warren, 1907

protmeta eurychoma Prout, 1938

osiris protmeta (Prout), 1938

postica (Walker), 1854

radiata Warren, 1906

roxana (Druce), 1885

Figure 3-1. Holotype of Atyria albifrons. Scale bar graduated at 1 mm above specimen.

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BIOGRAPHICAL SKETCH

Delano St. Aubyn Lewis received his Bachelor of Science in zoology from the

University of the West Indies in 1998. His area of specialty during his undergraduate studies was entomology. After completing his undergraduate studies, he entered graduate school at the University of the West Indies where his research led to the completion of a

Masters of Philosophy in 2001. The topic of his thesis was “Contributions to the

Conservation of the Hellshire Hills Herpetofauna: Analysis of Mongoose Diet and

Radiotelemetry of Headstarted Jamaican Iguanas”. He also started part-time instructing at

Northern Caribbean University while he was still in graduate school and then accepted a full time position as an instructor upon his completion, a position he held until his acceptance at the University of Florida in 2004. At the University of Florida, he pursued a Master of Science in entomology, focusing on Lepidoptera taxonomy.