Classification, Phylogeny, and

Zoogeography of the Genus Penlypus

(Coleoptera: Cleridae)

GINTER EKIS

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 227
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SMITHSONIA N CONTRIBUTION S TO ZOOLOGY • NUMBE R 227

Classification, Phylogeny, and

Zoogeography of the Genus Perilypus

(Coleoptera: Cleridae)

Ginter Ekis

MAR 3119771i

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Ekis, Ginter. Classification, Phylogeny, and Zoogeography of the Genus Perilypus

(Coleoptera: Cleridae). Smithsonian Contributions to Zoology, number 227, 138

pages, 386 figures, 2 tables, 1977.—New World Genus Perilypus is redefined,

and its natural history summarized. Species occur in at least three macrohabitats defined as oak, liana, and herbaceous assemblages. Beetles with

rectangular bodies inhabit oak and liana assemblages whereas oval-bodied beetles

are found among herbaceous plants; mimetic interactions are thought to be

the basis for these habitat-body form correlations. Perilypus mimics are regarded as generalists having evolved in appearance and/or behavior characteristics that simulate more than one distasteful model. Altitudinally the genus

ranges from sea level to 3400 meters; however, most specimens were collected

between 1000 and 2000 meters. Laboratory observations indicate that Perilypus

beetles are highly predatory, with size and rigidity of victims being important

limiting factors in prey acceptability. The criteria used for recognition of species

and of infra- and suprageneric groupings are discussed and techniques for dissections, illustrations, and measurements are described, a morphological analysis

of the major external and internal organs is presented, and a key to the species

groups and species based on adult characteristics is provided. Larval and pupal

stages of Perilypus are described for the first time. The species groups are characterized, and given for each species are, as appropriate, synonymic list, diagnostic combination, description, discussion of structural and chromatic variation,

discussion of natural history, geographic distribution, etymological derivation,

locality records, general notes, and illustrations. Distribution maps with symbols visually convey the geographic range of species and species groups.

There are 49 species presently recognized in Perilypus and 30 of these are

described for the first time. Twelve new synonymies are established. A phylogeny

of Perilypus is postulated using the Hennigian method of phylogenetic analysis.

The zoogeography of the genus is discussed in terms of faunal limits and vicariance. Concepts of historical zoogeography ("faunal tracks", "center of origin and

dispersal", and "forest refugia") are briefly discussed and two of them are used

to explain distributions of extant Perilypus taxa. Relationship discussions within

the species groups vary in completeness according to availability of data. A

fairly complete treatise is presented for the larger limbatus, quadrilineatus, and

ornaticollis groups, but the treatment of the reventazon group, the largest group

in the genus, is very fragmentary. Ancestral Perilypus probably evolved in northern Central America, during mid-Tertiary, and ultimately evolved into eight

species groups. Progenitors of the frontalis, criocerides, chaletoides, and quadrilineatus groups remained in northern Central America as did all of their

descendants. Ancestors of the limbatus and viridipennis groups dispersed to

southern Central America; some members of both groups ultimately penetrated

South America. The ornaticollis group, the most derived species group in the

genus, and the reventazon group are especially widespread geographically.

[Abstract in Spanish on p. 135]

OFFICIAL PUBLICATION DATF is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Ekis, Ginter

Classification, phylogeny, and zoogeography of the genus Perilypus (Coleoptera, Cleridae)

(Smithsonian contributions to zoology ; no. 227)

Bibliography: p.

1. Perilypus. 2. Insects—Classification. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 227.

QL1.S54 no. 227 [QL596.C62] 591'.08s [595.7'64] 76-18705
Contents

Page

Introduction 1

Natural History S

Macrohabi tats 4

Mimicry ; 4

Seasonal and Altitudinal Distribution 4

Laboratory Observations 4

Criteria for Species and for Infra- and Supraspecinc Groupings 5

Materials, Methods, and Terminology 6

Dissecting Techniques 6

Illustrations 6

Measurements 7

Types 7

List of Abbreviations 7

Morphological Analysis 8

External Morphology 8

Head 8

Prothorax 9

Mesothorax 12

Metathorax IS

Wings IS

Legs IS

Abdomen 16

Reproductive Organs 18

Internal Morphology 18

Digestive System 18

Nervous System 20

Reproductive Systems 20

Genus Perilypus 20

Key to the Species of Perilypus 23

The frontalis Group 26

1. Perilypus frontalis (Gorhani), new combination 28

2. PerUypus sensilis, new species SO

The criocerides Group SS

3. Perilypus criocerides (Gorham), new combination SS

4. Perilypus insectus, new species 35

The viridipennis Group 36

5. Perilypus viridipennis (Spinola), new combination 37

6. Perilypus relucens (Gorham), new combination 39

7. Perilypus levis, new species 42

The limbatus Group 45

8. Perilypus limbatus (Gorham), new combination 45

9. Perilypus columbicus (Spinoia), new combination 48

10. Perilypus apocopatus, new species 50

11. Perilypus iris, new species 52

12. Perilypus acus, new species 52

IS. Perilypus buga, new species 54

14. Perilypus latilira, new species 55

15. Perilypus prolixipenis, new species 58

iii
IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

16. Perilypus testaceicornis (Pic), new combination 60

17. Perilypus decoris, new species 60

The reventazon Group 61

18. Perilypus reventazon, new species 63

19. Perilypus nigriventris (Gorham), new combination 64

20. Perilypus sapientis, new species 65

21. Perilypus claudus (Wolcott) 66

22. Perilypus cultratus, new species 69

23. Perilypus bilineatus (Gorham), new combination 71

24. Perilypus cerroazul, new species 73

25. Perilypus ordinatus, new species 73

26. Perilypus calcaris, new species 74

27. Perilypus orthopleuridus (Thomson), new combination 76

28. Perilypus exilis, new species 78

29. Perilypus latilobus, new species 79

30. Perilypus dislinctus (Chevrolat), new combination 80

31. Perilypus crassus, new species 82

32. Perilypus immitis, new species 83

33. Perilypus caliculus, new species 83

34. Perilypus carbonarius Spinola 86

The chaletoides Group 88

35. Perilypus chaletoides, new species 88

36. Perilypus bicolor (Chevrolat), new combination 91

The quadrilineatus Group 92

37. Perilypus quadrilineatus (Chevrolat), new combination 92

38. Perilypus telephoroides (Gorham), new combination 94

39. Perilypus coroniformis, new species 96

40. Perilypus fluctus, new species 98

41. Perilypus bicristatus, new species 99

The ornnticollis Group 99

42. Perilypus ventralis (Gorham), new combination 102

43. Perilypus ornaticollis (LeConte), new combination 105

44. Perilypus galbeus, new species 107

45. Perilypus prolixicornis, new species 108

46. Perilypus pilatus, new species Ill

47. Perilypus antarius, new species Ill

48. Perilypus sinuapicis, new species Ill

49. Perilypus floralis (Gorham), new combination 113

Immature Specimens 114

Phylogeny and Zoogeography 116

Phylogenedc Methods 116

Zoogeographic Methods 125

Phylogenetics and Zoogeography of Perilypus 126

Intergeneric Relationships 126

Relationships among, the Species Groups 126

Relationships within the Species Groups ISO

Abstract in Spanish 135

literature Cited 136
Classification, Phylogeny, and

Zoogeography of the Genus Perilypus

(Coleoptera: Gleridae)

Ginter Ekis

Introduction

Checkered beetles of the genus Perilypus are exclusively New World and predominantly tropical;

they are predaceous surface dwellers with great

agility in gait and flight. Structurally, the beetles

are either rectangular or oval in body form and

their integumental color ranges from dull black to

floricolorous patterns of yellow, red, and blue.

Woody vegetation in dry montane forests seems to

be the macrohabitat for many of the species, but

lampyroid mimics occur on herbaceous plants located in the more open highland habitats. Intraand interpopulation color polymorphism (often

correlated with sex) and mimetic life styles are some

interesting traits of most of the species.

I first acquired an interest for these beetles in

1970 when William F. Barr of the University of

Idaho conveyed the following notes about specimens of Perilypus (heretofore in the genus Colyphus Spinola) sent to him for identification: "I'm

at a loss as to what to call these Colyphus specimens. The names available in the literature really

are meaningless until we can establish what constitutes a species in this group. The possibility of a

considerable amount of variation within a species

including sexual dimorphism should not be overlooked." From this reply evolved my objectives for

Ginter Ekis, Research Fellow, Department of Entomology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

the beginning stages of this work; to resolve the

species concept in Perilypus, clarify nomenclatural

problems, and provide diagnostic aids for species

recognition. At first there was little more to be done

with this group because only a handful of specimens were available for study. Perilypus specimens

are not abundantly collected in nature, as is the

case with most mimics; consequently, the beetles

were unfamiliar to most keepers of collections and

essentially unavailable for loan shipments. In time,

additional specimens became available, particularly

during my tenure as a research fellow of the Smithsonian Institution. During that period (June 1973

to June 1974) two research grants provided financial

support for an extensive field expedition to Central

and South America and a visit to various museums

in Europe. These expeditions nearly doubled the

available study material. Particularly important was

the trip to Latin America, during which I obtained

data about the natural history of the group, collected specimens of several species in fluid for

analyses of internal anatomy, and discovered the

larval and pupal stages of the genus.

The procurement of additional specimens and

biological data prompted an expansion of the scope

of this study to include generalizations about

Perilypus natural history, and analyses of internal

and external anatomy, and of phylogeny and zoogeography. The reader may find some of these

sections less than complete; for example, in the sections of phylogeny and zoogeography, relationships
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 1-2.—Perilypus decoris: 1, forebody: 2. habitus.
NUMBER 227

within the largest species group, the reventazon

group, remain undeciphered. Even a terse and/or

provisional treatment of some of the aforementioned subjects, however, may positively influence a

group's systematic progress. The purpose of this

paper is twofold. First, it is offered as a unit of

information consisting of all the current knowledge

about the genus Perilypus, including a resynthesis

of the data in sections of morphology, practical

taxonomy, and evolutionary relationships. Second,

this work is intended to establish a ground plan for

revisions involving genera of the subfamily Clerinae.

LITERATURE REVIEW.—Perilypus was validated by

Spinola in 1841; he included the name in a diagnostic key of the clerid genera of the world. The

genus was subsequently more fully described by

Spinola in 1844a on the basis of P. carbonarius

which until recently was classified under the subfamily Tillinae. In a previous paper, I (1975:25)

discussed the classification of the genus and transferred two other Spinola species to Perilypus from

Colyphus. In this work, the majority of Colyphus

species described by other authors are also transferred to Perilypus. There is little else to say about

the history of the group, except that the species

were never studied collectively, and that species

descriptions are scattered throughout the literature,

are very brief, and rarely include illustrations. Besides Spinola, the following authors published one

or more names of species now considered valid and

currently placed in Perilypus: Chevrolat (1843,

1874), Gorham (1878, 1882, 1886), LeConte (1880),

Pic (1941), Thomson (1860), and Wolcott (1927a).

ACKNOWLEDGMENTS.—During the course of this

study, I was very fortunate to work at the National

Museum of Natural History, Smithsonian Institution, with many competent persons who were always willing to discuss concepts or assist in material

needs. For practical reasons, only a few can be mentioned herein. I am particularly indebted to Terry

L. Erwin, who provided among other things, museum space, equipment, travel funds for fieldwork

in Latin America, a review of the manuscript, and,

most important, guidance and encouragement during a research fellowship at the Smithsonian Institution. This manuscript was also reviewed by

Donald R. Whitehead of the University of Michigan and Lee H. Herman, Jr., of the American

Museum of Natural History, with whom I also

enjoyed occasional discussions about systematics

theory. John M. Kingsolver kindly made available a microscope during the initial stages of

this work. I also must acknowledge his patience

during repeated visits to his office, the first point

of inquiry whenever I needed equipment relevant

to taxonomic work. As usual, William F. Barr of

the University of Idaho was very helpful in making

available his notes and photographic transparencies

of type specimens, and George C. Steyskal rendered

guidance on matters relevant to nomenclature. I am

indebted to George L. Venable for drawing the

habitus and forebody illustrations of P. decoris,

new species (Figures 1, 2) and to Michael W.

Druckenbrod for illustrating P. limbatus larva and

pupa (Figures 370, 380,381).

I extend my gratitude to the museum curators

and private collectors (see "Materials") who entrusted me with their specimens. For lending type

specimens I am grateful to Christine M. F. von

Hayek (British Museum of Natural History), Mme.

A. Bons (Museum d'Histoire Naturelle de Paris),

F. Heike (Museum fur Naturkunde on der Humbolt-Universitat zu Berlin), Marco Spinola (Tassarolo Castle in Novi Ligure, Italy), Henry Dybas

(Field Museum of Natural History), and John F.

Lawrence (Museum of Comparative Zoology).

This study was supported in part by a one year

Smithsonian research fellowship in the Institution's

program in Evolutionary and Systematic Biology, a

travel grant from the American Philosophical Society (Penrose Fund #6658) for type studies abroad,

and a Walter Rathbone Bacon Scholarship from the

Smithsonian Institution (awarded to T. L. Erwin)

for field studies in Latin America. For typing various sections of the manuscript I am grateful to Jo

Whitehead and Nancy Stefansson. Jos6 Cuatrecasas

kindly translated the abstract into Spanish.

Natural History

Although little is known about the natural history of Perilypus beetles, some general comments

can be made about their macrohabitats, approximate seasonal and altitudinal distribution, and

apparent involvement in several mimetic complexes. More specific notes relevant to mating behavior and prey acceptance are based on laboratory

observations. In addition, on page 47 I briefly

allude to the larval and pupal microhabitat of P.

limbatus.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MACROHABITATS

Field observations (June, July) in Mexico, Guatemala, Costa Rica, and Venezuela indicate that species of Perilypus frequent at least three kinds of

vegetation assemblages, here broadly denned as the

oak assemblage, the liana assemblage, and the herbaceous assemblage. Specimens were collected from

the oak assemblage, essentially a montane oak-pine

forest, by beating low hanging branches of Quercus.

The liana assemblage consists of an assortment

of lianas intertwined with branches, twigs, and low

canopy shrubby vegetation. A substantial part of

the woody vegetation is dead. The most productive

method of collecting specimens from the liana assemblage is by beating between 10:00 AM and 1:00

PM. This macrohabitat, usually found in the drier

regions of montane forests (about 1500 m), also is

frequented by wood-boring beetles. This suggests,

at least superficially, a potential biological relationship between the wood borers and the species of

Perilypus found among them; under laboratory

conditions, bark bettles were the preferred food of

P. limbatus.

The herbaceous assemblage involves broad-leaf

herbaceous plants (including such members of Compositae as Vernonia deppeana Lessing), which were

particularly evident along hedgerows adjacent to

cultivated fields and near roadsides. Adult Perilypus

on these plants were collected easily by hand during late afternoon hours.

There is an interesting correlation between the

body shape of the beetles and two of the vegetation

types on which they were collected. Beetles with

rectangular bodies were found in liana assemblages,

whereas those with oval bodies were found on

broadleaf herbaceous plants. Consequently, unlike

the slender beetles from the liana plants, the more

ovate beetles always were exposed and conspicuously visible on broad-surface foliage. This is not

surprising, however, as oval Perilypus are lampyrid

mimics whose success is dependent on their "inconspicuous" exposure among models.

MIMICRY

Even a cursory survey of Neotropical clerids reveals how extensively mimicry and warning coloration have influenced clerid evolution. These traits

certainly are conspicuous in Perilypus whose members may simulate lampyrids, cantharids, lycids, or

chrysomelids. Characteristics of the mimic important in the perfection of the mimetic pattern may

involve body form, integumental color, and behavior, or a combination of all three as in P.

ventralis. The mimetic relationships within Perilypus will be discussed in more detail. For the present

suffice to say that there is a tendency among some

species of Perilypus to simulate more than one

distasteful model; that is, the mimic has evolved a

generalized appearance and/or behavior that simulates members of several model species with more or

less similar color patterns. Membership in such a

generalized mimetic complex requires phenotypic

plasticity on the part of the mimic. Perhaps the

extensive color polymorphism present in this genus

is a manifestation of that requirement.

SEASONAL AND ALTITUDINAL DISTRIBUTION

Seasonal and altitudinal distribution records were

taken from collector labels and from data recorded

by me during field work. Most adult specimens were

collected from June to August; very few were obtained before June or after August, and none during December or January. Three immatures (two

larvae and one pupa) were captured in June. The

composite altitudinal range of the members of

Perilypus extends from sea level to about 3400

meters, most specimens being collected between

1000 to 2000 meters.

LABORATORY OBSERVATIONS

Specimens of P. limbatus were collected alive,

sexed, and maintained in petri dishes for general

observation. Longevity of captive specimens ranged

from five to ten days after capture; males perished

first and became a meal for survivors if not quickly

removed from the enclosures (dried specimens were

not eaten).

Interphenon matings occurred between members

of all phena (pp. 5, 47). The mating position is

dorsoventral. The male lifts his forebody onto the

female while maintaining his hind legs in contact

with the substrate; copulation lasts approximately

45 seconds. Precopulation events are rapid and uncomplicated. The male overtakes the female from

behind and with his hind legs on the substrate

clasps the female epipleural margin with his pro-
NUMBER 227

and mesotarsal claws. I suspect that the convex

female epipleuron and the conspicuously asymmetrical male protarsal claws function as copulatory

anchorage structures.

Field and laboratory observations suggest that

only one egg is released during each oviposition.

The habit of depositing a single egg at each oviposition site is possibly a safeguard against larval

cannibalism. One captive female devoured her

singly laid egg immediately after it was deposited

on the surface of a small twig.

My observations pertinent to prey acceptability,

admittedly based on superficial tests, show that

specimens of P. limbatus feed on a wide array of

insects as most checkered beetles probably do. Apparently two important limiting factors in prey

acceptance are size and rigidity of the exoskeleton

of the potential victim. Smaller prey, such as

scolytid beetles, are held with the middle legs (usually around the abdomen) and simultaneously

oriented with the front legs. The predator follows

a distinct sequence when devouring its catch: first

eaten are the contents of the cervical membranes,

followed by the soft tissues at the promesothoracic

junction, and finally the contents of the abdominal

cavity. Some larger beetles such as elaterids, chrysomelids, and cerambycids also were eaten, but

these were acceptable only when freshly killed and

with some exposure of internal tissue.

Criteria for Species and for Infraand Supraspecific Groupings

I accept the biological species concept (Mayr,

1969:26) as the theoretical premise for recognition

of species despite the fact that conjectures about

reproductive isolation are usually based on indirect

evidence (in part indicative of our inability to determine extent of gene flow). No species concept

alternative to Mayr's is more consistent with evolutionary theory.

The nominal species included in this work represent my best estimates (hypotheses) of what the biological species of Perilypus are. Each estimate is

based largely on anatomical criteria, but structural,

distributional, and biological characteristics are

integrated in species delimitations whenever possible. In Perilypus the genital organs, and particularly the aedeagus, provide the most consistent

characteristics for species recognition. Aedeagal

characteristics are exceptionally homogeneous at

the infraspecific level but otherwise they are highly

variable. Characteristics of the ovipositor are specifically distinct within some species groups.

I consider stable gaps (morphological or otherwise) not correlated with sex between sympatric

forms to indicate nonconspecificity; I assume such

gaps are manifestations of disrupted gene flow.

Allopatric samples are regarded as conspecific if

they show no appreciable morphological or biological differences. Within this definition, minor variations are interpreted as due to geographic variation.

When allopatric samples are notably different, however, the magnitude of their difference is compared

with that observed between sympatric species. In

absence of contrary evidence and when such differences between the allopatric samples approximate

those between the sympatric species, I judged allopatric forms not conspecific.

I generally oppose formal naming of species subunits, particularly in the absence of nonanatomical

information. Even when such information is available, it usually consists of gross approximations of

environmental limits and of distribution, and therefore is deficient for in-depth assessments of population dynamics. The recent work of Ehrlich, et al.

(1975) and of authors cited therein demonstrates

the level of experimentation necessary to gain

meaningful information about interpopulation gene

flow.

In addition to avoiding the introduction of ambiguous trivial names into the literature, nonusage

of the subspecific category necessitates a more thorough discussion of interpopulation differences. Too

often, and after minimal assessments, taxonomic

entities are named with justifications for their formal recognition ill-defined or hidden under the

cloak of theory.

I believe that most differences among infraspecific

populations can be conveyed effectively and emphasized through discussions, and if necessary, variant

populations may be identified by informal designations such as "morph" and "phenon." In this

work, noteworthy infraspecific variations are discussed under "Variation" in the species descriptions.

When variation is noted at the interpopulation

level, specimens exhibiting the atypical characteristics are assigned under a "morph" name (e.g., redlegged morph). I assigned groups of specimens

exhibiting a common intrapopulational color varia-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tion to a "phenon"; observed color polymorphism is

often sex dependent.

I do not formally recognize supraspecific taxa

within Perilypus. Until relationships among the

various New World clerine groups (of dubious

generic rank) closely related to Perilypus are better

understood, I will recognize the "species group"

category and, when necessary, the "species subgroup" category. Informal species groups and species subgroups will identify adequately the respective major and minor monophyletic lineages of these

genera until such time when a balanced generic

classification in the Clerinae is established. Lindroth (1969:xxiv) and Whitehead (1972:140) also

have alluded to the pitfalls of indiscriminate use of

the subgenus category.

Materials, Methods, and Terminology

I examined more than 1200 adult specimens, one

pupa, and two larvae of North, Central, and South

American Perilypus, and approximately 300 adults

and 12 larvae of other genera of Clerinae. The majority of these specimens were loaned to me by

museums but material from private collectors is

also well represented. I collected these insects from

Mexico southward to Venezuela and Charles W.

O'Brien, Terry L. Erwin, and Henry P. Stockwell

made special efforts to collect material in Pampel's

fluid especially for this study.

DISSECTING TECHNIQUES

Fluid-preserved specimens were killed and fixed

in Pampel's fluid (glacial acetic acid, 4 parts; 40%

formaldehyde, 6 parts; 95% alcohol, 15 parts; water,

30 parts). I found this preservative superior to any

other that I have used to date (Ekis and Gupta,

1971:52); the beetles are not discolored, do not become brittle internally, and can remain in fluid

for several years without notable damage. Several

structures described under "Morphological Analysis" would not have been noticed had I only studied

dried-preserved specimens. Indeed, I found that by

examining fluid-preserved specimens in fluid (usually water) integumental regions, and particularly

the soft membranous structures, were more clearly

visible and easier to maneuver for detailed study.

Another advantage to collecting specimens in Pampel's fluid is that specimens collected in that fluid

may be dry mounted and subsequently rehydrated

to an extent that delicate internal tissue can be

easily dissected. The quality of these dissections was

often equivalent to those made with specimens

retained in fluid.

Fluid-preserved specimens were dissected in water

as follows: First, the hind pair of wings was removed with microscissors, submerged in absolute

ethanol, unfolded between two glass slides, dried,

removed from the slide, mounted on paper point,

and pinned beneath the specimen and at right

angles to it. The second step involved removal of

the abdominal contents including the copulatory

organs. With an eye knife the abdomen was incised

from the third spiracle to the anus; with a watchmaker's forceps the digesting tract was severed at

the base of the abdomen; and the intact mass of

internal organs was lifted out of the abdominal

cavity. The organs were subsequently separated and

reintroduced into Pampel's fluid until they could

be studied and illustrated. Once dissected, beetles

were rinsed in water, dried on tissue paper, and

pointed. Whenever possible, at least four fluidpreserved specimens (two males and two females)

were dissected as described above, but the genital

organs were extracted from all male specimens irrespective of preservation technique used.

To remove membranous wings from a dry specimen, it was boiled for five minutes in water containing five drops of 409 detergent. After dissection

the beetle was washed in an ultrasonic cleaner,

dried, and pinned. Extraction of genital organs

from dry-preserved beetles involved severing the

abdomen from the specimen, boiling in 10 percent

KOH, and dissection procedures as described in the

previous paragraph. Occasionally internal reproductive organs from dry-preserved beetles were

sufficiently rehydrated so that a detail study of

them was possible. Genitalia and portions of internal organs were stored in glycerine and introduced into plastic genitalia vials, which were

sealed with polyethylene stoppers and pinned below

appropriate specimens.

ILLUSTRATIONS

All drawings were prepared with a camera

lucida mounted on a Wild stereoscopic microscope.

Sometimes line drawings of small organs (e.g.,

mouthparts) had to be optically magnified so that
NUMBER 227

structural details, examined through a compound

microscope, could be presented in adequate scale.

In the drawings of the male internal reproductive

organs only one-half of the accessory glands, testes

and associated tubes were drawn. The posterior

accessory glands are stippled for contrast. Unless

indicated otherwise the scale line accompanying a

drawing equals 1 mm. The scales for electron

micrograph figures are presented in the legends.

Distribution maps, usually by species groups, are

provided; dots and various other symbols represent

locality records. I used a question-mark symbol

whenever the locality records referred to a country

only. All illustrated morphological structures discussed in the text are indicated by italics at first

mention.

MEASUREMENTS

The meristic data provide a close approximation

of the size and proportions of the beetles and of

some of their parts. They also provide a quantitative basis for substantiating descriptive terms, such

as "feebly transverse," "boldly transverse." When

available, I measured at least 10 males and 10

females of each species. All measurements were

made with an ocular micrometer through a Spencer

stereoscopic microscope. Measurements and their

abbreviations (when relevant) found in the descriptions are: length = body length, sum of lengths of

head, pronotum, and elytron (head length is the

linear distance from the antennal carina to the

most posterolateral limit of the eye; lengths of

pronotum and elytron are explained below);

width = body width, maximum width across

elytra; AL = antennal length, sum of lengths of all

articles; EL = elytral length, linear distance from

the humeral margin, proximal to the suture, to the

elytral apex; EW = maximum width across the

elytron; HW = maximum width of the head across

eyes; IOW = maximum width between ocular sutures; PL = maximum dorsal pronotal length;

PLS = maximum lateral pronotal length; PW =

maximum dorsal pronotal width.

TYPES

I examined the primary type-specimen (holotype

or lectotype) of each Perilypus species except

Derestenus columbicus var. reynoldsi Wolcott and

Colyphus melanopterus Dury. Because male genital

characteristics provide the most reliable criteria for

diagnosing species, whenever possible I selected

male specimens to establish holotypes or designate

lectotypes. Procedures pertinent to selection of primary or secondary types were carried out in accordance with the International Rules of Zoological

Nomenclature (1964).

Data relevant to type-specimens and to typelocalities are noted at the beginning of each species

description.

LIST OF ABBREVIATIONS

The following abbreviations are used in the text.

They indicate the repository of specimens and

collections from which material was borrowed. Addresses and the names of curators within the institution, and private collectors are supplied to

facilitate inquiries regarding the collections.

AMNH American Museum of Natural History, New York,

New York 10024; L. H. Herman, Jr.

BMNH British Museum (Natural History), Entomology,

SW7 5BD, London, England; Christine M. F. von

Hayek

CASC California Academy of Sciences, San Francisco,

California 94118; D. H. Kavanaugh

CBAZ C. Bordon Azzali, Avenida Lima, Edificio Taormina, Apt. 8, Los Caobos, Venezuela; private

collection

CISC California Insect Survey, Entomology, University

of California, Berkeley, California 94720; J. A.

Chemsak

CMPP Carnegie Museum, Insects and Spiders, 4400

Forbes Avenue, Pittsburgh, Pennsylvania 15212;

G. E. Wallace

CMNH Cincinnati Museum of Natural History, 1720

Gilbert Ave., Cincinnati, Ohio 45202; C. Oehler

CNCC Canadian National Collection of Insects, Entomology Research Institute, Ottawa, Ontario, Canada;

D. Bright, E. C. Becker

CUNY Cornell University, Entomology, Ithaca, New York

14850; L. L. Pechuman

DEIE Deutsches Entomologisches Institute, 13 Eberswalde, D. D. R.; H. Morge

FMNH Field Museum of Natural History, Chicago, Illinois 60605; H. Dybas

GEKI G. Ekis, c/o National Museum of Natural History,

Smithsonian Institution, Entomology, Washington,

D.C. 20560; private collection

GHNE G. H. Nelson, Kansas City College of Osteopathy

and Surgery, Anatomy, 2105 Independence Avenue,

Kansas City, Missouri 64124; private collection

HFHO H. F. Howden, Carleton University, Biology,

Ottawa, Ontario, Canada; private collection
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HPST H. P. Stockwell, Gorgas Memorial Hospital, Canal

Zone; private collection

ISNB Institute Royal des Sciences Naturelles de Belgique, Bruxelles

LACM Los Angeles County Museum of Natural History,

900 Exposition Boulevard, Los Angeles, California

90007; J. P. Donahue

LMDR L. M. Drukenbrod, 5519 Decatur St., Bladensburg, Maryland 20710; private collection

MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138; J. F.

Lawrence

MHNM Museu de Historia Natural de la ciudad de

Mexico, Apartado 18-845, Mexico, D.F. 18; D.F.A.

Barrera

MNHB Museum fur Naturkunde on der HumboltUniversitat zu Berlin, 104 Berlin, Invalidenstrasse

43, DDR; F. Hieke

MNHP Museum National d'histore Naturelle, Entomologie, 45 bis, Rue de Button, Paris (Ve), France;

A Bons, M. Villiers, J. J. Menier

MSPI Massimiliano Spinola Collection, Tassarolo Castle,

Novi Ligure, Italy; M. Spinola

MZSP Museu de Zoologia da Universidade de Sao Paulo,

Caixa Postal 7172, 04263, Sao Paulo, Brazil; H.

Reichardt

MZSI Museo ed Instituto di Zoologia Sistematica della

Universita di Torino, 10123 Torino, Via Giovanni

Giolitti, 34 Italy; M. Zunino

NMNH National Museum of National History, Smithsonian Institution, Entomology, Washington, D.C.

20560; R. E. White, P. J. Spangler

RNHN Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden, Netherlands; J. Krikken

SEMK Snow Entomological Museum, University of Kansas, Lawrence, Kansas 66044; G. W. Byers

TAMT Texas A & M University, Entomology, College

Station, Texas 77843; H. R. Burke

TTUT Texas Tech University, Entomology, P. O. Box

4169, Lubbock, Texas 79409; D. Foster

UCMV Universidad Central de Venezuela, Instituto de

Zoologia, Facultad de Agronomia, Agricola, Maracay, Venezuela; G. Yepes T.

UZMD Universiteits Zoologiske Museum, Universitetsparken 15, Copenhagen, Denmark; Sv. G. Larsson

WBIV W. Bivin, Box 1854 Balboa, Canal Zone; private

collection

WFBA W. F. Barr, University of Idaho, Entomology,

Moscow, Idaho 83843; private collection

ZMAN Zoologisch Museum der Universiteit van Amsterdam, Entomologie, Plantage Middenlaan 64, Amsterdam 1004, Netherlands; J. P. Duffels

Morphological Analysis

In studies of insect systematics, anatomy often

provides most if not all of the data accessible for

relationship discussions, hence the importance of

comprehensive morphological analyses in revisionary works. By investigating more than the convenient integumental and internal characteristics,

the basis for evolutionary inferences is maximized.

With this in mind I decided to present a reasonably

thorough morphological analysis of Perilypus and

to accomplish this I chose P. reventazon since its

members exhibit the "typical" exoskeletal characteristics of most Perilypus species. Herein, I treat

most of these characteristics and those of the alimentary canal, ventral nerve cord, and internal

reproductive organs of both sexes. The characters

and their states are defined and illustrated, not only

to facilitate the descriptions and discussion included

in this work, but also to provide a foundation of

terminology for future works.

EXTERNAL MORPHOLOGY

Head

CRANIUM.—Snodgrass (1935:111) used head sutures to delimit six major areas of the generalized

insect cranium. Unfortunately, comparable areas

cannot always be so rigidly denned; in beetle adults

some of the sutures described by Snodgrass are

obscured or absent. An alternative used herein is

to subdivide the cranial surface on a more general

basis.

The major structures and areas on the front of

the head are illustrated in Figure 3. The frontoclypeal region is bounded laterally by the antennal

carina and interocular portion of the ocular suture,

anteriorly by the labral base, and posteriorly by the

hind limit of the interocular depressions. The epistomal suture traverses the frontodypeal area, dividing it into the clypeus and the broader frons. The

latter is marked by a pair of interocular depressions

that partially encircle the frontal umbo. The region

from the posterior limit of the frons to the postoccipital suture (Figure 7) is the epicranium (Figure

3) which extends from the most dorsal aspect of the

ocular suture to the base of the epicranial acumination (Figure 4).

The gena (Figure 5) is the postocular region of

the cranial wall, bounded posteriorly by the postoccipital suture, dorsally by the epicranium, and

ventrally by the gular suture and obscured lateral

borders of the submentum. The gena is conspicuously sculptured with shallow wrinkles at the sides
NUMBER 211 9

proximal to the ocular suture. The transverse gula

(Figure 5) is prolonged posteriorly into an oblong,

feebly elongate gular process that is microsetiferous

on the hind margin. The free borders of the process

unite with the ventral portion of the inflexed

postocciput. The latter borders the occipital foramen and is widely splayed proximal to the epicranial acumination. The compound eyes are setose,

boldly convex, have a well-developed ocular notch

(Figure 6) in front, and consist of ommatidia that

are about as wide as the ocular suture.

ANTENNA.—The antenna (Figure 8) consists of 11

articles, is serrate along the anterior border, and

via the articulatory process of the scape pivots on

the antennifer (Figure 6) of the antennal sclerite.

The scape is feebly arcuate and slightly concave

ventrally, and is about twice as long as the subspherical pedicel. The articles of the flagellum are

compressed dorsoventrally but as a group become

progressively more subcylindrical toward the antennal apex. The distal margins of articles 1 through

10 are truncate whereas article 11 is acuminate; all

articles are densely setose, particularly the distal

five. The macrosetae (Figure 19) are long, acuminate, and grooved longitudinally, whereas the microsetae are short, smooth, and blunt apically.

Microsetae occur only on the last five articles, being concentrated on the dorsodistal region of articles 7 to 10, and are particularly prominent on the

dorsal surface of article 11. The ventral surface of

the antenna is more sparsely setaceous, particularly

at the distal third where microturbercles are present. These tubercles are most abundant on the

venter of article 11, and are very similar to the

sensilla basiconica of Tenebrio molitor Linnaeus

reported by Doyen (1966:110).

MOUTHPARTS.—The labrum (Figure 9) is a transverse, strongly emarginated preoral flap attached to

the frontoclypeal inflection. The labrum is vested

dorsally with long setae and has an anterior fringe

of short setae. The torma projects from the hind

angles of the labrum and bifurcates distally into a

medial tormal process and a lateral tormal process.

The tormal sclerite is similarly bifurcated in other

clerids (Solervicens, 1973a: 165; 1973b:235, 238) and

in other beetles (Evans, 1961:307; Doyen, 1966:111).

Anteroventrally, the labrum has a U-shaped ridge

that partially outlines a thin, feebly concave sderotized region.

The mandible (Figures 10, 11) is subfalcifonn

and prominently visible in repose. Its outer facies is

broadly depressed and setose in the basal two-thirds.

Three dentes occupy most of the masticatory surface; the anterior dens is broadly arcuate and

acuminate; the medial dens is short, truncate, and

feebly arcuate; the posterior dens is broad, shallow,

and projects an asperate profile. Behind the

posterior dens is a small transparent lobe, which

appears continuous with the mandibular penicillus;

the latter extends anteriorly along the inner base

of the medial and posterior dentes. The condyle

and fossa of the mandibular base articulate with the

ventrolateral edge of the cranium.

The maxilla (Figure 12) connects to the cranium

behind the mandibular fossa. Its triangular base,

the cardo, is bifurcated into a posterior inner and

outer cardal process. Anteriorly the cardo connects

to the basistipes, a triangular sclerite flanked by the

palpifer and mediostipes. The latter is continuous

with the lacinia, whose base is distinguishable in

maxillary dorsal view only. Distally the lacinia is

divided into a triangular medial lobe (the mediolacinia) and smaller basolateral lobe (the

laterolacinia). A short, transverse inflection divides

the galea into a small basigalea and vastly larger

distigalea. The maxillary palpus consists of four

setose articles; the first (the basal) is the shortest,

the second and third together are slightly longer

than the fourth, and the fourth is cylindrical and

membranous apically.

The labium (Figure 13) connects to the submental

region of the cranium (Figure 5). Its mentum is

predominantly membranous, feebly sclerotized

basally and laterally. The palpigers also are feebly

sclerotized, subcylindrical, and partially underlie

the mentum. The ligula has a setose submarginal

fringe anteriorly, some scattered setae medially,

and one exceptionally long setae proximal to each

bisetose palpiger. The labial palpus consists of three

articles; the first is conical; the second slightly

swollen apically and nearly twice as long as the first;

and the third, the largest, is securiform, membranous along its distal border, and densely vested with

short recumbent setae.

Prothorax

The prothorax (Figures 14-16) is a squat moderately transverse cylinder flexibly connected to the

cranium by the cervical membrane. The cervical
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

COMPOUND

IYES

FRONS

ANTENNAL

CARINA

CLYPIUS

EPICRANIUM

GENA

OCULAR SUTURE

FRONTAL UMIO

INTEROCULAR

DEPRESSION

EPISTOMAL

SUTURE

SUBMENTAL

REGION

GULA

GULAR SUTURE

GULAR PROCESS

GENA

EPICRANIAL ACUMINATION

OCULAR

NOTCH

POSTOCCIPITAL

SUTURE

OCCIPITAL

FORAMEN

POSTOCCIPUT

ANTENNIFER

FIGURES Z-8.~Perilypus reventazon: 3, head, anterior; 4, head, dorsal; 5, head, ventral;

6, head, lateral; 7, head, posterior; 8, antenna. (Soles = 1 mm)
NUMBER %Z 11

MEDIAL TORMAL

PROCESS

LATERAL TORMAL

PROCESS

^LATEROLACINIA

J LAONIA

LABIAL PALPUS

PRONOTAL ARCH

15

UTUM

ANTERIOR

FORAMINAL FLANGE

PROSTERNUM

CRYPTOSTERNUM

PUURAl APOPHYSIS

TROCHANTIN

INTERCOXAL PROCESS

ANTEROVENTRAL

MEMBRANEOUS

TUBERCLE

POSTEROVENTRAL

MEMBRANEOUS

TUBERCLE

POSTERIOR

FORAMINAL FLANGE

PROSTERNAL APOPHYSIS

FIGURES 9-18.—Perilypus reventazon: 9, labium; 10-11, mandible; 12, maxilla; 13, labium; 14,

prothorax, dorsal view; 15, prothorax, lateral view; 16, prothorax, ventral view; 17, mesoscutellum, dorsal view; 18, mesoscutellum, lateral view. (Scale = 1 mm)
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sclerites are slender, feebly pigmented plates that

do not articulate with the head or prothorax. The

dorsal and lateral wall of the prothorax comprise

the pronotum; the ventral wall, or prosternum, is

greatly reduced externally but substantially

inflected.

The pronotum (Figures 14, 15) is traversed by

anterior and posterior depressions that delimit

three regions, the pronotal arch, the pronotum

proper, and the pronotal collar. At about the

anterior fourth, the subapical depression marks the

hind limit of the convex pronotal arch that extends

over most of the epicranial region when the head

is reposed. To minimize the potential friction between the epicranial surface and the inner wall of

the pronotal arch, the latter is lined with a transparent inflection, the anterior foraminal flange

(Figure 16). This thin, almost membranous flange

extends posteriorly to the inner ridge of the subapical depression where it connects to the cervical

membrane.

In dorsal view the pronotum proper is subspherical, boldly convex laterally, sinuous anteriorly,

and truncate posteriorly. Near each hind angle

there is a spheroid muscle-attachment depression,

the pronotal fovea (Figure 14). Behind this fovea,

the prebasal depression precedes the narrow pronotal collar, which widens at the sides and is

broadly inflected along its posterior margin. This

inflection, the posterior foraminal flange (Figure

16), is a broad, rigid lining that combines with

the pronotal collar to provide additional rigidity to

the posterior region of prothorax. Anteriorly, the

flange connects to the ensuing intersegmental

membrane.

The prosternum (Figure 16) is a narrow transverse plate adjoined to the anterior half of the

pronotal deflection. Posteriorly, the plate is broadly

inflected, particularly so laterad to the prosternal

intercoxal process. The prosternal inflection, the

cryptosternum of Hlavac (1972:126), supports the

prosternal apophysis and extends laterally to the

pleural apophysis. The latter is short, broadly

bilobed distally, and continuous with the trochantin. (In the prothorax of the clerid Necrobia rufipes

DeGeer, the distal end of the pleural apophysis is

also broad but not bilobed (Tremblay, 1958:98).)

The pronotum is densely clothed by short vertical

setae; such setae line the posterior margin of the

pronotal postcoxal process. The prosternum is predominantly glabrous; setae are present on the

anterior margin and on the feebly expanded distal

region of the intercoxal prosternal process.

Mesothorax

The pro-mesothoracic intersegmental membrane

contains one dorsal and two ventral pairs of

transverse membranous tubercles (Figure 16). The

dorsal pair is similar in form and vestiture to the

anteroventral pair. The posteroventral tubercles

differ from the others by being more conical and

by bearing stout vertical setae; they are probably

sensory organs as suggested by their connection

with a mesothoracic nerve.

The mesothorax (Figures 21, 22) is approximately

one-half the size of the metathorax. Its thick,

strongly sclerotized notal plate (Figures 17, 18) may

be divided conveniently into scutum and scutellum;

the former is transverse, the latter subquadrate. A

longitudinal mesonotal suture divides equally the

rugose scutum, which partially projects into the

pronotum. Side margins of the mesonotum are

broadly inflexed, particularly so below the scutellum where the inflection serves as a shelf on which

rests the mesobasal angle of the elytron.

The mesopleuron (Figures 21, 22) is divided into

a large triangular mesepisternum and a smaller,

tapering mesepimeron. The first is coarsely rugose

in the dorsal half, conspicuously ridged at the

anterior margin, and is copiously vested with recumbent setae, as are most of the metathoracic pleural

and sternal plates. The substantially narrower

mesepimeron is finely rugose and emarginated at

the pleurocoxal articulation.

The mesosternum is sagittal in outline and

prolonged medially. The anterior prolongation is

short and tumid. The posterior one, the mesothoracic intercoxal process, is long and slender, and

is invaginated apically to accommodate the apex

of the metathoracic intercoxal process with which it

forms a strong but flexible connection. Posteriorly,

the mesosternum is deeply inflexed, particularly

beneath its intercoxal process where it forms a

nearly imperforated septum. Posteroventrally, the

mesosterneal inflection adjoins a feebly sclerotized

plate located anteriad to a metasternal transverse

ridge. (A comparable area in the cantharid Chanliognatus pennsylvanicus (DeGeer) was interpreted
NUMBER 227 13

by Matsuda (1970:213) to consist of "mesospinasternal" and "metapresternal" elements.)

Metathorax

The metathorax (Figures 21, 22, 24) is feebly

sclerotized dorsally, strongly sclerotized and boldly

convex ventrally. The metanotum is typically complex, being partitioned into various sclerites,

sutures, and/or other structures correlated with the

development of flight. A lengthy description of

the metanotal components is not necessary because

they have no immediate relevance to the taxonomic

data herein; but an illustration (Figure 24) of some

of the more important regions might be useful for

subsequent comparative studies. The metapleuron

(Figures 21, 22) consists primarly of the slender

metepisternum; the metepimeron is largely membranous, being feebly sclerotized along the mesopleural suture and at the pleural wing processes.

The metasternum is trapezoidal, about half as

long as wide, and is divided into two halves by a

midlongitudinal suture (the discriminal line of

Ferris, 1940:37). Ferris apparently was also the

first to suggest that the beetle metathoracic venter,

commonly called the "metasternum," consists of

subcoxal elements. The discriminal line is also

well-defined in the clerids Enoclerus sphegeus

Fabricius and Necrobia rufipes DeGeer (Campau,

1940:78).

Internally, the discriminal line is expressed by a

bold inflection referred to as the discrimen by

Ferris (1940:36) and the median ridge by Doyen

(1966:123). The inflection is a keel-like structure

that nearly spans the length of the metasternum

and increases in depth posteriorly where it becomes

continuous with the internal ridge of the antecoxal

suture. Posteriad to this suture is a narrow tapered

antecoxal piece ("Katepisternum 3" of Campau,

1940:78) whose ventral extremity forms the ventral

coxal condyle.

To my knowledge a metafurcal sclerite, or metasternellum, is not distinguishable in the Cleridae;

however, a well-formed Hylecoetoid metendosternite (Crowson, 1944:274) does not vary appreciably

in Perilypus. The metendosternite (Figure 25) of

P. reventazon consists of a broad laterally arcuate

stalk, oval lamina, and feebly arcuate jurcal arms

that project anterolaterally. Solervicens (1973a, fig.

5b; 1973b, figs. 4a,b) studied the metendosternites

of the clerid genera Epiclines Blanchard and Natalts

Laporte. He noted a substantial difference between

the metendosternites of N. laplacii Laporte and

N. impressus (Spinola).

Wings

MESOTHORACIC WINGS.—The elytron (Figures 26,

28, 29) is an oblong protective cover that overlays

the thorax and all abdominal segments; its posterior

deflection, the apical slope (Figure 29), is acute.

The dorsal surface is copiously indented with

coarse favus (or honeycombed-shaped) punctations

and has a sparsely microrugose microsculpture

(note arrow in Figure 307). The latter gives the

punctate and interpunctate surface a glittering

arenose appearance that becomes particularly apparent under intensive light. The elytral disc also

has a dense vesture of short reclinate and longer

vertical setae.

In dorsal view, the elytron is outlined by three

margins (Figure 26). Two of these, the humeral

and sutural, are defined according to their morphological origin. The third, the posthumeral, is

defined in accord with its position and/or function.

Below the posthumeral margin is the epipleural

margin (Figure 28) that consists of lateral epipleural

fold and ventral epipleuron.

The humeral margin is feebly sinuous, the posthumeral margin is linear in basal two-thirds and

acutely arcuate in apical third. The sutural margin

is linear and has a prominent sutural ridge that

extends from the sutural cleft (as in Figure 27) to

the elytral apex. The sutural margin has a sutural

groove and sutural flange that interlock the reposed

elytra.

METATHORACIC WINGS.—A membranous wing is

illustrated in Figure 23. Wing venation characteristics of JP. reventazon are not described because

they are of no consequence in delimitation of

perilypine species and because terminology of clerid

wing venation is currently being reevaluated by

other workers.

Legs

Close apposition of the intrasegmental leg bases,

as found in the more successful surface predatory

beetles, is a primary structural adaptation for an

efficient mechanism of cursorial locomotion. Of
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MESOSTERNUM

MESEPIMERON

MITAPUURAL WING PROCESS

M f SOTHORACIC INTERCOXAL PROCESS

METATHORACIC INTERCOXAL PROCESS

METEPISTERNUM

METEPIMERON

MESOPIEURAL SUTURE

DISCRIMINAL LINE

ANTECOXAl SUTURE

FIRST VISIBLE ABDOMINAL STERNUM

STERNAL DEPRESSION

FIGURES 19-23.—Perilypus reventazon: 19, antennal articles 5 to 8 (arrows indicate microsetae);

20, abdominal sterna n, in, ventral view; 21, mesothorax, metathorax, and abdomen, ventral

view; 22, mesothorax, metathorax, and abdomen, lateral view; 23, metathoracic wing. (Sale =

I mm)
NUMBER 227 15

AXILLARY I

AXILLARY 3

LAMINA

HUMERAL MARGIN

POSTHUMERAl*-

MARGIN

.SUTURAl

'RIDGE

.SUTURAL

\ MARGIN

FIGURES 24-31.—Perilypus reventazon: 24, metanotum and axillary sclerites; 25, metendosternite;

26, elytron, dorsal view; 28, elytron, cross section; 29, elytron, lateral view. P. orthopleuridus:

27, mesobasal region (arrow indicates sutural cleft). P. limbatus: 30, elytron, lateral view. P.

quadrilineatus: 31, elytron, lateral view. (Scales = 1 mm)
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

secondary importance in this scheme are modifications of structures and/or proportions of the leg

components, particularly those of the tarsal articles.

All of these considerations are to some degree

pertinent to a discussion of the leg structure of

clerid beetles which, as a group, are almost exclusively cursorial predators.

In P. reventazon the intrasegmental coxae are

separated by a thin intercoxal process, which in

the meso- and metathorax communicate with a

midlongitudinal extension of the subsequent segmental venter. The procoxa (Figure 32) is conical

and more protrusive than the pyrifonn mesocoxa

(Figure 33) and transverse metacoxa (Figure 35).

The pro- and mesotrochanter are triangular and the

metatrochanter is oblong. In general, the femora

are very similar but the front femur is shorter and

slightly more robust than the rest. The tibiae are

also very similar, but protibia differs by having one

tibial spur, not two (Figure 32). All tibiae are conspicuously carinate (the tibial carina) on their

anterior and posterior fades, and each is particularly densely setose at its inner apical third.

The tarsus, the most variable leg region, consists of five conspicuous tarsomeres, which differ

with respect to length and pulvillar shape on each

leg (Figure 34). The basitarsus (Figure 33) is always

the shortest; the second and fifth tarsomere are

about equal in length; and the third is slightly

longer than the fourth. The emargination of the

distal border of the pulvillus of each tarsus becomes

more pronounced from basitarsus to tarsomere four.

Another structural progression involves abridgement of the first two pulvilli from pro- to metatarsus. The basitarsal pulvillus, which along with

pulvillus 2 is conspicuously asymmetrical on the

metatarsus (note left arrow in Figure 36), is reduced

and obscured by a narrow setal cluster (note right

arrow in Figure 36).

Protarsal claws are sexually dimorphic with respect to symmetry. In male specimens the inner

claw is conspicuously longer than the outer claw

(Figure 34). In female specimens the protarsal claws

are symmetrical as are the claws of the middle and

hind legs of both sexes.

Abdomen

The abdomen (Figures 21, 22) is soft, flexible,

with eight apparent segments. In males, rudiments

of a ninth segment may be present in the form of

a feebly sclerotized crescentic plate (the interspicular

plate) located beneath the aedeagus and between

the posterior explanation of the spiculi (Figures 37,

38). Doyen (1966:138) found a dorsal interspicular

plate in Tenebrio and considered it the tenth

tergum. Tanner (1927, pi. 6: figs. 56, 57), in his

coverage of female genitalia of Tillinae clerids,

identified the membrane between segment 8 and

the ovipositor as the ninth segment.

In general the abdominal dorsum (Figure 22) is

feebly sclerotized. However, tergal sclerotization

becomes more intense from tergum 1, which is

entirely membranous, to tergum 8; the sclerotization of sterna 3 to 8 is about equivalent. The first

five terga are glabrous, whereas the sixth and

seventh have bands of microsetae at their posterior

margins, which are probably instrumental in the

folding mechanism of the membranous wings.

Tergum 8, the pygidium, is vested with long

reclinate setae, and is subquadrate in males (Figure

39) and transverse in females (Figure 40). The

lateral area of the abdomen is broadly membranous.

In more advanced coleopterans it is often difficult to identify components of the anteroventral

region of the abodmen. This region was modified

substantially during the posterior extension of the

metathoracic ventral plate, an adaptation yielding

greater surface area for attachment of flight muscles.

In general, sternum i is considered absent in most

polyphagan beetles but a vestige of this plate is

present in Tenebrio (Doyen, 1966:135). In Perilypus, sternum i appears to be entirely suppressed.

The second abdominal sternum (Figure 20) is narrow and feebly sclerotized and is fused with an

anterior depression of the third abdominal sternum

to form the metacoxal cavity. Medially however,

the two sterna form a keel-like intercoxal process

(Figure 20). Forbes (1925:290) described a similar

development of the abdominal venter in the

anobiid Trypopitys sericeus (Say) (now in the genus

Priobium).

The posterior six sterna, which comprise the

visible abdominal venter, are well-sclerotized,

transverse plates, the lateral and posterior margins

of which are narrowly and palely indicated. Sterna

in to VII each have a pair of sternal depressions

(Figure 22). To the inner surface of these are attached short longitudinal muscles that connect to

the antecostal ridge. Doyen (1966:136) found similar
NUMBER 227 17

FIGURES 32-42.—Perilypus reventazon: 32, front leg; 33 middle leg; 34, front tarsus; 35, hind

leg; 36, hind tarsus (top arrow indicates pulvillus 2, bottom arrow indicates basitarsai pulvillus);

37, spicular fork, ventral view; 38, spicular fork, lateral view; 39, male pygidium; 40, female

pygidium; 41, male abdominal sternum vm; 42, female abdominal sternum vm, (Scales = 1 mm)
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

muscle attachment regions on exposed sterna of

Tenebrio molitor. The posterior margin of sternum

VII (Figure 21) is broadly emarginated in both sexes,

whereas that of sternum vm is sexually dimorphic,

being emarginated in males (Figure 41) and arcuate

in females (Figure 42). The exposed sterna are

vested with long, reclinate setae.

Reproductive Organs

MALE.—The aedeagus (sensu Lindroth, 1957:242)

consists of two major structures, the tegmen (Figure

43) and the phallus (Figure 46). The tegmen is a

dorsoventrally compressed cylinder that consists of

a pair of parameres, the phallobase, two phallobasic

struts, and the phallobasic apodeme. Each paramere

has a basolateral depression, is hollow, and has two

mesal margins (the mesodorsal and the mesoventral

margins). Together, the parameres enclose a dorsal

and ventral sinus (Figure 44). Anteriorly, the parameres are continuous with the phallobase, whose

anterior limit projects as the mesoventral phallobasic apodeme and the lateral phallobasic struts.

The phallus, which when reposed lies in the

tegmen, consists of one dorsal and one ventral

phallic plate (Figure 46) and two lateral membranes. Each phallic plate is set with a row of

marginal denticles aligned along the margin

proximal to the phallic plicae. Also, the posterior

ends of these plates fuse to form the phallic apex;

anteriorly each plate projects as a phallic strut.

The phallic plicae are distended during copulation

(as in Figure 47). Their numerous phallic papillae

(as in Figure 155; top arrow) may be stimulatory

organs or anchorage structures; the latter function

is unlikely because the inner lining of the bursa

copulatrix does not have structures that might be

construed as elements of an interlocking mechanism.

When reposed the aedeagus rests on the spicular

fork (Figure 37), which provides an attachment

surface for the aedeagal protractor and retractor

muscles, and guides aedeagal movements. The

spicular fork consists of two slender spicules that

are broadened posteriorly and fused at the anterior

third. Between the spicules and attached to them is

the spicular sac, which is opened proximal to the

interspicular plate. To the ventrolateral walls of

this sac connects a spheroid gland (the spicular

gland), whose function has not been determined.

Cholodkovsky (1913:530) described and figured

similar glands in the clerid Necrobia ruficollis

Latreille and indicated that the duct of each gland

opened on either side of the anus. Scott (1919:108,

footnote) suggested that the glands reported by

Cholodkovsky might be neotenic structures that

in mature larvae produce secretions important to

construction of the pupal cocoon. During studies

of clerid internal anatomy I found the spicular

gland present in both sexes of species from all

suprageneric taxa. In females, the spicular gland

connects to the membranous sac of the spiculum

ventrale (Figure 42).

FEMALE.—The ovipositor (Figures 48, 49), a predominantly membranous tube, connects to the

eighth segment via an expansive connecting membrane (the ninth segment of Tanner, 1927, figs. 56,

57); three pairs of slender sclerites provide structural rigidity to the genital tube. Two pairs of these

sclerites, the oblique and ventral bacculi (Figure

48) are remnants of the first and second valvifers,

respectively (Tanner, 1927:23). The mesoventral

extremity of the oblique bacculus articulates with

the posterior limit of the ventral bacculus, which

at the posterior fourth projects as a short lateral

extension (the baccular acumination). The third

pair of supportive sclerites, proctigeral bacculi (Figure 49) widen on the proctiger, are fused at posterior third, and are strongly divergent anteriorly.

The coxites (the hemisternites of Lindroth, 1957:

250) are lobate structures that show sclerotized

vestiges (the coxital plates) of valvifer two. The

outer margin of the coxite is broadly vested with

setae and the inner margin is asetose and strongly

sclerotized in the posterior half. Between the dorsal

and ventral bases of the coxites extend membranous

and acuminate laminae. The dorsal laminal incision

divides the posterior half of the dorsal lamina

(Figure 49) into two equal lobes; this lamina is

substantially larger than the unilobed ventral

lamina (Figure 48). Posteriorly, the coxite projects

as a well-developed coxital stylus that is multisetose

distally.

INTERNAL MORPHOLOGY

Digestive System

The male and female alimentary canal of P.

reventazon were described by Ekis and Gupta

(1971:56, figs. 28, 29, 76, 77) under C. signaticollis
NUMBER 227 19

PARAMERE

ESOVENTRAL MARGIN

ESODORSAL MARGIN

VENTRAL SINUS

DORSAL SINU

PHALLOBASE

43

-_PHAUOBASIC

STRUTS

PHAILOBASIC

APODEME

PHALLIC

PUCAE

PHALLUS

TEGMEN

45

COXITAL

STYLUS

COXITE

COXITAL

PLATE

DORSAL LAMII

LAMINAL INCISIO

VENTRAL LAMINA

OBLIQUE

BACCULUS

PROCTIGER

r

BACCULAR

ACUMINATION

__VENTRAL

^ ^ BACCULUS

PROCTIGERAl

BACCULUS

FIGURES 45-50.—Perilypus reventazon: 43, tegmen, ventral view, 44, tegmen, dorsal view, 45,

tegmen, lateral view, 46, phallus. P. chaletoides: 47, aedeagus, dorsal view; 48, ovipositor,

ventral view; 49, ovipositor, dorsal view; 50, brain and ventral nerve cord, dorsal view. (Scales =

1 mm)
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ANTERIOR

ACCESSORY GLAND

POSTERIOR

ACCESSORY GLAND

OUTER BRANCH

INNER BRANCH

SPERMATHECAL CAPSULE

PERMATHECAL GLAND

DUCT

SPERMATHECAL

GLAND

SPERMATHECAL

DUCT

BURSA

COPULATRIX

OVARY

MEDIAN OVIDUCT

LATERAL OVIDUCT

CALYX

VAGINA

56

FIGURES 51-56.—Perilypus reventazon: 51, male internal reproductive organs; 52, anterior accessory gland; 53, posterior accessory gland; 54, ejaculatory duct; 55, female internal reproductive organs. P. xtcntralis: 56, hind leg. (Scales = 1 mm)
NUMBER 227 21

Spinola and C. bilineatus Gorham, respectively.

The descriptions and illustrations in that work

adequately characterize perilypine digestive organs,

which have no appreciable intrageneric variation.

Nervous System

An illustration of the ventral nerve cord (Figure

50) is provided to establish a frame of reference for

subsequent intergeneric comparisons. The nerve

cord was not notably varied among the species in

which it was studied.

Reproductive Systems

MALE.—The more important male organs (Figures 51-54) are the testes, vas deferens, accessory

glands, and ejaculatory duct.

Each of two oblong testes consists of 12 tubular

testicular follicles that are encapsulated by the

peritoneal membrane. The follicles unite with the

tubular vas deferens via the vasa efferentia, which

join the ejaculatory duct. An oblong swelling on

the vas deferens proximal to the ejaculatory duct

is the seminal vesicle. To the anterior limit of the

ejaculatory duct there also are attached two pairs

of accessory glands which according to their position of insertion are designated as the posterior

accessory glands and the anterior accessory glands.

The anterior accessory gland (Figures 51, 52) is

very long, undivided, and coiled; its narrower distal

third is tightly convoluted. The inner walls of this

gland has two shallow, longitudinal ridges that

delimit at least two different secretional regions.

The products of one of these regions are externally

visible as a dark band along the inner half of the

gland. The posterior accessory gland (Figures 51, 53)

divides near the base into two variously convoluted

branches. The outer branch is about one-third

longer than the inner branch; the texture and other

visible properties of the secretions of both branches

are identical. The ejaculatory duct (Figure 54) is

bulbous and sinuous in its anterior half, slender

and linear in the remainder.

FEMALE.—The major female organs (Figure 55)

are the ovaries, lateral and median oviducts, bursa

copulatrix, spermathecal capsule, spermathecal

gland, and vagina. The shape and external appearance of these organs are generally homogeneous at

infra- or interspecific level. However, the shape of

the bursa copulatrix is correlated with the amount

of semen in its lumen.

The ovaries consist of 12 acrotrophic ovarioles

that stem from a well-developed calyx. The latter

connects to short, lateral oviducts that communicate with a narrower more elongate median oviduct.

The bursa copulatrix, a saccular extension of the

anterior region of the vagina, is recurved basally.

The bursa copulatrix is usually spatulate, but in

gravid females it is swollen substantially by seminal

fluid. Near the base of the bursa copulatrix the

spermathecal capsule, a sclerotized ovoid structure,

communicates with the vagina via the spermathecal

duct; the latter is distended basally, narrowed and

convoluted distally. The spermathecal gland connects to the spermathecal capsule via a short brunneous spermathecal gland duct. The vagina is flexed

near the middle and is tapered.

Genus Perilypus

Perilypus Spinola, 1841:72 [type-species: Perilypus carbonarius

Spinola, 1844a: 105, by monotypy].—Spinola, 1844a: 103.—

Lacordaire, 1857:430.—Desmarest, [1860]:235.—Schenkling,

1903:13.—Ekis, 1975:26.

Derestenus Chevrolat, 1843:13 [type-species: Derestenus quadrilineatus Chevrolat, 1843:13, by monotypy].—Desmarest,

1860:238.—Lacordaire, 1857:443.—Gorham, 1878:162.— Corporaal, 1948:243.

DIAGNOSIS.—The combination of serrate antenna

(Figure 8) and intricate elytral punctation (Figure

2; punctations are usually honeycomb-shaped) distinguish specimens of most species of Perilypus

from those of other New World genera of Clerinae

(sensu Crowson, 1964). Specimens of the frontalis

and criocerides groups (involving four species) have

clubbed antenna. Members of the first group are

superficially similar to some specimens of the genus

Placopterus, but are separated easily from them

(p. 26). Specimens of P. criocerides (Gorham) might

be confused with those of Systenoderes amaenus

Spinola, which differ by having the elytron boldly

convex, sparsely setose, and impunctate.

DESCRIPTION.—Form: Variable, commonly rectangulate (Figure 2) with elytron plane in basal

two-thirds and deflexed in apical third. Members of

the quadrilineatus and ornaticollis groups are variously oval (Figures 279, 293) and usually compressed

dorsoventrally, whereas those of the chaletoides and

viridipennis groups are respectively conspicuously

robust, and short and variously convex.
22

Size: Length 5.4 to 14.4 mm, width 1.6 to 5.2 mm.

Females usually more robust than males.

Color: Head (including antennae), thoracic

venter, elytron, legs, and abdomen usually flavotestaceous or black, or both. Pronotum concolorous,

or lateral regions flavotestaceous to rufous and disc

with black vitta; latter percurrent or interrupted

at middle; sometimes pronotum predominantly

flavotestaceous with dark brown or black midapical

macula. Abdomen usually entirely black in males,

black and flavotestaceous in females. Elytron concolorous, fasciate, or vittate (usually vittate or

fasciate in females only).

Integumental Setae: Head, pronotum, elytron,

and legs densely setose; setae particularly long and

copious on antenna and tibia. Antennal setal length

inversely related to elytral setal length; in lampyriform specimens (where antenna most serrate)

antennal setae exceptionally long, elytral setae

exceptionally short.

Head: Interocular depressions deeply impressed

or shallow, confluent posteriorly or not. Eyes usually

boldly convex; ommatidia fine, as wide as ocular

suture. Mouthparts, see pages 9, 11. Antenna serrate (Figure 8); rarely clubbed (Figure 67); articles

usually increasing in width from scape to article

11; funicular articles filiform (Figure 67) to boldly

serrate (Figure 284).

Thorax: For discussion of prothoracic venter,

metathorax, and metathoracic wings and legs,

see pp. 9-16. Pronotum usually conspicuously

transverse, length rarely equals width; subapical

depression deeply impressed throughout, or deeply

impressed laterally and shallow medially, sinuous

or linear; pronotal foveae shallow or deeply impressed, punctiform or elongate; side margins of

pronotum proper variously convex; prebasal depression and pronotal collar usually well developed.

Mesoscutellum subquadrate, transverse (Figure 312),

or triangular (Figure 300); rarely stalked (Figure

310). Elytron variable; outer margins (epipleural

or posthumeral) broadly arcuate, or straight in

basal two-thirds (or more); arcuate in apical third

(or more); surface microsculptured (see arrow in

Figure 307) or not; punctation usually favus, deeply

impressed or shallow, and profusely distributed

throughout elytral surface, or punctations small

and round and restricted to posthumeral region.

Epipleural fold plane or convex; apical slope

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

deflection gradual (Figure 30), very gradual (Figure

31), or acute (Figure 29). Front tarsal claws boldly

(Figure 69) or feebly (Figure 34) asymmetrical, or

symmetrical.

Abdomen: The structures of the abdomen are

discussed on page 16. Male pygidium varied in size

and shape, particularly robust in some species

(Figure 217); posterior margin usually entire, rarely

emarginated (Figure 94).

Male Genitalia: For a general description, see

page 18. Usually strongly pigmented throughout;

in members of the ornaticollis group phallobase

marked with two dorsal and two ventral longitudinal bands. Combined length of phallobase and

phallobasic apodeme 1.5 to 10 times longer than

paramere. Combined length of paramere and phallobase 1.0 to 4.8 times longer than phallobasic apodeme. Paramere: well developed; dorsum usually

convex, explanate medially or not; mesodorsal margin denticulated or not; venter usually concave;

dorsal and ventral sinuses well defined. Phallus:

marginal denticles present or not; phallic plicae

usually well developed.

Female Genitalia: For a general description see

page 18. Dorsal lamina bilobed (Figure 101), or

trilobed (Figure 71), rarely heptalobed (Figure 318);

ventral lamina usually trilobed (Figure 85), rarely

unilobed (Figure 48), or pentalobed (Figure 70);

laminae often deeply incised; proctigeral bacculi

fused (Figure 49) or not (Figure 71); ventral

bacculi never fused, acuminate at posterior twothirds or near middle; coxital plates present or

absent.

Internal Organs: The treatise of the alimentary

canal, ventral nerve cord, and the reproductive

organs of P. reventazon, (p. 18) adequately serves

as a generic characterization of the internal organs

considered. The accessory glands of the male reproductive organs, however, are somewhat variable

interspecifically. The most notable variation involves the anterior glands, which are feebly coiled

in P. ventralis (Figure 316), and the posterior

glands which are exceptionally long in P. chaletoides (Figure 277).

DISTRIBUTION.—The species of this genus occur

in the New World only. Except for P. ornaticollis,

whose northern limit is recorded as Ohio (USA),

the genus is exclusively Neotropical and ranges

from Mexico to central Brazil. Most of the species

occur in Central America, from Mexico to Panama.
NUMBER 227 23

Key to the Species of Perilypus

I. Antenna (Figure 67) loosely clubbed, funicular articles usually filiform, rarely serrate 2

Antenna (Figure 8) not clubbed, serrate 3

2.(1) Elytron copiously punctate (Mexico to Nicaragua) {frontalis group) 8

Elytral punctations absent, or few punctations in posthumeral region (Mexico) (criocerides group) 9

3.(1) Elytra (Figures 279, 293) ovate, and/or elytral surface copiously impressed with small

shallow punctations; punctate and interpunctate elytral surfaces arenose, rarely

smooth and shiny; pronotum (Figure 279) boldly transverse; antenna (Figure 284)

usually boldly serrate 4

Elytra (Figure 2) rectangulate; elytral surface impressed with large deeply impressed

punctations (Figure 2), punctations profusely distributed (Figure 2), or concentrated

in posthumeral region (Figure 99); punctate and interpunctate elytral surface usually smooth and shiny; pronotum (Figure 110) feebly transverse, length rarely equals

width; antenna rarely boldly serrate 6

4.(3) Mesoscutellum (Figure 312) transverse; femora usually dark brown or black and with

subapical annulus (Figure 56), rarely entirely dark brown or flavotestaceous (USA

to Venezuela) (ornaticollis group) 10

Mesoscutellum subquadrate or trigonal; femora predominantly flavotestaceous or entirely dark brown or black, rarely annulated subapically 5

5.(4) Dorsal fades entirely stramineous or predominantly castaneous (Mexico) (chaletoides

group) , 17

Dorsal fades not as above; elytral surface entirely black, rarely predominantly

piceous and vittate discally (Mexico to Guatemala) (quadrilineattis group) 18

6.(3) Epipleural fold convex, and/or elytral surface impunctate at apical fourth or more 7

Epipleural fold plane, concave or grooved longitudinally; elytral surface coarsely punctate throughout (Mexico to Peru) (reventazon group) 22

7.(6) Elytral apical fourth impunctate or with few very small punctations (but see variation

under P. levis) (Panama to Brazil) (viridipennis group) 56

Elytral surface coarsely punctate throughout (Coast Rica to Peru) (limbatus group) 58

8.(2) Front tarsal claws asymmetric; epipleural fold grooved longitudinally (males), or

shifted dorsally (females) by boldly convex epipleuron at elytral middle half (Figure

61) (Mexico to El Salvador) 1. P. frontalis (Gorham), new combination

Front tarsal claws symmetric; epipleural fold convex from elytral base to elytral apex

(El Salvador to Nicaragua) 2. P. setuUis, new spedes

9.(2) Antennal club compact (Figure 80), funicular articles filiform; elytron predominantly

flavotestaceous; posterior margin of pygidium feebly emarginated in males (Figure

86), entire in females (Mexico) 3. P. criocerides (Gorham), new combination

Antennal club lax (Figure 96), funicular articles serrate; posterior margin of pygidium

deeply emarginated in males (Figure 94), feebly emarginated in females (Figure 95);

elytron dark blue or purpurescent (Mexico) 4. P. insectus, new spedes

10.(4) Femora entirely dark brown or entirely flavotestaceous, not annulated 11

Femora predominantly black, annulated (Figure 56) 12

11.(10) Pronotal discal vitta furcate (Figures 322-324); elytral discal vitta (when present) not

strongly diverged from elytral suture at elytral apical fifth; femora entirely dark

brown (USA to Mexico) 43. P. ornaticolUs (LeConte), new combination

Pronotal discal vitta reduced to a midbasal macula (Figure 342); elytral discal vitta

strongly diverged from elytral suture at elytral apical fifth (Figure 342); femora

entirely flavotestaceous (Panama) 49. P. floralis (Gorham), new combination

12.(10) Fourth and fifth visible abdominal sternum flavous; mesoscutellar disc concave; average size about 13 mm (Mexico to Guatemala)

42. P. vtntralu (Gorham), new combination

Fourth and fifth visible sternum black; mesoscutellar disc plane; average size from 6

to 8 mm IS

13.(12) Epipleural fold plane 14

Epipleural fold obliquely convex 16
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

14.(13) Elytron sinuate apically (Figure 354), surface with stout vertical setae, sutural margin

diverged at apical fifth (Figure 354) (Panama) 46. P. pilatus, new species

Elytron truncate apically, vertical setae and sutural margin normal 15

15.(14) Antennal article 11 as long as preceding three combined (Figure 331) (Mexico)

45. P. prolixicornis, new species

Antennal article 11 as long as preceding two combined (Figure 332) (Mexico to Guatemala) 44. P. galbeus, new species

16.(12) Elytral apex uplifted (Figure 353), surface dull, epipleural fold obliquely convex (Costa

Rica) 47. P. tmtarius, new species

Elytral apex not uplifted but sinuatotruncate. surface shiny, epipleural fold evenly

convex (Venezuela) 48. P. sinuapicis, new species

17.(5) Elytral surface entirely stramineous (Mexico)

36. P. bicolor (Chevrolat), new combination

Elytral surface predominantly rufous, apex and sutural margin usually black (Figure

266), rarely only faintly infuscated at suture (Mexico) 35. P. chaletoides, new species

18.(5) Epipleural fold obliquely convex, visible in specimen dorsal view; elytral vitta as in

Figure 279 (Mexico to Guatemala)

37. P. quadrittneatus (Chevrolat), new combination

Epipleural fold not obliquely convex and not visible in specimen dorsal view; elytral

disc avittate 19

19.(18) Elytral surface undulated longitudinally (Figure 305); pronotum not coarsely macrosculptured 20

Elytral surface not undulated; pronotum coarsely macrosculptured 21

20.(19) Pronotal vitta exceptionally broad, with oblong roseous macula at center; femur

testaceous ventroapically (Mexico) 39. P. coronijormis, new species

Pronotal vitta constricted medially, not maculated centrally, femora annulated subapically (Guatemala) 40. P. fiuctus, new species

21.(19) Elytral disc bicarinate (Figure 300); pronotum concave centrally and paralaterally.

coarsely punctate (Mexico) 41. P. bicristatus, new species

Elytral disc not carinate; pronotum plane, transversely wrinkled; posthumeral margin

briefly vitiate (Figure 287) (Mexico to Guatemala)

38. P. telephoroides (Gorham), new combination

22.(6) Posterior margin of sixth visible sternum emarginated (Figure 41) (males) 23

Posterior margin of sixth visible sternum not emarginated (Figure 42) (females) 39

23.(22) Front tarsal claws boldly asymmetrical 24

Front tarsal claws feebly asymmetrical, or symmetrical 28

24.(23) Length about 6 mm; elytron olive green or blue green, apex usually testaceous

(Mexico) 32. P. immitis, new species

Length more than 8 mm; elytral coloration not as above 25

25.(24) Antenna testaceous (Mexico to Costa Rica)

27. P. orthoplettridus (Thomson), new combination

Antenna entirely black or dark brown 26

26.(25) Pygidium transverse (Mexico) 29. P. latilobus, new species

Pygidium oblong 27

27.(26) Antennal articles 3-5 moderately serrate (Figure 8); procoxae infuscated (Guatemala) ....

26. P. calcaris, new species

Antennal articles 3-5 feebly serrate, nearly filiform; procoxae immaculate (Mexico) ...

28. P. exttis, new species

28.(23) Elytral apex testaceous (Panama) 23. P. bilineatus (Gorham)

Elytral apex not testaceous 29

29.(28) Antenna boldly serrated (Figure 244) SO

Antenna moderately serrated 31

30.(20) Femora infuscated at apex only; elytra not truncated (Panama)

24. P. cerroaxul, new species

Femora infuscated at distal third or more; elytra truncated (Peru)

31. P. crassut, new species

31.(29) Specimens north of Nicaragua 32

Specimens south of Nicaragua 35
NUMBER 227 25

32.(31) Body form slender (Figure 239); elytra feebly convex; elytral vertical setae about onethird longer than elytral reclinate setae; pronotal vitta constricted at subapical

depression (Mexico to Guatemala) 30. P. distinctvu (Chevrolat), new combination

Body form squat; elytra plane; elytral vertical setae about one-half longer than elytral

reclinate setae; pronotal vitta not as above S3

33.(32) Pronotal subapical depression shallow at middle; pronotal arch feebly convex, approaching plane (Guatemala) 19. P. nigriventris (Gorham)

Pronotal subapical depression deeply impressed throughout; pronotal arch boldly convex 34

34.(33) Parameres short, strongly arcuate (Figure 255); parameres with combined venter deeply

concave (Figure 255) (Mexico) 33. P. caliculus, new species

Parameres not as above, but as in Figure 258 (Mexico) 34. P. carbonarius Spinola

35.(31) Profemur black in ventroapical third or entirely flavotestaceous (Costa Rica)

21. P. claudut (Wolcott)

Profemur flavotestaceous in ventroapical third 36

36.(35) Metafemur entirely flavotestaceous or infuscated dorsoapically S7

Metafemur black in apical half or less (Costa Rica) 20. P. sapient is, new species

37.(36) Antenna more serrate than in Figure 8; metafemur flavotestaceous (Costa Rica)

22. P. cultratus, new species

Antennal serration as in Figure 8 38

38.(37) Metafemur infuscated dorsopreapically (infuscation narrowed to femoral apex dorso

laterally or not narrowed) (Costa Rica) 18. P. rrventazon, new species

Metafemur infuscated dorsoapically (Panama) 25. P. ordinatus, new species

39.(22) Epipleuron broadly explanate in elytral middle third (Figure 230) (Mexico)

27. P. orthopUuridus (Thomson)

Epipleuron not broadly explanate 40

40.(39) Sixth visible abdominal sternum conspicuously arenose and with broad shallow depressions; fourth and fifth visible sternum flavous (Guatemala)

26. P. calcaris, new species

Abdominal sterna not as above 41

41.(40) Elytral disc vittate (sometimes vitta only faintly visible) 42

Elytral disc avittate 51

42.(41) Elytral discal vitta closer to posthumeral margin than to sutural margin (Figure

186) 43

Elytral discal vitta about equidistant from side margins (Figure 193) 46

43.(42) Elytral discal vitta gradually incurved posteriorly, terminating at elytral apex 44

Elytral discal vitta acutely incurved posteriorly, terminating at sutural margin before

elytral apex 45

44.(43) Elytral sutural margin flavotestaceous at basal third or more (Mexico)

34. P. carbonarius Spinola

Elytral sutural margin piceous (Panama) ... 23. P. bilineatus (Gorham), new combination

45.(43) Femora black in distal half or more, not flavotestaceous ventroapically (Costa Rica)

21. P. claudus (Wolcott)

Femora infuscated in dorsodistal half, flavotestaceous ventroapically (Costa Rica)

18. P. reventaum, new species

46.(42) Anterior limit of elytral discal vitta before humeral margin (Figure 239); first five

visible abdominal sterna flavotestaceous 47

Anterior limit of elytral discal vitta at humeral margin (Figure 186); abdominal sternum

not as above 48

47.(46) Elytron feebly convex; metafemur infuscated at dorsoapical third (Mexico)

30. P. distinctTM (Chevrolat)

Elytron plane; metafemur infuscated at apex only (Costa Rica)

22. P. cultratus, new species

48.(46) Femora predominantly black (Mexico) 29. P. latilobus, new species

Femora predominantly flavotestaceous 49

49.(48) Antenna flavotestaceous (Costa Rica) 21. P. claudm (Wolcott)

Antenna black 50

50.(49) Meso- and metafemur black in apical third or more (Costa Rica)

20. P. sapientis, new species
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Meso- and metafemur black at apex only (Guatemala)

19. P. nigriuentris (Gorham), new combination

51.(41) Elytron broadly flavotestaceous at or near margin, fasciate (Figures 262-265), or entirely

Bavotestaceous (Mexico) 34. P. carbonarhu Spinola

Elytron not as above, disc entirely black 52

52.(51) Antenna boldly serrate (Figure 244); elytra feebly convex (Peru)

31. P. crassus, new species

Antenna moderately serrate (Figure 8); elytra plane 53

53.(52) Elytral apex testaceous and/or elytra olive green or blue green (Mexico)

32. P. unmitu, new species

Elytral apex not testaceous 54

54.(53) Femora flavotestaceous in basal third; pronotal vitta abruptly narrowed at subapical

depression (Mexico) 29. P. latUobus, new species

Femora entirely black; pronotal vitta not as above 55

55.(54) Pronotum boldly transverse (Mexico) 34. P. carbonarhu Spinola

Pronotum feebly transverse (Mexico) 33. P. caliculut, new species

56.(7) Antenna boldly serrate (Figure 244) and densely setose (Canal Zone to Brazil)

7. P. levis, new species

Antenna only moderately serrate and moderately setose 57

57.(56) Antenna uniformly black; elytral apical fourth impunctate or with very small punctations; epipleural fold more collapsed than convex (Panama to Colombia)

6. P. relucens (Gorham), new combination

Antenna predominantly dark brown, flavotestaceous apically; elytral apical half impunctate; epipleural fold convex (Panama to Venezuela)

5. P. viridipennis (Spinola), new combination

58.(7) Legs testaceous, except hind femur black in distal half (Panama)

16. P. tettaceicornis (Pic), new combination

Combination of leg colors not as above 59

59.(58) Epipleural fold wider than antennal pedicel (north of Colombia) 60

Epipleural fold not wider than antennal pedicel (South America) 62

60.(59) Elytral apex black; elytral discal vitta sharply incurved and broadly outcurved posteriorly (Figure 2) (Panama) 17. P. decoris, new species

Elytral apex flavotestaceous; elytral disc avittate, or vitta not curvate posteriorly 61

61.(60) Specimens from north of Panama; male genitalia as in Figures 172-174

15. P. prolixipenis, new species

Specimens from Panama; male genitalia as in Figures 167-169

14. P. latilira, new species

62.(59) Distal half of hind femur predominantly or entirely black; frons partially or totally

black (Figures 58, 59) 63

Distal half of hind femur predominantly flavotestaceous, only apex infuscated; frons

flavotestaceous (Figure 57) 66

63.(62) Inner margins and venter of antennal articles 9 to 11 testaceous (Colombia)

10. P. apocopates, new species

Inner margins and venter of antennal articles 9 to 11 black 64

64.(63) Femur entirely black 11. P. Ms, new species

Femur flavotestaceous basally 65

65.(64) Labrum and frons black (Figure 59) (Colombia)

9. P. cohmtbicus (Spinola), new combination

Labrum testaceous; frons testaceous anteriorly, black posteriorly (Figure 58) (Colombia)

13. P. buga, new species

66.(62) Antennal articles 9 to 11 testaceous ventrally (Venezuela to Bolivia)

8. P. Umbatus (Gorham), new combination

Antennal articles 9 to 11 black ventrally (Peru) 12. P. acus, new species

The frantalis Group

The members of this group have clubbed antennae, a characteristic that separates them from all

other species of Perilypus except the two species

that constitute the criocerides group. Specimens of

the frontalis group differ from those of the criocerides group by having the elytral surface copiously
NUMBER 227 27

FIGURES 57-61.—Anterior limit of black color of frons: 57, Perilypus limbatus, 58, P. buga,

59, P. columbicus. P. frontalis: 60, antenna, dorsal view, X 60 (arrows indicate apical border

of antennal articles); 61, female elytron, lateral view, X 27 (arrow indicates convex epipleuron).

(Scale = 1 nun)
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and deeply punctated; the paramere (Figures 73,

78) long, feebly pigmented, and without lateral

depression; and the phallic plate (Figures 74, 79)

devoid of marginal denticles.

Specimens of the frontalis group superficially

resemble those of the genus Placopterus (Wolcott,

1910b: 363) from which they can easily be distinguished as follows: in frontalis group specimens the

antenna extends beyond the pronotal collar, antennal article 11 is oblong, body form is rectangulate,

and pronotal subapical depression is sinuous. In

Placopterus species the antenna does not reach the

pronotal collar, antennal article 11 is quadrate or

transverse, body form is more oval, and the pronotal

subapical depression is broadly arcuate.

The distribution of this group extends from

Chiapas, Mexico, to Rivas, Nicaragua.

1. Perilypus frontalis (Gorham), new combination

FIGURES 60-76, 356

Poecilochroa frontalis Gorham, 1886:338 [holotype: female,

deposited in BMNH; type-locality: Ciudad Guatemala,

Departamento de Guatemala, Guatemala].

Derestenus apicalis Pic, 1941:10 [lectotype: female, deposited

in MNHP, here designated; type-locality: Guatemala; new

synonymy].

Pic's type-specimen does not differ significantly

from Gorham's type-specimen.

DIAGNOSIS.—The extraordinary degree of asymmetry between the inner and outer front tarsal

claws (Figure 68) will distinguish males of this

species from those of any other congeneric species.

Female specimens are easily recognized by the

boldly convex epipleuron, which is most developed

in elytral middle half. The modification of the

female epipleuron causes the epipleural fold to be

shifted upward and the confluence of both structures gives the epipleural margin a tubular appearance (Figure 61).

DESCRIPTION.—Form: As in Figure 62.

Size: Length: males, average about 6.3 mm, range

5.2-7.4 mm; females, average about 6.7 mm, range

5.7-7.8 mm. Width: males, average about 1.9 mm,

range 1.8-2.3 mm; females, average about 2.2 mm,

range 1.8-2.7 mm. Ten males and 10 females

measured.

Color: Cranium either flavotestaceous, pale

orange, flavotestaceous with midepicranial black

macula, or clypeus, frons, and cranial venter flavotestaceous and epicranium and gena black; antenna

black; prothorax either flavotestaceous at sides and

with percurrent or medially interrupted black discal

vitta, or flavotestaceous or pale orange and with or

without dark brown midapical macula and dark

brown collar; femur either flavotestaceous in basal

four-fifths and black in apical fifth, flavotestaceous

in basal fifth and black in apical four-fifths, or

entirely flavotestaceous; tibia black or yellow; mesoand metasternum piceous; elytron either shiny

black but apex flavotestaceous (Figure 62), black

with sutural and marginal flavotestaceous vitta

(Figure 63), as in Figure 64, or stramineous and

infuscated subapically (Figure 66); abdomen black.

Head: Interocular depressions broad and shallow; eyes shallow; frons (Figure 65) conspicuously

broad (HW/IOW, average about 1.9, range 1.8-2-1;

10 males and 10 females measured); antenna

clubbed (Figures 60, 67); funicular articles feebly

serrate; apical border and disc of antennal articles 7

and 8 not microsetiferous; AL/PLS, about 1.7;

length /width ratio of each male antennal article

2.5:1.4:2.0:2.0:2.0:1.7:1.8:1.7:1.3:1.1:1.7.

Pronotum: PL/PW, average about 0.86, range

7.9-9.2 (10 males and 10 females measured); subapical and prebasal depressions deeply impressed;

side margin of pronotal proper boldly convex.

Mesoscutellum: Subquadrate.

Elytron: Epipleural margin straight in basal

three-fourths, broadly arcuate in apical fourth; slope

of apical deflection gradual; discal punctations well

impressed and evenly distributed; punctate and

interpunctate surfaces smooth, shiny, and densely

covered with pale reclinate setae; vertical setae

piceous; epipleuron and epipleural fold variable

(see "Variation"); EL/EW, average about 4.3, range

3.9-4.9 (10 males and 10 females measured).

Front Tarsal Claws: Strongly asymmetrical in

both sexes; inner claw abom *— e times longer

than outer claw in males (tifc-.e 69), and about

one and a half times longer in females (Figure 68).

Male Genitalia (Figures 72-74): Feebly pigmented. Phallobase very broad. Combined length

of phallobase and phallobasic apodeme 1.5 times

longer than paramere. Combined length of paramere and phallobase 3.0 times longer than phallobasic apodeme. Paramere: apex obtuse, feebly

declinate in lateral view; lateral depression absent;

dorsum convex, explanate medially; venter narrow,
NUMBER 227 29

62

63

64

66

FIGUKES 62-66.—Perilypus frontalis: 62-63, habitus; 64, elytra; 65, head, front view, X 30;

66, elytra. (Scale = I mm)
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mesoventral margin broadly arcuate; mesodorsal

margin sinuous. Sinus: dorsal narrow, lanceolate;

ventral elliptical. Phallus: marginal denticles absent; phallic plicae well developed (47 specimens

examined).

Female Genitalia (Figures 70, 71): Laminae very

short; dorsal lamina trilobed, lobes equal in length;

ventral lamina pentalobed, paralateral pair of lobes

shorter than medial lobe; proctigeral bacculi widely

separated; ventral bacculus acuminate at posterior

two-thirds; coxital plates absent (2 specimens

examined).

Internal Reproductive Organs: Male (Figures 75,

76); anterior accessory gland uniramous, tightly

coiled; posterior accessory gland biramous, outer

branch little less than twice as long as inner branch;

testis composed of 12 follicles. Female, as in Figure

55 (5 males and 2 females examined).

VARIATION.—Structural: The degree of asymmetry of the front tarsal claws varies with sex (see

"Front Tarsal Claws") as does development of the

epipleuron and the epipleural fold. In the female

elytron the epipleuron and epipleural fold are

boldly convex (Figure 61); in males the epipleuron

is not convex and the epipleural fold is grooved

longitudinally.

The degree of asymmetry of the front tarsal claws

also varies slightly independent of sex. The epipleuron and epipleural fold is collapsed in some female

specimens; however, this variation is probably due

to incomplete cuticular expansion.

Color: Variation in color, not correlated with

sex or geography, is extreme (Figures 62-66). In

general, color of various parts of the integument

not normally black or brown ranges from stramineous to fulgidus.

NATURAL HISTORY.—In June, in Sumpango,

Guatemala, I collected four specimens at 2012

meters and in Zinacatan, Mexico, three specimens at

1830 meters. The Guatemalan sample was collected

by beating roadside shrubs while those from Mexico

were collected by beating oak branches (Quercus

sp.). In May, on Cerro Verde, El Salvador, H. F.

Howden collected 13 specimens at 2000 meters.

Adult specimens were also collected in July, in

Mexico.

DISTRIBUTION (Figure 356).—The known range

of this species extends from southern Mexico to

southern £1 Salvador and southwestern Guatemala.

LOCALITY RECORDS (Figure 356).—I examined 110 adult

specimens from Central America. MEXICO: Estado de Chiapas: Chiapas (ZMAN, 1 female) ; 2 miles [3.2 km] northwest of Pueblo Nuevo (GHNE, 1 female); 20 miles [32 km]

north of Bochil, Yerba Buena (CNCC, 1 female; GEKI, 2

females); San Crist6bal de las Casas (GEKI, 1 male and

1 female); Huistan area, 27 miles [43 km] northeast of San

Cristobal de las Casas (GEKI, 1 male); 11 miles [18 km]

northeast of San Cristobal de las Casas (GEKI, 1 female);

7 miles [11 km] southeast of San Cristobal de las Casas

(HFHO, 1 male); Teopisca (MNHB, 1 male and 5 females; GEKI, 2 males and 5 females); Highway 24, 8 to 10

miles [13 to 16 km] southeast of Teopisca (CNCC, 4 males

and 3 females; GEKI, 2 males and 2 females; HFHO, 1

male and 3 females) ; Comitan (GEKI, 7 males and 5 females; MNHB, 7 males and 13 females); El Rincon, route

17 (HFHO, 1 female); Chincultic, near El Rincon (GEKI,

1 female); Zinacatan (GEKI, 1 male and 2 females; NMNH,

1 female). GUATEMALA: Departamento de Guatemala:

Ciudad Guatemala (BMNH, 1 female, holotype); Departamento de Sacatepequez: Duerias (BMNH, 1 male and 1 female; GEKI, 1 female); Capetillo (BMNH, 2 males and 1

female; GEKI, 4 males and 1 female; MNHB, 1 male; MNHP,

2 females; NMNH, 1 male). Departamento de Chimaltenango: Calderas (BMNH, 1 female; GEKI, 1 female).

"Guatemala" (MNHP, 1 female). EL SALVADOR: Departamento de Santa Ana: 23 kilometers north of Metapan

(GEKI, 1 female). Departamento de San Miguel: Cerro

Verde (CASC, 1 male; GEKI, 4 males and 2 females; HFHO,

6 males and 2 females).

2. Perilypus sensilis, new species

FIGURES 77-79, 356

TYPE-LOCALITY.—Cerro San Jer6nimo, Departamento de La Paz, El Salvador.

TYPE-SPECIMENS.—The holotype male is deposited in MNHP, the allotype in BMNH. Both

specimens were collected by G. C. Champion. Two

paratypes: GEKI, 1 male: HFHO, 1 male.

DIAGNOSIS.—Distinguishable from P. frontalis,

the only other member of the frontalis group, by

symmetry of the front tarsal claws, by the evenly

convex epipleural fold (male and female), and by

the less convex side margins of the pronotum

proper.

DESCRIPTION.—Form: As in P. frontalis (Figure

62).

Size: Length: males, average about 6.8 mm, range

5.6-8.0 mm; females 7.6 mm. Width: males, average about 2.2 mm, range 1.8-2.9 mm; females 2.2

mm. Three males and 1 female measured.

Color: Clypeus, from, and cranial venter flavotestaceous, epicranium and gena black; antenna
NUMBER 227 31

67

70

FIGURES 67-74.—Perilypus frontalis: 67, antenna; 68, female front tarsal claw, X 140; 69, male

front tarsal claw, X 140; 70, ovipositor, ventral view; 71, ovipositor, dorsal view; 72, tegmen.

lateral view; 73, tegmen, ventral view; 74, phallus. (Scale = 1 mm)
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

75

79

FICURFS 75-80.—Perilypus frontalis: 75, male internal reproductive organs; 76, posterior accessory

glands. P. sensilis: 77, tegmen, lateral view; 78, tegmen, ventral view; 79, phallus. P. criocerides: 80, antenna. (Scales = 1 mm)
NUMBER 227

black; prosternum and side margin of pronotum

flavotestaceous; pronotal disc broadly vittate;

femora flavotestaceous in basal four-fifths, dark

brown in apical fifth; tibia and tarsus dark brown;

meso- and metasternum black; elytron flavotestaceous along epipleural margin and at apex, black

in remainder; abdomen black.

Head: Cranium and antenna as in P. frontalis

except eyes more convex and AL/PLS 1.9.

Pronotum: As in P. frontalis except less transverse (PL/PW, average about 0.93, range 0.87-0.98;

4 specimens measured) and side margin of pronotum proper less convex.

Elytron: As in P. frontalis except surface less

shiny, punctations more conspicuous, and epipleural fold (of both sexes) evenly convex from

elytral base to elytral apex.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 77-79): Feebly pigmented. Phallobase very broad. Combined length

of phallobase and phallobasic apodeme 3 times

longer than paramere. Combined length of paramere and phallobase 2.8 times longer than phallobasic apodeme. Peramere: apex broadly incurved

and obtuse; apical third declinate in lateral view;

lateral depressions absent; dorsum convex, broadly

explanate medially; venter narrow and feebly concave; mesoventral margin broadly concave; mesodorsal margin sinuous. Sinus: dorsal narrowly

lanceolate; ventral elliptical. Phallus: as in P.

frontalis except phallic plates broader (3 specimens examined).

Female Genitalia: Not studied.

VARIATION.—Structural: No structural variation

was noted.

Color: The only noticeable variation is in the

intensity of the vitta on the elytral epipleural margin; the vitta is most pronounced in specimens

from Guatemala.

NATURAL HISTORY.—G. C. Champion collected

the holotype from Cerro San Jer6nimo, Guatemala,

at 914 meters. In May, in Boquer6n, El Salvador,

H. F. Howden collected one specimen at 1800

meters.

DISTRIBUTION (Figure 356).—The known range

extends from Alta Verapaz, Guatemala, to San Juan

del Sur, Nicaragua.

ETYMOLOGY.—Latin, adjective sensilis (sensitive),

with reference to the inferred sensory function of

the microsetae on antennal articles 9 to 11.

LOCALITY RECORDS (Figure 356).—I examined 4 adult

specimens from Central America. GUATEMALA: Departamento de Alta Verapaz: Purulha (BMNH, 1 female). EL

SALVADOR: Departamento de San Salvador: Boquer6n, near

Santa Tecla (HFHO, 1 male) ; Departamento de La Paz:

Cerro San Jeronimo (MNHP, 1 male, holotype). NICARAGUA: Departamento de Rivas: San Juan del Sur (GEK.I,

1 male).

The criocerides Group

The group consists of two rather dissimilar species. In both of these the antenna is clubbed,

elytral surface smooth and shiny and only sparcely

vested with small punctations behind the humerus,

and the male pygidium is conspicuously emarginated posteriorly (Figures 89, 94). Components of the

male genitalia are strongly pigmented and phallic

marginal denticles are present. In P. criocerides the

phallic plates are denticulated on both margins

(Figure 84), a characteristic not present elsewhere in

Perilypus.

Members of this group are known only from

southern Mexico.

3. Perilypus criocerides (Gorham),

new combination

FIGURES 80-87, 356

Colyphus criocerides Gorham, 1882:144 [holotype: male, deposited in BMNH; type-locality: Cerro de Plumas, Estado

de Veracruz, Mexico].

DIAGNOSIS.—The compact yellow antennal club

(Figure 80) and nearly impunctate and predominantly flavotestaceous elytron (Figure 81) will distinguish members of this species from other species

occurring in Mexico.

DESCRIPTION.—Form: As in Figure 81.

Size: Length: male, 8.0 mm; females, average

about 7.7 mm, range 7.4-8.1 mm. Width: male 2.2

mm; females, average about 4.4 mm, range 4.3-4.5

mm. One male and 3 females measured.

Color: Cranium flavotestaceous; first 8 antennal

articles dark brown, article 9 to 11 flavotestaceous;

prothorax flavotestaceous, pronotum with or without midapical macula or percurrent discal vitta;

meso- and metasternum black; femur predominantly flavotestaceous, infuscated apically; tibia and

tarsus dark brown; elytron entirely flavotestaceous,

or predominantly flavotestaceous and with one
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

basosutural and one discal black macula (Figure

81); abdomen dark brown.

Head: Frons broad (HW/IOW, 1.9; 4 specimens

measured); interocular depressions broad, deeply

impressed; eyes boldly convex; antenna (Figure 80)

distinctly clubbed, funicular articles filiform, articles 7 and 8 without microsetae; length/width ratio

of each male antennal article 2.0:1.0:2.2:2.2:2.2:

2.0:1.6:1.4:0.85:0.60:1.1.

Pronotum: Feebly transverse (PL/PW, average

about 0.91, range 0.88-0.93; 4 specimens measured),

subapical depression deeply impressed and strongly

sinuous, side margin of pronotum proper boldly

convex.

Mesoscutellum: Subquadrate.

Elytron: Posthumeral margin straight in basal

half, broadly arcuate in apical half; slope of apical

deflection gradual; surface punctations faintly visible posthumerally and near epipleural margin;

punctate and interpunctate surfaces smooth and

shiny; epipleural fold grooved longitudinally proximal to epipleural margin, feebly convex more dorFIGURFS 81-87.—Penllypus criocerides: 81, habitus; 82, tegmen, lateral view; 83, tegmen, ventral

view; 84, phallus; 85, ovipositor, ventral view; 86, male pygidium; 87, ovipositor, dorsal view.

(Scales = 1 mm)
NUMBER 227 35

sad; EL/EW, average about 4.4, range 4.3-4.5 (4

specimens measured).

Abdomen: Posterior margin of pygidium emarginated in males (Figure 86), arcuate in females.

Front Tarsal Claws: Asymmetrical in both sexes.

Male Genitalia (Figures 82-84): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5 times longer than paramere.

Combined length of paramere and phallobase 2.7

times longer than phallobasic apodeme. Paramere:

lateral depression present; dorsum feebly convex;

venter feebly concave; mesoventral margin concave;

mesodorsal margin irregular, feebly explanate

medially, and boldly braced basally. Sinus: dorsal

lanceolate; ventral broadly elliptical. Phallus: Phallic plates denticulated on both margins; phallic

plicae well developed (1 specimen examined).

Female Genitalia (Figures 85, 87): Dorsal and

ventral lamina trilobed, lateral lobes shorter than

median lobe; proctigeral bacculi widely separated;

ventral bacculus acuminate near posterior twothirds; coxital plates absent (1 specimen examined).

VARIATION.—Structural: The available specimens

are structurally homogeneous.

Color: The dorsum of one female is entirely

flavotestaceous. The black markings on the pronotum and the elytron vary in size.

NATURAL HISTORY.—J. M. Campbell collected

three adults during July, in Chiapas, Mexico.

DISTRIBUTION (Figure 356).—Known only from

Veracruz and Chiapas, Mexico.

LOCALITY RECORDS (Figure 356).—I examined 4 adult

specimens from Central America. MEXICO: F.stado de

Chiapas: 17 kilometers north of Tuxtla Gutierrez (GEKI,

3 females) . Estado de Veracruz: Cerro de Plumas (BMNH,

1 male, holotype).

4. Perilypus insectus, new species

FICURES 8&-96, 356

TYPE LOCALITY.—Mexico.

TYPE-SPECIMENS.—The holotype male and the

allotype, deposited in MNHB, were collected by J.

Flohr G. One paratype: GEKI, 1 male.

DIAGNOSIS.—The extent of emargination of the

pygidial posterior margin (Figure 94) distinguishes

male specimens. In female specimens the pygidium

is feebly emarginated (Figure 95). Small size and

metallic luster of the elytron are diagnostic characteristics for both sexes.

DESCRIPTION.—Form: As in Figure 88.

Size: Length: male, average about 6.0 mm, range

5.8-6.2 mm; female, 6.0 mm. Width: male, average

about 1.8 mm, range 1.7-1.9 mm; female 1.8 mm.

Two males and 1 female measured.

Color: Cranium flavotestaceous; antenna flavotestaceous to light brown; prothorax flavotestaceous,

except pronotum with brown midapical macula;

mesoscutellum flavotestaceous; pro- and mesofemur

flavotestaceous, except infuscated dorsoapically;

metafemur flavotestaceous in basal half, brown in

apical half; tibia and tarsus brown; mesosternum

flavotestaceous, metasternum brown; elytron shiny,

metallic dark blue or purple; abdomen black.

Head: As in P. criocerides except frons broader;

HW/IOW, 1.7 (3 specimens measured); antennal

club less compact, funicular articles serrate (Figure

96); and length/width ratio of each male antennal

article 2.1:1.3:1.4:1.7:1.3:1.7:1.4:1.4:1.4:1.2:2.0.

Pronotum: As in P. criocerides except side margins of pronotum proper less convex.

Elytron: As in P. criocerides except basal half behind humerus finely punctate and distal half impunctate.

Abdomen: Posterior margin of pygidium deeply

emarginated in male (Figure 94), feebly emarginated in female (Figure 95).

Front Tarsal Claws: Asymmetrical in both sexes.

Male Genitalia (Figures 89-91): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 3.5 times longer than paramere.

Combined length of paramere and phallobase 2.6

times longer than phallobasic apodeme. Paramere:

carinate lateroapically; attenuate; lateral depression

well impressed and localized medially; dorsum

feebly convex; venter concave; mesoventral margin

strongly sinuous; mesodorsal margin feebly sinuous

and boldly braced basally. Sinus: dorsal subparallel,

ventral broadly elliptical. Phallus: phallic plate

broad to phallic plicae; marginal denticles extended

from basal third to phallic plicae (2 specimens examined).

Female Genitalia (Figures 92, 93): Laminae and

coxites very short; dorsal lamina trilobed, lobes

about equal in length; ventral lamina pentalobed,

paralateral pair of lobes shorter than medial lobe;

proctigeral bacculi widely separated; coxital plates

absent.

VARIATION.—Structural: Emargination of the py-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 88-95.—Perilypus insectus: 88. habitus; 89, tegmen, lateral view; 90, tegmen, ventral

view; 91, phallus; 92, ovipositor, ventral view; 93, ovipositor, dorsal view; 94, male pygidium;

95, female pygidium. (Scales = 1 mm)

gidium is sex dimorphic (Figures 94, 95) as is the

degree of asymmetry of the front tarsal claws; the

claws are more asymmetrical in males than in the

female.

Color: Elytron may be dark blue or purple. The

antenna varies in extent of paleness.

ETYMOLOGY.—Latin, insectus (to cut into; masculine past participle of inseco). I refer to the emargination of the pygidial posterior margin.

DISTRIBUTION (Figure 356).—Known only from

"Mexico."

LOCALITY RECORDS (Figure 356).—I examined 3 adult

specimens from Central America. "Mexico" (GEKI, 1 male;

MNHB, 1 male, holotype, and 1 female).

The vtridipennis Group

The conspicuous absence of deeply impressed

punctations from the apical third (or more) of the
NUMBER 227 37

elytron is the major diagnostic characteristic for the

group. Others are antenna pronouncely serrate, and

elytral punctate and interpunctate surface smooth

and shiny. Elytral coloration (Figures 103, 110) is

unique in Perilypus.

Geographically, the members of this group are

distributed from northern Panama to northeastern

Brazil with distribution records from northern

Colombia and Venezuela.

5. Perilypus viridipennis (Spinola),

new combination

FIGURES 97-108, 357, 358

Systenoderes viridipennis Spinola 1844a: 132 [holotype: sex

not determined, deposited in MSPI; type-locality: Colombia].— Desmarest, [1860]:239.—Ekis, 1975:27.

Cleronomus amaenus Schenkling, 1906:263 [lectotype: male,

deposited in DEIE, and 1 paralectotype, female, deposited

in ZMAN, here designated; type-locality: "Magdalenstrom" = Rio Magdalena, Colombia; new synonymy].

Schenkling's type-specimen does not differ significantly from the type of P. viridipennis.

DIAGNOSIS.—The gradual increase of flavotestaceous color in the distal half of the antenna, and the

lack of deeply impressed punctations at the elytral

apical half easily distinguishes specimens of this

species from those of the other two species in the

viridipennis group.

DESCRIPTION.—Form: As in Figure 98.

Size: Length: males, average about 5.8 mm, range

5.7-7.0 mm; females, average about 6.5 mm, range

5.9-7.6 mm. Width: males, average about 1.8 mm,

range 1.6-2.1 mm; females, average about 2.2 mm,

range 1.8-2.4 mm. Ten males and 10 females measured.

Color: Cranium rufescent; antenna bicolorous,

basal half dark brown, distal half increases in paleness; thorax rufescent except female pronotum (and

usually of males) with dark brown midapical macula; mesoscutellum rufescent; femora flavotestaceous, usually infuscated dorsally; elytron variable

(see "Variation"); abdominal venter rufescent.

Head: Interocular depressions shallow; frontal

umbo shallow; eyes moderately convex; HW/IOW,

average about 2.2, range 2.0-2.3 (10 males and 10

females measured); antenna (Figure 97) serrate;

AL/PLS, 1.4; length/width ratio of each male antennal article 2.4:1.4:1.8:1.8:1.5:1.3:1.2:1.1:1.0:0.93:1.5.

Pronotum: Only slightly transverse, rarely length

equals width (PL/PW, average about 0.95, range

0.90-1.0; 10 males and 10 females measured); subapical and prebasal depressions deeply impressed;

side margin of pronotum proper moderately arcuate.

Elytron: Epipleural margin straight in basal half,

broadly arcuate in apical half; elytral convexity

varied (see "Variation); large punctations localized

behind humerus (Figure 99); punctate and interpunctate surfaces smooth and shiny, not arenose;

epipleural fold feebly convex, but lower region

slightly collapsed; slope of apical deflection gradual;

EL/EW, average about 4.2, range 3.8-5.0 (20 specimens measured).

Front Tarsal Claws: Male, feebly asymmetrical.

Female, symmetrical.

Male Genitalia (Figures 104-106): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 6 times longer than paramere.

Combined length of paramere and phallobase 4.0

times longer than phallobasic apodeme. Paramere:

apical region papilliform in lateral view; lateral

depression broad and well depressed, clearly visible

in ventral view; dorsum convex and explanate

medially; venter concave; mesoventral margin concave; mesodorsal margin sinuous and denticulate.

Sinus: dorsal constricted at basal third, spheroid

behind constriction, narrowly elliptical in front of

constriction. Phallus: marginal denticles of phallic

plate broad, shallow, and extended to phallic plicae

(9 specimens examined).

Female genitalia (Figures 100, 101): Dorsal lamina

bilobed, deeply incised; ventral lamina trilobed,

lateral lobes slightly shorter than medial lobe;

proctiger robust; proctigeral bacculi not fused;

ventral bacculi acuminate at posterior third; coxital

plates absent (1 specimen examined).

Internal Reproductive Organs: Male (Figures 107,

108); anterior accessory gland uniramous, tightly

coiled; posterior accessory gland biramous, outer

branch one-third longer than inner branch; testis

composed of 12 follicles; basal third of ejaculatory

duct exceptionally bulbous. Female as in Figure 55

(2 males and 1 female examined).

VARIATION.—Structural: The convexity of the

elytron seems to correlate geographically, the elytra

being more convex in specimens from Panama than

in those from Venezuela.

Color: In some Panamanian specimens the midapical macula of the pronotum extends posteriorly
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

98

FIGURES 96-103.—Perilypus insectus: 96, antenna. P. viridipennis: 97, antenna; 98, habitus; 99,

elytral posthumeral region; 100, ovipositor, ventral view; 101, ovipositor, dorsal view; 102-103,

elytra. (Scale = 1 mm)
NUMBER 227 39

to form a narrow discal streak. Variation of elytral

coloration suggests an east to west cline, but it may

also correlate with altitude as the few elevation

records indicate. The lighter specimens were collected at low elevations in Panama, the darker

ones at higher Andean elevations in Colombia and

Venezuela. Panamanian specimens are predominantly rufescent and have a black to violaceous

basal macula on the elytron (Figure 103). This

macula may be confined to the humeral angle or

extend from the post humeral margin to the sutured margin. In specimens from Colombia the

elytron may be entirely blue, or blue in basal and

apical thirds and rufescent in middle third (Figure

102). Venezuelan specimens have the elytron shiny

blue or shiny blue-green.

NATURAL HISTORY.—In May, on Cerro Campana,

Panama, I collected a female by beating woody

shrubs and lianas at an altitude of 460 meters. At

the same site and by the same collecting method,

H. Stockwell collected two male adults in January.

Other specimens were collected in Panama in

February, March, May, June, July, and November,

and in Colombia in May. The specimens from

Venezuela were collected in June by J. and B.

Bechyne\ one at 850 meters, the other at 1000

meters.

DISTRIBUTION (Figures 357, 358).—Specimens were

collected in central Panama, northern Colombia,

and northern Venezuela.

LOCALITY RECORDS (Figures 357, 358).—I examined 21 adult

specimens from Central and South America. PANAMA:

Provincia de Code: El Valle (NMNH, 1 male and 1 female).

Provincia de Panama: Rio Diablo (GEKI, 1 male) ; Cerro

Campana (GEKI, 1 male and 1 female; HPST, 1 male).

Cacahualito (CMPP, 1 male). Provincia de Colon: Portobelo

(GEKI, 1 male). Canal Zone: Summit (GEKI, 1 female);

3 miles [4.8 km] west of Paraiso (GEKI, 1 female); Barro Colorado Island (WBIV, 1 male). COLOMBIA: Departamento de

Magdalena: Sevilla (MCZC, 1 male and 1 female); Aracataca

(GEKI, 2 females; MCZC, 1 female); Rio Magdalena (DEIE,

1 male; ZMAN, 1 female). Colombia (MSPI, 1 specimen, sex

not known). VENEZUELA: Estado de Carabobo: Las

Trincheras, Hacienda El Palmar (CMPP, 1 female); Cerro

Aquirre (GEKI, 1 male) .

6. Perilypus relucens (Gorham), new combination

FICURES 109-114,357

Colyphus relucens Gorham, 1886:336 [lectotype: female, deposited in BMNH, and 4 paralectotypes, females, 2 deposited in BMNH, 1 in GEKI, and 1 in MNHP. here

designated; type-locality: Volcan de Chiriquf, Provincia de

Chiriquf, Panama].

DIAGNOSIS.—Specimens of this species can be

easily confused with Panamanian specimens of P.

viridipennis, primarily because their elytral coloration is very similar (compare Figures 103, 110).

Anatomically, however, the two species are quite

distinct. Specimens of P. relucens are generally

broader, their elytra less convex posteriorly and

feebly depressed at middle third. The convexity of

their epipleural fold is almost obliterated; the

fold is collapsed and appears grooved longitudinally. Also their elytral punctations are slightly

smaller, extend to the suture, and terminate at

elytral posterior fourth.

DESCRIPTION.—Form: As in Figure 110.

Size; Length: males, average about 6.9 mm, range

5.9-7.7 mm; females, average about 7.4 mm, range

6.1-8.0 mm. Width: males, average about 2.2 mm,

range 2.0-2.4 mm; females, average about 2.5 mm,

range 2.2-2.8 mm. Ten males and 10 females

measured.

Color: Cranium refescent; antenna refescent or

brown; prothorax rufescent, with or without dark

brown midapical macula; scutellum, meso- and

metasternum, legs, and abdomen rufescent or dark

brown; elytron either blue in basal fourth and

rufescent in apical three-fourths, or entirely blue

and with a violaceous or virescent tinge.

Head: As in P. viridipennis except interocular

width proportionally wider (HW/IOW, average

about 1.9, range 1.9-2.1; 20 specimens measured);

eyes shallower; antenna (Figure 111) proportionally

longer (AL/PLS, 1.6); and length/width ratio of

each male antennal article 2.0:1.2:1.8:1.6:1.4:1.4:

1.4:1.4:1.3:1.2:1.8.

Pronotum: As in P. viridipennis except more

transverse (PL/PW, average about 0.87, range 0.77-

0.98; 20 specimens measured), less convex, and

subapical depression more sinuous.

Elytron: As in P. viridipennis except less convex,

feebly depressed at middle third, punctations larger

(Figure 109) and extended to posterior threefourths, epipleural fold collapsed, and epipleural

margin less convex at posterior third.

Front Tarsal Claws: As in P. viridipennis.

Male Genitalia (Figures 112-114): Strongly pigmented. Combined length phallobase and phal-
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 104-110.—Perilypus viridipennis: 104, tegmen, lateral view; 105, tegmen, ventral view;

106, phallus; 107, male internal reproductive organs; 108, posterior accessory glands. P.

relucens: 109, elytral posthumeral region; 110, habitus. (Scale = 1 mm)
NUMBER 227

41

Y--- -

113

114

FIGURES 111-117.—Perilypus relucens: 111, antenna; 112, tegmen, lateral view; 113, tegmen,

ventral view; 114, phallus. P. levis: 115, tegmen, lateral view; 116, tegmen, ventral view;

117, phallus. (Scale = 1 mm)
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lobasic apodeme 7 times longer than paramere.

Combined length of paramere and phallobase 3.6

times longer than phallobasic apodeme. Paramere:

apex uncinate in ventral view; lateral depression

broad, more basoventral than basolateral; dorsum

feebly convex and not explanate medially; venter

narrowly concave; mesoventral margin straight in

basal two-thirds, slightly arcuate in apical third;

mesodorsal margin sinuous. Sinus: dorsal and ventral sagittate, ventral slightly narrower than dorsal.

Phallus: phallic plate narrowing from middle half;

marginal denticles restricted to apical half of phallic plate (10 specimens examined).

Female Genitalia: As in P. viridipennis except

proctiger broader and posterior margin more arcuate (1 specimen examined).

Internal Reproductive Organs: Female, as in

Figure 55 (1 specimen examined).

VARIATION.—Structural: No conspicuous structural variation was noted, except that the epipleural fold is more collapsed in some specimens

than in others.

Color: Specimens from the type-locality are partially sex dimorphic in elytral and abdominal

coloration. In male specimens, the elytron is entirely blue and may have a purpurescent or viriscent

tinge, and the abdomen is dark brown. In females

the elytron is blue in the basal fourth and rufescent

in the apical three-fourths, and the abdomen is

rufescent. In one female the elytron is entirely

blue and the abdomen predominantly brown.

In one of two other females from Panama (exact

locality not known) the elytron is entirely blue and

has a purpurescent tinge, and the abdomen is entirely brown. In the second specimen the elytron is

entirely rufescent except for a small blue macula on

the humeral angle. The abdomen is entirely rufescent.

NATURAL HISTORY.—In May, in lakeside forest at

Las Lagunas, Panama, I collected one female specimen by beating tree foliage. G. E. Champion collected adults on Volcan de Chiriquf at elevations

from 610 to 1829 meters.

DISTRIBUTION (Figure 358).—The known range

extends from northern Panama to western Colombia.

LOCALITY RECORDS (Figure 358) .—I examined 21 adult

specimens from Central and South America. PANAMA:

Provincia de Chiriquf: Volcan de Chiriquf (BMNH, 4 males

and 3 females, lectotype; GEKI, 5 males and 2 females;

MNHP, 2 males and 1 female); Las Lagunas (GEKI, 1 female). "Panama" (ZMAN, 2 females). COLOMBIA: Departamento de Valle del Cauca (GEKI, 1 male).

REMARKS.—Gorham (1886:336), unaware of this

species' sex linked color polymorphism, identified

the males as Colyphus signaticollis Spinola. He did,

however, note the anatomical similarity between the

two sexes. In discussing the females he states, "Allied to the blue variety [= P. relucens males] of

C. signaticollis figured in the earlier part of this

volume."

7. Perilyptis levis, new species

FIGURES 115-123,358

TYPE-LOCALITY—Santarem, Estado de Brasil.

TYPE-SPECIMENS.—The male holotype and allotype are deposited in CMPP. Five paratypes: GEKI,

2 males and 1 female; CMPP, 1 male and 1 female.

DIAGNOSIS.—Within the viridipennis group specimens of this species are easily recognized by their

boldly serrate antenna (Figure 123).

DESCRIPTION.—Form: As in Figure 120.

Size: Length: Males, average about 6.7 mm, range

6.1-7.1 mm; females, average about 6.8 mm, range

6.5-7.8 mm. Width: males, average about 1.9 mm,

range 1.8-2.2 mm; females, average about 1.9 mm,

range 1.8-2.0 mm. Eight males and 9 females

measured.

Color: Cranium usually flavotestaceous, sometimes dark brown or black; antenna dark brown or

black, rarely flavous apically; pronotum ranging

from predominantly flavotestaceous to predominantly black (see "Variation" and Figures 118-122);

femur flavotestaceous, infuscated apically; tibia and

tarsus dark brown; elytron violaceous, cupreus, or

blue-black, and with or without broad flavotestaceous vitta on epipleural margin, rarely with faint

discal vitta (epipleural-margin vitta extended to

apex or not); abdominal venter dark brown or

black.

Head: As in P. viridipennis except antenna

(Figure 123) broader, more serrate (see "Variation"), more setose, and proportionally longer

(AL/PLS, 1.6); and length/width ratio of each

male antennal article 2.0:1.3:1.4:1.5:1.0:1.0:1.0:0.9:

0.8:0.8:1.3.

Pronotum: As in P. viridipennis except length

equals width (7 specimens measured).
NUMBER 227 43

Elytron: As in P. viridipennis except more slender (EL/EW, average about 4.7, range 4.4-5.2; 17

specimens measured), less convex, and punctation

extended to sutural margin and to elytral apical

third (but see "Variation").

Front Tarsal Claws: As in P. viridipennis.

Male Genitalia (Figures 115-117): Feebly pigmented; tegmen spatulate. Combined length of

phallobase and phallobasic apodeme 4.7 times

longer than paramere. Combined length of paramere and phallobase 2.5 times longer than phallobasic apodeme. Paramere: very broad in lateral

view; dorsum boldly convex, explanate medially;

venter convex broad at base and tapered to apex;

mesoventral margin sinuous; mesodorsal margin

arcuate. Sinus: dorsal digitform but widened between paramere apices; ventral abconic, slightly

shorter than dorsal, and feebly constricted basally.

Phallus: phallic plates without marginal denticles

(8 specimens examined).

Female Genitalia: As in P. viridipennis except

lobes of ventral lamina equal in length (1 specimen examined).

Internal Reproductive Organs: Female, as in

Figure 55 (1 specimen examined).

VARIATION.—Structural: Degree of antennal serration, and apportionment of elytral punctations

vary substantially. Antennal serration varies geographically, which is particularly evident among

male specimens; serration of the clavola is more

prominent in eastern than in western South American specimens.

There is some indication that development and

distribution of elytral punctations is also correlated

with geography; the. punctations are notably reduced or absent at elytral apical third in specimens

from Santarem (Brazil), Colombia, or Venezuela.

Elytral punctation is conspicuously coarser and

more densely distributed among the geographically

disjunct specimens from the Canal Zone and

Amapa, Brazil (the two specimens from Amapa are

particularly coarsely punctated). I assign these

specimens to a deep-punctated morph.

Color: Color polymorphism appears to correlate

geographically and possibly altitudinally. Specimens

from lower elevations (Canal Zone and Santarem,

in Brazil) have a lighter pronotum than those collected from the more mountainous regions of Colombia and Venezuela.

The pronotum is predominantly flavotestaceous

in specimens from Santarem, Brazil, and in females

from Colombia and the Canal Zone (Figure 122).

The pronotum (Figures 118-120) is predominantly

black in specimens from Amapa, Brazil, and in

males from Venezuela and Colombia.

The male from Bogota, Colombia, has a short

and narrow flavotestaceous vitta on the elytral disc

and one on the epipleural margin. In specimens

from Colombia, and in one of three females from

the Canal Zone, the epipleural vitta is broader than

the epipleural fold and extends to the elytral apex;

in other specimens the marginal vitta is as broad or

narrower than the epipleural fold and terminates

before the elytral apex. The antennae are uniformly dark brown or black except in three females

from the Canal Zone, in which they are increasingly

flavotestaceous towards the apex.

NATURAL HISTORY.—From Barro Colorado Island, Canal Zone, T. L. Erwin collected one adult

specimen in June. P. J. Darlington collected two

additional adults from the same locality in May.

Other specimens were collected from Colombia in

September by M. de Ma than, from Brazil between

October and November by Thieme, and from Venezuela in August by J. and B. Bechyn6, and between

September and October by E. A. Klages.

DISTRIBUTION (Figure 358).—This widely distributed species ranges from the Canal Zone to Brazil

and is recorded from highlands of Colombia and

Venezuela.

ETYMOLOGY.—Latin, the adjective lews (smooth).

I refer to the smooth surface at the sutural and

apical region of the elytron.

LOCALITY RECORDS (Figure S58).—I examined 17 adult

specimens from Central and South America. CANAL ZONE:

Barro Colorado Island (GEK.I, 1 female; MCZC, 1 female;

NMNH, 1 female). COLOMBIA: Departamento de Cundi

namarca; Bogota (GEKI, 1 male); Conanche (NMNH, 1

female). Departamento de Boyaca; Muzco (GEKI, 1 male

and 1 female). VENEZUELA: Distrito Bolivar: Suapure,

Rio Cauca (CUNY, 1 male). BRAZIL: Territorio de

Amapa: Sena Lombarda (USMV, 1 female); Rio Calcoene

(GEKI, 1 male). Estado de Para: Santarem (GEKI, 2

males and 1 female; CMPP, 2 males, holotype, and 2 females) .

REMARKS.—The magnitude of infraspecific variation involving elytral punctation is quite exceptional. Perhaps the specimens from the Canal Zone

and Amapa, Brazil, are not conspecific with the

other specimens examined, but the genitalic char-
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURES 118-124.—Perilypus levis: 118-119, 121-122, pronotum; 120, habitus;

123, antenna. P. limbatus: 124, antenna. (Scale = 1 mm)
NUMBER 227 45

acteristics in the male from Amapa are identical

with males whose elytral punctation is typical. I

assume that within the viridipennis group homogeneity of genitalic characteristics indicates conspecificity, as they do in other species groups of the

genus; perhaps the observed differences are early

indications of peripheral isolation.

Although I did not study male specimens from

the Canal Zone, I tentatively consider the three

female specimens from that region as conspecific. I

labeled them "Perilypus levis Ekis?"

The limbatus Group

The following combination of characteristics distinguish members of the limbatus group from other

Perilypus: body form rectangulate; antenna moderately serrate (flagellar articles subfiliform in males);

elytron with convex epipleural fold, deeply impressed and widely distributed punctations, punctate and interpunctate surfaces smooth and shiny,

and with a gradual apical slope.

The composite distribution of the 10 species of

this group extends from Chiapas, Mexico, to San

Martin, Peru; the range north of Panama is attributed only to locality records of P. prolixipenis.

8. Perilypus limbatus (Gorham), new combination

FIGURES 124-132, 136, 137, 362

Colyphus limbatus Gorham, 1878:161 [lectotypc: male, here

designated, deposited in MNHP; type-locality: Caracas,

Venezuela]; 1886:336.

Derestenus limbatus var. latesuturalis Pic, 1941:11 [lectotype:

female, here designated, deposited in MNHP; type-locality:

Bolivia; new synonymy].

Derestenus limbatus var. notaticollis Pic, 1941:11 [lectotype:

female, here designated, deposited in MNHP; type-locality:

Venezuela; new synonymy].

Derestenus limbatus var. suturalis Pic, 1941:11 [lectotype:

male, here designated, deposited in MNHP; type-locality:

Venezuela; new synonymy].

Derestenus columbicus var. reynoldsi Wolcott, 1927b: 106

[holotype not studied; see "Remarks," one paratype in

FMNH; type-locality: El Valle, Venezuela; new synonymy].

Derestenus latefasciatus Pic, 1941:10 [lectotype: female, here

designated, deposited in MNHP; type-locality: Bolivia; new

synonymy].

The names in synonymy represent intrapopulation color variants.

DIAGNOSIS.—Within the limbatus group only

specimens of P. limbatus, P. apocopatus, and P.

decoris, have the anterior margin of antennal article

9 to 11 flavotestaceous. Perilypus limbatus differs

from P. apocopatus by having the femora predominantly flavotestaceous, and from P. decoris by the

narrower epipleural fold and absence of the elytral

discal vitta.

DESCRIPTION.—Form: As in Figure 125.

Size: Length: males, average about 6.9 mm, range

6.3-7.6 mm; females, average about 8.2 mm, range

7.2-9.2 mm. Width: males, average about 1.9 mm,

range 1.8-2.0 mm; females, average about 2.4 mm,

range 2.1-2.8 mm. Ten males and 10 females

measured.

Color: Clypeus, frons (Figure 57), and venter of

cranium flavotestaceous; epicranium and gena shiny

black; antenna predominantly black, anterior margin of articles 9 to 11 flavotestaceous, pronotum

variable (see "Variation"); legs bicolorous; femur

flavotestaceous, with dorsoapical infuscation diminishing from front to hind femur; tibia and tarsus

black to brown; prosternum flavotestaceous, mesoand metasternum black to brown; elytron variable

(see "Variation"); abdomen black.

Head: Interocular depressions shallow; frontal

umbo indistinct; eyes boldly convex (HW/IOW,

average about 2.2, range 2.1-2.4; 20 specimens

measured); antenna (Figure 124) moderately serrate;

AL/PLS, 1.7; length/width ratio of each male antennal article 2.5:1.4-1.7:1.7:2.0:1.5:1.4:1.3:1.1:1.0

1.9.

Pronotum: Only slightly transverse; PL/PW,

average about 0.93, range 0.89-0.96 (20 specimens

measured); subapical depression deeply impressed

and strongly sinuous; pronotal arch impressed with

few shallow punctations; side margin of pronotum

proper moderately arcuate.

Mesoscutellum: Subquadrate.

Elytron: Epipleural margin straight in basal

three-fourths, arcuate in apical fourth; apical slope

gradual; disc impressed with large punctations at

basal two-thirds, punctations are smaller at apical

third; punctate and interpunctate surfaces smooth

and shiny; epipleural fold convex; EL/EW, average

about 5.1 range 4.6-5.7 (20 specimens measured).

Front Tarsal Claws: Male, strongly asymmetrical.

Female, feebly asymmetrical.

Male Genitalia (Figures 128-130): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 4 times longer than paramere.

Combined length of paramere and phallobase 4.8
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 125-130.—Perilypus limbatus: 125, habitus; 126, pronotum; 127, elytra; 128, aedeagus,

ventral view; 129, phallus, ventral view, X 130 (arrow indicates marginal denticles); 130,

phallus, lateral view, X 130 (arrow indicates phallic plicae. (Scale = 1 mm)
NUMBER 227 47

times longer than phallobasic apodeme. Paramere:

Apex obtuse; lateral depression present; dorsum

convex, explanate medially; venter concave; mesoventral margin straight in basal half, faintly sinuous

in distal half; mesodorsal margin sinuous. Sinus:

dorsal lanceolate, constricted subapically; ventral

obconic. Phallus: apex papilliform; phallic plates

contiguous dorsally; marginal denticles stout and

extended beyond limit of phallic plicae (20 specimens examined).

Female Genitalia (Figures 131, 132): Dorsal

lamina bilobed; ventral lamina trilobed, lateral

lobes shorter than medial lobe; proctiger reduced;

proctigeral bacculi not fused; ventral bacculus

acuminate at posterior fourth; coxital plates prominent; coxital stylus bulbous distally (2 specimens

examined).

Internal Reproductive Organs: Male (Figures

136, 137), anterior accessory gland uniramous,

tightly coiled; posterior accessory gland biramous,

outer branch about one-fourth longer than inner

branch; testis composed of 12 follicles. Female, as

in Figure 55; ovary composed of 12 ovarioles (5

males and 3 females examined).

VARIATION.—Structural: The degree of concavity

of the elytral epipleural margin is notably variable.

Females from El Valle, Venezuela, have the epipleural margin conspicuously more concave than

do females from other localities; El Valle females

are also more robust in body form.

Color: The available specimens, although generally homogeneous in external structure, are

chromatically polymorphic. Pronotal and elytral

pigmentation segregate specimens into four color

phena: the bimaculate phenon (with reference to

pronotal lateroapical maculae), the bivittate

phenon (with reference to bivittate elytron), the

punctate phenon (with reference to pronotal and

apical macula), and the fasciate phenon (with reference to fasciate elytron). Both sexes are represented by the first two phena, the last two only by

females.

To the bimaculate phenon I assign individuals

that have a flavotestaceous lateroapical macula on

each anterior angle of the pronotum (as in Figure

157) and whose elytron is vittate at the epipleural

margin. In two female specimens of this phenon

the pronotal disk is flavotestaceous. Specimens of

the bivittate phenon have a bimaculate pronotum,

and each elytron (Figure 127) has a yellow vitta on

the epipleural margin and one on the sutural

margin. In some specimens of the bimaculate and

bivittate phena the pronotum has a fuscous testaceous macula anteriad to the fovea. In one specimen of the bivittate phenon, the marginal vitta

extends only to the elytral basal half. In specimens

of the punctate phenon, the pronotum is fulvous to

fulvescent and has a black midapical macula and

black collar (Figure 125); the pronotal macula

varies in width. Members of the fasciate phenon

have a punctate pronotum and a broad flavous

discal fascia on the midelytron (Figure 125). The

entire spectrum of color polymorphism described

above is present in one population sample from

Rancho Grande, Venezuela.

Seven specimens from El Lim6n, Venezuela, are

more melanistic than the other specimens examined. They exhibit the following characteristics:

posterior region of clypeus black, femur black

distally, and elytral vitta absent or indistinct.

NATURAL HISTORY.—In May, in the vicinity of

the Rancho Grande Biological Station in Aragua,

Venezuela, I collected 53 adults, 2 larvae, and 1

pupa. The macrohabitat of the site can be described

briefly as a predominantly deciduous, subtropical

montane forest. The topographical location of this

Andean region, generally considered a cloud forest,

affords some protection from the nubilous conditions generated by the Caribbean Ocean. Consequently the collection site (at about 1100 meters)

is considerably drier than adjacent mountainous

areas, particularly those at higher elevations. I did

not find P. limbatus specimens at 1200 meters

where clouds maintained moisture level at constant saturation.

Woody lianas and low canopy verdure are two

of the site's most characteristic types of vegetation.

Adults and one larva were collected by beating

vegetation assemblages rich in lianas of about 7-13

cm in diameter. I found a second larva and one

pupa after examining numerous sections of dead

lianas that housed the immatures in a shallow

cavity beneath thin bark. On the same kind of

liana I observed a female oviposit into a crevice in

which I subsequently found one egg. Although I

failed to rear the aforementioned immatures to

imago stage I am confident that they are members

of P. limbatus. The pupa (Figures 380, 381) albeit

not mature, clearly exhibits typical characteristics

of adult Perilypus. The color markings of the
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pupal abdomen are identical to that found in the

larvae; and I do not know of any other species of

Perilypus collected from Rancho Grande vicinity.

At Rancho Grande I also collected adults of this

species by black light; however, the species apparently is only mildly phototropic since only three

specimens were collected.

The altitudinal range of P. limbatus, as indicated by label data, is from 900 to 1500 meters.

DISTRIBUTION (Figure 362).—I studied specimens

from northern Venezuela and Bolivia. The seemingly disjunct distribution is probably a consequence of coincidental collecting. The affinity of

this species for subtropical montane forests suggests that they occur along the Atlantic slopes of

the Andes, which would make a transamazonian distribution unlikely.

LOCALITY RECORDS (Figure 362).—I examined 122 adults,

2 larvae, and 1 pupa from South America. VENEZUELA:

Distrito Federal: Caracas (ZMAN, 1 male and 1 female;

FMNH, 1 male and 1 female; GEKI, 2 males; MNHP, 1

male, lectotype; NMNH, 1 male); Quebrada Avila (CBAZ,

1 male); Rio Gurimare (GEKI, 1 female); El Lim6n, Cordillera Del Litoral (CBAZ, 2 males and 1 female; GEKI, 2

males and 2 females); El Valle (FMNH, 2 females; GEKI,

1 female). Estado de Carabobo: Cerro de Caffe (GEKI, 1

female); Santa Clara (UCMV, 1 male); Trincheras (GEKI, 1

female, UCMV, 1 female). Estado de Aragua: Maracay,

Rancho Grande (AMNH, 2 males and 1 female; ZMAN, 3

males; BMNH, 3 males and 1 female; CBAZ, 3 males and

1 female; FMNH, 4 males; GEKI, 18 males, 19 females, 1

larva and 1 pupa; CMPP, 2 males; UCMV, 2 males and 2

females; MNHP, 2 males and 1 female; MCZC, 2 males and

1 female; MZSP, 2 males; NMNH, 13 males, 4 females and

1 larva). "Venezuela" (FMNH, 1 female, MNHP, 5 females) . BOLIVIA: Departamento de Cochabamba: Cochabamba (GEKI, 1 female). "Bolivia" (GEKI, 1 male; MNHP,

3 males and 1 female).

REMARKS.—The new synonymies are based on

comparisons of the appropriate type-specimens

and on available biological data; I did not study

the type of Wolcott's variety. The color variants,

herein assigned to phena, were given varietal or

species status by previous authors. The varietal

names of Pic (1941), latesuturalis, notaticollis,

suturalis, and that of Wolcott (1927b), reynoldsi,

correspond to the bimaculate, bivittate, punctate,

and fasciate phenon, respectively. Further, Derestenus latefasciatus Pic, corresponds to my fasciate

phenon and to Wolcott's reynoldsi variety. I did not

study the holotype specimen of D. columbicus var.

reynoldsi Wolcott although a para type and

Wolcott's description leaves little doubt that the

synonymy is correct; the holotype of this variety

is assumed lost (Henry H. Dybas, of the Field

Museum of Natural History, pers. comm.).

9. Perilypus columbicus (Spinola), new combination

FIGURES 133-135, 138, 139, 362

Thanasimus columbicus Spinola, 1844a: 195 [lectotype: female, and one paralectotype, male, subsequent designation by Elds, 1975:27, lectotype deposited in MNHP, paralectotype in MSPI; type-locality: Bogota, Colombia].—

Chevrolat, 1876:46,—Schenkling, 1906:263,—Ekis, 1975:27.

Clerus colotnbiae Spinola, 1844b:135 [lectotype: female, here

designated, deposited in MNHP; type-locality: Colombia;

new synonymy].—Gemminger and Harold, 1869:1735.—

Chevrolat, 1876:46.

Spinola's (1844b: 135) lectotype represents a

color phenon of P. columbicus.

DIAGNOSIS.—In addition to the characteristics

given in the key, specimens of this species are distinguished from other members of the limbatus

group by the proportionally shorter antenna

(AL/PLS, 1.5) and by the length/width ratio of

pronotum (0.99). Also, the lateral explanation of

the phallobase and the basal region of the lateral

lobes (Figure 134) is a modification of the male

genitalia unique in Perilypus.

DESCRIPTION.—Form: As in P. limbatus (Figure

125).

Size: Length: male, average about 7.6 mm, range

7.4-7.9 mm; female, average about 7.7 mm, range

7.1-8.3 mm. Width: male, average about 2.2 mm,

range 2.1-2.4 mm; female, average about 2.4 mm,

range 2.0-2.6 mm. Seven specimens measured.

Color: Clypeus, gula, prosternum, and basal half

of femur flavotestaceous; labrum, frons, cranium,

distal half of femur, tibia, tarsus, meso- and metasternum, and abdomen black; pronotum and elytron variable (see "Variation").

Head: As in P. limbatus.

Pronotum: As in P. limbatus except feebly transverse (PL/PW, average about 0.99, range 0.95-1.0

(7 specimens measured).

Elytron: As in P. limbatus except EL/EW, average about 4.5, range 4.3-4.9 (7 specimens measured).

Front Tarsal Claws: As in P. limbatus.

Male Genitalia (Figures 133-135): Strongly pigmented. Combined length of phallobase and phallo-
NUMBER 227 49

FIGURES 131-141.—Perilypus limbatus: 131, ovipositor, ventral view; 132, ovipositor, dorsal

view. 136, male internal reproductive organs; 137, posterior accessory glands. P. columbicus:

133, tegraen, lateral view; 134, tegmen, ventral view; 135, phallus; 138, ovipositor, ventral

view; 139, ovipositor, dorsal view. P. iris: 140, ovipositor, ventral view; 141, ovipositor, dorsal

view. (Scale = 1 mm)
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

basic apodeme 4 times longer than paramere.

Combined length of paramere and phallobase 3.1

times longer than phallobasic apodeme. Paramere:

explanate basally; apical region feebly arcuate in

lateral view; lateral depression small but well impressed; dorsum convex, explanate medially; venter

concave; mesoventral margin concave; mesodorsal

margin sinuous. Sinus: dorsal narrow and almost

twice as long as ventral, ventral elliptical. Phallobase: explanate in apical half. Phallus: acuminate;

marginal denticles as in P. limbatus (3 specimens

examined).

Female Genitalia (Figures 138, 139): As in P.

limbatus except lateral lobes of ventral lamina as

long as medial lobe and proctiger more prominent.

VARIATION.—Structural: The beetles examined

are structurally homogeneous externally.

Color: The male pronotum and elytron are

black, except the former is fuscous lateroapically,

the latter feebly vittate on the epipleural margin.

Two color patterns of the female elytron distinguish a vittate phenon and a fasciate phenon.

Vittate females have the elytron black; a faint

vitta on the epipleural margin may be present. In

fasciate females, the elytron has a broad rufescent

fascia at the midelytron which extends to the

sutural margin, and a broad rufescent macula is

present on the elytral apex.

NATURAL HISTORY.—Unknown, except that two

specimens were collected from Bogota, a locality

at about 4000 meters on the Cordillera Oriental

of the Colombian Andes.

DISTRIBUTION (Figure 362).—Known only from

Colombia.

LOCALITY RECORDS (Figure 362).—I examined 6 adult specimens from South America. COLOMBIA: Departamento de

Cundinamarka: Bogota (MNHP, 2 females, lectotype; MSPI,

1 male). "Colombia" (GEK.I, 2 males and 1 female).

REMARKS.—I (Ekis, 1975:25) erroneously concluded that the specimen in MSPI was the holotype.

With regard to Clerus colombiae Spinola, Chevrolat (1876:46) poses the question "C'est un

Colyphus?" He was quite correct in doubting the

placement of this species under Clerus (now Enoclerus). The width of the distal articles of colombiae antennae is exaggerated in the habitus illustration published by Spinola (1844b, pi. 6; fig. 6).

10. Perilypus apocopatus, new species

Figures 143-148, 362

TYPE-LOCALITY.—Ocana, Departamento de None

de Santander, Colombia.

TYPE-SPECIMENS.—The holotype male is deposited in MNHP, the allotype in GEKI.

DIAGNOSIS.—Distal half of hind femur predominantly black; inner margins and venter of antennal

articles 9 to 11 testaceous. Elytral coloration of the

P. apocopatus female specimen is similar to that of

fasciate females of P. limbatus and P. columbicus,

except that the elytral fascia of P. apocopatus does

not reach the sutural margin.

DESCRIPTION.—Form: As in P. limbatus (Figure

125).

Size: Length: male, 7.4 mm; female, 7.0 mm.

Width: male, 2.2 mm; female, 2.2 mm. One male

and one female measured.

Color: As in P. limbatus except posterior half of

frons black; antennal articles 9 to 11 predominantly

flavotestaceous; and hind femur black in distal

half. Pronotal and elytral coloration varies with

sex (see "Variation").

Head: As in P. limbatus.

Pronotum: As in P. limbatus.

Elytron: As in P. limbatus except EL/EW, average about 4.5, range 4.4-4.6 (2 specimens

measured).

Front Tarsal Claws: As in P. limbatus.

Male Genitalia (Figures 146-148): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 3 times longer than paramere.

Combined length of paramere and phallobase 4.0

times longer than phallobasic apodeme. Paramere:

apical region arcuate and acuminate in lateral

view; lateral depression feebly developed; dorsum

convex, broadly explanate medially; venter concave; mesoventral and mesodorsal margin concave.

Sinus: dorsal narrow, digitiform; ventral exceptionally long and elliptical. Phallus: apical region

acutely narrowed; phallic plates narrowed in

apical half; marginal denticles as in P. limbatus.

Female Genitalia (Figures 143, 144): Dorsal

lamina broadly incised; ventral lamina reduced,

lateral lobes longer than very shallow medial lobe;

oblique bacculus incorporated into large anterior

coxital plate; proctiger short; proctigeral and ventral bacculi as in P. limbatus.
NUMBER 227 51

VARIATION.—Structural: The specimens examined

are structurally homogeneous externally.

Color: The two specimens studied differ in pronotal and elytral coloration. In the male pronotum

is predominantly black and fuscous lateroapically;

the elytron is black, with a purpurescent tinge, and

is broadly flavotestaceous apically. In the female

the pronotum is predominantly flavotestaceous and

has a black midapical macula and black pronotal

collar; the elytron has a broad flavotestaceous fascia

at the middle and an epipleural vitta that expands

apically (Figure 145); and the nonflavotestaceous

portions of the elytron are more purpurescent than

black.

DISTRIBUTION (Figure 362).—Known only from

Ocana, located on the northern latitude of the

Colombian Cordillera Oriental.

ETYMOLOGY.—A Latin derived adjective from

the noun apocope (a cutting off) + the suffix

-atus (cut off). The name refers to the abbreviated

length of the dorsal sinus of the male genitalia.

142

FIGURES 142-145.—Perilypus iris: 142, habitus. P. apocopatus: 143, ovipositor, ventral view;

144, ovipositor, dorsal view; 145, elytra. (Scale = 1 mm)
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

LOCALITY RECORDS (Figure 362).—I examined 2 adult

specimens from South America. COLOMBIA: Departamento

de Norte de Santander: Ocaiia (GEKI, 1 female; MNHP, 1

male, holotype).

11. Perilypus iris, new species

FIGURES 140-142, 362

TYPE-LOCALITY.—Tocota, Departamento de Valle

del Cauca, Colombia.

TYPE-SPECIMENS.—The holotype female is deposited in ZMAN.

DIAGNOSIS.—Within the limbatus group specimens belong to this species if their femora are

entirely black.

DESCRIPTION.—Form: As in Figure 142; forebody

narrow; elytra convex.

Size: Female: length, 8.3 mm; width, 2.5 mm.

Color: Head, meso- and metasternum, legs, and

abdominal venter black; prothorax flavotestaceous,

except pronotal arch with transverse black macula;

elytron with broad sharply outlined fascia at middle, and at apex.

Head: As in P. limbatus.

Pronotum: As in P. limbatus except feebly transverse (PL/PW, 0.99).

Elytron: As in P. limbatus except more convex

and with more depth.

Front Tarsal Claws: Female, feebly asymmetrical.

Female Genitalia (Figures 140, 141): Ventral

lamina with lateral lobes shorter than medial lobe;

oblique bacculus incorporated into small anterior

coxital plate; coxite well sclerotized throughout;

dorsal lamina and proctigeral and ventral bacculi

as in P. limbatus.

ETYMOLOGY.—Latin, the noun iris (rainbow);

with reference to the dorsal coloration of this

beetle.

LOCALITY RECORDS (Figure 362).—I examined 1 adult specimen from South America. COLOMBIA: Departamento de

Valle del Cauca: Tocota (ZMAN, 1 female, holotype).

12. Perilypus acus, new species

FICURES 149-151,154, 155, 362

TYPE-LOCALITY.—Callanga, Provincia de Cusco,

Peru.

TYPE-SPECIMENS.—The holotype male is deposited in MNHB, the allotype in FMNH. Fourteen

paratypes are deposited in repositories as noted

under "Locality Records."

DIAGNOSIS.—Distinguishable from the very similar specimens of P. limbatus by the following combination of characteristics: front tarsal claws symmetrical, antennal articles 9 to 11 not testaceous,

lateroapical maculae of pronotum (when present)

extended posteriorly beyond subapical depression.

DESCRIPTION.—Form: As in P. limbatus (Figure

125).

Size: Length: males, average about 7.2 mm, range

6.1-7.9 mm; females, average about 7.5 mm, range

6.4-8.4 mm. Width: males, average about 2.0 mm,

range 1.8-2.1 mm; females, average about 2.2 mm,

range 2.0-2.3 mm. Twelve males and 5 females

measured.

Color: Head, thoracic sterna, legs, and abdomen

as in P. limbatus, or postocular region of cranium

flavotestaceous; antennal articles 9 to 11 entirely

black; pronotum predominantly black, lateroapical

flavotestaceous maculae (when present) extended

beyond subapical depression, and discal vitta percurrent or not (Figure 149); and vitta on

epipleural margin broader.

Head: As in P. limbatus.

Pronotum: As in P. limbatus.

Elytron: As in P. limbatus.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 154, 155): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.5 times longer than paramere.

Combined length of paramere and phallobase 4.5

times longer than phallobasic apodeme. Paramere:

acuminate; dorsum plane, explanate medially;

venter concave; mesoventral margin arcuate; mesodorsal margin feebly convex and denticulate. Sinus:

dorsal narrowly obconic; ventral broadly elliptical.

Phallus: apex lobifonn; phallic plates widely

separated dorsally with marginal scales near apex

(note arrow in Figures 154, 155) (12 specimens

examined).

Female Genitalia (Figures 150, 151): As in P.

limbatus; except coxites more robust; dorsal lamina

deltoid, with short broad incision; lateral lobes of

ventral lamina very short; and proctiger more

prominent (2 specimens examined).

VARIATION.—Structural: The specimens examined

are structurally homogeneous externally.

Color: Pronotal coloration is sex dimorphic. The
NUMBER 227 53

FIGURES 146-155.—Perilypus apocopatus: 146, tegmen, lateral view; 147, tegmen, ventral view;

148, phallus. P. acus: 149, pronotum; 150, ovipositor, ventral view; 151, ovipositor, dorsal

view; 154, phallus, lateral views, X 130 (arrow indicates phallic scales); 155, phallus, lateral

view, X 1000 (upper arrow indicates phallic papillae; lower arrow indicates phallic scale). P.

buga: 152, ovipositor, ventral view; 153, ovipositor, dorsal view. (Scale = 1 mm)
54

female pronotum is predominantly flavotestaceous

and has a black midapical macula and black

pronotal collar. However, in three female specimens the black basal coloration of the pronotum

extends anteriorly (Figure 149). In males, the

pronotum is always predominantly black and the

vitta on the elytral epipleural margin is narrower

than in female specimens.

NATURAL HISTORY.—Adults were collected in

Peru in May and August.

DISTRIBUTION (Figure 362).—The known distribution extends from Tarapoto to Marcapata, Peru.

ETYMOLOGY.—Latin, the noun acus (husk). I

refer to the scales on the apical region of the

phallic plates.

LOCALITY RECORDS (Figure 362).—I examined 16 adult specimens from South America. PERU: Provincia de Cusco:

Callanga (FMNH, 1 male and 1 female; GEKI, 2 males and

1 female; MNHB, 1 male, holotype); Marcapata (ZMAN, 1

male). Provincia de San Martin: Tarapoto (GEKI, 1 male

and 1 female; MNHP, 2 males and 1 female). "Peru"

(BMNH, 1 male; GEKI, 2 males; MNHP, 1 female; ZMAN,

1 male).

13. Perilypus buga, new species

FICURES 152,153, 156-162, 362

TYPE-LOCALITY.—Buga, Valle del Cauca, Departamento de Valle del Cauca, Colombia.

TYPE-SPECIMENS.—The holotype male and allotype are deposited in NMNH. These and three

paratypes were collected by me in 1973. Three paratypes: GEKI, 3 females.

DIAGNOSIS.—In addition to characteristics given

in the key, specimens of this species are distinguished from specimens of other species in the

limbatus group by the longer antenna (AL/PL, 1.9),

more conspicuous elytral costae, and by the preponderance of pale erect setae on the elytral

surface.

DESCRIPTION.—Form: As in Figure 156.

Size: Length: male, 5.9 mm; female, average

about 7.6 mm, range 7.3-7.8 mm. Width: male

1.5 mm; female, average about 2.2 mm, range 2.0-

2.3 mm. One male and 4 females measured.

Color: As in P. limbatus except venter of antennal articles 9 to 11 black, venter of cranium and

posterior region of frons totally or predominantly

black (Figure 58), and femur black in apical half or

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

more. Color of pronotum and elytron variable (see

"Variation").

Head: As in P. limbatus except frontal umbo

more prominent; antenna proportionally longer

(AL/PL, 1.9); and length/width ratio of each male

antennal article, 2.5:1.1:1.7:1.6:1.7:1.7:1.5:1.6:1.6:

1.3:2.0.

Pronotum: As in P. limbatus except PL/PW,

average about 0.91, range 0.88-0.95 (5 specimens

measured).

Elytron: As in P. limbatus except elytral costae

prominent; vertical setae pale; and EL/EW, average about 5.0, range 4.7 to 5.4 (5 specimens

measured).

Front Tarsal Claws: As in P. limbatus.

Male Genitalia (Figures 158-160): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 6 times longer than paramere.

Combined length of paramere and phallobase 3.4

times longer than phallobasic apodeme. Paramere:

apex uncinate; lateral depression broad, conspicuously long and most prominent near parameral

base; dorsum plane, only slightly explanate

medially; venter concave; mesoventral margin concave; mesodorsal margin straight. Sinus: dorsal

elliptical; ventral abovate; both sinuses equal in

length. Phallus: marginal denticles small, barely

extended to phallic plicae (1 specimen examined).

Female Genitalia (Figures 152, 153): As in P.

limbatus except coxites shorter, coxital stylus not

bulbous distally; proctiger more prominent; and

dorsal lamina with very short medial lobe (2 specimens examined).

Internal Reproductive Organs: Male (Figures

161, 162); anterior accessory gland uniramous,

tightly coiled; posterior accessory gland biramous,

outer branch only slightly longer than inner

branch; testis composed of 12 follicles; ejaculatory

duct conspicuously bulbous in proximal half. Female as in Figure 55; ovary composed of 12 ovarioles (1 male and 4 females examined).

VARIATION.—Structural: Conspicuous structural

variation appears to be restricted to convexity of

the epipleural fold, which is most developed in the

vittate female.

Color: Sexual dimorphism with respect to pronotal and elytral color is present. Color variation

groups females into a punctate or bivittate phenon.

These phena are generally similar to their P. limbatus counterparts, but in this species the pronotum
NUMBER 227

55

156

FIGURES 156-163.—Perilypus buga: 156, habitus; 157, pronotum; 158, tegmen, lateral view;

159, tegmen, ventral view; 160, phallus; 161, male internal reproductive organs; 162, posterior accessory gland. P. latilira: 163, elytron (arrow indicates epipleural fold). (Scale = 1 mm)
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of punctate specimens (Figure 156) is black along

the lower sides and at the base anteriad to the

pronotal collar. The posthumeral elytral vitta is

faintly visible or absent in punctate females. The

elytron of the bivittate phenon (Figure 156) has a

broad discal vitta that does not join the marginal

vitta at the elytral apex. In the male, color of the

pronotum is similar to Figure 157; the elytron is

black and avittate.

NATURAL HISTORY.—In May, in Buga, Colombia,

at an altitude of 1700 meters, 1 collected 5 specimens of this species by beating tree branches intertwined with small woody lianas.

DISTRIBUTION (Figure 362).—Specimens were collected from Buga, a location between Cordillera

Occidental and Cordillera Central of the Colombian Andes.

ETYMOLOGY.—"Buga" is the name of the locality

from which these beetles were collected. The

epithet is a noun in apposition.

LOCALITY RECORDS (Figure 362).—I examined 5 adult

specimens from South America. COLOMBIA: Departamento de Valle del Cauca: Valle del Cauca, Buga (GEKI, 3

females; NMNH, 1 male, holotype, and 1 female).

REMARKS.—The avittate females are superficially

very similar to the darker P. limbatus specimens

from El Limon, Venezuela. However, in P. buga

specimens the elytral vertical setae are pale, not

black as in El Limon specimens. The black coloration on the pronotal lower sides and at the base

beyond the pronotal collar (Figure 156) further

distinguish P. buga specimens from the more melanistic specimens of P. limbatus.

14. Perilypus latilira, new species

FIGURES 164-171, 361

TYPE-LOCALITY.—Bugaba, Provincia de Chiriqui,

Panama.

TYPE-SPECIMENS.—The holotype male and the

allotype are deposited in BMNH. These and 14

paratypes were collected from the type-locality by

G. C. Champion. The 14 paratypes are deposited

in repositories as noted under "Locality Records."

DIAGNOSIS.—The combination of robust body,

epipleural fold wider than pedicel, and epicranium

with black subquadrate macula separate specimens

of this species from those of other species in the

limbatus group except P. prolixipenis. Specimens of

P. latilira are distinguished from specimens of

P. prolixipenis by male genitalic characteristics

(compare Figures 168, 173) and probably by geographic distribution.

DESCRIPTION.—Form (Figure 165): A broad and

deep-bodied species.

Size: Length: males, average about 7.6 mm, range

6.4-8.9 mm; females, average about 8.6 mm, range

6.5-9.8 mm. Width: males, average about 2.0 mm;

range 1.9-2.5 mm; females, average about 2.6 mm,

range 1.9-3.0 mm. Ten males and 3 females

measured.

Color: Frons and venter of cranium flavotestaceous; gena black; epicranium with black subquadrate macula; antenna black; pronotal disc usually

with black vitta that extends to hind angles of

pronotum proper; lower sides of pronotum black;

femur flavotestaceous but black or infuscated dorsoapically; tibia and tarsus dark brown; prosternum

flavotestaceous; meso- and metasternum black;

elytron piceous, with purpurescent tinge and with

broad flavotestaceous marginal vitta that expands

apically; elytron in male with or without short

and narrow flavotestaceous discal vitta, in female

vitta (Figure 166) is broad and long and not confluent with vitta on epipleural margin; abdominal

venter in male black, in female flavotestaceous except visible sterna 5 and 6 black.

Head: As in P. limbatus except eyes less convex

(HW/IOW, average about 2.2, range 1.8-2.4, 13

specimens measured) and antenna (Figure 164)

proportionally shorter (AL/PL, 1.4).

Pronotum: As in P. limbatus except more convex,

subapical depression more sinuous, and pronotal

arch shallower and more prolonged medially.

Elytron: As in P. limbatus, except broader and

deeper (EL/EW, average about 4.9 range 4.5-5.4;

13 specimens measured) and epipleural fold (Figure 163) wider.

Front Tarsal Claws: As in P. limbatus.

Male Genitalia (Figures 167-169): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 6 times longer than paramere.

Combined length of paramere and phallobase 3.4

times longer than phallobasic apodeme. Paramere;

feebly arcuate in lateral view; dorsum narrow and

feebly convex; venter conspicously broader than

dorsum, feebly concave; mesoventral and mesodorsal margin concave. Sinus: dorsal and ventral

elliptical, former broader. Phallus: acuminate;
NUMBER 227 57

FIGURES 164-171.—Perilypus latilira: 164, antenna; 165, habitus; 166, elytra; 167, tegmen,

lateral view; 168, tegmen, ventral view; 169, phallus; 170, ovipositor, ventral view; 171, ovipositor, dorsal view. (Scales = 1 mm)
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

marginal denticles broad, shallow, and restricted

in middle region of phallic plate (13 specimens

examined).

Female Genitalia (Figures 170, 171): Dorsal

lamina deeply incised and acuminate; lateral lobes

of ventral lamina only slightly shorter than medial

lobe; proctigeral bacculi fused as in Figure 171;

inner margin of coxite well sclerotized; coxital

plates not visible (1 specimen examined).

VARIATION.—Structural: The epipleural fold is

conspicuously broader and more convex in females

(Figure 163) than in males. The terminal article of

the male antenna varies in length and in extent of

acumination.

Color: Color variation is evident in both males

and females. One male specimen from Bugaba,

Panama, is particularly atypical; its cranium, antenna, prothorax, and mesosternum, and legs are

entirely flavotestaceous. Definition of the elytral

discal vitta is correlated with sex. Two of three

females examined have broad, clearly defined vittae

on the elytral disc (Figure 166). The elytral discs

of the other female and some males show a faint

abbreviated streak (Figure 165).

DISTRIBUTION (Figure 361).—Known only from

northern Panama.

ETYMOLOGY.—The trivial name is a Latin compound noun formed from the adjective latus (wide)

+ the noun lira (slide). I refer to the width of the

epipleural fold.

LOCALITY RECORDS (Figure 361).—I examined 16 specimens

from Central America. PANAMA: Provincia de Chiriqui:

Bugaba (BMNH, 3 males, holotype, and 1 female; GEKI,

5 males and 1 female; MNHP, 4 males and 1 female). Corrigimiento de Caldera (GEKI, 1 male).

REMARKS.—Specimens of this species were mistaken for the superficially similar P. distinctus

(Figure 239), and P. nigriventris. Elytral color

characteristics of female P. latilira accentuate the

similarity between that species and the aforementioned congeners, but the broad and convex

epipleural fold easily segregates P. latilira females

from females outside the limbatus group.

15. Perilypus prolixipenis, new species

FIGURES 172-174, 360, 361

TYPE-LOCALITY.—Turrialba, Provincia de Cartago, Costa Rica.

TYPE-SPECIMENS.—The holotype male, deposited

in FMNH, was collected by A. Hayne in 1922. Four

paratypes: deposited in repositories as noted under

"Locality Records."

DIAGNOSIS.—Within the limbatus group specimens of this species are reliably separated from the

externally similar specimens of P. latilira, by male

genital characteristics (compare Figures 168 and

174).

DESCRIPTION.—Form: As in P. latilira (Figure

165).

Size: Length: males, average about 8.0 mm,

range 7.4-8.9 mm; female, 9.8 mm. Width: males,

average about 2.2 mm, range 2.0-2.5 mm; female,

3.0 mm. Four males and 1 female measured.

Color: As in P. latilira.

Head: As in P. latilira.

Pronotum: As in P. latilira.

Elytron: As in P. latilira.

Front Tarsal Claws: As in P. latilira.

Male Genitalia (Figures 172-174): As in P. latilira except combined length of phallobase and

phallobasic apodeme 7 times longer than paramere;

ventral sinus inversely acuminate; and marginal

denticles less conspicuous (4 specimens examined).

Female Genitalia: As in P. latilira.

VARIATION.—There is no conspicuous variation

other than that noted for P. latilira.

DISTRIBUTION (Figures 360, 361).—The known

range extends from Chiapas, Mexico, to Cartago,

Costa Rica.

ETYMOLOGY.—The epithet is a Latin compound

noun formed from the adjective prolixus (long) +

the noun penis (penis). I refer to the length of the

aedeagus.

LOCALITY RECORDS (Figures 360, 361) .—I examined 5 adult

specimens from Central America. MEXICO: Estado de

Chiapas (GEKI, 1 male). COSTA RICA: Provincia de

Cartago: Turrialba (FMNH, 2 males, holotype, and 1 female; GEKI, 1 female; ZMAN, 1 male).

REMARKS.—As mentioned in the diagnosis, only

the differences in male genitalia reliably distinguish

P. prolixipenis from P. latilira. I am confident that

the noted differences reflect reproductive isolation.

If the differences were geographic, one would expect

a less than identical appearance of genital characteristics between the P. prolixipenis specimen from

Mexico and those from Costa Rica.

Three locality labels have the abbreviation
NUMBER 227 59

172

FIGURES 172-180.—Perilypus prolixipennis: 172, tegmen, lateral view; 173, tegmen, ventral

view; 174, phallus. P. testaceicornis: 175, habitus; 176, tegmen, lateral view; 177, tegmen,

ventral view; 178, antenna; 179, ovipositor, ventral view; 180, ovipositor, dorsal view. (Scale =

1 mm)
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

"acq," which, I suspect, refers to the Latin word

acquisitus (to add to, to obtain). I suspect that

Hayne used this abbreviation to identify the collector in the same way that one would use "leg"

(abbreviation for legulns meaning a gatherer, a

collector). One specimen has two different locality

records. One of these indicates that the specimen

originates from "Guatemala, Chiapas," the other

label confines the locality to "Chiapas," which was

once part of Guatemala.

16. Perilypus testaceicornis (Pic),

new combination

FIGURES 175-180, 361

Colyphus testaceicornis Pic, 1941:11 [holotype: female, deposited in MNHP; type-locality: Panama].

DIAGNOSIS.—Specimens of this species are easily

recognized by the following combination of femoral

color; front and middle femora testaceous, hind

femur black in distal half, testaceous in proximal

half.

DESCRIPTION.—Form (Figure 175): Shallow

bodied; elytra feebly concave postmedially.

Size: Length: males, average about 7.5 mm, range

6.6-8.2 mm; females, average about 7.3 mm, range

6.0-8.7 mm. Width: males, average about 2.0 mm,

range 1.8-2.1 mm; females, average about 2.1 mm,

range 1.8-2.5 mm. Three males and 10 females

measured.

Color: Cranium, antenna, and prothorax flavotestaceous, except pronotum sometimes dusky midapically and pronotal collar black at sides; legs

flavotestaceous, except hind femur black in distal

half; meso- and metasternum mostly black, former

fulvous medially; elytron and abdominal sterna

variable (see "Variation").

Head: As in P. limbatus except antenna (Figure

178) proportionately longer (AL/PL, 1.9); antennal

articles 9 and 10 not as wide as long; and length/

width ratio of each male antennal article 2.2:1.4:

1.9:1.9:1.8:1.9:1.7:1.5:1.4:1.3:1.9.

Pronotum: As in P. limbatus except less convex,

subapical depression more sinuous and feebly impressed medially, and side margins of pronotum

proper less arcuate.

Elytron: Posthumeral margin, surface, and

epipleural fold as in P. limbatus; otherwise shallower, feebly concave postmedially, and generally

more slender.

Front Tarsal Claws: Feebly asymmetrical in

both sexes.

Male Genitalia (Figures 176, 177): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 6.5 times longer than paramere.

Combined length of paramere and phallobase 3.4

times longer than phallobasic apodeme. Paramere:

apex slightly declinate; dorsum narrow, feebly

convex; venter feebly concave; mesoventral margin

sinuous; mesodorsal margin concave. Sinus: elliptical dorsal narrower than lanceolate ventral.

Phallus: phallic plates acuminate; marginal denticles shallow, broad, and restricted to middle

region of phallic plates (3 specimens examined).

Female Genitalia (Figures 179, 180): Dorsal

lamina convex laterally, incision short; lateral

lobes of ventral lamina shorter than medial lobe;

proctigeral bacculi fused as in Figure 180, coxite

with small coxital plate (1 specimen examined).

VARIATION.—Structural: The specimens examined

do not vary appreciably in external characteristics.

Color: The sutural, apical, and epipleural

regions of the elytron may be dark brown or black.

The paler, nearly fulvous coloration of the elytral

disc suggests a dimly defined vitta that varies in

width. In general, lighter elytra are reddish, darker

ones purpurescent.

NATURAL HISTORY.—In May, near Las Lagunas,

Panama, at an altitude of 1400 meters, I collected

one female specimen by beating tree foliage. G. C.

Champion collected 23 adult specimens on Volcan

de Chiriqui at elevations of from 920 to 1227

meters.

DISTRIBUTION (Figure 361).—To my knowledge

these beetles have only been collected on the

Pacific slope of the Cordillera de Talamanca, in

northern Panama.

LOCALITY RECORDS (Figure 361).—I examined 25 adult

specimens from Central America. PANAMA: Provincia de

Chiriqui: Volcan de Chiriqui (AMNH, 1 female; BMNH, 1

male and 6 females; GEKI, 1 male and 6 females; ISNB, 1

female; MNHP, 3 females; MNHB, 1 female; RNHN, 1

female; NMNH, 1 male and 1 female); Las Lagunas (GEKI,

1 female). "Panama" (MNHP, 1 female, holotype).

17. Perilypus decoris, new species

FIGURES 1, 2, 181-185, 187, 188, 361

TYPE-LOCALITY.—Cerro Campana, Provincia de

Panama, Panama.
NUMBER Zil 61

TYPE-SPECIMENS.—The holotype male, deposited

in NMNH, was collected by T. L. Erwin in 1974.

The allotype is deposited in GEKI. Three paratypes deposited in repositories noted under "Locality Records."

DIAGNOSIS.—The right-angled incurvature of the

elytral discal vitta (Figure 2), near the elytral apex,

easily distinguishes members of this species from

congeners. The general shape of the elytral discal

vittae in P. decoris is roughly similar in specimens

of P. floralis (Figure 342) of the ornaticollis group,

but in P. floralis specimens the vitta is not acutely

incurved and the elytra is more oval than rectangulate.

DESCRIPTION.—Form: As in Figure 2.

Size: Length: males, 7.6; females, average about

8.8, range 8.3-9.3 mm. Width: males, 2.2; females,

average about 2.3, range 2.5-2.8 mm. Two males

and 3 females measured.

Color: Labrum, clypeus, frons, and cranial venter

flavotestaceous, remainder of cranium black; antenna black, except articles 9 to 11 flavotestaceous;

prothorax predominantly flavotestaceous, with midapical, midbasal, and basolateral black maculae;

femur predominantly flavotestaceous, dark brown at

dorsopreapical half; tiba flavotestaceous or dark

brown; meso- and metasternum black; elytron predominantly black, with two white vittae that converge near elytral apex; discal vitta acutely

incurved to sutural margin apically; abdominal

venter black in male, predominantly black in

female (first five visible sterna yellow along posterior margin).

Head: As in P. limbatus except interocular impressions (Figure 1) broader and more deeply

impressed, frontal umbo more prominent, antenna

as in Figure 181.

Pronotum: As in P. limbatus except more transverse (PL/PW, average about 0.86, range 0.88-

0.90; 5 specimens measured).

Elytron: As in P. limbatus except epipleural fold

broader and more convex, and EL/EW 4.6 (5 specimens measured).

Abdomen: Male pygidium as in Figure 185.

Front Tarsal Claws: As in P. limbatus.

Male Genitalia (Figures 181-184): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.1 times longer than paramere.

Combined length of paramere and phallobase 2.3

times longer than phallobasic apodeme. Paramere:

apex uncinate; lateral depression shallow and

elongate; dorsum feebly convex, explanate mesally;

venter narrow and concave; mesoventral margin

feebly arcuate except apically where strongly recurved; mesodorsal margin feebly sinuous and more

sclerotized at base. Sinus: dorsal narrow, slightly

expanded at basal half and longer than lanceolate

ventral sinus. Phallus: phallic plate feebly sinuous;

marginal denticles conspicuous, not extended beyond anterior limit of phallic plicae (1 male examined).

Female Genitalia (Figures 189, 190): Dorsal lamina deeply incised; ventral lamina reduced to one

very short triangular lobe; proctigeral bacculi fused

as in Figure 190; baccular accumination arcuate

and projecting anteriorly; coxites slender; coxital

plates prominent (2 specimens examined).

Internal Reproductive Organs: Male (Figures

187, 188), anterior accessory glands uniramous,

tightly coiled; posterior accessory glands biramous,

outer branch almost twice as long as inner branch;

testis composed of 12 follicles. Female as in Figure

55 (1 male and 2 females examined).

VARIATION.—Structural: The specimens studied

showed no noteworthy structural variation.

Color: The posterior junction between the elytral vittae is less distinct in male specimens.

NATURAL HISTORY.—Specimens were collected

from the type-locality (850 meters) in May, June,

and July. Three adults were collected during night

by beating "slash and burn" vegetation; T. L. Erwin, C. W. O'Brien, and B. Marshall.

DISTRIBUTION.—Known only from the typelocality.

ETYMOLOGY.—Latin, the adjective decoris (ornamented); with regard to the striking coloration of

the elytra.

LOCALITY RECORDS (Figure 361).—I examined 5 adult specimens from Central America. PANAMA: Provincia de Panama: Cerro Campana (GEKI, 1 male and 2 females; NMNH,

1 male, holotype; WBIV, 1 female).

REMARK.—The male paratype is without abdomen.

The reventazon Group

Specimens of the reventazon group are characterized by having the elytral apical slope acute (Figures 29-31) and the curvature of the epipleural
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 181-192.—Perilypus decoris: 181, antenna; 182, tegmen, lateral view; 183, tegmen,

ventral view; 184, phallus; 185, male pygidium; 187, male internal reproductive organs, inner

lateral view; 188, male internal reproductive organs, outer lateral view; 189, ovipositor, ventral

view; 190, ovipositor, dorsal view. P. reventazon: 186, habitus. P. nigriventris: 191, tegmen,

lateral view; 192, tegmen, ventral view. (Scale = 1 mm)
NUMBER 227 63

margin at elytral apical fourth acutely arcuate (except in P. orthopleurid.es where both characteristics

are secondarily lost). Within the group there is

little nongenitalic structural variation, but color

polymorphism is abundant at intraspecific level.

Color differences usually pertain to the elytral surface and are most often correlated with sex.

The composite distribution of the 17 species that

currently comprise this group extends from Mexico

to Peru.

18. Perilypus reventazon, new species

FIGURES 3-26, 28, 29, 32-46, 48-55, 186, 363

TYPE-LOCALITY.—Rio Reventazon, Turrialba,

Estado de Cartago, Costa Rica.

TYPE-SPECIMENS.—The holotype male and the

allotype, deposited in NMNH, were collected by

me in 1973. Sixty-eight paratypes deposited in repositories as noted under "Locality Records."

DIAGNOSIS.—Similar to P. claudus, P. sapientis,

and P. cultratus. Specimens of P. reventazon are

substantially larger than members of P. claudus

and P. sapientis. From P. cultratus, P. reventazon

differs by the less-developed antennal serration,

more convex pronotum, and shallower interocular

depressions.

DESCRIPTION.—Form: As in Figure 186.

Size: Length: males, average about 8.4 mm, range

7.2-10.0 mm; females, average about 9.0 mm, range

8.0-10.5 mm. Width: males, average about 2.3 mm,

range 2.0-2-7 mm; females, average about 2.4 mm,

range 1.6-3.0 mm. Ten males and 10 females measured.

Color: Cranium, pronotum, elytron, and abdomen variable (see "Variation"); antenna black;

meso- and metasternum dark brown; femur predominantly flavotestaceous, infuscated dorsoapically; tibia and tarsus brown.

Head: Interocular depressions deeply impressed,

feebly oblique, and not confluent posteriorly; frontal umbo prominent; eyes boldly convex; HW/

IOW, average about 2.2, range 2.1-2.3 (20 specimens measured); antenna (Figure 8) serrate; AL/

PLS about 1.9; length/width ratio of each male

antennal article 2.2:1.1:1.6:1.8:1.7:1.3:1.5:1.3:1.1:

1.2:1.7.

Pronotum: Moderately transverse (PL/PW, average about 0.88, range 0.83-0.93; 20 specimens measured); subapical and prebasal depressions deeply

impressed, former strongly sinuous; pronotal arch

with few coarse punctations; side margins of pronotum proper boldly convex; fovea small, spheroid,

and deeply impressed; collar prominent.

Mesoscutellum: Subquadrate.

Elytron: Epipleural margin straight in basal

three-fourths, acutely arcuate in apical fourth;

apical slope acute; surface arenose; disc deeply impressed with large punctations; epipleural fold

plane; EL/EW, average about 4.8, range 4.4-5.3 (20

specimens measured).

Front Tarsal Claws: Males, distinctly asymmetrical. Female, feebly asymmetrical.

Male Genitalia (Figures 43-46): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.5 times longer than paramere. Combined length of paramere and phallobase 2.4 times

longer than phallobasic apodeme. Paramere: apex

feebly uncinate; lateral depression feebly impressed;

dorsum plane; venter slightly concave; mesoventral

margin concave; mesodorsal margin sinuous and

conspicuously more sclerotized in basal third than

in apical two-thirds. Sinus: dorsal digitiform; ventral lanceolate. Phallus: marginal denticles extended

from basal half to plicae (30 specimens examined).

Female Genitalia: See page 18.

Internal Reproductive Organs: See page 20.

VARIATION.—Structural: Except for the sex dimorphism present in the front tarsal claws, the

specimens studied are structurally homogeneous

externally.

Color: There is substantial color variation and

except for the elytral and abdominal coloration,

color polymorphism is not sex correlated. The entire spectrum of color variation presented below is

evident in a series of 45 specimens collected from

the type-locality.

The cranium may "be variously bicolored, but

usually is predominantly black. If predominantly

black the cranium is flavotestaceous in the postocular epicranial region. When the flavotestaceous

macula of this region is prolonged anteriorly (along

the interocular margin of the eyes) it marks the

epicranium with a subquadrate black macula; the

latter combination of cranial color is most often

found in males with vittate pronotum. In some

males, the black epicranial macula is reduced to

the extent that the cranium is predominantly flavotestaceous to rufescent.
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The pronotum is either flavotestaceous or refuscent, and vittate or avittate; its lower sides may

be infuscated or entirely black. The pronotal disc

is either marked by a midapical black macula or

by a black vitta.

Elytral coloration distinguishes an avittate phenon, a univittate phenon, and a bivittate phenon.

In an avittate beetle the elytron is entirely black,

except for a small pale macula at the base. This

macula corresponds, in position, to the basal limit

of a fully developed vitta. In beetles of the univittate phenon the elytron has a discal vitta that begins at the base, gradually obliques toward the suture, and incurves sharply at its apical limit; the

incurved portion may or may not reach the sutural

margin. In members of the bivittate phenon the

elytron has two vitta, one on the epipleural margin

and one on the disc. The marginal vitta, which is

narrower than the discal one, usually terminates

at the elytral apical eight. Of 32 male specimens

examined only three have avittate elytra; in one of

these the discal vitta diminishes in the basal half.

Female specimens whose pronotum is vittate have

univittate elytra whereas those that have the pronotum maculated have bivattate elytra; in one female

specimen the elytron is entirely flavotestaceous.

The male abdomen is black. In females, visible

abdominal sterna i to v are either entirely flavotestaceous or predominantly brown (when mostly

brown their middle and posterior margin is pale),

and vi is entirely brown.

NATURAL HISTORY.—I collected 45 specimens of

this species, from May to July, in Turrialba, Costa

Rica. Thirty-eight of these were collected by beating tree branches intertwined with lianas, seven by

sweeping herbaceous hedge growth. All specimens

were collected from a forested ravine adjacent to

Rio Reventaz6n; the altitude of the site is 610 meters. A. Hayne collected two specimens from Turrialba at 900 meters.

In-captivity feeding behavior of adult specimens

is similar to that observed in P. limbatus (Gorham).

DISTRIBUTION (Figure 363).—Known only from

the type-locality.

ETYMOLOGY.—The trivial name, reventazon, constitutes a noun in apposition and refers to the

type-locality.

LOCALITY RECORDS (Figure 363).—I examined 71 adult

specimens from Central America. COSTA RICA: Estado

de Cartago: Turrialba, at Rio Reventazon (AMNH, 1 male

and 1 female; BMNH, 1 male and 1 female; CASC, 1 male

and 1 female; CISC, 1 male; CNCC, 1 male; FMNH, 1 male;

GEKI, 18 males and 14 females; CMPP, 1 male and 1 female; UCMV, 1 male; MCZC, 1 male and 1 female; MNHB,

1 male; MNHP, 1 male and 1 female; MZSP, 1 male and 1

female; NMNH, 2 males, holotype, and 1 female; ZMAN, 1

male); Turrialba (CNCC, 1 female; FMNH, 8 females;

MNHB, 1 female; NMNH, 1 male and 1 female; ZMAN,

1 female. Locality not known: FMNH, 1 male and 1 female.

19. Perilypus nigriventris (Gorham),

new combination

FIGURES 191, 192, 198, 199, 364

Colyphus nigriventris Gorham, 1886:336 [lectotype: male, deposited in BMNH, here designated; type-locality: El Zapote,

Departamento de Escuintla, Guatemala].—Wolcott, 1927a:

S3.

DIAGNOSIS.—Within the reventazon group male

specimens of P. nigriventris are superficially very

similar to those of P. latilobus, P. calcaris, and P.

exilis. Among the four species, however, only P.

nigriventris has symmetrical male front tarsal

claws. Females of these species are more difficult to

separate.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except more robust.

Size: Length: males, average about 7.5 mm, range

6.6-8.5 mm; females, average about 8.3 mm, range

7.6-9.4 mm. Width: males, average about 2.0 mm,

range 1.8-2.2 mm; females, average about 2.3 mm,

range 2.2-2.6 mm. Four males and five females

measured.

Color: Clypeus, frons, cranium venter, and postocular region of epicranium flavotestaceous; remainder of epicranium and gena black; antenna

black; pronotum broadly flavotestaceous dorsolaterally, piceous ventrolaterally and discally; pronotal collar piceous; prosternum flavotestaceous, mesoand metasternum piceous; femora flavotestaceous,

except distal region infuscated; tibia and tarsus

piceous; elytron piceous, with or without testaceous

discal vitta; abdomen entirely black, or partially

dark brown and partially flavotestaceous.

Head: As in P. reventazon except antenna sex

dimorphic in length (AL/PLS, male 2.1; female

1.8) and in proportion of length/width ratio of

each antennal article, male 2.0:1.0; 1.5:1.7:1.6:

1.7:1.7:1.7:1.6:1.3:1.9; female 2.0:1.0:1.0:1.0:1.0:1.0:

1.0:1.0:1.2:1.0:1.7.
NUMBER 227 65

Pronotum: As in P. reventazon except subapical

depression shallower, pronotal arch less convex, and

pronotum proper side margins less arcuate.

Elytron: As in P. reventazon.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 191, 192): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 6 times longer than paramere. Combined length of paramere and phallobase 2.8 times

longer than phallobasic apodeme. Paramere: arcuate and acuminate; lateral depression shallow

and elongate; dorsum plane; venter feebly concave; mesoventral and mesodorsal margin concave.

Sinus: dorsal and ventral equal in length and

lanceolate. Phallus: feebly sinuous; marginal denticles distributed from basal fourth to apical fourth

of phallic plate (18 specimens examined).

Female Genitalia (Figures 198, 199): Dorsal lamina bilobed, outer margins arcuate, ventral lamina trilobed, lateral lobes about equal to median

lobe; dorsal bacculus not fused; ventral bacculus

acuminate at about posterior third; coxite robust;

coxital plates absent (1 specimen examined).

VARIATION.—Structural: The male antenna is

more slender and proportionally shorter than the

female antenna (see "Head"). Also, in one female

the side margin of the pronotum proper is more

conspicuously convex and the elytral apical curvature is less arcuate than in other specimens.

Color: The female elytron has a testaceous discal

vitta that is closer to the posthumeral margin than

to the sutural margin. The discal vitta begins posteriad to the humeral margin, ends anteriad to the

apex, and varies in width. The male elytron is

avittate. Also in males, the flavotestaceous postocular macula of the epicranium extends anteriorly,

marking the epicranium with a black subquadrate

macula; in females, the epicranial postocular macula is reduced or absent. Color of abdominal venter is also correlated with sex, being entirely black

in males and predominantly flavotestaceous in females.

DISTRIBUTION (Figure 364).—The single locality

record from Costa Rica extends the known range

of this species from southwestern Guatemala to

central Costa Rica.

LOCALITY RECORDS (Figure 364).—I examined 9 adult specimens from Central America. GUATEMALA: Departamento

de Escuintla: El Zapote (BMNH, 1 male, lectotype, and 2

females; GEKI, 2 males and 2 females; MNHP, 1 male and

1 female). COSTA RICA: Provincia de Cartago: Turrialba

(GEKI, 1 male).

REMARKS.—Two P. nigriventris females formed

part of the syntype series of P. bilineatus, a third

was identified as P. distinctus. Wolcott's (1927a:33)

treatise of this species refers to specimens of P.

reventazon and P. acus.

20. Perilypus sapientis, new species

FICURES 195-196, 363

TYPE-LOCALITY.—Montes de Oca, San Pedro,

Provincia de San Jos£, Costa Rica.

TYPE-SPECIMENS.—The holotype male and the

allotype are deposited in NMNH. These and 11

paratypes were collected by C. H. Ballou, 10 in

1936 and one in 1937. Twenty-six paratypes deposited in repositories as noted under "Locality

Records."

DIAGNOSIS.—Within the reventazon group specimens of this species are superficially most similar

to specimens of P. claudus Wolcott. No one external characteristic will easily separate all members

of these two species, but males are separable by

genital differences (compare Figures 195, 201), females by elytral form; female specimens of P.

claudus have shorter and more quadrate elytra.

DESCRIPTION.—Form: As in Figure 193.

Size: Length: males, average about 6.8 mm, range

6.0-7.8 mm; females, average about 7.2 mm, range

6.7-7.9 mm. Width: males, average about 2.0 mm,

range 1.8-2.3 mm; females, average about 2.2 mm,

range 2.0-2.4 mm. Ten males and 10 females measured.

Color: Cranium entirely testaceous, or clypeus

frons and cranium venter testaceous and epicranium

and gena black; antenna black; venter and sides of

pronotum testaceous; pronotal disc testaceous, with

or without midapical infuscation or with broad

piceous vitta that narrows abruptly at subapical depression (Figure 193); pronotal collar black; prosternum testaceous; meso- and metasternum black

or piceous; femur flavotestaceous in basal half,

piceous in distal half; tibia piceous, distal region

pale or not; tarsus piceous; elytron piceous, with

or without discal vitta; elytral discal vitta begins at

midbase extends posteriorly equally distant from

suture and posthumeral margin and ends at elytral

apical eighth, where feebly explanate (Figure 193)
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

or not; abdominal venter entirely black, or predominantly dark brown having middle region of visible

sternum i to iv flavotestaceous.

Head: As in P. reventazon except antenna proportionally shorter (AL/PLS, 1.7) and length/width

ratio of each antennal article 2.0:1.2:1.3:1.2:1.3:

1.0:1.3:1.3:1.0:1.0:1.8.

Pronotum: As in P. reventazon.

Elytron: As in P. reventazon except proportionally shorter (EL/EW, average about 4.2, range

4.1-5.2; 20 specimens measured) and apical curvature more acute.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 194-196): As in P. nigriventris except dorsal and ventral sinus broader and

paramere more acutely arcuate in apical region (3

specimens examined).

Female Genitalia: As in P. nigriventris.

Internal Reproductive Organs: Male and female

as in P. reventazon (1 male and 1 female examined).

VARIATION.—Structural: A male specimen from

Turrialba, Costa Rica, is more robust and has a

larger more convex pronotal arch; the side margins

of the pronotum proper are broadly arcuate and

the body form is generally more robust. Female

specimens are shorter; their elytral apical curvature is more acute than it is in males. Depth of

the pronotal subapical depression varies independent of sex.

Color: All males examined have the cranium

testaceous. Partial sex-linked color variation is apparent in coloration of the pronotal disc: vittate

in 8 of 11 females studied, avittate in all males examined. Color of elytron and abdominal venter is

invariably sex linked: always vittate discally in females (in one female, from the type-locality, this

vitta is faintly visible), avittate in males. Male abdominal sterna are black; those of females are part

piceous and part testaceous, except for the last

visible sternum, which is totally black.

NATURAL HISTORY.—Adult specimens were collected in Costa Rica from February to July. I collected one female specimen in May by beating tree

foliage in Alajuela. In June, in Montes de Oca,

once a forested environ of San Jos6 and now occupied by the campus of the University of Costa Rica,

C. H. Ballon collected 10 specimens on Hamelia

erecta Jacquin (probably equals H. patens Jacquin),

one specimen on Trichilia havanensis Jacquin, and

one on Mirabilis jalapa Linnaeus. Ballou collected

one other specimen in July on Veronica brachiata

Bentham. C. Fernandez collected one specimen

from San Jose\ in May, between 1000 to 2000 meters.

DISTRIBUTION (Figure 363).—The known range

of this species extends from central Guatemala to

central Costa Rica.

ETYMOLOGY.—The epithet represents a genitive

case of the latin adjective sapiens (wise). I dedicate

the name of this species to my learned friend

George C. Steyskal.

LOCALITY RECORDS (Figure 363).—I examined 28 adult

specimens from Central America. GUATEMALA: Departamento de Guatemala: northwestern outskirt of Guatemala

City (GEKI, 1 male). COSTA RICA: Provincia de Alajuela:

Alajuela (GEKI, 1 female), Provincia de San Jose; San Ignacio

(GEKI, 1 male); Farm La Caja, 8 kilometers, west of San

Jose (ZMAN, 1 male); San Pedro, Montes de Oca (GEKI, 5

males and 2 females; MHNM, 2 males; NMNH, 2 males,

holotype, and 2 females); San Jose (FMNH, 1 male and 1

female; GEKI, 1 male; NMNH, 2 males and 1 female).

Provincia de Cartago: Turrialba (GEKI, 1 male; NMNH, 1

female; ZMAN, 1 male and 2 females). "Cost Rica" (MNHP,

1 female).

REMARKS.—The more robust male specimen from

Turrialba, heretofore identified as Colyphus nigriventris, may not be conspecific; but without additional data I cannot justify its specific distinction

from other specimens of P. sapientes or its conspecificity with P. nigriventris.

21. Perilypus claudus (Wolcott)

FIGURES 197, 200-203, 364

Derestenus claudus Wolcott, 1927a:32 [holotype: male, deposited in FMNH; type-locality: Turrialba, Estado de

Cartago, Costa Rica.].

Cleronomus nigrifrons Chevrolat var. flavicornis Wolcott,

1927a:36 [holotype: female, deposited in FMNH; new

synonymy].

Chevrolat's type-specimen is a color variant of

P. claudus.

DIAGNOSIS.—Males of this species can be reliably

distinguished from males of other species of Perilypus only by genitalic characteristics as described

(see "Male Genitalia") and depicted in Figures

200-202. No one characteristic will diagnose P.

claudus females but the combination of small size,

slender body, and distal confluence of the elytral

discal vittae will generally separate them from females of other Costa Rican congeners.
NUMBER 227 67

The shape and position of the discal vitta in P.

claudus specimens from Volcan Turrialba is very

similar to that in females of P. sapientis. However,

female P. sapientis is more robust and more quadrate in body form than female P. claudus. The

configuration of the discal elytral vitta in P. claudus

females from Monteverde is the same as in females

of P. reventazon (Figure 186), but specimens of the

latter species are considerably larger than those of

the former.

DESCRIPTION.—Form: As in P. sapientis (Figure

193) except more slender.

Size: Length: males, average about 6.2 mm, range

5.3-7.3 mm; females, average about 6.8 mm, range

5.7-8.0 mm. Width: males, average about 1.7 mm,

range 1.4-1.9 mm; females, average about 1.8,

range 1.7-2.3 mm. Ten males and 10 females

measured.

Color: Color characteristics are exceptionally

variable and because of this a composite description of this species' integumental color is not provided. Instead, the general color patterns and the

deviations from them are described under "Variation."

Head: As in P. reventazon except antenna proportionally longer (AL/PLS, 2.0) and length/width

ratio of each male antennal article 2.8:1.4:1.8:1.8:

1.6:1.6:1.4:1.4:1.2:1.0:2.3.

Pronotum: As in P. reventazon.

Elytron: As in P. reventazon.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 200-202): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5 times longer than paramere.

Combined length of paramere and phallobase 3

times longer than phallobasic apodeme. Paramere:

feebly arcuate in lateral view; parameral septum

prominent (note arrow in Figure 201); lateral depression indistinct; dorsum tumid midbasally, feebly

convex medially; venter narrow and feebly convex;

mesoventral margin concave; mesodorsal margin

sinuous. Sinus: dorsal irregular; ventral broadly

elliptical. Phallus: as in Figure 202 (29 specimens

examined).

Female Genitalia: As in P. nigriventris (2 specimens examined).

Internal Reproductive Organs: Male (Figures

197-203), anterior accessory gland uniramous.

tightly coiled; posterior accessory gland biramous,

outer branch one-fourth longer than inner branch;

testis composed of 12 follicles. Female (Figure 55),

ovary composed of 12 ovarioles (3 males and 2

females examined).

VARIATION.—Structural: Elytral punctations are

slightly shallower in specimen from Volcan Irazu

and in those from Quebrada Florida.

Color: On the basis of leg color, specimens from

Volcan Irazu and those from Quebrada Florida are

assigned to a brown-legged morph. These specimens

are considerably homogeneous in color unlike in

other population samples of this species. Color

characteristics of specimens of the dark-legged

phenon are as follows: Head: cranium testaceous,

immaculate; antenna piceous with articles 9 to 11

not testaceous beneath. Thorax: prothorax predominantly testaceous; pronotum with midapical

macula, or with or without discal vitta, latter interrupted at middle or not; pronotal collar piceous;

meso- and metasternum piceous; legs piceous; elytron piceous, with cupreous or purpurescent luster.

Abdomen: sternum piceous in both sexes.

Leg color can also be used to distinguish a yellowlegged morph. In specimens of this morph the head,

antenna, pronotum, mesoscutellum and legs are

entirely flavotestaceous, and the elytra purpurescent

and avittate.

All other specimens examined exhibit the following possible color characteristics: Head: cranium flavotestaceous, with or without midepicranial

macula, or clypeus and frons flavotestaceous, epicranium black (except postocular region flavotestaceous), gena piceous, and venter piceous or

flavotestaceous; antenna flavotestaceous or black,

when piceous articles 9 to 11 testaceous or dark

brown. Thorax: pronotum predominantly black or

predominantly flavotestaceous; when predominantly

black vittate discally and flavotestaceous laterally;

when predominantly flavotestaceous discal vitta

reduced; pronotal discal vitta (when present) may

be entire or interrupted at middle; when pronotum

predominantly flavotestaceous midapical macula

present; pronotal collar piceous; prosternum

flavotestaceous; meso- and metasternum piceous;

profemur flavotestaceous in basal half, piceous in

apical half, meso- and metafemur flavotestaceous in

basal two-thirds and piceous in apical third, or all

femora flavotestaceous in basal half and piceous in

apical half; tibia entirely piceous, or piceous in

basal half and testaceous in apical half (usually only

hind tibia testaceous in apical half); tarsus piceous;
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 193-203.—Perilypus sapientis: 193, habitus; 194, tegmen, lateral view; 195, tegmen,

ventral view; 196, phallus. P. claudus: 197, male internal reproductive organs; 200, tegmen,

lateral view; 201, tegmen, ventral view (arrow indicates parameral septum); 202, phallus; 203,

posterior accessory gland. P. nigriventris: 198, ovipositor, ventral view; 199, ovipositor, dorsal

view. (Scale = 1 mm)
NUMBER 227 69

elytron piceous, with or without purpurescent luster

and vittate or avittate discally. Abdomen: venter

entirely black, or visible sternum i to v flavotestaceous infuscated discally and laterally, sternum vi

piceous.

Specimens from Monte Verde are sex dimorphic

with respect to cranial, pronotal, and possibly

elytral color. Male specimens have the cranium

entirely flavotestaceous, pronotal vitta absent, and

elytron avittate. Female specimens have the cranium

predominantly piceous, the pronotal vitta is always

entire, and of 14 specimens eight have vittate

elytra. The elytral discal vitta, which is nearer to

the posthumeral margin than to the sutural margin,

is incurved apically; the incurvature reaches the

sutural margin.

I studied too few specimens from Volcan Turrialfa to determine relationships between color and

sex. The following color characteristics are present,

however. The legs are entirely flavotestaceous in 5

of 11 males; in the other males the legs are partly

piceous and partly flavotestaceous. Among all specimens studied, 2 males and the only female from this

locality have univittate elytra while in another

male the elytral vitta is reduced to a short basal

streak.

NATURAL HISTORY.—In June, on the northeastern

slope of Volcan Turrialba, Costa Rica, at an altitude of 1450 meters, I collected a series of 12 specimens by beating live branches of Psidium guajava

Linnaeus.

I collected 26 additional specimens, in June, from

Monte Verde, Costa Rica, by beating dense understory composed mostly of live and dead tree

branches and liana vines. The macrohabitat of the

locality may be described as a dry section of montane forest situated at 1600 meters on the Pacific

slope of the Cordillera de Tilaran. Holdridge, et al.

(1971), classified this region as "lower montane rain

forest."

In April and May, F. Nevermann collected four

specimens of the brown-legged phenon from the

western slope of Volcan Irazii (at 1500 meters);

four others were collected during May near Quebrada Florida (at 1400 meters). A single specimen,

with an F. Nevermann label but apparently collected by "Schild" (the writing is not clear), was

collected in October from Turrialba.

DISTRIBUTION (Figure 364).—The single specimen

from Mexico extends the known range of this

species from Mexico to Costa Rica.

LOCALITY RECORDS (Figure 364).—I examined 52 adult

specimens of this species from Central America. MEXICO:

Estado de Morelos: Yautepec (GEKI, 1 male). COSTA RICA:

Provincia de Puntarenas: Monteverde (AMNH, 1 male and 1

female; BMNH, 1 male and 1 female; CASC, 1 male and 1

female; FMNH, 1 male and 1 female; GEKI, S males and 6

females; CMPP, 1 male; MNHB, 1 female; MNHP, 1 male;

NMNH, 2 males and 2 females; ZMAN, 1 male and 1 female).

Provincia de Cartago; Turrialba (FMNH, 2 males, holotype,

and 2 females; GEKI, 1 male); Volcan Turrialba (GEKI, 10

males and 1 female); Volcan Irazii (FMNH, 1 female; GEKI,

1 female; NMNH, 3 females). Provincia de Limdn: Quebrada Florida (GEKI, 2 males; NMNH, 1 male and 1 female).

REMARKS.—The vast color variability present in

the available samples is evident at intra- and interpopulational levels. Diversity of integumental color

is particularly evident in coloration of the elytron.

The elytral discal vitta may be linear or incurved

at the apex. Elytra are avittate in the beetles from

Volcan Irazu and Quebrada Florida.

While intrapopulation color polymorphism is

not extraordinary in Perilypus the extent reported

here is more than usual. Accordingly, one might

suspect that the beetles from Monteverde, Volcan

Turrialba, Volcan Irazu, and Quebrada Florida are

not conspecific, as indeed they may not be. However, for the present I assume them conspecific in

view of their structural similarity including characteristics of the genital organs of both sexes.

Wolcott's (1927a:36) description of Derestenus

nigrifrons Chevrolat var. flavicornis is based on one

female specimen from Turrialba. Integumental

color (Wolcott's criteria for recognizing var. flavicornis) of a male specimen from Monte Verde,

Costa Rica, is identical to that of Wolcott's holotype. There is no noticeable anatomical difference

between these two specimens except for size and

structures related to sex. I believe the specimens

under consideration are conspecific, not only to each

other, but also with the other specimens included

herein. I assign two male specimens and Wolcott's

var. flavicornis to a yellow-legged phenon (see

"Variation)."

22. Perilypus cultratus, new species

FIGURES 204-206, 211, 216, 363

TYPE-LOCALITY.—Rio Reventazdn, Turrialba,
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Estado de Cartago, Costa Rica.

TYPE-SPECIMENS.—The holotype male is deposited in NMNH, the allotype in GEKI; both were

collected by me in 1971. Five paratypes are deposited in the repositories noted under "Locality

Records."

DIAGNOSIS.—Within the reventazon group, specimens from Costa Rica belong to this species if

their pronotal disc is faintly tumid behind the

subapical depression and in front of the prebasal

depression, if the pronotal foveae are deeply impressed and not particularly spheroid (sometimes

they approach a crescentic shape), and if their

antenna is more serrate than in P. reventazon

(Figure 8). Specimens of this species also can be

easily separated from the very similar sympatric

specimens of P. reventazon by differences in the

male pygidium (compare Figures 211, 39) and by

the lineate female elytral vitta.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except pronotum less convex.

Size: Length: males, average about 7.6 mm, range

8.0-6.6 mm; females, average 8.0 mm. Width: males,

average about 2.2 mm; range 2.0-2.6 mm; females:

2.4 mm. Five males and 2 females examined.

Color: Clypeus, frons, postocular region of epicranium, and cranial venter testaceous; gena and

remainder of epicranium black; antenna black; side

margin of pronotum proper flavotestaceous; pronotal disc broadly vittate; pro-, meso-, and metasternum dark brown; femora predominantly

flavotestaceous, infuscated dorsoapically; elytron

piceous, with or without short flavotestaceous discal

vitta; abdomen entirely black or predominantly

flavotestaceous.

Head: As in P. reventazon except: interocular

depressions more deeply impressed, rugose or not;

antenna more serrate; and length/width ratio of

each male antennal article 2.4:1.2:1.4:1.3:1.3:1.3:

1.2:1.0:1.2:1.1:2.2.

Pronotum: As in P. reventazon except foveae

more deeply impressed, more elongate than spheroid; and disc behind subapical depression and

before prebasal depression feebly tumid.

Elytron: As in P. reventazon.

Front Tarsal Claws: Male and female feebly

asymmetrical.

Abdomen: Male pygideal apodeme as long as

pygidial plate (Figure 211).

Male Genitalia (Figures 204-206, 216): Strongly

pigmented. Combined length of phallobase and

phallobasic apodeme 7 times longer than paramere.

Combined length or paramere and phallobase 3

times longer than phallobasic apodeme. Paramere:

acuminate in lateral view; lateral depression not

conspicuous; dorsum feebly convex; venter concave;

mesoventral and mesodorsal margin sinuous, margins confluent in apical half, not in distal half.

Sinus: dorsal narrow, as long as inversely acuminate

ventral. Phallus: attenuate; marginal denticles

prominent (4 specimens examined).

Female Genitalia: As in P. nigriventris.

Internal Reproductive Organs: Male and female

as in P. reventazon (1 male and 2 females examined).

VARIATION.—Structural: The interocular depressions are less depressed and abbreviated in the

female specimen from Hamburg Farm; they are

conspicuously rugose in the male from Tuis and in

one female from Turrialba.

Color: The pronotal vitta is deltoid in the female

specimen from Hamburg Farm, and in one from

Turrialba. Except for the male from Hamburg

Farm, male elytra are avittate; female elytra are

vittate discally.

NATURAL HISTORY.—In June and July, within a

forested ravine adjacent to Rio Reventaz6n, I collected a male and two females by sweeping herbaceous hedge growth.

In June, within the Rio Reventaz6n drainage

system, in Limon, F. Nevermann collected one

female in "troknem holz" (probably translated as

fallen timber, or from a wood pile). He also collected a male in August from Hamburg Farm.

DISTRIBUTION (Figure 363).—The few locality

records suggest that this species is primarily distributed along the Costa Rican Atlantic drainage

system.

ETYMOLOGY.—Latin, the adjective cultratus

(knife-shaped); the trivial name refers to the knifeshaped phallus.

LOCALITY RECORDS (Figure 363).—I examined 7 adult specimens from Central America: COSTA RICA: Provincia de

Cartago: Turrialba at Rio Reventaz6n (GEKI, 2 females;

NMNH, 1 male, holotype); Tuis (GMNH, 1 male). Provinda

de Lim6n: Hamburg Farm at Rio Reventaz6n (GEKI 1 male;

NMNH, 1 female). Locality not known (FMNH, 1 male).

REMARKS.—Females of this species have been

confused with females of P. bilineatus; however,

the conspicuous convexity of the elytron and the
NUMBER 227 71

distinct confluence of the epipleural and discal

vittae at the elytral apex in P. bilineatus should

easily separate specimens of that species from those

of P. cultratus.

23. Perilypus bilineatus (Gorham),

new combination

FIGURES 207-209, 364

Colyphus bilineatus Gorham, 1886:335 [lectotype: female, deposited in BMNH, two paralectotypes, females, one deposited in BMNH and one in ZMAN, here designated;

type-locality: Volcan de Chiriqui, Provincia de Chiriquf,

Panama].—Wolcott, 1927a:37.

DIAGNOSIS.—The slender, shallow body combined

with the following characteristics will distinguish

specimens of this species within the reventazon

group; elytral apex testaceous, elytral disc vittate

or not, pronotum as long as wide, and pronotal

subapical depression shallow and wrinkled transversely midbasally.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except shallower and more slender.

Size: Length: male, average 6.7 mm; females,

average about 7.3 mm, range 6.4-8.5 mm. Width:

males 2.0; females, average about 2.1 mm, range

2.0-2.4 mm. One male and 5 females measured.

Color: Cranium entirely testaceous, or clypeus

frons and postocular region of epicranium testaceous and remainder of cranium black; antenna

black; pronotum predominantly testaceous and

infuscated midapically, or predominantly black and

pronotum proper with side margins and anterior

angles testaceous; prosternum testaceous; meso- and

metasternum black; femur predominantly testaceous, infuscated apically; tibia and tarsus dark

brown; elytron predominantly piceous, epipleuron

and apex faintly testaceous and flavotestaceous discal vitta closer to posthumeral margin than to

sutural margin (as in Figure 264); discal vitta may

or may not be confluent with testaceous epipleural

vitta; abdominal venter black, or visible sterna i to

iv testaceous and infuscated anteriorly, v predominantly testaceous or predominantly dark brown,

and vi black.

Head: As in P. reventazon except cranium sometimes conspicuously macrosculptured; antenna less

serrate; and length/width ratio of each male antennal article 2.5:1.0:1.1:1.4:1.2:1.1:1.1:1.1:1.0:1.0:1.8.

Pronotum: As in P. reventazon except as long as

wide, subapical depression more sinuous, and pronotal arch transversely wrinkled midapically.

Elytron: As in P. reventazon except shallower and

more convex.

Front Tarsal Claws: Male, damaged. Female,

symmetrical.

Male Genitalia (Figures 207-209): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 6 times longer than paramere.

Combined length of paramere and phallobase 2.5

times longer than phallobasic apodeme. Paramere:

feebly uncinate in ventral view; lateral depression

deeply impressed; dorsum convex; venter concave;

mesoventral margin concave; mesodorsal margin

sinuous. Sinus: dorsal ovate; ventral lanceolate.

Phallus: phallic plates tapered; marginal denticles

stout (1 specimen examined).

Female Genitalia: As in P. nigriventris.

VARIATION.—Structural: Antenna is proportionally shorter and more cylindrical in one female

specimen, particularly articles 8 to 10.

Color: In the male specimen the pronotum is

predominantly testaceous, the abdomen black, and

the elytral disc avittate. Females have the pronotum

predominantly black, abdomen part black and part

testaceous, and the elytral disc distinctly vittate. The

elytral epipleural vitta varies in extent of expression, and is absent in the female specimen in which

antennal articles 8 to 10 are more cylindrical.

NATURAL HISTORY.—G. C. Champion collected

the available specimens from Volcan de Chiriquf at

various altitudes ranging from 610 to 1829 meters.

DISTRIBUTION.—Presently known only from the

type-locality.

LOCALITY RECORDS (Figure 364).—I examined 6 adult specimens from Central America. PANAMA: Provincia de Chriqui: Volcan de Chiriqui (BMNH, 1 male and 2 females, lectotype; GEKI, 1 female; MNHP, 1 female; ZMAN, 1 female).

REMARKS.—The name of this species has been

applied to members of P. nigriventris, P. reventazon,

and P. copiscolis. In view of the substantial similarity in elytral color among females of these species,

particularly with regard to the form of elytral

vittae, it is not surprising that the aforementioned

misidentincations involved only female specimens.
72

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 204-216.—Perilypus cultratus: 204, tegmen, lateral view; 205, tegmen, ventral view;

206, phallus; 211, male pygidium; 216, tegmen (arrow indicates confluency between mesoventral and mesodorsal margins). P. bilineatus: 207, tegmen, lateral view; 208, tegmen, ventral view; 209, phallus. P. cerroazul: 210, posterior accessory gland; 212, tegmen, lateral view;

213, tegmen, ventral view; 214, tegmen, dorsal view (arrow indicates dorsal brace); 215, anterior

accessory gland. (Scale = 1 mm)
NUMBER 227 73

24. Perilypus cerroazul, new species

FIGURES 210, 212-215, 217, S64

TYPE-LOCALITY.—Cerro Azul, Provincia de Panamd, Panama.

TYPE-SPECIMENS.—The holotype male, deposited

in NMNH, was collected by W. Bivin in 1971.

DIAGNOSIS.—The only specimen studied is superficially very similar to the sympatric specimens of

P. ordinatus. However, the antenna is more distinctly serrate and the pygidium more robust

(compare Figures 217, 245) in male specimens

of P. cerroazul.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except proportionally longer.

Size: Length: 9.2 mm. Width: 2.4 mm.

Color: As in P. reventazon except postocular

region of epicranium broadly flavotestaceous, femur

black at apex only, and elytron with faint discal

vitta visible only at elytral basal half.

Head: As in P. reventazon except antenna more

serrate.

Pronotum: As in P. reventazon except more

convex and side margins of pronotum proper less

arcuate.

Elytron: As in P. reventazon except proportionally longer (EL/EW, 5.2) and more densely

pubescent.

Abdomen: Male pygidium (Figure 217) particularly robust.

Front Tarsal Claws: Feebly asymmetrical.

Male Genitalia (Figures 212-214): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 9 times longer than paramere.

Combined length of paramere and phallobase 4

times longer than phallobasic apodeme. Paramere:

acuminate and arcuate in lateral view; lateral

depression indistinct; dorsum narrow and feebly

convex; venter broadly concave; mesodorsal margin

feebly concave, strongly braced, and margins

contiguous at basal fifth (note arrow in Figure

214). Sinus: dorsal and ventral broadly lanceolate.

Phallus: acuminate.

Internal Reproductive Organs: Male (Figures 210,

215), anterior accessory gland uniramous, coiled;

posterior accessory gland biramous, outer branch

about twice as long as inner branch; testis composed

of 12 follicles.

NATURAL HISTORY.—In June, on Cerro Azul,

Panama, W. Bivin collected the holotype specimen

by beating vegetation at 700 meters.

ETYMOLOGY.—The epithet is a noun in apposition

and refers to the type-locality.

LOCALITY RECORDS (Figure 364).—I examined 1 adult specimen from Central America. PANAMA: Provincia de Panama: Cerro Azul (NMNH, 1 male, holotype).

25. Perilypus ordinatus, new species

FIGURES 218-221, 245, 364

TYPE-LOCALITY.—Cerro Campana, Provincia de

Panama, Panama.

TYPE-SPECIMENS.—The holotype male, deposited

in NMNH, was collected by W. Bivin in 1971. Five

paratypes deposited in the repositories noted under

"Locality Records."

DIAGNOSIS.—See P. cerroazul.

DESCRIPTION.—Form: As in P. reventazon (Figure

186).

Size: Length: Average about 7.5 mm, range

6.8-8.5 mm. Width: Average about 2.1 mm, range

1.8-2.4 mm. Three males measured.

Color: Clypeus, frons, cranium venter, and

postocular region of epicranium flavotestaceous;

remainder of epicranium and gena black (see

"Variation"); antenna black; pronotum with side

margin broadly flavotestaceous, lower sides and

disc black; pronotal collar black (see "Variation");

prosternum testaceous; meso- and metasternum

black; profemur black dorsally, flavotestaceous

ventrally; meso- and metafemur black in dorsoapical fourth, flavotestaceous in remainder; proand mesotibia black, metatibia at least partially

testaceous; elytron piceous; abdomen black.

Head: As in P. reventazon except antenna more

serrate, proportionally shorter (AL/PL, 1.7) and

frontal umbo more prominent.

Pronotum: As in P. reventazon except less transverse (PL/PW, average about 0.94, range 0.92-0.96;

3 specimens measured), more convex, and side

margins of pronotum proper less arcuate.

Elytron: As in P. reventazon except more densely

pubescent.

Abdomen: Male pygidium as in Figure 245.

Front Tarsal Claws: Feebly asymmetrical.

Male Genitalia (Figures 218-220): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 3.5 times longer than para-
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mere. Combined length of paramere and phallobase 3.6 times longer than phallobasic apodeme.

Paramere: lateral depression broad, extended beyond basal limit of paramere; dorsum narrow and

convex; venter narrow and concave; mesodorsal

margin concave, strongly incised at basal third,

mesoventral margin strongly sinuous, feebly incised

at basal third and uncinate preapically (note arrow

in Figure 219). Sinus: as in Figure 219. Phallus: as

in Figure 220 (5 specimens examined).

Internal Reproductive Organs: Male (Figure 221);

anterior accessory gland uniramous, tightly coiled;

posterior accessory gland biramous, outer branch

longer than inner branch; testis composed of 12

follicles (5 specimens examined).

VARIATION.—Structural: I found no noteworthy

structural variation among the specimens studied.

Color: In one specimen the head, pronotum, and

mesoscutellum is entirely flavotestaceous.

NATURAL HISTORY.—On Cerro Campana (850

meters) W. Bivin collected two specimens by beating, one during March and one during May. From

the same locality additional specimens were

collected by S. S. and W. D. Duckworth (in April

by sweeping) and by H. P. Stockwell (in May by

beating).

ETYMOLOGY.—Latin, the adjective ordinatus (well

ordered). I dedicate the name of this species to my

colleague and friend Terry L. Erwin.

LOCALITY RECORDS (Figure 364).—I examined 6 adult specimens from Central America. PANAMA: Provincia de Panama: Cerro Campana (GEKI, 2 males; HPST, 1 male;

NMNH, 2 males, holotype; WBIV, 1 male).

26. Perilypus calcaris, new species

FIGURES 222-226, 363

TYPE-LOCALITY.—Finca Recreo, Municipio de

Yepocapa, Departamento de Chimaltenango, Guatemala.

TYPE-SPECIMENS.—The holotype male and the

allotype, deposited in FMNH, were collected by

R. D. Mitchell in 1948. Two paratypes: BMNH,

1 male; GEKI, 1 male.

DIAGNOSIS.—Specimens of P. calcaris are superfically very similar to specimens of P. nigriventris

and of P. lobulatus. The pygidium, however, is

substantially larger in males of P. calcaris, being

about equal in size to the one illustrated in Figure

185 (compare with Figure 245). Female specimens

of these three species are difficult to separate, but

color of the abdominal venter seems to provide

some basis for distinguishing them. The abdominal

venter is dark brown in the anterior half and

flavotestaceous in the posterior half (except the

fifth visible sternum is infuscated) in females of

P. calcaris, entirely dark brown in females of P.

lobulatus, and predominantly flavotestaceous (sternum vi is dark brown) in females of P. nigriventris.

DESCRIPTION.—Form: As in P. reventazon (Figure

186).

Size: Length: males, average about 7.6 mm, range

7.2-7.8 mm; female 7.9 mm. Width: males, average

about 2.3 mm, range 2.0-2.4 mm; female, 2.2 mm.

Three males and one female measured.

Color: As in P. nigriventris except female elytron

faintly pale along posthumeral margin, femora dark

brown in apical half (except testaceous ventroapically), coxae strongly infuscated, and female abdominal venter not predominantly flavotestaceous.

Head: As in P. reventazon except antenna more

slender.

Pronotum: As in P. reventazon except pronotal

arch and pronotum proper with shallow rugosities.

Elytron: As in P. reventazon except punctations

smaller and shallower and surface more arenose.

Front Tarsal Claws: Male, distinctly asymmetrical; female, feebly asymmetrical.

Male Genitalia (Figures 222-224): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.3 times longer than paramere.

Combined length of paramere and phallobase 4

times longer than phallobasic apodeme. Paramere:

apex feebly uncinate in ventral view; lateral

depression narrow, more dorsad than ventrad;

dorsum convex and broadly explanate medially;

venter narrow and feebly concave; mesoventral

margin emarginated and biacuminate; mesodorsal

margin sinuous, with stout brace at base. Sinus:

dorsal much narrower than elliptical ventral.

Phallus: acuminate; outer margin inflexed; stout

marginal denticles centralized at middle half of

phallic plate (3 specimens examined).

Female Genitalia (Figures 225, 226): As in P.

orthopleuridus except dorsal lamina reduced.

VARIATION.—Structural: The pronotal rugosities

vary in extent of expression.

Color: Color of the abdominal venter varies with
NUMBER 227 75

FIGURES 217-228.—Perilypus cerroazul: 217, male pygidium. P. ordinatus: 218, tegmen, lateral

view; 219, tegmen, ventral view (arrow indicates preapical uncination of mesoventral margin);

220, phallus; 221, male internal reproductive organs. P. calcaris: 222, tegmen, lateral view;

223, tegmen, ventral view (arrow indicates acumination of mesoventral margin); 224, phallus;

225, ovipositor, ventral view; 226, ovipositor, dorsal view. P. orthopleuridus: 221, ovipositor,

ventral view; 228, ovipositor, dorsal view. (Scale = 1 mm)
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sex; in male specimens entirely dark brown, in

females dark brown in basal half and mostly

flavotestaceous in apical half.

NATURAL HISTORY.—R. D. Mitchell collected

three adult specimens (at 1341 meters) in May from

Finca Recreo, Guatemala, by beating "dead

branches." G. C. Champion collected one specimen

from Cerro Zunil, Guatemala, at an elevation

between 1219 and 1832 meters.

DISTRIBUTION (Figure 363).—Known only from

the southwestern highlands of Guatemala.

ETYMOLOGY.—The trivial name is the genetive

case of calcar (a spur) and refers to the spur-like

acumination on the mesoventral margin of parameres.

LOCALITY RECORDS (Figure 363).—I examined 4 adult specimens from Central America. GUATEMALA: Departamento

de Quetzaltenango: Cerro Zunil (BMNH, 1 male). Departamento de Chimaltenango: Finca Recreo (GEKI, 1 male;

FMNH, 1 male holotype, and 1 female).

27. Perilypus orthopleuridus (Thomson),

new combination

FIGURES 27, 227-234, 364

Derestenus orthopleuridus Thomson, 1860:57 [holotype, female, deposited in MNHP; type-locality: Mexico].—Gorham, 1882:144—Schenkling, 1906:263.

Derestenus vittipennis Chevrolat, 1874:290 [lectotype: male,

deposited in BMNH, here designated; type-locality: Mexico; new synonmy].—Gorham, 1882:143.

Derestenus collaris Chevrolat, 1876:12 [lectotype: female, deposited in MNHP, here designated; type-locality: Mexico].

—Schenkling, 1906:263.

The new synonmy is based on the discovery that

D. vittipennis Chevrolat is the heretofore undescribed male of P. orthopleuridus.

DIAGNOSIS.—The combination of front tarsal

claws boldly asymmetrical, and antenna partially or

totally flavotestaceous will easily separate males of

this species from other males within the reventazon

group. The epipleuron (Figure 230) is explanate

and plane ventrally in P. orthopleuridus females

only.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except elytral apical slope gradual.

Size: Length: males, average about 8.2 mm, range

5.8-9.9 mm; females, average about 8.9 mm, range

7.6-10.4 mm. Width: males, average about 2.1 mm,

range 2.0-2.8 mm; females average about 2.3 mm,

range 2.0-3.4 mm. Ten males and 10 females

measured.

Color: Cranium unicolorous or bicolorous; unicolorous cranium is flavotestaceous; bicolorous

cranium has clypeus, frons, and cranial venter

flavotestaceous, and epicranium and gena piceous;

antenna entirely flavotestaceous, or articles 1 to 8

brunneus and 9 to 11 flavotestaceous; pronotum

predominantly flavotestaceous and with midapical

piceous macula, or flavotestaceous at sides and

piceous discally; pronotal collar piceous or predominatly flavotestaceous; prosternum flavotestaceous; mesoscutellum and meso- and metasternum

flavotestaceous or piceous; femur entirely flavotestaceous, flavotestaceous in basal half and ventroapical half and piceous in dorsoapical half, or

predominantly flavotestaceous and infuscated dorsoapically; tibia and tarsus testaceous or piceous;

elytron piceous, with or without flavous discal

vitta; visible abdominal sterna I to v entirely

flavotestaceous, predominantly flavotestaceous and

infuscated basally or laterally, or piceous; sixth

visible abdominal sternum piceous.

Head: As in P. reventazon except antenna less

serrate.

Pronotum: As in P. reventazon except more

convex.

Elytron: As in P. reventazon, except apical slope

gradual, punctations shallower, surface not arenose,

and female epipleuron explanate and plane ventrally (Figures 230, 231).

Abdomen: Male pygidium (Figure 229) conspicuously large and feebly convex.

Front Tarsal Claws: Male, boldly asymmetrical;

female, feebly asymmetrical.

Male Genitalia (Figures 232-234): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 4.7 times longer than paramere. Combined length of paramere and phallobase

3.0 times longer than phallobasic apodeme. Paramere: apex uncinate; lateral depressions extended

beyond base of ventral sinus; dorsum particularly

narrow and broadly braced basally; venter narrow,

concave; mesoventral margin emarginated (note

arrow in Figure 233); mesodorsal margin sinuous.

Sinus: dorsal about twice as broad as ventral.

Phallus: acuminate; outer margin inflexed in basal

half and denticulated from middle to basal three-
NUMBER 227 77

fourths; plicae feebly developed (45 specimens

examined).

Female Genitalia (Figures 227, 228): Dorsal and

ventral lamina and coxites very long; dorsal lamina

FIGURE 229-238.—Perilypus orthopleuridus: 229, male pygidium, 230, female elytron, ventral

view, X 25 (arrow indicates epipleuron); 231, female elytron, lateral view, X 25 (arrow indicates epipleuron); 232, tegmen, lateral view; 233, tegmen, ventral view (arrow indicates

acumination of mesoventral margin); 234, phallus, lateral view. P. exilis: 235, tegmen, lateral

view; 236, tegmen, ventral view. P. latilobus: 237, tegmen, lateral view; 238, tegmen, ventral

view. (Scale = 1 mm)
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slender, deeply incised; lateral lobes of ventral

lamina shorter than medial lobe; inner margin of

coxites well sclerotized; remainder as in P. reventazon (2 specimens examined).

Internal Reproductive Organs: Male, anterior

accessory gland uniramous tightly coiled, posterior

accessory gland biramous (3 specimens examined).

VARIATION.—Structural: The epipleuron is explanate in female elytra only; epipleural explanation varies but is always prominent. The cranium,

femora, pronotum, and abdominal sterna are

conspicuously rugose and the elytral surface is more

arenose in the female specimen from Temazcal,

Mexico.

Color: Coloration of cranium, pronotum, femur,

meso- and metasternum, and of abdominal sterna

is correlated with sex.

Pronotal color distinguishes 2 male phena, a

maculate phenon and a vittate phenon. Among the

45 male specimens studied, 35 belong to the

maculate phenon (the pronotum is predominantly

flavotestaceous and has a piceous midapical macula)

and 10 to the vittate phenon (the pronotal disc

has a piceous vitta that is narrowly percurrent or

interrupted at the middle).

Among the maculate males, 11 have the elytron

distinctly vittate, in the remaining specimens the

elytral vitta is faintly visible or absent. In all

maculate males the cranium, femur, and meso- and

metasternum are entirely flavotestaceous. In two of

the 10 vittate males the elytra are distinctly vittate,

while in the remaining 8 the elytral discal vitta

are either faintly visible or absent. Vittate males

have the cranium and the femur bicolorous, and

the meso- and metasternum piceous. In all males

the abdomen is entirely piceous.

The female pronotum is vittate, the elytron

either avittate, faintly vittate, or distinctly vittate.

The female cranium, femur, and the first five

visible abdominal sterna are partly testaceous and

dark brown; sternum vi is always black.

DISTRIBUTION (Figure 364).—The known range

extends from Mexico to Costa Rica with locality

records from Guatemala and Honduras.

LOCALITY RECORDS (Figure 364).—I examined 98 adult specimens from Central America. MEXICO: Estado de Nayarit:

Tepic (MNHB, 1 male); Estado de Mexico; Toluca (ZMAN, 1

female). Estado de Veracruz: Atoyac (BMNH, 1 female); 49

miles [78 km] south of Acayucan (GEKI, 1 female); Orizaba

(BMNH, 1 male; CASC, 4 males and 4 females; GEKI, 3

males and 3 females; GEKI, 2 males; MNHB, 1 female,

LACM, 1 male; MNHP, 3 females, holotype; ZMAN, 1 male);

1 mile [1.6 km] west of Fortin de las Flores (CISC, 1 male);

Cordoba (BMNH, 2 males; CASC, 4 males and 2 females;

FMNH, 1 male; GEKI, 4 males and 2 females; NMNH, 2

males); Jalapa (CASC, 1 male; FMNH 1 male); Mirador

(MNHB, 1 male); Lago de Catemaco (CNCC, 1 male; GEKI,

2 males and 1 female); San Andres Tuxtla (GEKI, 1 male);

Playa Vincente (BMNH, 1 female); Cerro de Palmas (BMNH,

1 female). Estado de Oaxaca: Oaxaca (BMNH, 1 female,

GEKI, 1 female); Temazcal (GEKI, 1 female; GHNE, 1 female). Estado de Chiapas: 8 miles [13 km] south of Simojovel

(CNCC, 1 male); 27 kilometers southwest of Simojovel (GEKI,

1 female); 2 miles [3 km] northeast of Bochil (GEKI, 1 male);

2 kilometers north of Bochil (GEKI, 1 female); 12 miles [19

km] north of Tuxtla Gutierrez (GEKI, 1 male); 11 miles [18

km] north of Ocozocoautla (GEKI, 1 female; TAMT, 1 female); 22 kilometers north of Ocozocoautla (CNCC, 1 female);

El Suspiro, Berriozabal (GEKI, 1 male). "Mexico" (BMNH, 1

male and 2 females; FMNH, 1 male; GEKI, 3 males and 7

females; MNHB, 1 female; MNHP, 1 male and 7 females;

ZMAN, 2 females). GUATEMALA: Departamento de Alta

Verapaz: San Juan (BMNH, 1 male). Tamahu (AMNH, 1

female); Senahii (BMNH, 1 female). HONDURAS: Departamento de Choluteca: Cantarranas, Rio Choluteca (GEKI, 1

female.) COSTA RICA: Provincia de Cartago: Turrialba

(GEKI, 1 male).

REMARKS.—Specimens of this species were identified as P. quadrilineatus, P. distinctus, P. nigrifrons, and Colyphus signoticollis Spinola.

28. Perilypus exilis, new species

FIGURES 235, 236, 363

TYPE-LOCALITY.—Thirty-two kilometers north of

Huixtla, Estado de Chiapas, Mexico.

TYPE-SPECIMENS.—The holotype male, deposited

in CNCC, was collected by W. R. M. Mason in

1969.

DIAGNOSIS.—The combination of conspicuously

slender antenna (with article 11 strongly acuminate),

front tarsal claws conspicuously asymmetrical, and

pygidium exceptionally robust and truncate will

distinguish males of this species from other males

within the reventazon group.

DESCRIPTION.—Form: As in P. reventazon (Figure

186) except more slender.

Size: Length: male, 8.0 mm. Width: male, 2.4 mm.

Color: As in males of P. calcaris except coxae not

infuscated.

Head: As in P. reventazon except frons proportionally narrower (HW/IOW, 2.5), antenna more

slender, antennal article 11 more acuminate, and
NUMBER 227 79

length/width ratio of each antennal article 2.0:

1.3:1.7:1.7:1.7:1.9:1.9:1.9:1.6:1.4:2.1.

Pronotum: As in P. reventazon except less transverse (PW/PLS, 0.91).

Elytron: As in P. reventazon.

Abdomen: Male pygidium conspicuously robust

and truncate.

Front Tarsal Claws: Boldly asymmetrical.

Male Genitalia (Figures 235, 256): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 6.3 times longer than paramere. Combined length of paramere and pallobase

3 times longer than phallobasic epodeme. Paramere: outer margin compressed at middle; feebly

curvate in apical fourth in lateral view; lateral

depressions well depressed; dorsum convex and

venter concave at their respective apical half, basal

half dorsum and venter plane; mesoventral and

mesodorsal margin sinuous. Sinus: dorsal narrow in

apical half, broad at basal half; ventral narrow,

constricted medially. Phallus: as in P. claudus

except more slender.

NATURAL HISTORY.—W. R. M. Mason collected

the only available specimen of this species in June,

in Chiapas, Mexico, at an altitude of 914 meters.

ETYMOLOGY.—Latin, the adjective exilis (thin). I

refer to the slender form of the antenna.

LOCALITY RECORDS (Figure 363).—I examined 1 adult specimen from Central America. MEXICO: Estado de Chiapas:

20 miles [32 km] north of Huixtla (CNCC, 1 male, holotype).

29. Perilypus latilobus, new species

FIGURES 237, 238, 363

TYPE-LOCALITY.—Six kilometers northwest of

Pueblo Nuevo, Rio Bahada, Estado de Chiapas,

Mexico.

TYPE-SPECIMENS.—The holotype male is deposited in NMNH, the allotype in CNCC. The first

of these specimens was collected by G. H. Nelson in

1965, the second by J. M. Campbell in 1969. One

paratype: GEKI, 1 female.

DIAGNOSIS.—The combination of femora predominantly black to dark brown and pronotal

discal vitta abruptly constricted at subapical depression will generally distinguish members of this

species from the very similar specimens of P. nigriventris, P. calcaris, and P. exilis.

DESCRIPTION.—Form: As in P. reventazon (Figure

186).

Size: Length: male, 8.0 mm; females, average

about 8.5 mm, range 7.9-9.2 mm. Width: Male,

2.2 mm; females, average about 2.4 mm, range 2.0-

2.8 mm. One male and 2 females measured.

Color: As in P. calcaris.

Head: As in P. reventazon except antenna proportionally shorter (AL/PLS, 1.7).

Pronotum: As in P. reventazon except disc more

convex and side margin of pronotum proper less

arcuate.

Elytron: As in P. reventazon.

Abdomen: Male pygidium subquadrate; pygidial

apodeme as long as pygidial plate.

Front Tarsal Claws: Male, boldly asymmetrical;

female, feebly asymmetrical.

Male Genitalia (Figures 237, 238): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 4.3 times longer than paramere.

Combined length of paramere and phallobase 5

times longer than phallobasic apodeme. Paramere:

conspicuously broad in basal four-fifths, abruptly

narrow apical fifth, lateral depression inconspicuous in lateral view; dorsum broadly convex,

explanate medially; venter broadly concave; mesoventral margin concave; mesodorsal margin feebly

sinuous and briefly denticulated preapically. Sinus:

dorsal narrow in apical half, slightly expanded in

studied (1 specimen examined).

Female Genitalia: As in P. nigriventris.

Internal Reproductive Organs: Female, as in

Figure 55; ovary composed of 12 ovarioles (2 specibasal half; ventral subelliptical. Phallus: not

mens examined).

VARIATION.—Structural: The female specimen

from Tuxtla Gutierrez is more robust than the one

from Pueblo Nuevo.

Color: The specimen from Tuxtla Gutierrez has

a short narrow vitta on the elytral, disc which is not

the case in the other two specimens studied.

NATURAL HISTORY.—In June in Tuxtla Gutierrez,

Mexico, at 1280 meters, I collected one adult specimen by beating roadside vegetation. G. H. Nelson

collected the holotype in July, by sweeping roadside vegetation near Rio Bahada, in Pueblo Nuevo,

Mexico.

DISTRIBUTION (Figure 363).—Known only from

southern Mexico.

ETYMOLOGY.—The trivial name is a Latin com-
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pound noun formed from the adjective latus (wide)

+ the noun lobus (lobe). I refer to the lobe-like

appearance of the paramere.

LOCALITY RECORDS (Figure 363).—I examined 3 adult specimens from Central America. MEXICO: Estado de Chiapas:

four miles [6 km] northwest of Pueblo Nuevo, Rio Bahada

(NMNH, 1 male, holotype); Tuxtla Gutierrez (GEKI, 1 female): junction between routes 190 and 195 (CNCC, 1 female).

30. Perilypus distinctTM (Chevrolat),

new combination

FIGURES 239-242, 363

Derestenus distinctus Chevrolat, 1874:289 [lectotype: female,

deposited in MNHP, here designated; type-locality: Yucatan, Estado de Yucatan, Mexico].—Gorham, 1882:143.—

Schenkling, 1906:262.

DIAGNOSIS.—Within the reventazon group specimens from between the southern half of Mexico

and north of Honduras belong to this species if

they are particularly densely pubescent, have the

pronotal sides broadly flavotestaceous and often

roseate, and have a short but robust body form.

DESCRIPTION.—Form: As in Figure 239.

Size: Length: males, average about 8.5 mm, range

6.3-9.4 mm; females, average about 8.1 mm, range

6.5-9.8 mm. Width: males, average about 2.0 mm,

range 1.8-2.6 mm; females, average about 2.5 mm,

range 2.0-2.8 mm. Ten males and 10 females

measured.

Color: Clypeus, frons, and venter of cranium

flavotestaceous; epicranium predominantly black or

predominantly flavotestaceous, often marked by

subquadrate black macula; gena black or flavotestaceous; antenna black; pronotum with black

discal vitta that narrows at middle; pronotal sides

broadly flavotestaceous or roseate; pronotal collar

black; prosternum flavotestaceous; meso- and

metasternum dark brown or black; femora

predominantly flavotestaceous, infuscated dorsoapically; tibia and tarsus dark brown or black;

elytral disc with or without narrow flavotestaceous

vitta that when present begins at midbase, gradually oblique toward suture, and usually not incurved apically; abdominal venter entirely black, or

first five visible sterna flavotestaceous, sixth black.

Head: As in P. reventazon except antenna more

serrate and proportionally shorter (AL/PLS, 1.6).

Pronotum: As in P. reventazon except more convex, subapical depression more sinuous, and side

margins less arcuate.

Elytron: As in P. reventazon except elytron proportionally shorter (EL/EW, average about 4.4,

range 4.3-5.6; 20 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 240-242): Strongly pigmented. Combined length of phallobase and phallobasic apodeme variable in relation to length of

paramere (see "Variation"). Combined length of

paramere and phallobase 3.0 times longer than

phallobasic apodeme. Paramere: apex boldly

uncinate; lateral depression broad and deeply impressed; dorsum convex, strongly braced at base;

venter narrow, concave; mesoventral and mesodorsal

margins sinuous. Sinus: dorsal broadly lanceolate;

ventral elliptical, constricted basally. Phallus:

abruptly narrowed near apex; marginal denticles

prominent (18 specimens examined).

Female Genitalia: As in P. nigriventris.

Internal Reproductive Organs: Male, anterior

accessory gland uniramous; coiled posterior accessory gland biramous, divided into inner and outer

branch (3 specimens examined).

VARIATION.—Structural: The ratio between the

combined length of the phallobase and the phallobasic apodeme over the length of the paramere

is highly variable (6.5 to 9.0). The variation may

be geographic but the available data are inconclusive. In general, the aforementioned ratio is lower

in males from Oaxaca than in those from Veracruz.

Color: Sex dimorphism is evident in the coloration of the abdominal venter and in the elytron. In

females, the abdominal venter is bicolorous and the

elytron vittate. In males, the abdominal venter is

concolorous and the elytron avittate. Of 17 females

examined 2 have the elytral vitta feebly incurved

apically.

NATURAL HISTORY.—Adult specimens were collected in Mexico from May to August. H. F.

Howden collected 9 specimens at 609 meters by

beating tree foliage in Oaxaca, Mexico.

DISTRIBUTION (Figure 363).—The known range of

this species extends from Veracruz, Mexico, to Alta

Verapaz, Guatemala.

LOCALITY RECORDS (Figure 363).—I examined 45 adult specimens from Central America. MEXICO: Estado de Veracruz:

Cordoba (BMNH, 1 female, CASC, 1 male; GEKI, 1 male and

1 female; FMNH, 1 female); Montepio, 8 miles [13 km] north

of Sontecomapan (GEKI, 1 female); Lago de Catemaco
NUMBER TZ1 81

i

FIGURES 239-245.—Perilypus distinctus: 239, habitus; 240, tegmen, lateral view; 241, tegmen,

ventral view; 242, phallus. P. crassus: 243, habitus; 244, antenna. P. ordinatus: 245, male

pygidium. (Scale = 1 mm)
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(CNCC, 5 males and 1 female; GEKI, 2 males and 2 females);

Cerro de Palma (BMNH, 1 female). Estado de Oaxaca: 6

miles [10 km] south of Valle Nacional (CNCC, 1 male;

GEKI, 4 males and 1 female; HFHO, 4 males and 1 female).

Estado de Chiapas: 20 to 25 miles [32 to 40 km] north of

Huixtla (CNCC, 1 female); Rio de Salvador, 4 kilometers

northwest of Ixhautan (GHNE, 1 female). Estado de Yucatan: Yucatan (MNHP, 1 female, lectotype). "Mexico"

(MNHP, 1 male and 3 females; MNHB. 1 female; ZMAN,

1 male). GUATEMALA: Departamento de Alta Verapaz:

Teleman (BMNH. 2 males). "Guatemala" (GEKI, 1 male).

BELIZE: Distrito de Corozal: Rio Hondo (GEKI, 1 male).

Distrito de Belize: Belize (GEKI, 1 male). "M-tee Dist"

(GEKI, 1 female; MCZC, 1 male and 1 female).

REMARKS.—A male specimen formed part of the

syntype series of P. nigriventris. Other specimens

were identified as P. quadrilineatus, P. orthopleuridus, P. nigrifrons, and Colyphus signaticollis

Spinola.

31. PerUypus crassus, new species

FIGURES 243, 244, 246-248, 365

TYPE-LOCALITY.—Rio Huallaga, Yurimaguas,

Chambireyacu, Provincia de Huallaga, Peru.

TYPE-SPECIMENS.—The holotype male and the

allotype, deposited in MNHP, were collected by

M. de Mathan in 1885. One paratype: GEKI, 1

male.

DIAGNOSIS.—The combination of robust body that

is densely pubescent and truncated, boldly serrate

antenna (Figure 244), and South American distribution will easily identify members of this species

within the reventazon group.

DESCRIPTION.—Form: As in Figure 243.

Size: Length: males, 7.4 mm; female, 7.3 mm.

Width: males, 4.6 mm; female, 4.3 mm. Two males

and 1 female measured.

Color: Clypeus, anterior half of frons, and venter

of cranium navotestaceous; posterior half of frons

black; epicranium predominantly black, fulvous in

postocular region; prosternum flavotestaceous; pronotum flavotestaceous dorsolaterally, black ventrolaterally and discally; pronotal collar black; front

femur flavotestaceous in proximal half, dark brown

in distal half; middle and hind femur flavotestaceous in basal two-thirds, dark brown in distal third;

elytron piceous but epipleural fold slightly paler;

abdominal venter black.

Head: As in P. reventazon except antennal ridge

more prominent; antenna (Figure 244) more serrate, flattened, and profusely pubescent; scrape

shorter than article 11; and length/width ratio of

each male antennal article 2.0:1.0:1.0:1.1:1.1:1.0:

1.0:1.0:1.0:1.0:1.8.

Pronotum: As in P. reventazon except only feebly

transverse (PL/PW, average about 0.93, range 0.91-

0.95; 3 specimens measured), integumental setae

more numerous and longer, disc more convex, subapical depression more sinuous, and side margins

of pronotum proper more arcuate.

Elytron: As in P. reventazon except with more

depth, prominently convex and truncated, surface

more arenose and more densely pubescent, epipleural fold explanate dorsally, EL/EW, average about

4.4, range 4.3-4.6 (3 specimens measured).

Front Tarsal Claws: Feebly asymmetrical in both

sexes.

Male Genitalia: Strongly pigmented. Combined

length of phallobase and phallobasic apodeme 5.7

times longer than paramere. Combined length of

paramere and phallobase 3.4 times longer than

phallobasic apodeme. Paramere: apex feebly upcurved; lateral depression particularly prominent;

dorsum convex; explanate medially; venter concave; mesoventral margin arcuate; mesodorsal margin feebly arcuate in most of posterior two-thirds,

strongly arcuate in anterior third. Sinus: dorsal

feebly elliptical at middle, oval basally; ventral elliptical. Phallus: apex feebly digitiform; marginal

denticles prominent (2 specimens examined).

Female Genitalia: As in P. nigriventris.

VARIATION.—Structural: The female antenna is

slightly more serrate than male antenna.

Color: The pronotal discal vitta is narrow medially in the females and broad throughout in

males.

NATURAL HISTORY.—Unknown except M. de

Mathan collected two specimens near Rio Huallaga,

Peru, one in June and one in August.

DISTRIBUTION (Figure 365).—Known only from

Huallaga to Loreto, Peru.

ETYMOLOGY.—Latin, the adjective crassus (stout),

with reference to the short stout body of this insect.

LOCALITY RECORDS (Figure 365).—I examined 3 adult specimens from South America. PERU: Provincia de Huallaga:

Chambireyacu, Yurimaguas, Rio Huallaga (MNHP, 1 male,

holotype, and 1 female). Provincia de Loreto: Rio Amazonas

(GEKI, 1 male).
NUMBER 227

83

32. Perilypus immitis, new species

FICURES 249-253, 364

TYPE-LOCALITY.—Atlixco, Estado de Puebla,

Mexico.

TYPE-SPECIMENS.—The holotype male and allotype are deposited in CASC. Five paratypes deposited in repositories as noted under "Locality Records."

DIAGNOSIS.—Within the reventazon group, specimens from Mexico belong to this species if their

elytra are predominantly or entirely blue-green or

olive green (the elytral apex is usually testaceous),

and if their elytral surface is densely vested with

gray reclinate setae.

DESCRIPTION.—Form: rectangulate; these beetles

are exceptionally small and shallow-bodied.

Size: Length: males, average about 6.2 mm, range

5.4-6.9 mm; females, average about 6.9 mm, range

5.9-8.0 mm. Width: males, average about 2.2 mm,

range 1.6-3.0 mm; females, average about 2.3 mm,

range 2.0-2.6 mm. Five males and 2 females measured.

Color: Cranium entirely testaceous, or clypeus

frons and venter of cranium testaceous and epicranium and gena black; antenna dark brown;

prothorax testaceous, except pronotum with midapical dark brown macula; mesosternum predominantly testaceous; metasternum black; femora

becoming more dark brown from front to middle

femur; elytron olive green or blue-green; abdomen

testaceous or dark brown.

Head: As in P. reventazon except eyes less convex and cranium more coarsely punctated (particularly in males).

Pronotum: As in P. reventazon.

Elytron: As in P. reventazon except elytron proportionally shorter (EL/EW, average about 4.2,

range 3.2-4.7; 11 specimens measured) and more

densely vested with reclinate setae.

Front Tarsal Claws: Male, boldly asymmetrical;

female, feebly symmetrical.

Male Genitalia (Figures 249-251): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5 times longer than paramere. Combined length of paramere and phallobase 2.7 times

longer than phallobasic apodeme. Paramere: very

broad basally; lateral depression deeply impressed

at middle third; dorsum convex, explanate medially

to extent that mesodorsal margins overlap; venter

concave; mesoventral margin and mesodorsal margin sinuous, latter denticulated in apical half only.

Sinus: dorsal indistinct, ventral obovate and acuminate basally. Phallus: phallic plate abruptly narrowed in apical fourth; marginal denticles localized

at middle third of plate (5 specimens examined).

Female Genitalia (Figures 252, 253): Dorsal and

ventral lamina short; proctiger particularly broad;

remainder as in P. nigriventris (1 specimen examined).

VARIATION.—Structural: The epicranium and

frons are more coarsely punctated in males than in

females.

Color: The male specimen from Yecora does not

have the elytral apex testaceous. Also, in this beetle

the elytron has a more bluish tinge; in most other

specimens, the elytron is olive green.

NATURAL HISTORY.—In May, in Yecora, Mexico,

H. F. Howden collected one adult specimen at

2134 meters by beating oak foliage (Quercus sp.).

Specimens were also collected in June and July.

ETYMOLOGY.—Latin, the adjective immitis

(rough). The name refers to the coarse dentitions

along the mesodorsal margin of the parameres.

DISTRIBUTION (Figure 364).—From Sonora to

Puebla, in Mexico.

LOCALITY RECORDS (Figure 364).—I examined 7 adult specimens from Central America. MEXICO: Estado de Sonora:

Yecora (GEKI, 1 male). Estado de Queretaro: San Juan del

Rio (GEKI, 1 female; MNHB, 1 male). Estado de Puebla:

Atlixco (CASC, 1 male, holotype, and 1 female; GEKI, 1

male). Estado de Morelos: Tepoztlan (MHNM, 1 male).

33. Perilypus caliculus, new species

FIGURES 254-256, 363

TYPE-LOCALITY.—Tejupilco, Estado de Mexico,

Mexico.

TYPE-SPECIMENS.—The holotype male, deposited

in FMNH, and 73 paratypes were collected by

H. E. Hinton and R. L. Usinger in 1933. Paratypes

are deposited in repositories as noted under "Locality Records."

DIAGNOSIS.—The male genitalia (Figures 254-

256) provides the only reliable characteristics for

diagnosing members of this species.

DESCRIPTION.—Form: As in P. reventazon (Figure

186).
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 246-259.—Perilypus crassus: 246, tegmen, lateral view; 247, tegmen, ventral view; 248,

phallus. P. immitis: 249, tegmen, lateral view; 250, tegmen, ventral view; 251, phallus; 252,

ovipositor, ventral view; 253, ovipositor, dorsal view. P. caliculus: 254, tegmen, lateral view;

255, tegmen, ventral view; 256, phallus. P. carbonarius: 257, tegmen, lateral view; 258, tegmen, ventral view; 259, phallus. (Scale = 1 mm)
NUMBER 227 85

Size: Length: males, average about 7.8 mm, range

5.9-9.0 mm; females, average about 7.6 mm, range

ti.4-9.2 mm. Width: males, average about 2.3 mm,

range 2.1-2.6 mm; females, average about 2.5 mm,

range 2.3-3.0 mm. Ten males and 10 females measured.

Color: Cranium entirely flavotestaceous, or clypeus postocular region of epicranium and cranial

venter flavotestaceous and frons and epicranium

black; prothorax flavotestaceous, with or without

piceous macula at pronotal midapex, or prosternum

and upper sides of pronotum and pronotal disc

black; mesoscutellum flavotestaceous or black;

meso- and metasternum from flavotestaceous to

black; legs predominantly flavotestaceous or predominantly black; elytron usually piceous or black

and avittate, rarely flavotestaceous or vittate; vittate

elytron with flavotestaceous vitta on epipleural margin, disc, and sutural margin.

Head: As in P. reventazon except setiferous punctations on frons and epicranium larger and antenna

more serrate.

Pronotum: As in P. reventazon except subapical

depression more deeply impressed.

Elytron: As in P. reventazon except elytral surface more arenose.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 254-256): Strongly pigmented. Combined length of phallobase and phallobasic apodeme about 7 times longer than paramere.

Combined length of paramere and phallobase about

3.0 times longer than phallobasic apodeme. Paramere: arcuate in lateral view; dorsum convex; venter broadly concave; mesoventral and mesodorsal

margin concave. Sinus: dorsal and ventral elliptical.

Phallus: phallic plates broad, with marginal denticles faintly visible (74 males examined).

Female Genitalia: As in P. nigriventris except

lateral lobes of ventral lamina proportionally

shorter.

Internal Reproductive Organs: Male, anterior accessory gland uniramous, coiled; posterior accessory

gland biramous (5 specimens examined).

VARIATION.—Structural: The female pronotum is

usually more transverse than the male pronotum.

The setiferous punctations on the head and pronotum are particularly robust in some specimens

from Valle De Bravo, Mexico; also in some of

these specimens the interocular depressions are

coarsely rugose.

Color: Among the males studied 60 have the

cranium entirely flavotestaceous and the prothorax

predominantly flavotestaceous; in other specimens

the cranium and prothorax is predominantly black.

In females, the cranium is always mostly black; the

pronotum either predominantly flavotestaceous and

maculated midapically or predominantly black, in

which case it has a black discal vitta.

I assign 14 specimens from Tejupilco, Mexico, to

a trivittate phenon. These have the elytron trivittate; the vitta on the epipleural margin extends

from the elytral base to the elytral apical half or

more; the discal vitta begins at the elytral base

(nearer the posthumeral margin than to the sutural

margin) curves inward at the basal fifth and gradually outcurves to the elytral apical fifth where it

curves inwards towards the elytral apex; the sutural vitta is faintly developed and extends from

the elytral base to the elytral basal third.

NATURAL HISTORY.—Adult specimens were collected from Mexico in May; in June by H. E.

Evans from Morelos at 1829 meters and by G. H.

Nelson on Mimosa sp. in Valle de Bravo; in July

by H. E. Hinton from Tejupilco at 1164 meters;

in August by G. H. Dieke from Michoacdn at 1378

meters; and in October by H. H. Smith from Chilpancingo at 378 and 914 meters.

DISTRIBUTION (Figure 363).—A specimen from

Turrialba, Costa Rica, extends the known range of

this species from Mexico to Costa Rica.

ETYMOLOGY.—Latin, a masculine diminutive

noun caliculus (cup). I refer to the cup-shaped

contour of the combined venter of the parameres.

LOCALITY RECORDS (Figure 363).—I examined 168 adult

specimens from Central America. MEXICO: Estado de

Hidalgo: Metamoros (CASC, 2 males and 2 females; GEKI, 1

male). Estado de Mexico: Tejupilco (FMNH, 36 males, holotype, and 41 females; GEKI, 6 males and 6 females); Real de

Arriba (FMNH, 1 male and 4 females; GEKI, 1 male and 1

females); Valle de Bravo (GEKI, 3 males and 2 females;

GHNE, 10 males and 8 females); Cuernavaca (CASC, 2 males

and 3 females; CNCC, 2 males and 1 female; CUNY, 1 male

and 1 female; GEKI, 2 males and 2 females). Estado de

Morelos: Yautepec (BMNH, 1 male and 1 female). Estado

de Guerrero: Tepetlapa (BMNH, 1 male and 1 female; GEKI,

1 male); Chilpancingo (AMNH, 1 female; BMNH, 5 males

and 8 females; CNCC, 1 female; GEKI, 1 male; Iguala

(CNCC, 1 male and 1 female); Soledad (BMNH, 1 male);

Tasco (BMNH, 1 female; NMNH, 1 male and 1 female).

"Mexico" (BMNH, 1 male). "Guatemala" (GEKI, 1 male).

COSTA RICA: Provincia de Cartago: Turrialba (GEKI, 1

male).
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—At first glance specimens of the trivittate phenon are very different from the others

included herein, but they do not differ appreciably

in any structural characteristic studied.

34. Perilypus carbonarius Spinola

FIGURES 257-265, 36S

Perilypus carbonarius Spinola, 1844a: 105 [holotype: male,

deposited in MSPI; type-locality: "Mexico"].—Ekis, 1975:26.

Derestenus lateralis Chevrolat, 1874:289 [lectotype: female,

and 1 female paralectotype, here designated, deposited in

BMNH; type-locality: "Mexico"; new synonymy].—

Gorham, 1882:142.

Derestenus mutabilis Chevrolat, 1874:289 [lectotype: female,

and S female paralectotypes, here designated, deposited in

MNHP; type-locality: "Mexico"; new synonymy].—

Gorham, 1882:142.

Derestenus nigrifrons Chevrolat, 1874:289 [lectotype: male,

here designated, deposited in MNHP; type-locality:

Oaxaca, Estado de Oaxaca, Mexico; new synonymy].—

Gorham, 1882:142.—Wolcott, 1927a:35.

Colyphus flammeus Gorham, 1878:162 [lectotype, female,

here designated, deposited in BMNH; type-locality:

"Mexico"; Gorham, 1882:142, placed this nominal species

in synonymy with Derestenus mutabilis Chevrolat].

Colyphus marginatus Gorham, 1878:162 [lectotype: female,

here designated, deposited in MNHP; type-locality: "Mexico"; Gorham, 1882:142, placed this nominal species in

synonymy with Derestenus lateralis Chevrolat].

See "Remarks" for the justification of the new

synonymies.

DIAGNOSIS.—Although somewhat variable, genital

characteristics provide the most reliable clues for

identifying male members of this species (Figures

257-259). In most cases Mexican specimens belong

to this species if they possess the following combination of characteristics: body deep, pronotum

boldly transverse, elytral surface densely arenose,

from rugose, and cranium and pronotum impressed

with large setiferous punctations. Color patterns of

the elytron are also diagnostic in some individuals

(Figures 262-265), particularly females.

DESCRIPTION.—Form: As in Figures 260, 261.

Size: Length: males, average about 8.0 mm, range

6.8-10.0 mm; females, average about 8.6 mm, range

6.8-10.3 mm. Width: males, average about 2.3 mm,

range 2.0-3.2 mm; females, average about 2.7 mm,

range 2.4-3.3 mm. Ten males and 10 females measured.

Color: Cranium entirely flavotestaceous or entirely black, or clypeus, cranial venter, and postocular region of epicranium flavotestaceous and

remainder of cranium black; prothorax entirely

flavotestaceous except for midapical piceous macula, or predominantly piceous and pronotal disc

vittate; meso- and metasternum flavotestaceous, or

from dark brown to black; legs flavotestaceous or

black, or both; elytron variable (see "Variation");

abdomen from dark brown to black.

Head: As in P. reventazon except antenna (Figure 260) more serrate; epicranial setiferous punctations larger; frons often rugose; and length/width

ratio of each male antennal article 2.0:1.0:1.4:1.3:

1.3:1.2:1.2:1.1:1.1:1.1:1.9.

Pronotum: As in P. reventazon except more transverse in females, subapical depression more deeply

impressed, setiferous punctations larger, and side

margin of pronotum proper more boldly arcuate.

Elytron: As in P. reventazon except surface more

arenose.

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 257-259): Strongly pigmented. Combined length of phallobase and phallobasic apodeme about 5 times longer than paramere. Combined length of paramere and phallobasic

apodeme about 3.6 times longer than phallobasic

apodeme. Paramere: feebly arcuate in lateral view;

dorsum feebly convex, boldly braced basally; venter

concave; mesodorsal and mesoventral margin concave. Sinus: dorsal and ventral lanceolate; dorsal

slightly broader and longer than ventral. Phallus:

as in Figure 259 (38 specimens examined).

Female Genitalia: As in P. nigriventris.

Internal Reproductive Organs: Male; anterior accessory glands uniramous, coiled; posterior accessory glands biramous, divided into an inner and

outer branch (1 specimen examined).

VARIATION.—Structural: There is considerable

variation in the degree of concavity of the parameral mesodorsal margin, and in width of the

paramere and parameral dorsobasal brace. The elytral apex is sometimes acuminate at the suture as

is the case in all females from El Palmito. The

pronotum is generally more transverse in females

than in males.

Color: Elytra are mostly concolorous, usually

black, rarely testaceous. When bicolorous the elytron may have one of three color patterns, corresponding to the marginate phenon, the trivittate

phenon, or the subfaciate phenon.

In specimens of the marginate phenon the ely-
NUMBER 227 87

tron is broadly vittate at the posthumeral margin; elytral apex. In males, the same vitta may extend

the vitta extends from the humeral angle to the from the humeral angle to the elytral apex, ter261

262

FIGURES 260-265.—Perilypus carbonarius: 260, habitus, lateral view; 261, habitus, dorsal view;

262-265, elytras. (Scale = 1 mm)
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

minate at midelytron, or be faintly visible at the

elytral middle half. In one female specimen the

posthumeral vitta gradually extends towards the

elytral apex (Figure 265). I studied 5 males and 11

females of this phenon.

The elytral color pattern of trivittate specimens

is as described for the trivittate phenon of P. caliculus (see "Variation"). However, in specimens of

this species the discal vitta is less incurved at the

elytral basal fifth and is closer to the posthumeral

margin; in two male specimens the vittae are only

faintly visible at the elytral base. I studied 5 males

and 2 females of this phenon.

The elytra of subfasciate specimens are illustrated in Figures 262 and 263. The nonpiceous portion of the elytral disc ranges from stramineous to

rufotestaceous. I studied 1 male and 7 females

of this phenon.

NATURAL HISTORY.—Specimens were collected

from Mexico in June, July, and August. H. F. Howden collected 8 adult specimens, by beating, from

El Palmito at 1251 meters. J. Laue collected 1

adult from Volcan Colima at 1000 meters.

DISTRIBUTION (Figure 363).—From Sinaloa to

Oaxaca, Mexico.

LOCALITY RECORDS (Figure 363).—I examined 91 adult

specimens from Central America. MEXICO: Estado de

Sonora: Arroyo del Juchujaqui, Alamos (GEKI, 1 male);

Guirocoba (GEKI, 1 male). Estado de Sinaloa: 15 miles [24

km] west of El Palmito (CNCC, 4 males and 2 females;

GEKI, 1 male and 2 females); about 38 miles [61 km] east

of Villa Union (CNCC, 1 female; GEKI, 1 male and 1 female).

Estado de Durango: Ventanas (GEKI, 1 male). Estado de

Nayarit: El Pich6n (CISC, 1 female; GEKI, 1 male); Tepic

(MNHB, 1 female). Estado de Jalisco: Guadalahara (CMPP,

3 males and 1 female; GEKI, 1 male and 1 female). Estado de

Colima: Volcan Colima (GEKI, 1 male). Estado de Morelos:

Yautepec (MNHB, 1 male). Estado de Veracruz: Cerro de

Palmas (MNHB, 1 female); Playa Vicente (BMNH, 1 male

and 1 female). Estado de Mexico: Bejucos (FMNH, 1 male);

Temascaltepec (BMNH, 1 female). Estado de Guerrero:

Acapulco (WFBA, 1 female); 8 miles [13 km] north of

Iguala (GEKI, 1 male); 5 miles [8 km] south and 2.5 miles

[4 km] east of Chilpancingo (GEKI, 1 female). Estado de

Oaxaca: Jugila (BMNH, 5 males and 7 females; GEKI, 3

males and 3 females; MNHP, 5 females; MNHB, 1 male and

4 females); Panixtlahuaca (BMNH, 3 males and 2 females);

GEKI, 3 males); Oaxaca (MNHP, 2 males and 1 female).

"Mexico" (BMNH, 2 females; GEKI, 4 males and 3 females;

MNHP, 3 males and 5 females; MNHB, 1 female; MSPI, 1

male, holotype).

REMARKS.—Some specimens, here assigned to the

subfasciate phenon, were described as Derestenus

mutabilis Chevrolat or Colyphus flammeus Gorham,

while some of the marginate phenon were described as Derestenus lateralis Chevrolat or Colyphus marginatus Gorham. I believe the specimens

representing these phena are conspecific primarily

because their elytral color characteristics intergrade

to some extent as do characteristics of the male

genitalia.

As a group, specimens of P. carbonarius are

variable in color and in male genitalia, but the

variation is unequivalent to interspecific variations

within Perilypus. Perhaps P. carbonarius is undergoing more rapid differentiation than congeneric

species and perhaps the observed interpopulation

variations are manifestations of restricted gene flow

and possibly of incipient speciation. Alternatively,

P. carbonarius, as defined herein, may consist of

several biological species in which stable external

gaps have not yet materialized.

The nominal species placed in synonymy by me

(see above) were considered "good" species by other

authors, primarily on the basis of color characteristics. I found such characteristics unsuitable for

inferring reproductive isolation within Perilypus.

The chaletoides Group

The members of this group are characterized by

their stout body; small, shallow, and profusely distributed elytral punctations; very arenose elytral

surface; light coloration (predominantly castaneous

or stramineous); and by the unusually long male

posterior accessory glands.

The two species that comprise the group occur

in Mexico from Durango to Chiapas.

35. Perilypus chaletoides, new species

FIGURES 266-271, 274, 277, 278, 359

TYPE-LOCALITY.—Twenty-four kilometers north

of LaVentosa, at junction with highway 185, Estado

de Oaxaca, Mexico.

TYPE-SPECIMENS.—The holotype male is deposited in NMNH, the allotype in GHNE. Both specimens were collected by G. H. Nelson in 1965.

DIAGNOSIS.—The color pattern of the elytron as

described and figured (Figure 266), will distinguish

members of this species from congeneric species.

DESCRIPTION.—Form (Figure 266): The specimens
NUMBER 227 89

are broad and deep bodied, and conspicuously convex in lateral view.

Size: Length: males, average about 8.6 mm, range

7.6-9.9 mm; females, average about 9.7 mm, range

8.0-10.6 mm. Width: males, average about 2.7 mm,

range 2.4-3.2 mm; females, average about 3.2 mm,

range 2.6-3.6 mm. Nine males and 6 females measured.

Color: Cranium castaneous, with or without midepicranial black macula; antenna black; prothorax

castaneous, except lower sides of pronotum and pronotal discal vitta black; mesoscutellum castaneous,

black, or castaneous but infuscated distally; mesoand metasternum entirely black, or middle region

castaneous and lateral regions black; femur variable (see "Variation"); tibia and tarsus black; elytron narrowly black along sutural margin, black

at apical fifth, castaneous in remainder; abdomen

black.

Head: Interocular depressions shallow; frontal

umbo prominent; eyes moderately convex; HW/

IOW, average about 2.3, range 2.0-2.3 (15 specimens measured); antenna (Figure 274) short; AL/

PLS, 1.4, boldly serrate; length/width of each antennal article sex dimorphic, male 2.7:1.4:1.6:1.5:

1.2:1.2:1.2:1.0:0.88:0.72:1.6, female 1.8:1.2:1.3:1.1:

1.1:1.0:1.0:0.89:0.72:0.67:1.5.

Pronotum: Boldly convex and only slightly transverse (PL/PW, average about 0.96, range 0.92-1.0;

15 specimens measured); subapical and prebasal depressions deeply impressed; pronotal arch coarsely

punctate; side margins of pronotum proper boldly

convex.

Mesoscutellum: Subquadrate.

Elytron: Conspicuously convex and with exceptional depth; epipleural margin straight in basal

half, arcuate in apical half; punctations very small

and profusely distributed; punctate and interpunctate surfaces conspicuously arenose; setal densely

distributed; EL/EW, average about 4.2, range 3.3-

4.7 (15 specimens measured); slope of apical deflexion gradual.

Front Tarsal Claws: Male and female feebly

asymmetrical.

Male Genitalia (Figures 269-271): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.4 times longer than paramere. Combined length of paramere and phallobase 3.0 times

longer than phallobasic apodeme. Paramere: apical

fourth abruptly narrowed in dorsal or ventral view,

feebly declinate in lateral view; lateral depression

feebly impressed; dorsum plane, with narrow brace

at basal third (Figure 271); venter feebly concave;

mesoventral margin faintly arcuate; mesodorsal

margin sinuous in basal half, linear in apical half.

Sinus: dorsal and ventral lanceolate. Phallus: acuminate; marginal denticles on middle half of phallic

plate (12 specimens examined).

Female Genitalia (Figures 267, 268): Dorsal lamina bilobed, incision short; ventral lamina trilobed,

lateral lobes shorter than medial lobe; dorsal bacculi not fused; ventral bacculus acuminate near

middle; coxital plates absent (2 specimens examined).

Internal Reproductive Organs: Male (Figures

277, 278); anterior accessory glands uniramous,

coiled throughout; posterior accessory glands biramous, outer branch about 1.5 longer than inner

branch, both branches exceptionally long and extensively convoluted; testis composed of 12 follicles. Female; spermathecal capsule bulbous distally

and spermathecal gland duct exceptionally long (2

males and 2 females examined).

VARIATION.—Structural: The front tarsal claws

are particularly asymmetrical in the male specimen

from El Camer6n, Oaxaca, Mexico. The interocular depressions and disc of the pronotal arch are

more coarsely punctated in male specimens than

in female specimens. Also correlated with sex is

the degree of concavity of the epipleural margin

(best seen in the elytral lateral view), which is

conspicuously more concave in males than in females.

Color: The pronotal vitta may be percurrent, interrupted at the middle, or reduced to a midapical

macula. In three male specimens the dark regions

of the elytron are confined to an elongate sutural

infuscation. The elytron of some specimens is predominantly flavocastaneous.

NATURAL HISTORY.—In August in Chiapas, Mexico, C. W. O'Brien collected three specimens, by

beating, at 885 meters. G. H. Nelson collected

specimens in July by sweeping roadside vegetation

in Oaxaca, Mexico. One specimen was collected

at light in July by Mastro and Schaffner. Other

specimens were collected from Mexico in June.

DISTRIBUTION (Figure 359).—Known only from

southwestern Mexico.
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

266

273

FIGURES 266-274.—Perilypus chaletoides: 266, habitus; 267, ovipositor, ventral view; 268, ovipositor, dorsal view; 269, tegmen, lateral view; 270, tegmen, ventral view; 271, tegmen, dorsal

view; 274, antenna. P. bicolor: 272, tegmen, lateral view; 273, tegmen, ventral view. (Scale =

1 mm)
NUMBER 227 91

ETYMOLOGY.—Latin compound adjective formed

from Chaletus (a chrysomelid generic name) + the

suffix -oides (like). I refer to the similarity in the

elytral color pattern between members of this species and those of Chaletus.

LOCALITY RECORDS (Figure 359).—I examined 34 adult specimens from Central America. MEXICO: Estado de Oaxaca:

9 miles [15 km] east of El Cameron (TTUT, 1 male); 15

miles [24 km] north of La Ventosa, at junction with highway

185 (NMNH, 1 male, holotype; GEKI, 1 male and 1 female;

GHNE, 2 females); 8 miles [13 km] north of La Ventosa

(TAMT, 1 female); 14 miles [23 km] south of Matias Romero

(GEKI, 1 female; TAMT, 2 females); Almoloya (NMNH, 1

male). Isthmus of Tehuantepec (MCZC, 1 female). Estado de

Chiapas: 4 miles [7 km] northwest of Pueblo Nuevo, Rio

Bajada (GHNE, 1 male); At junction between highways 190

and 195 (HFHO, 1 male); Tuxtla Gutierrez (CNCC, 1 female); Suchiapa (CISC, 1 male); Los Amates (AMNH, 1

male); 21 miles [34 km] west of Lazaro Cardenas (CNCC, 1

lemale); 2.3 miles [4 km] west of Las Cruces (GEKI, 2

males); 31 miles [50 km] southeast of Comitan (GEKI, 1

male and 1 female; TAMT, 3 males and 1 female); 7 miles [11

km] southeast of Chiapa de Corzo (GEKI, 1 male); 19 miles [31

km] west of Ocozocoautla (TAMT, 1 female); 7 miles [11 km]

southwest of Ocozocoautla (GEKI, 1 male and 1 female); 13

miles [21 km] northeast Cintalapa (GEKI, 2 males and 1

lemale); 2 miles [3 km] east Rizo de Oro (GEKI. 1 male).

36. Perilypus bicolor (Chevrolat),

new combination

FIGURES 272, 273, 275, 359

Sallea bicolor Chevrolat, 1874:287 [lectotype, female, deposited

in MNHP, here designated; type-locality: Veracruz, Estado

de Veracruz, Mexico].—Gorham, 1882:144.—Schenkling,

1898:364.

DIAGNOSIS.—The dorsal fascies of these specimens

is entirely testaceous.

DESCRIPTION.—Form: As in P. chaletoides (Figure 266).

Size: Length: males, average about 8.2 mm, range

7.1-8.9 mm; females, average about 8.6 mm, range

7.3-10.2 mm. Width: males, average about 2.7 mm,

range 2.4-3.0 mm; females, average about 2.8 mm,

range 2.3-3.2 mm. Six males and 10 females

measured.

Color: Cranium, prothorax, and elytron testaceous; antenna, meso- and metasternum, tibia, tarsus, and abdomen dark brown; femora variable (see

"Variation").

Head: As in P. chaletoides except antenna (Figure 275) slightly less serrate, and length/width ratio of each male antennal article 2.3:1.2:1.3:1.3:1.5:

1.3:1.1:1.0:1.0:1.0:1.5.

Pronotum: As in P. chaletoides.

Elytron: As in P. chaletoides.

Front Tarsal Claws: As in P. chaletoides.

Male Genitalia (Figures 272, 273): Strongly pigmented. Combined length of phallobase and phallobasic apodeme 5.7 times longer than paramere.

Combined length of paramere and phallobase 3.2

times longer than phallobasic apodeme. Paramere:

acuminate, abruptly narrowed at apical third; lateral depression narrow, deep, and projecting beyond

basal limit of ventral sinus; dorsum plane; venter

feebly convex; mesoventral margin concave; mesodorsal margin feebly convex in basal third, linear in

apical two-thirds. Sinus: dorsal and ventral lanceolate, dorsal slightly longer than ventral. Phallus:

phallic plates broad in basal three-fourths, gradually narrowed to apex in apical fourth; marginal

denticles small, extended from basal fourth to

apical three-fourths of phallic plate (7 specimens

examined).

Female Genitalia: As in P. chaletoides.

VARIATION.—Structural: No appreciable structural variation was noted.

Color: The dorsal fascies is concolorous in the

specimens studied, but femoral coloration is variable and not correlated geographically or with regard to sex. In four specimens, the front and middle femora are entirely testaceous, the hindfemur

is predominantly testaceous and infuscated apically.

In 6 specimens, the front and middle femur is

testaceous and infuscated apically, the hind femur

entirely dark brown; in 14 specimens the front femur is testaceous and infuscated apically, the middle and hind femur entirely dark brown.

Additional color variation involves the mesosternum which is lighter, as dark, or paler than the

metasternum. Also, in one female specimen the

epipleural fold is faintly infuscated at the midelytron.

NATURAL HISTORY.—Specimens were collected in

southern Mexico from June through August.

DISTRIBUTION (Figure 359).—This Mexican species

ranges from Durango to Oaxaca.

LOCALITY RECORDS (Figure 359).—I examined 24 adult

specimens from Central America. MEXICO: Estado de

Veracruz: Veracruz (MNHP, 1 female, lectotype); Cotaxtla

Experiment Station, Cotaxtla (CISC, I female; GEKI, 1 male);

Puente Nacional (CISC, 1 male); Motzorongo (MNHB, 1
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

female). Estado de Durango: Canelas (GEKI, 1 male). Estado

de Guerrero: Taxco (GEKI, 1 female); Almolonga (GEKI, 1

male). Estado de Oaxaca: Oaxaca (BMNH, 1 female; GEKI,

1 male and 1 female; MNHP. 1 female). "Mexico" (FMNH, 1

male and 1 female; GEKI, 1 male; MNHB, 1 male and 2

females; RNHN, 1 male; UZMD, 1 female; ZMAN, 2 females).

The quadrilineatus Group

The most striking characteristic of the species

included herein is their lampyroid body. Members

of this group are also characterized by antenna

boldly serrate; elytron with arenose surface, small

punctations, very gradual apical slope, and deflexed

posthumeral region deeply concave; and mesodorsal margin of paramere denticulated.

The quadrilineatus group exhibits more interspecific structural variation than the other species

groups do. Perilypus bicristatus, for example, has

some particularly discordant characteristics including, elytron bicarinate, mesoscutellum triangular,

and pronotum concave medially and paralaterally.

The combined distribution of the five species

included in this group extends from northeastern

Mexico to central Guatemala. However, only P.

quadrilineatus extends north of the Isthmus of

Tehuantepec where it occurs primarily along the

Atlantic drainage systems.

37. Perilypus quadrilineatus (Chevrolat),

new combination

FIGURES 276, 279-283, 285, 286, 366

Derestenus quadrilineatus Chevrolat, 1843:14 [holotype: female, deposited in MNHP; type-locality: Veracruz, Estado

de Veracruz, Mexico].—Lacordaire, 1857:443.—Gorham,

1882:141.

Derestenus similis Thomson, 1860:57 [lectotype: male, here

designated, deposited in MNHP; type-locality: Mexico].

DIAGNOSIS.—The bold, obliquely convex epipleural fold and the narrow vitta of the elytral disc

(Figure 279) easily separate specimens of this species

from congeners within the quadrilineatus group.

DESCRIPTION.—Form: As in Figure 279.

Size: Length: males, average about 10.1 mm,

range 9.0-10.9 mm; females, average about 10.6

mm, range 8.2-13.2 mm. Width: males, average

about 3.5 mm, range 3.0-4.4 mm; females, average

about 3.5 mm, range 2.6-4.6 mm. Ten males and

10 females measured.

Color: Clypeus, frons, and postocular region of

epicranium flavotestaceous; remainder of cranium

dark brown; pronotal sides flavotestaceous, sometimes roseous; pronotal disc with piceous vitta;

thoracic sterna brown; legs variable (see "Variation"); elytron predominantly dark brown, with

marginal and discal flavotestaceous vittae; abdominal sternum variable (see "Variation").

Head: Interocular depression broad, shallow, not

crescentic; eyes boldly convex (HW/IOW, average

about 2.2, range 2.0-2.4; 10 males and 10 females

measured); antenna (Figure 276) serrate, not particularly flattened, length/width ratio of each male

antennal article 1.7:1.4:1.5:1.7:1.4:1.5:1.4:1.4:1.4:

1.4:2.3; AL/PLS, 1.7.

Pronotum: Boldly transverse (PL/PW, average

about 0.90, range 0.82-0.96; 20 specimens measured); sides of pronotum proper boldly arcuate;

dorsum feebly convex; pronotal arch coarsely punctate; subapical depression well impressed at sides,

shallow at middle; pronotal foveae and prebasal

depression deeply impressed; pronotal collar normal, not particularly narrow.

Mesoscutellum: Subquadrate.

Elytron: Posthumeral margin arcuate in outline

and contour; narrow longitudinal region proximal

to epipleural fold gradually deflected, deflected

region concave; punctations small, shallow, and

evenly distributed throughout elytral disc although

diminished near suture; surface arenose and profusely vested with short reclinate setae; epipleural

fold obliquely convex; epipleuron convex, explanate internally; apical slope very gradual; EL/EW,

average about 4.4 range 3.2-4.9 (20 specimens

measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 281-283): Strongly

sclerotized. Combined length of phallobase and

phallobasic apodeme 5 times longer than paramere.

Combined length of paramere and phallobase 2.3

times longer than phallobasic apodeme. Paramere:

outer margin acutely inflected at apical third; apex

incurved; lateral depression elongate and deeply

impressed; dorsum convex, strongly explanate

medially; venter narrow and concave; mesoventral

margin concave; mesodorsal margin sinuous and

denticulated at middle third. Sinus: dorsal narrow

in apical half, expanded in basal half; ventral elliptical. Phallus: phallic plates broad, feebly sclerotized subapically; marginal denticles extended from
NUMBER 227

93

283

FIGURES 275-283.—Perilypus bicolor: 275, antenna. P. chaletoides: 277, posterior accessory gland;

278, male internal reproductive organs. P. quadrilineatus: 276, antenna; 279, habitus; 280,

male internal reproductive organs; 281, tegmen, lateral view; 282, tegmen, ventral view; 283,

phallus. (Scale = 1 mm)
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

middle of phallic plate to phallic plicae (17 specimens examined).

Female Genitalia (Figure 285, 286): Dorsal lamina

bilobed; ventral lamina trilobed; proctiger short,

acute; proctigeral bacculus acuminate at posterior

fifth; coxital plates absent (2 specimens examined).

Internal Reproductive Organs: Male (Figure 280);

anterior accessory gland uniramous, coiled throughout; posterior accessory gland biramous, outer

branch nearly twice as long as inner branch (1

specimen studied).

VARIATION.—Structural: The angularity of the

convex epipleural fold is more pronounced in some

specimens; in others it is the extent of its inner

and outer explanation that is striking. In general,

female antennae are more serrate than male antennae.

The geographically disjunct female specimen

from Volcan Colima, Mexico, differs from all other

specimens studied as follows: pronotum more transverse (PL/PW, 0.81), with subapical depression

more deeply impressed, and with arch narrower and

more convex; elytral form more rectangulate elytral

disc avittate; and legs entirely dark brown. Whether

these differences can be inferred to represent genetic discontinuity or geographic variation will depend on study of additional material from Colima

and its environs. Should this specimen (which I

labeled P. quadrilineatus (Chevrolat)) be judged

distinct from P. quadrilineatus it would undoubtedly be a member of the sister of that species.

Color: The flavous femoral annulus varies in

width, its definition is obliterated when the femur

is predominantly flavous. Coloration of the abdominal venter varies with sex and is dark brown in

males. In females, visible sterna I, n, and vi are

brown, sterna in to v are flavotestaceous. Sometimes

the female sternum m is partially flavotestaceous

and partially brown.

NATURAL HISTORY.—Specimens were collected

in Mexico in June by G. H. Nelson (by beating

roadside vegetation in Oaxaca) and by H. S. Dybas

from Veracruz at 920 meters. A female specimen

was intercepted in New Orleans (U.S.A.) on banana

debris apparently transported from Mexico. Other

adult specimens were collected in Mexico in May,

June, and August.

DISTRIBUTION (Figure 366).—The center of distribution of this Mexican species appears to be the

Isthmus of Tehuantepec: most of the specimens

studied were collected from the Atlantic side of the

Isthmus. Two specimens were collected from northern Guatemala.

LOCALITY RECORDS (Figure 366).—I examined 68 adult

specimens from Central America. MEXICO: Estado de

Tamaulipas: 5 miles [8 km] west of Gomez Farias (GEKI, 1

female). Estado de San Luis Potosi: El Salto de Agua (GEKI,

1 female; 25 miles [40 km] north of Tamazunchale (CNCC,

2 females; GEKI, 2 females). Estado de Puebla: 15 miles

[24 km] east of Teziutlan (CNCC, 1 male and 1 female;

GEKI, 1 female). Estado de Colima: Volcan Colima (GEKI,

I female). Estado de Veracruz: Jalapa (GEKI, 1 female);

Fortin (FMNH, 2 males and 1 female); Los Amates (CNCC, 1

female); Tezonapa (FMNH, 1 female); Veracruz (BMNH, 1

female; FMNH, 1 male; GEKI, 1 male; MNHP, 1 female,

holotype); Cordoba (CASC, 3 males; GEKI, 1 male and 1

female; MNHP, 1 female); San Rafael, Jicaltepec (MCZC, 2

females; GEKI, 1 female); Cotaxtla Experiment Station,

Cotaxtla (CISC, 1 male and 1 female); Puente Nacional

(GEKI, 1 female); Lago de Catemaco (CISC, 1 female; NMNH,

1 female); 2 miles [3 km] north of Santiago Tuxtla (GEKI,

1 female); Cerro de Palmas (BMNH, 2 females; GEKI, 2

females); Santiago Tuxtla (BMNH, 2 females); Playa Vicente

(BMNH, 2 males); Cosamaloapan (BMNH, 1 female); Atoyac

(MNHB, 1 female). Estado de Oaxaca: Oaxaca (GEKI, 1

female); Temescal (GEKI, 1 female; GHNE, 2 males). Estado

de Chiapas: Portugal, 7 miles [11 km] southeast of Simojovel

(GEKI, 1 female); 8 miles [13 km] south of Simojovel (GEKI,

1 female); Santo Domingo, 15 miles [24 km] southeast of

Simojovel (CISC, 1 female); Simojovel (CISC, 1 female); 10

miles [16 km] south of Malpaso (GEKI, 1 female). "Mexico"

(BMNH, 1 female; GEKI, 3 females; MNHP, 2 males and

1 female; MNHB, 4 females; MZST, 1 female; NMNH, 2

females; ZMAN, 1 female). GUATEMALA: Departamento

de Alta Verapaz: Teleman (BMNH, I female).

38. Perilypus telephoroides (Gorham),

new combination

FIGURES 284, 287-291, 366

Colyphtis telephoroides Gorham, 1882:142 [lectotype; male,

and two paralectotypes, male and female, here designated,

deposited in BMNH; type-locality: Capetillo, Departamento

de Sacatepequez, Guatemala].—Gorham, 1886:335.—Schenkling, 1906:262.

DIAGNOSIS.—The short testaceous vitta on the

elytral humeral angle (Figure 287) identifies specimens of this species.

DESCRIPTION.—Form: As in Figure 287.

Size: Length: males, average about 8.6 mm, range

6.2-10.4 mm; females, average about 9.4 mm, range

7.9-10.1 mm. Width: males, average about 2.9 mm,

range 1.9-3.6 mm; females average about 3.2, range

2.6-3.6 mm. Eight males and 9 females measured.
NUMBER 227

95

FIGURES 284-291.—Perilypus telephoroides: 284, antenna; 287, habitus; 288, ovipositor, ventral

view; 289, ovipositor, dorsal view; 290, tegmen, lateral view; 291, tegmen, ventral view. P.

quadrilineatus: 285, ovipositor, ventral view, 286, ovipositor, dorsal view. (Scale = 1 mm)
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Color: Clypeus and postocular region of epicranium testaceous, remainder of cranium black; antenna black; pronotum testaceous dorsolaterally,

black ventrolaterally and discally; pro-, meso-, and

metasternum black; legs black; elytron black except

with short testaceous vitta at basal third of posthumeral margin (Figure 287); abdominal venter

color variable (see "Variation").

Head: Interocular depressions broad and well

impressed; interocular surface coarsely rugose; epicranium densely punctate; eyes boldly convex;

HW/1OW, average about 2.4, range 2.1-2.7 (20

specimens measured); antenna (Figure 284) conspicuously serrate and sex dimorphic with respect

to length (AL/PLS, male 2.0, female 2.3); length/

width ratio of each male antennal article 2.5:1.0:

1.1:1.4:1.2:1.1:1.1:1.1:1.0:1.0:1.8.

Pronotum: Distinctly transverse and feebly trapezoidal; subapical depression shallow; pronotal arch

feebly convex and coarsely punctate; disc of

pronotum proper transversely wrinkled; PL/PW,

average about 0.83, range 0.79-0.90 (17 specimens

measured).

Elytron: Posthumeral margin straight in basal

two-thirds, arcuate in apical third; narrow longitudinal region proximal to epipleural fold acutely

deflexed; concavity of deflexed posthumeral region

elongate and deeply impressed; apical slope very

gradual; surface copiously impressed with small

shallow punctations; punctate and interpunctate

surfaces arenose and profusely vested with short

reclinate setae; epipleural fold unusually narrow;

EL/EW, average about 4.1, range 3.9-4.9 (17 specimens measured).

Front Tar sal Claws: Male, feebly asymmetrical;

female, symmetrical.

Male Genitalia (Figures 290, 291): Strongly pigmented. Combined length of phallobase and

phallobasic apodeme 5 times longer than paramere.

Combined length of paramere and phallobase 3.4

times longer than phallobasic apodeme. Paramere:

apex obtuse; outer margin depressed at middle;

lateral depression deep, extended from paramere

base to paramere apical fourth; dorsum convex,

strongly explanate medially; venter narrow and

concave; mesoventral margin concave in basal twothirds, sinuous in apical third; mesodorsal margin

sinuous, denticulated in apical third only, and with

conspicuous brace at base. Sinus: dorsal narrowly

lanceolate; ventral elliptical and feebly constricted

apically. Phallus: as in P. quadrilineatus (10

specimens examined).

Female Genitalia (Figures 288, 289): As in P.

quadrilineatus except dorsal lamina and its incision

broader and proctiger more robust (1 specimen

examined).

Internal Reproductive Organs: Male, anterior

accessory gland uniramous, coiled throughout.

VARIATION.—Structural: Notable structural variation is evident in the length and degree of

antennal serration. Male antennae are proportionally longer and less serrate than female antennae.

Color: The size of the postocular epicranial

macula varies as does the length of the testaceous

vitta on the elytral humeral angle; in one specimen

the vitta is broadened and extends from the humeral

angle to the elytral apex. Coloration of abdominal

venter is sex dimorphic being black in males; in

females, visible sterna i-m are black, sterna IV-VI

testaceous.

NATURAL HISTORY.—In October in the interior

valley of Sierra de las Minas, Guatemala, R. D.

Mitchell collected one adult specimen at 1550

meters. Other adult specimens were collected from

Mexico in May and June.

DISTRIBUTION (Figure 366).—The known range

extends from southern Mexico to southern

Guatemala.

LOCALITY RECORDS (Figure 366).—I examined 20 adult

specimens from Central America. MEXICO: Estado de

Chiapas: Chilcultic, near El Rinc6n (CNCC, 1 male; GEKI,

1 male); Lagos des Calores, route 17 (HFHO, 1 male); El

Rincrtn, route 7 (GEKI, 2 females); 17 miles [27 km] southeast of Teopisca, route 24 (CNCC, 1 female). "Mexico"

(MNHP, 1 male, ZMAN, 1 female). GUATEMALA: Departamento de Sacatepequez: Capetillo (BMNH, 1 male,

lectotype, and 2 females; GEKI, 1 male and 1 female; MNHP,

1 female). Departamento de Zacapa: Santa Clara, north of

Cabanas (FMNH. 1 female).

39. Perilypus coroniformis, new species

FICURES 292-296. S66

TYPE-LOCALITY.—Eight kilometers west of San

Cristobal de las Casas, Estado de Chiapas, Mexico.

TYPE-SPECIMENS.—The holotype male and the

allotype are deposited in HFHO. These and 4

paratypes were collected by H. F. Howden in 1969.

Four paratypes: GEKI, 1 male and 1 female;

HFHO, 2 females.
NUMBER 227 97

FIGURES 292-297.—Perilypus coroniformis: 292, antenna; 293, habitus; 294, tegmen, lateral view;

295, tegmen, ventral view; 296, posterior accessory glands. P. fluctns: 297, habitus. (Scales =

1 mm)
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—The configuration of the pronotal

discal vitta, as illustrated (Figure 293) and described

is unique in Perilypus.

DESCRIPTION.—Form: As in Figure 293.

Size: Length: males, average about 8.1 mm, range

8.5-10.7 mm; females, 10.0 mm. Width: males,

average about 3.8 mm, range 3.6-4.0 mm; females,

3.6 mm. Two males and 2 females measured.

Color: Clypeus testaceous, remainder of cranium,

and antenna black; prosternum and pronotal sides

roseous; pronotal disc broadly vittate, vitta with

oblong roseous macula at center; meso- and

metathorax, legs, elytron, and abdomen black,

except femora testaceous ventroapically.

Head: As in P. teleph.oroid.es except HW/IOW,

average about 2.4, range 2.3-2.5 (4 specimens

measured); antenna (Figure 292) boldly serrate;

AL/PLS, sex dimorphic, male 2.7, female 2.3; and

length/width ratio of each antennal article sex

dimorphic, male 2.3:1.4:1.2:1.4:1.4:1.3:1.4:1.4:1.4:

1.4:1.9, female 2.4:1.0:1.2:1.3:1.2:1.2:1.1:1.0:1.0:1.0:

1.5.

Pronotum: As in P. telephoroides except more

transverse (PL/PW, average about 0.80, range

0.77-0.84; 4 specimens measured), more trapezoidal,

and disc smooth.

Elytron: As in P. telephoroides except posthumeral margin arcuate, disc undulated longitudinally, and EL/EW, average about 4.0, range

3.8-4.2 (4 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 294, 295): Strongly pigmen ted. Combined length of phallobase and

phallobasic apodeme 5 times longer than paramere.

Combined length of paramere and phallobase 3

times longer than phallobasic apodeme. Paramere:

outer margin broadly arcuate; apex subacute;

lateral depression well impressed, extended from

parameral base to parameral apical two-thirds;

dorsum convex, strongly explanate mesally; venter

concave; mesoventral margin concave; mesodorsal

margin convex and finely denticulated at middle

third. Sinus: dorsal indistinct; ventral elliptical.

Phallus: as in P. telephoroides (2 males examined).

Female Genitalia: As in P. telephoroides.

Internal Reproductive Organs: Male; as in Figure

296 (1 specimen examined).

VARIATION.—Structural: Antennal length varies

with sex; the male antenna is proportionally longer

than the female antenna. In general, the antennal

articles are more transverse in females.

Color: There is no conspicuous color variation

among the 4 specimens examined.

NATURAL HISTORY.—During June near San

Crist6bal de las Casas, Mexico, H. F. Howden

collected 6 adult specimens by beating oak (Quercus

sp.) foliage.

DISTRIBUTION (Figure 366).—Known only from the

type-locality.

ETYMOLOGY.—Latin noun corona (crown) +

noun forma (shape), and adjectival ending -is. The

name refers to the peculiar shape of the pronotal

discal vitta.

LOCALITY RECORDS (Figure 366).—I examined 6 adult specimens from Central America. MEXICO: Estado de Chiapas: 5

miles [8 km] west of San Cristobal de las Casas (GEKI, 1 male

and 1 female; HFHO. 1 male, holotype, and 3 females).

40. Perilypus fiuctus, new species

FIGURES 297, 301-303, 366

TYPE-LOCALITY.—Sinanja, Departamento de Baja

Verapaz, Guatemala.

TYPE-SPECIMENS.—The holotype male and allotype, deposited in BMNH, were collected by G. C.

Champion.

DIAGNOSIS.—Distinguishable from P. coroniformis, the only other species in the quadrilineatus

group with elytral surface undulated longitudinally,

by the testaceous annulation of the femora and by

the sagittate pronotal vitta.

DESCRIPTION.—Form: As in Figure 297.

Size: Length: male, 8.7 mm; female, 9.0 mm.

Width: male, 3.2 mm; female 3.5 mm. One male and

1 female measured.

Color: As in P. telephoroides, except posthumeral

margin avittate, pronotal vitta strongly constricted

at subapical depression, and femora annulated.

Head: As in P. telephoroides.

Pronotum: As in P. telephoroides, except not

transversely wrinkled, margins of pronotum proper

boldly arcuate, and fovae more spheroid.

Elytron: As in P. telephoroides except posthumeral margin arcuate from base to apex, surface

distinctly undulated longitudinally, and EL/EW

3.9 (2 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 301-303): Strongly pig-
NUMBER 227

99

mented. Combined length of phallobase and phallobasic apodeme 9 times longer than paramere.

Combined length of paramere and phallobase 3.2

times longer than phallobasic apodeme. Paramere:

apical region broadly uncinate in lateral view;

lateral depression shallow and localized at middle

third; dorsum boldly explanate in apical half, feebly

explanate in basal half; venter narrowly concave;

mesoventral margin sinuous; mesodorsal margin

convex and coarsely denticulated in middle half.

Sinus: dorsal indistinct; ventral broadly lanceolate.

Phallus: as in Figure 303.

Female Genitalia: Not studied.

VARIATION.—Structural: The two specimens are

structurally homogeneous.

Color: The male abdominal venter is black; in

female specimens visible sterna i-m are black and

IV-VI testaceous.

DISTRIBUTION (Figure 366).—Known only from

Baja Verapaz, Guatemala.

ETYMOLOGY.—Latin noun fluctus (wave), referring to the longitudinal undulation of the elytral

disc.

LOCALITY RECORDS (Figure 366).—I examined 2 adult specimens from Central America. GUATEMALA: Departamento

de Baja Verapaz: Sinanja (BMNH, 1 male, holotype); San

Jeronimo (BMNH, 1 female).

REMARKS.—The holotype was part of the syntype

series of P. telephoroides and can be easily distinguished from members of that species by the more

convex margins of the pronotum and the elytron,

undulation of the elytral surface, testaceous annulus

of femur, and by the absence of an elytral humeral

vitta.

The allotype was part of the syntype series of

JP. ventralis. While there is considerable habitus

similarity between female members of the two

species, orientation of the epipleural fold easily

separates them. In specimens of P. ventralis the

epipleural fold is in the ventral position while in

specimens of the quadrilineatus group the fold is

lateral.

41. Perilypus bicristatus, new species

FIGURES 298-300, 366

TYPE-LOCALITY.—Twenty-four kilometers west

of El Palmito, Estado de Sinaloa, Mexico.

TYPE-SPECIMENS.—The holotype female, deposited in CNCC, was collected by H. F. Howden in

1964.

DIAGNOSIS.—Elytron bicristate (Figure 300), pronotum concave medially and paralaterally and

with paramedial longitudinal elevations, mesoscutellum triangular, and frontal umbo particularly

prominent (Figure 299). The frontal umbo is the

only region of body not black.

DESCRIPTION.—Form: As in Figure 300.

Size: Length 9.4 mm; width 2.3 mm.

Color: Frontal umbo flavous, remainder of body

black.

Head: Cranium coarsely punctate; interocular

depressions deeply impressed and rugose posteriorly;

frontal umbo prominent, conspicuously convex,

and narrowly prolonged posteriorly; antenna

(Figure 298) boldly serrate and very flat, article 1

and 11 oblong, 2 to 9 transverse, 10 as long as

broad; length/width ratio of each antennal article

1.2:0.8:0.9:0.8:0.7:0.8:0.9:0.9:0.8:1.0:1.9; AL/PLS,

2.4.

Pronotum: Transverse (PL/PW, 0.83), coarsely

punctate; surface concave centrally and paralaterally, and with paramedial longitudinal elevations; subapical depression deeply impressed at

side margins, feebly impressed discally.

Mesoscutellum: Triangular.

Elytron: Posthumeral margin arcuate; narrow

longitudinal region proximal to epipleural fold

acutely deflexed; concavity of deflexed posthumeral

region long and deeply impressed; apical slope very

gradual; surface with small coarse punctations,

prominently arenose; disc bicarinate (Figure 300);

epipleural fold conspicuously narrow; EL/EW, 3.7.

Front Tarsal Claws: Symmetrical.

NATURAL HISTORY.—The holotype was collected

from El Palmito, Mexico in July.

ETYMOLOGY.—Latin compound noun formed

from the prefix bi- (two) + noun crista (ridge),

referring to the two conspicuous longitudinal

ridges on the elytral disc.

LOCALITY RECORDS (Figure 366).—I examined 1 adult specimen from Central America. MEXICO: Estado de Sinaloa:

15 miles [24 km] west of El Palmito (CNCC, 1 female, holotype).

The ornaticollis Group

The species included herein are substantially

diverse anatomically, but as a group are charac-
100

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

299

302

FIGURES 298-303.—Perilypus bicristatus: 298, antenna; 299, head; 300, habitus. P. fluctus: 301,

tegmen, lateral view; 302, tegmen; ventral view; 303, phallus. (Scale = 1 mm)
NUMBER 227 101

FIGURES 304-307.—Perilypus ventralis: 304, antenna; 305, habitus; 306, elytron, X 16;

307, elytron, X 65 (arrow indicates microsculpture). (Scale = 1 mm)
102

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terized by mesoscutellum transverse, body form

moderately to strongly oval, interocular depressions

crescentic, clytral sutural notch right angle, elytral

punctations small, and elytral surface usually conspicuously arenose. Male genital characteristics

include, combined length of paramere and phallobase less than twice as long as phallobasic apodeme,

phallobase marked with two dorsal and two ventral

longitudinal bands, and first connecting membrane

vested with serrulated scales.

Perilypus ventralis is a particularly aberrant

member in the group. Only specimens of that

species have the mesoscutellum stalked, postscutellar

disc concave, epipleural fold in ventral position,

elytron undulated longitudinally, and anterior

accessory gland of male reproductive organs not

completely coiled. In general, there is more habitus

concordance between specimens of P. ventralis and

lampyriform members of the quadrilineatus group

than with the other members of the ornaticollis

group. The similarity, however, is due to convergence and is not an expression of immediate common ancestry.

The composite distribution of the eight species

in this group extends from Ohio, U.S.A., to

Distrito Puerto Caballo, Venezuela.

42. Perilypus ventralis (Gorham), new combination

FIC.URKS 304-319, 368

Colyphus ventralis Gorham, 1882:141 [lectotype: female, and

3 paralectotypes, 2 males and 1 female, here designated,

deposited in BMNH; type-locality: Duenas, Departamento

de Sacatepequez, Guatemala].

DIAGNOSIS.—In addition to the characteristics

given in the key, specimens of this species are easily

diagnosed by their large size (average length about

13 mm), concave mesoscutellar disc (Figure 312),

and by the narrow and longitudinal pronotal

foveae.

DESCRIPTIONS.—Form: As in Figure 305.

Size: Length: males, average about 12.9 mm,

range 10.4-14.1 mm; females, average about 13.0

mm, range 10.0-14.4 mm. Width: males, average

4.6 mm, range 4.0-5.0 mm; females, average about

4.4 mm, range 3.6-5.2 mm. Eight males and 10

females measured.

Color: Head including antenna black; pronotum

flavous at sides, with black triangular discal vitta

(Figure 305); prosternum black; legs black except

femur with testaceous annulus (Figure 56); mesoand metasternum black; elytron color variable (see

"Variation"); abdomen bicolorous, visible sterna

i-m and vi black, iv-v flavous.

Head: Gular process (Figure 308) filiform; interocular depressions broad, shallow, conspicuously

rugose and only feebly crescentic; frontal umbo

prominent, narrowed from clypeus to epicranium;

antennal carina coarsely punctate; frons broad;

eyes not particularly convex; HW/IOW, average

about 2.2, range 2.0-2.3 (18 specimens measured);

antenna (Figure 304) serrate; length/width ratio of

antennal articles sex dimorphic, male 2.2:1.5:1.5:

1.7:1.3:1.2:1.2:1.2:1.2:1.0:1.9, female 2.3:1.5:1.7:2.0:

1.9:1.8:1.6:1.5:1.3:1.3:2.0; AL/PLS, 2.2.

Pronotum (Figures 305): Transverse (PL/PW,

average about 0.88, range 0.84-0.93; 18 specimens

measured); densely setose; pronotal arch poorly

denned; subapical depression feebly impressed at

sides, nearly obliterated medially; pronotal foveae

longitudinal and transversely rugose; prebasal

depression shallow; pronotal collar narrow.

Mesoscutellum (Figures 310, 312): Transverse,

stalked, and concave discally.

Elytron: Shallow, undulated longitudinally and

copiously vested with short setae; posthumeral

margin arcuate; slope of apical deflection very

gradual; punctations small and profusely distributed

throughout surface (Figure 306); punctate and

interpunctate surface arenose (Fig. 307); epipleural

fold broad, concave, and deflected internally;

epipleural fold (Fig. 311) as wide as antennal

article 11; sutural notch right angle; apical slope

gradual; EL/EW, average about 4.5, range 4.3-5.0

(18 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 313, 314): Feebly sclerotized; phallobase with 2 dorsal and 2 ventral

longitudinal bands, latter more sclerotized than

remainder of phallobase. Combined length of

phallobase and phallobasic apodeme 8 times longer

than paramere. Combined length of paramere and

phallobase 1.4 times longer than phallobasic

apodeme. Paramere: apex truncate; lateral depression absent; dorsum and venter plane; mesoventral

and mesodorsal margin broadly concave. Sinus:

dorsal indistinct, half as long as lanceolate ventral.

First connecting membrane with serrulate scales

(Figure 315). Phallus: phallic plate very slender and
NUMBER 227

10S

FIGURES 308-314.—Perilypus ventralts: 306, head venter and cervical membrane, X 90 (arrow

indicates gular process); 309, elytron, X 41 (arrow indicates sutural cleft); 310, mesoscutellum,

lateral view, X 45; 311, elytron, X 41; 312, mesoscutellum, dorsal view, X 45; 313, aedeagus,

ventral view, X 60; 314, phallus. X 290.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ficintzs 315-321.—Perilypus ventralis: 315, aedeagus, X 350 (arrow indicates first connecting

membrane); 316, male internal reproductive organs; 317, ovipositor, ventral view; 318, ovipositor, dorsal view; 319, posterior accessory gland. P. ornaticottis: 320, aedeagus, ventral

view, X 117; 321, aedeagus, ventral view, X 325 (arrow indicates first connecting membrane).

(Scale - 1 mm)
NUMBER 227 105

strongly sderotized; marginal denticles absent;

phallic plicae prolonged and set with very large

spines (Figure 314) (18 specimens examined).

Female Genitalia (Figures 317, 318): Dorsal lamina heptalobed; ventral lamina trilobed, laminal

incisions very short; proctiger semicircular; proctigeral bacculi not fused; ventral bacculus acuminate near middle; coxital plates absent (2 specimens

examined).

Internal Reproductive Organs: Male (Figures 316,

319): anterior accessory gland uniramous, coiled

distally, posterior accessory gland biramous, outer

branch slightly longer than inner branch, both

branches broadly arcuate; ejaculary duct not bulbous anteriorly; testis composed of 12 follicles.

Female: as in Figure 55 (4 males and 2 females

examined).

VARIATION.—Structural: A female specimen from

El Fortfn, Mexico, is more oval in body form.

Color: Intrapopulation variation of elytral color

not correlated with sex distinguishes a light phenon

and a dark phenon. Beetles of the light phenon

have flavous vitta on the sutural and epipleural

margins. The epipleural vittae may be slightly

expanded at midelytron. The elytron is entirely

black in beetles of the dark phenon. In general, the

elytral disc varies from brown to black.

NATURAL HISTORY.—In June Henry Dybas collected a female specimen from El Fortin, Mexico,

at 1000 meters. From Sumpango, Guatemala, at

2012 meters, I collected 15 adult specimens on the

compositae Vernonia deppeana Lessing. Fourteen

of these were of the dark phenon, one of the light

phenon. On the same plant I also collected

specimens of the lampyrid genus Photinus. The

striking similarity between these beetles was first

noted by Gorham (1882:141), ". . . this, together

with the color, gives this insect a singularly close

resemblance to some Lampyridae which the two

or three pale segments beneath heightens. The

variety from Oaxaca [my light phenon] equally

resembles those varieties of species of Photinus in

which the margins are pale."

DISTRIBUTION (Figure 368).—The known distribution of this species extends from Veracruz,

Mexico, to Sacatepequez, Guatemala. Most of the

specimens were collected at elevations from 1000

to 2012 meters.

LOCALITY RECORDS (Figure 368).—I have examined 28 adult

specimens from Central America: MEXICO: Estado de Veracruz: Fortfn (FMNH, 1 female). Jesus Carranza (NMNH, 1

female). Estado de Guerrero: Jalapa (BMNH, 2 males).

Estado de Oaxaca: Oaxaca (BMNH, 1 male). "Mexico"

(MNHP, 1 female). GUATEMALA: Departamento de Baja

Verapaz: San Jeronimo (GEKI, 1 female). Departamento de

Sacatepequez: Antiqua (GEKI, 1 female; NMNH, 1 female);

Sumpango (AMNH, 1 female; GEKI, 5 males and 4 females;

FMNH, 1 female; NMNH, 1 male and S females); Duenas

(BMNH, 4 females, lectotype; GEKI, 1 female).

43. Perilypus ornaticollis (LeConte),

new combination

FIGURES 520-330, 367

Cleronomous ornaticollis LeConte, 1880:194 [holotype: female,

deposited in MCZC; type-locality: Ohio, U.S.A.].

Colyphus melanopterus Dury, 1906:251 [type deposited in

CMNH; type-locality: Cincinnati, Ohio, U.S.A.; Synonymy

by Wolcott (1910a:851)].

Colyphus furcatus Schaeffer, 1904:218 [lectotype: male, and 7

paralectotypes, 1 male and 6 females, here designated, deposited in NMNH; type-locality: San Tomas, Esperanza

Ranch, Brownsville, Texas, U.S.A.; new synonymy].

According to Dury (1906:251) the type-specimen

of Colyphus melanopterus Dury (Cleronomus ornaticollis LeConte) is "longer, much less hairy, the

elytra are more shining and immaculate jet black"

than representatives of Colyphus furcatus Schaeffer.

Although I did not see the type-specimen of C.

melanopterus, two other specimens from Ohio were

studied. The characteristics mentioned by Dury

fall within the range of variation of this species.

DIAGNOSIS.—Distinguishable by the bifurcate

configuration of the pronotal discal vitta (Figures

322-324).

DESCRIPTION.—Form: As in Figure 322.

Size: Length: males, average about 6.7 mm, range

6.3-7.2 mm; females average about 7.9 mm, range

6.0-8.4 mm. Width: males, average about 2.0 mm,

range 1.6-2.4 mm; females, average about 2.3 mm,

range 1.5-2.8 mm. Six males and 15 females

measured.

Color: Clypeus and frons flavotestaceous; epicranium, gena, antenna, and legs dark brown;

pronotum and elytron variable (see "Variation");

prosternum flavotestaceous; legs, meso- and metasternum, and abdomen brown to black.

Head: Interocular depressions broad, well impressed, and crescentic; frontal umbo prominent;

frons broad; eyes moderately convex; HW/IOW,
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

325

FIGURES 322-330.—Perilypus ornaticollis: 322, habitus; 323-324, pronota; 325, antenna; 326,

tegmen, lateral view; 327, tegmen, ventral view; 328, phallus; 329, ovipositor, ventral view;

330, ovipositor, dorsal view. (Scale = 1 mm)
NUMBER 227

107

average about 1.8, range 1.3-1.9 (21 specimens

measured); antenna (Figure 325) boldly serrate;

length/width ratio of antennal articles sex dimorphic, male 2.0:1.1:1.3:1.2:1.1:1.1:1.2:0.9:0.9:0.7:1.3,

female 2.0:1.1:1.3:1.2:1.2:1.2:1.0:1.0:0.8:0.7:1.2; AL/

PLS, 1.7.

Pronotum: Strongly transverse (PL/PW, average

about 0.81, range 7.4-8.7; 21 specimens measured);

side margin of pronotum proper boldly convex;

subapical depression parallel with anterior margin,

pronotal arch narrow, feebly convex; prebasal

depression shallow; pronotal collar narrow.

Mesoscutellum: Transverse.

Elytron: Posthumeral margin straight in basal

third, arcuate in apical two-thirds; slope of apical

deflection gradual; punctations broad and shallow;

punctate and interpunctate surfaces with rugose

microsculpture, latter makes elytral surface appear

arenose; densely setose; EL/EW, average about 4.5,

range 4.1-5.6 (21 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 320, 321, 326-328): Feebly sclerotized; phallobase with 2 dorsal and 2 ventral longitudinal bands, latter more sclerotized than

remainder of phallobase. Combined length of

phallobase and phallobasic apodeme 7 times longer

than paramere. Combined length of paramere and

phallobase 1.4 times longer than pallobasic apodeme. Paramere: acuminate; inner walls of paramere densely setose. Sinus: dorsal, half as long as

ventral. First connecting membrane set with serrulate scales (Figure 321). Phallus: apex lobiform;

phallic plate abruptly narrowed at middle and at

junction with phallic apodeme; marginal denticles

absent; phallic plicae inconspicuous (6 specimens

examined).

Female Genitalia (Figures 329, 330): Dorsal and

ventral lamina trilobed, latter with very short

incisions; proctiger broad, nearly semicircular;

proctigeral bacculi not fused; ventral bacculus

accuminate near middle; coxital plates absent (2

specimens examined).

Internal Reproductive Organs: Male: lateral

accessory gland uniramous, coiled throughout (1

specimen examined).

VARIATION.—Structural: The antenna is more

serrate and body form more oval in three female

specimens from southeastern Mexico; two of these

also differ by their more transverse pronotum

(PL/PW, 0.76).

Color: In general, southern specimens are lighter

than northern ones. Beetles from the southern part

of the range (south of Louisiana) have an incomplete pronotal vitta (Figure 322), and the elytron is

broadly vittate (Figure 322). The pronotal vitta is

percurrent and the elytral disc a vittate and more

shiny in specimens north of Louisiana (Figure

324).

Two specimens, one from Tennessee and one

from Arkansas, have the pronotal vitta prolonged

unto the basal half of the pronotal arch (Figure

323).

NATURAL HISTORY.—Adult specimens were collected in March, May, June, and July. R. Kirkton

collected one specimen in Arkansas in June within

a "cotton alfalfa strip." Dury (1906:251) collected

"one specimen while sweeping low vegetation in

river bottom, July 5, 1905."

DISTRIBUTION (Figure 367).—This is the only

species of Perilypus prominently distributed north

of Mexico. The known range extends from Ohio,

U.S.A., to Cordoba, Mexico.

LOCALITY RECORDS (Figure 367).—I examined 46 adult specimens from North and Central America. UNITED STATES:

Ohio: Hamilton County (FMNH, 1 male); "Ohio" (MCZC. 1

female, holotype). Indiana: Tippecanoe County (GEKI, 1 female). Tennessee: Memphis (GEKI, 1 female). Louisiana:

Alexandria (GEKI, 1 female). Arkansas: Mississippi County

(WFBA, 1 female). Texas: Saint Thomas, Brownsville (GEKI,

1 female); Brownsville, Esperanza Ranch (AMNH, 1 female;

CASC, 1 female; BMNH, 1 female; FMNH, 3 females; GEKI,

2 males and 8 females; GHNE. 1 female; LMDR, 2 females;

MCZC, 1 male and 1 female; SEMK, 2 females; NMNH, 2

males and 7 females; UZMD, 1 female). MEXICO: Estado

de Veracruz: San Rafael, Jicaltepec (GEKI, I female; MCZC,

2 females); Cordoba (GEKI, 1 female); Orizaba (GEKI, 1 female); 17 miles [27 km] north of Nauxtla (GEKI, 1 female).

REMARKS.—The three female specimens discussed

under "Variation" are probably not conspecific,

but I cannot decide on the basis of the available information. I labeled these P. ornaticollis

(LeConte)?.

44. Perilypus galbeus, new species

FICURES 332, 337-339, 355, 368

TYPE-LOCALITY.—Guatemala.

TYPE-SPECIMENS.—The holotype male is depos-
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ited in MNHP and was collected by L. Conradt.

The allotype is in BMNH.

DIAGNOSIS.—There is considerable habitus similarity between specimens of this species and those

of P. prolixicornis and of P. pilatus. From the first

of these, P. galbeus differs by its shorter antennal

article 11 (compare Figures 331, 332), and from P.

pilatus by the more oval and more convex elytron.

Also, the elytral surface of P. pilatus is vested with

exceptionally long vertical setae.

DESCRIPTION.—Form: As in Figure 355.

Size: Length: male 7.5 mm, female 8.8 mm.

Width: male 2.4 mm, female 3.4 mm.

Color: Head black, except clypeus and midanterior region of frons flavotestaceous; antenna

black; prothorax predominantly flavotestaceous,

pronotal disc with tapering black vitta; legs black

except femur annulated distally; meso- and

metathorax, elytron, and abdomen black.

Head: Interocular depressions and frontal umbo

as in P. ornaticollis; frons broad, eyes moderately

convex; antenna (Figure 332) boldly serrate;

antennal length sex dimorphic (AL/PLS, male 2.5,

female 2.1) and article 11 as long as preceding two;

length/width ratio of each antennal article, male

2.2:1.4:1.5:1.3:1.2:1.2:1.0:1.0:1.0:1.0:2.0.

Pronotum: Strongly transverse (PL/PW, average

about 0.77, range 0.76-0.78; 2 specimens measured);

subapical depression feebly sinuous, shallow at

middle; pronotal arch narrow and shallow; foveae

more transverse than punctiform; pronotum proper

margin feebly arcuate; prebasal depression very

shallow; pronotal collar very narrow medially.

Mesoscutellum: Transverse.

Elytron: More convex and more arenose than

in P. ornaticollis; posthumeral margin arcuate

throughout; slope of apical deflection gradual;

epipleural fold feebly concave, as broad as antennal

article 11; apex truncate; EL/EW, average about

4.2, range 3.8-4.5 (2 specimens measured).

Front Tarsal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 337-339): As in P.

ornaticollis except paramere apex more gradually

acuminate; dorsal and ventral sinus very narrow,

latter sinus only slightly longer than dorsal sinus;

and phallic plates uniformly slender.

Female Genitalia: Not studied.

VARIATION.—Structural: Antennal length is sex

dimorphic; the male antenna is the longer. The two

specimens examined differ in shape of interocular

depression which is more crescentic in the male.

Color: The two specimens do not vary appreciably in color.

DISTRIBUTION (Figure 368).—The known range

extends from Veracruz, Mexico, to Guatemala.

ETYMOLOGY.—Latin noun galbeus (arm band),

in allusion to the testaceous annulus on the

femora.

LOCALITY RECORDS (Figure 368).—I examined 2 adult specimens from Central America. MEXICO: Estado de Veracruz,

Cordoba (BMNH, 1 female). GUATEMALA (MNHP, 1 male,

holotype).

45. Perilypus prolixicornis, new species

FIGURES 331, 334-336, 368

TYPE-LOCALITY.—Cordoba, Estado de Veracruz,

Mexico.

TYPE-SPECIMENS.—The holotype male, deposited

in CASC, was collected by A. Feyes.

DIAGNOSIS.—The only specimen studied can be

distinguished from congeneres by the length of

antennal article 11 (Figure 331), which is as long

as the preceding three.

DESCRIPTION.—Form: As in P. galbeus (Figure

355).

Size: Length, 7.5 mm; width, 2.4 mm.

Color: As in P. galbeus.

Head: As in P. galbeus except antennal article

11 (Figure 331) proportionally longer; length/width

ratio of each antennal article, 2.0:1.1:1.0:1.3:1.3:

1.1:1.0:0.9:0.8:0.9:3.5; and AL/PLS, 4.9.

Pronotum: As in P. galbeus except foveae

broader and shallower.

Elytron: As in P. galbeus.

Front Tarsal Claws: Symmetrical.

Mesoscutellum: Transverse.

Male Genitalia (Figures 334-336): Generally as

in P. ornaticollis (LeConte) but differs from that

species as follows: combined length of phallobase

and phallobasic apodeme 10 times longer than

paramere; paramere apices excurved, ventral sinus

only slightly longer and broader than dorsal sinus,

and phallic plate uniformly narrow.

ETYMOLOGY.—Latin adjective prolixus (stretched

out) + the noun cornu (horn) + 1 adjectival

ending -is; referring to the long antennal article 11.
NUMBER Til 109

337 338 339 340 341

1-341.—Perilypus proltxicomis: 331, antenna; 334, tegmen, lateral view; 335, teginen,

w; 336, phallus. P. galbeus: 332, antenna; 337, tegmen, lateral view; 338, tegmen,

w; 339, phallus. P. floralis: 333, antenna; 340, legmen, lateral view; 341, teemen.

t

1

 IGURES 331—JTI . M. c / HVLSIAO u i un.Mii// riu, JJ I , a«ii«_iuia, O-J~X, I«.^IIILII

ventral view; 336, phallus. P. galbeus: 332, antenna ; 337, legmen , _..„.*** ,*»„ . ..,...., ^.^ ,.

ventral view; 339, phallus. P. floralis: 333, antenna ; 340, tegmen , lateral view; 341, tegmen,

i_ ^ i mrn \ ventral view. (Scale = 1 mm)
110

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

342

FIGURES 342-350.—Perilypus floralis: 342, habitus; 343, tegmen, lateral view; 344, tegmen, ventral

view; 345, phallus; 346, ovipositor, ventral view; 347, ovipositor, dorsal view. P. pilatus: 348,

ovipositor, ventral view; 349, ovipositor, dorsal view. P. antarius: 350, antenna. (Scale = 1 mm)
NUMBER 227

111

LOCALITY RECORD (Figure 368).—I examined 1 adult specimen from Central America. MEXICO: Estado de Veracruz:

Cordoba (CASC, 1 male, holotype).

LOCALITY RKCORDS (Figure 368).—I examined 1 adult specimen from Central America. PANAMA: Provincia de Panama, Cerro Campana (NMNH, 1 female, holotype).

46. Perilypus pilatus, new species

FIGURES 348, 349, 351, 354, 368

TYPE-LOCALITY.—Cerro Campana, Provincia de

Panama, Panama.

TYPE-SPECIMENS.—The holotype female, deposited in NMNH, was collected by H. P. Stockwell

in 1970.

DIAGNOSIS.—The stout black setae on the elytral

surface and the conspicuous divergence of sutural

margins at the elytral apical fifth are diagnostic

characteristics of this species.

DESCRIPTION.—Form: As in Figure 354.

Size: Length: 8.4 mm. Width: 2.9 mm.

Color: As in P. ornaticollis except pronotal vitta

not furcate; pronotal sides roseate; apex and suture

of elytron black; and femora with subapical

testaceous annulus.

Head: As in P. ornaticollis except eyes more

convex (HW/IOW, 2.1); antenna (Figure 350)

more compressed, more distinctly serrate, and

proportionally longer (AL/PLS, 2.0); and length/

width of each antennal article 2.0:1.0:1.1:1.3:1.2:

1.2:1.2:1.0:0.9:0.8:1.3.

Pronotum: As in P. ornaticollis except subapical

depression more sinuous and foveae more deeply

impressed.

Mesoscutellum: Transverse.

Elytron: Posthumeral margin arcuate; punctations particularly small and shallow; punctate and

interpunctate surfaces densely arenose; surface

copiously vested with short pale reclinate setae and

with stout black vertical setae; sutural hinge

abruptly narrowed at apical fifth; elytral sutural

margin diverging in apical fifth; apex sinuatotruncate (Figure 354); EL/EW, 3.9.

Front Tarsal Claws: Symmetrical.

Female Genitalia (Figures 348, 349): As in P.

ornaticollis except dorsal laminal incisions shorter

and proctiger more slender.

NATURAL HISTORY.—In July H. P. Stockwell

collected the holotype from the type-locality (800

meters) by beating.

ETYMOLOGY.—Latin adjective pilatus (thick),

referring to the stout elytral vertical setae.

47. Perilypus antarius, new species

FIGURES 350, 352, 353, 368

TYPE-LOCALITY.—San Pedro, Montes de Oca,

Provincia de San Jose", Costa Rica.

TYPE-SPECIMEN.—The holotype female, deposited

in NMNH, was collected by C. H. Ballou in 1936.

DIAGNOSIS.—Elytral apex strongly upcurved

(Figure 353).

DESCRIPTION.—Form: As in Figure 352.

Size: Length 7.8 mm. Width 2.5 mm.

Color: As in P. ornaticollis except epipleural fold

flavotestaceous, pronotal vitta not furcate, elytral

disc avittate, femora annulated subapically, and

posterior margin of abdominal sterna flavotestaceous.

Head: As in P. ornaticollis except antenna

(Figure 350) proportionally longer (AL/PLS, 2.5);

length/width ratio of each antennal article 2.2:

1.1:1.2:1.9:1.7:1.5:1.5:1.3:1.3:1.1:1.9; and HW/

IOW, 2.2.

Pronotum: As in P. ornaticollis except subapical

depression more impressed and PL/PW, 0.73.

Mesoscutellum: Transverse.

Elytron: As in P. ornaticollis except apex uplifted

epipleural fold obliquely convex; and EL/EW, 4.8.

Front Tarsal Claws: Symmetrical.

NATURAL HISTORY.—In June, from the typelocality, G. H. Ballou collected the holotype on

Hemalia erecta Linnaeus.

ETYMOLOGY.—Latin adjective antarius (hoisting),

referring to the uplifted elytral apex.

LOCALITY RECORDS (Figure 368).—I examined 1 adult specimen from Central America. COSTA RICA: Provincia de San

Jos£: San Pedro, Montes de Oca (NMNH, 1 female, holotype).

48. Perilypus sinuapicis, new species

FIGURES 343-345, 369

TYPE-LOCALITY.—San Esteban, Estado de Carabobo, Venezuela.

TYPE-SPECIMENS.—The male holotype, deposited

in FMNH, was collected by P. J. Anduze in 1939.

DIAGNOSIS.—Easily distinguishable from speci-
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

354

FIGURES 351-354.—Perilypus pilatus: 351, antenna; 354, habitus. P. antarius: 352, habitus;

353, hind body. (Scale = 1 mm)
NUMBER 227 113

mens of P. fioralis, the only other member of the

ornaticollis group whose specimens have the elytral

surface very shiny, by differences in elytral color

(elytron is avittate in specimens of P. sinuapicis).

Also, in specimens of P. sinuapicis the antenna is

black and femora predominantly dark brown

whereas in those of P. fioralis the antenna and

the abdomen is entirely flavotestaceous.

DESCRIPTION.—Form: As in P. fioralis except

epipleural margin less convex.

Size: Length: 6.2 mm. Width: 1.9 mm.

355

FIGURE 355.—Perilypus galbeus, habitus. (Scale = 1 mm)

Color: As in P. ornaticollis except pronotal vitta

not furcate and femur with testaceous annulus.

Head: As in P. ornaticollis except eyes more convex (HW/IOW, 2.2), antenna proportionally longer

(AL/PLS, 2.1), and length/width ratio of each

antennal article 2.0:1.1:1.4:1.5:1.5:1.3:1.2:1.2:1.1:

1.2:1.5.

Pronotum: As in P. ornaticollis except less transverse and subapical depression more deeply

impressed.

Mesoscutellum: Transverse.

Elytron: As in P. ornaticollis except punctate and

interpunctate surfaces shiny, only feebly arenose,

epipleural fold convex, and apex sinuatotruncate.

Front Tarsal Claws: Symmetrical.

Male Genitalia (Figures 343-345): As in P. ornaticollis except combined length of paramere and

phallobase 1.6 times longer than phallobasic apodeme, paramere apex more acuminate, dorsal and

ventral sinus narrower, and phallic plates not

abruptly widened at middle.

NATURAL HISTORY.—Unknown except the holotype was collected in December.

ETYMOLOGY.—Latin noun sinus (pocket) + noun

apex (tip) + adjectival ending -is, referring to the

emargination of elytral apex.

LOCALITY RECORDS (Figure 369).—I examined 1 adult specimen from South America. VENEZUELA: Estado de Carabobo: San Esteban (FMNH, 1 male, holotype).

49. Perilypus fioralis (Gorham), new combination

FIGURES 333. 340-342, 346, 347, 368

Colyphus fioralis Gorham, 1882:142 [holotype: male, deposited

in BMNH; type-locality: Volcan de Chiriquf, Provincia de

Chiriqui, Panama].

DIAGNOSIS.—Distinguishable from all other congeneric species by the unique configuration of the

elytral vittae. The flavous discal vitta begins at the

elytral midbase and arches to join the epipleural

margin vitta at the elytral apical fifth (Figure 342).

DESCRIPTION.—Form: As in Figure 342.

Size: Length: males, average about 6.8 mm, range

6.6-6.9 mm; females, average about 6.7 mm, range

6.0-7.4 mm. Width: males, 2.3 mm; females, average

about 2.1 mm, range 1.9—2.4 mm. Two males and 3

females measured.

Color: Clypeus, frons, posterior epicranial region,

venter of head, and antenna flavotestaceous; an-
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terior epicranial region and gena black; prothorax

flavotestaceous, except pronotum with black midbasal macula; legs flavotestaceous; meso- and

metasternum black; elytron predominantly black,

with sinuous discal and linear marginal vitta converging at apical fifth; abdomen black.

Head: Interocular depression, frontal umbo, and

eye convexity as in P. ornaticollis; frons broad

(HW/1OW, average about 2.0, range 1.8-2.1; 5

specimens measured); antenna (Figure 333) distinctly serrate; length/width ratio of each male

antennal article 1.8:1.3:1.3:1.8:1.7:1.5:1.5:1.7:1.7:

1.2:2.0; AL/PLS sex dimorphic, male 2.6, female

2.3.

Pronotum: Boldly transverse (PL/PW, average

about 0.78, range 0.71-0.80; 5 specimens measured);

subapical depression deeply impressed; side margin

of pronotum proper boldly convex; fovea prominent, broad and shallow; prebasal depression shallow; pronotal collar narrow.

Mesoscutellurn: Transverse.

Elytron: Elytral surface as in P. ornaticollis

except surface smooth, shiny, less setose, and punctations reduced near suture and apex; epipleural

fold very broad and convex (as wide as antennal

article 11); apex feebly sinuatotruncate.

Front Tar sal Claws: Symmetrical in both sexes.

Male Genitalia (Figures 340, 341): As in P.

ornaticollis except phallobasic apodeme as long as

phallobase and paramere combined; dorsal and

ventral sinus broader, latter only slightly shorter

than ventral; and phallus uniformly slender (2

specimens examined).

Female Genitalia (Figures 346, 347): As in P.

ornaticollis except ventral lobes more prominent

and proctiger reduced (1 specimen examined).

Internal Reproductive Organs: Male, anterior

accessory gland uniramous, coiled throughout (1

specimen examined).

VARIATION.—Antennal length is sex dimorphic,

being longer in males than in females (AL/PL,

male 2.6, females 2.3). The elytral vitta varies in

width.

NATURAL HISTORY.—In August, at 915 meters on

Cerro Campana, H. and A. Howden collected 1

adult specimen by beating. G. C. Champion collected 4 specimens from Volcan de Chiriqui at

elevations from 915 and 1220 meters.

DISTRIBUTION (Figure 368).—The known distribution extends from Provincia de Chiriqui, to

Provincia de Panama, Panama.

LOCALITY RECORDS (Figure 368).—I examined 5 adult specimens from Central America. PANAMA: Provincia de Chiriqui: Volcan de Chiriqui (BMNH, 2 males, holotype; GEKI, 1

female; MNHP, 1 female). Provincia de Panama: Cerro

Campana (HFHO, 1 female).

Immature Specimens

I studied two larvae and one pupa of P. limbatus

(Gorham), the only known immature specimens of

Pcrilypus. Although the larval description is based

on one species, I tentatively consider the description

generic even though the specimens studied probably do not present adequately the full complement

of generic character states. Fortunately, the comprehensive publication of clerid larvae by Boving

and Champlain (1920) provides some basis for

making tentative decisions about generic characterizations, even when only a few larval specimens

are available.

Lack of sufficient data and specimens precludes

a precise age determination of the two larvae

examined. On the basis of size, the larger specimen

probably belongs to at least the penultimate larval

instar.

I do not provide a description of the pupa at

this time because the only available specimen exhibits most of the Perilypus imaginal characteristics.

The small uncinate urogomphi, and chaetotaxy,

may be taxonomically important, but I cannot

decide without comparative information. In lieu

of a written description, habitus figures of the

pupa (Figures 380, 381) are provided as preliminary diagnostic aids.

DIAGNOSIS.—Larval Perilypus are distinguished

from larval specimens of most other genera of

Clerinae studied to date by having only one seta on

the outer surface of the mandible. The only known

exceptions are Trichodes bicinctus Green and T.

nutalli (Kirby) (Foster, 1976); the conspicuously

concave hypostomal margin in Trichodes distinguish members of that genus from the available

larvae of Perilypus in which the hypostomal margin

is feebly oblique. Foster (1976) provided detailed

characterizations of Trichodes larvae in his revision

of North American Trichodes. The mandible of

other known clerine larvae has at least two mandibular setae. Also, the anterior margin of the labrum
NUMBER 25J7

115

is emarginated deeply in larvae of Perilypus; the

anterior labial margin in other known clerine

larvae is more transverse, and shallowly emarginated or feebly arcuate.

DESCRIPTION.—Form (Figure 370): Digitiform,

and cylindrical except head capsule flattened dorsoventrally; forebody more tapered than hindbody;

densely vested with pale setae.

Size: Length 8.8 mm, width 1.6 mm.

Color: Mandible, cranium, pronotal plate, and

ninth abdominal segment dark brown; remainder

of mouthparts, antennae, legs, prosternal plate, and

meso- and metanotal plates light brown; abdominal

terga with pair of transverse purpurescent maculae;

remainder of body white.

Head (Figures 371, 372): Prognathus and protruding. Head capsule transverse (length 0.9 mm,

width 1.0 mm) plane, venter feebly tumescent;

epicranial suture absent; frontal sutures feebly

arcuate at anterior third; frons triangular, with

shallow longitudinal rugosities; epicranium smooth;

maxillary hypostomal margin (Figure 371) feebly

oblique; gular sutures feebly sinuous; gular plate

narrow, tapering posteriorly. Five stemmata (Figure

372) present; stemmata aligned in two parallel

vertical rows, three in anterior row and two in

posterior row. Antenna (Figure 376) composed of

three articles and broad transparent basal membrane; article 1 largest, about twice as long as wide;

article 2 quadrate, ventroapical region projecting

a conical antennal appendix; article 3 about three

times as long as wide and with long stout apical

seta. Mandible (Figure 373) falciform, with one seta

on posterior outer surface; mandibular groove

deeply invaginated; apex unidentate; retinaculum

shallow, present on middle of masticatory surface;

lacinia mandibulae present, setiform, and only

feebly bifid apically. Maxilla (Figure 377) with

cardo and stipes trapezoidal, each with oblique

transverse plate at base, plate of former asetose, of

latter bisetose near outer margin; palpiger feebly

sclerotized, bisetose at inner margin; palpus with

three articles successively narrowing from basal

article, article 1 transverse, article 2 quadrate,

article 3 elongate; pedunculate seta well developed.

Labium (Figure 371) with rudimentary ligula; submentum and mentum faintly sclerotized, former

elongate, latter transverse; palpiger strongly sclerotized, with one stout seta on inner apical margin;

palpus with two articles, article 1 subquadrate,

article 2 elongate and feebly arcuate. Labrum (Figure 375) transparent and well developed; side

margins serrulated; anterior margin strongly emarginated; aboral surface bearing eight large setae

arranged into two anterolateral groups of four.

Epipharynx (Figure 375) with anterior row of 12

truncate epipharyngeal seta, and four epipharyngeal

sensilla; latter dorsad to tormal sclerite, which

extends into elongate processes posterolaterally.

Thorax (Figures 370, 378): Prothorax with notal

plate strongly sclerotized, scutiform, and divided

by a light midlongitudinal line; prosternum (Figure 378) with elliptical midlongitudinal prosternal

plate (the sternal plate of Boving and Champlain,

1920:581) that adjoins two transverse lateral prosternal plates (the presternal area of same authors).

Meso- and metathorax with pair of feebly sclerotized notal sclerotizations; sternomedial sclerotization elliptical in mesothorax and subovate in

metathorax, plates feebly sclerotized in both segments. Legs well sclerotized, composed of five

segments that taper to claw; coxa rectangulate;

femur as long as longitudinal diameter of coxa;

tibia narrow, about as long as femur; tarsus

unguiform.

Abdomen (Figure 370): Segments 1 to 8 without

ampullae or sclerotizations; segment 9 (Figure 379)

with strongly sclerotized notal plate, which projects

as pair of widely separated uncinate urogomphi;

segment 10 reduced, entirely covered by segment 9.

VARIATION.—In addition to size, the smaller

larva (length 8.0 mm, width 1.9 mm) differs from

the larger one most conspicuously by the bluntness

of the mandibular apex (Figure 374), shallower

mandibular groove, equilateral head capsule, and

by having shorter integumental setae.

DISCUSSION.—In addition to the immature specimens of Perilypus described in this paper, I studied

larvae of the genera Placopterus, Enoclerus, Thanasimus, Trichodes, Opilo, and Tillicera. Because

larvae of only a few genera of Clerinae are available, any discussion of relationships based on larval

character states is obviously tentative. Nevertheless,

such a discussion is presented to stimulate resyntheses as additional data become available.

The relative apotypy and plesiotypy of Clerinae

larval characters is beyond surmise at present,

because few clerine larvae are available for study.

However, the presence of the lacinia mandibulae in

the larvae of Perilypus (Figure 373) and Placopterus
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figure 382) suggest a potentially close relationship.

The lacinia mandibulae has heretofore not been

reported in Cleridae although Crowson (1964:277)

noted its occurrence in other families of the superfamily Cleroidea. Unfortunately, I have not been

able to substantiate the possible close relationship

between Perilypus and Placopterus on the basis of

adult synapotypy, but earlier (p. 20) I alluded to

the superficial similarities between some of their

members. Should the lacinia mandibulae be judged

a synapotypic characteristic, then it would be quite

legitimate to infer sister group relationship between

the aforementioned two genera since it is not

unusual for taxa to lose indicators of "immediate"

ancestry in the imago stage while retaining them

in the immature stages. This is precisely why

Hennig (1966:32) stressed the importance of the

holomorphological approach in phylogenetic

systematics.

The second item of interest regarding the aforementioned larvae is that they provide some hints of

relationships at suprageneric level. The configuration of the hypostomal margin separates all the

larvae examined into two groups. In Perilypus,

Placopterus, and Enoclerus the margin is feebly

oblique (as in Figure 371), whereas in the remaining

genera the hypostomal margin is distinctly concave

(Boving and Champlain, 1920, fig. 36). When the

closely related genera of each group are brought

into alignment an interesting pattern emerges, one

that may have considerable heuristic value. The

dichotomy, based on hypostomal margin configuration, essentially separates the more recently evolved

genera (Perilypus, Placopterus, Enoclerus) from the

aforementioned older generic groups (Trichodes,

etc). The former group of genera are exclusively

New World whereas the latter group is predominantly Old World except for Priocera, a New

World genus with apparent Old World affinities.

Phylogeny and Zoogeography

PHYLOGENETIC METHODS

The postulated phylogenetic relationships within

Perilypus are illustrated in Figures 383-386 and

are based on Hennigian principles (Hennig, 1965,

1966). Hennig's tenets for reconstructing phylogeny

were copiously reviewed elsewhere and I think it

unnecessary to rediscuss them or to restate their

logic as justification for their use herein. However,

I want to emphasize, an aspect of Hennig's

approach apparently not fully appreciated; that is,

systematic studies in which Hennigian methods are

applied result in a more comprehensive presentation

of available data about the organisms under study.

The reason for this should become evident in the

following paragraphs.

Fundamental to Hennig's methods of phylogenetic analysis are the breakdown of phenotypic

resemblance into symplesiomorphy, synapomorphy,

and convergence (for definitions see Hennig, 1965:

102), and the establishment of monophyletic groups

only on the basis of shared apomorphic character

states; that is, a taxon is monophyletic when it

includes all the known descendents of that taxon's

ancestor; the descendents are recognized by the

concept of synapomorphy.

Hennig's approach to phylogenetic analysis has

an inherent prerequisite for in-depth investigations

of characters from as many aspects of an animal's

gestalt as is available for study; hence, the possibility of discovering synapomorphies is maximized.

How readily such similarities are detected is, of

course, largely dependent on a group's evolutionary

history. Often it is dependent on a specialist's

willingness to conduct comprehensive analysis of

more than just the readily observable characters.

In-depth studies of phenotype (morphological or

otherwise) are essential to the holomorphological

approach of character analysis forwarded by Hennig

(1966:180). Results from such an approach are

presented in phylogenetic systems in which relationships are recognized by definable criteria

(autapomorphies or synapomorphies), which theoretically best approximate evolutionary history. The

concept of monophyly by autapomorphy only is

one of the crucial differences between the theory

of phylogenetic systematics advocated by Hennig

and that championed by Mayr (1969), or Sokal and

Sneath (1963). Works not mentioned elsewhere in

this paper that discuss some of Hennig's phylogenetic principles include Brundin (1966, 1968,

1972), Tuomikoski (1967), Darlington (1970), Schlee

(1971), Kavanaugh (1972), and Mayr (1974).

CHARACTER PHYLOGENY.—The phylogenetic status

of character states, whether apotypic (derived) or

plesiotypic (primitive), must be resolved before
NUMBER 227 117

P. insect US (exact loca

FICURE 356.—Distribution map of the frontalis and criocerides groups.

sister-group relationships can be established (the

prefixes apo- and plesio- are combined with the

neutral suffix -typic (cf. Ball, 1975), not with

-morphic which has strictly morphological implications). Criteria for inferring character phylogenies

were proposed and/or reviewed in detail by Hennig

(1965, 1966), Maslin (1952), Schlee (1969), Kluge

and Faris (1969), Marx and Rabb (1970, 1972),

Crowson (1970), Munroe (1974), Ross (1974), and

by Ball (1975). Of the criteria mentioned by these

authors those most relevant to this study are briefly

discussed below. The application of most of these

criteria require of the specialist a broad understanding of character states (particularly their distribution) among diverse groups relevant to the

taxa under investigation.

CRITERION OF FREQUENCY OF OCCURRENCE.—This

is the most conventional method of predicting a

characteristic's phylogenetic status, particularly

when two-state characters are involved. The objective is to determine how extensively a character

state is distributed within the taxon under study

(e.g., genus) and among taxa of at least the next

highest category (e.g., subtribe). Needless to say,
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

P. relucens

»P.viridipennis

P.levis

359

L^^S pjP.chaletoides

-^~Vij £ P. bicolor

'A 1o

FIGURES 357-359.—Distribution maps: 357-358, the viridipennis group;

359, the chaletoides group.

the more inclusive the survey of taxa the more

credible becomes the assessment of character phylogeny. Character states widespread among diverse

groups are classified as plesiotypic on the assumption their extensive distribution is a result of

inheritance, not due to independent evolution.

Conversely, character states with limited distributions are interpreted apotypic because their re-
NUMBER 227 119

O P.prolixipenis

v£) P.testaceicornis

0 P.decoris

X P.latilira

^ Rlimbatus

® Rbuga

® P.iris

© Rapocopatus

M P.columbicus

O P.acus

FIGURES 360-362.—Distribution maps of the limbatiis group.
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

.carbonarius

^yP.caliculus

P.distinctus

P.latilobus

P.calcaris

P.sapientis

P.cultratus

P. reventazon

P.exilis

P. immitis

P. orthopleuridus

P. nigriventris

P. claudus

P.bilineatus

P.ordinatus

P. cerroazul

P.crassus

FIGURES 363-365.—Distribution maps of the reventazon group.
NUMBER. TZ1 121

P.bicristatus

P.quadrilineatus

P.telepboroides

P. coroniformis

£) P.fluetus

P.ornaticollis

P. ventral is

P.antarius

P.floralis

P.galbeus

Rprolixicornis

P.pilatus

FIGURES 366-369.—Distribution maps: 366, the quadrilineatus group;

367-369, the ornaticollis group.

stricted occurrence is an assumed manifestation of

recent evolution. However, limited distribution of

characteristics may also indicate massive extinctions

in lineages once rich in taxa. Should the majority

of taxa of such a lineage become extinct, the few

survivors would exhibit sparsely distributed,

relictual characteristics; these could be mistaken as

recently evolved on the basis of the criterion under

consideration. While relictual characteristics, among

extant taxa, are generally considered the exception,

their potential presence must be carefully considered when the criterion of frequency of occurrence is used. The application of this criterion,

and of some that follow, requires the assumption

that in evolutionary progression character divergence is more common than character convergence.

CRITERION OF CORRELATION TO TRANSFORMATION

SERIES.—Morphoclines, or morphological transformation series, in which the order and direction of

change is known, provide a means of predicting

character phylogeny of less understood morphoclines, or of states of two-state character systems

whose phylogenetic value is unpredictable by the

criterion of frequency of occurrence. Maslin (1952)

and Hennig (1966) discussed various ways by which

the relative apotypic and plesiotypic extremes of

transformation series can be inferred. In general,

the most practical way is to establish the most

plesiotypic extreme by the criterion of frequency

of occurrence and infer the opposite extreme to be
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HYPOSTOMAL

MARGIN

GULA

372

STEMMA

374

MANDIBULAR

GROOVE

LACINIA

MANDIBULAE

0.5 mm

FICURES 370-374.—Perilypus limbatus larva: 370, habitus; 371, head, ventral view; 372, head,

lateral view; 373, mandible, older larva (see p. 114); 374, mandible, younger larva. (Scales =

1 mm)
TRUNCATE

EPIPHARYNGEAL

SETA

EPIPHARYNGEAL

SENSILLUM

376

^ANTENNA L

APPENDIX

PEDUNCULATE

SETA

STIPES

CARDO

LATERAL PROSTERNAL PLATE

MID-LONGITUDINAL

PROSTERNAL

PLATE

FIGURES 375-379.—Perilypus limbatus larvae: 375, epipharynx; 376, antenna; 377, maxilla; 378,

thorax, ventral view; abdominal segments vm-ix, lateral view; 379, abdominal segment ix.

(Scale = 1 mm)
124

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

K.D.

FIGURES 380-382.—Perilypus limbatus pupae: 380, lateral view; 381, ventral view.

Placopterus thoracicus: 382, mandible. (Scales = 1 mm)
NUMBER 227 125

apotypic. The state, or states, between the apotypic

and plesiotypic extremes are considered phylogenetic intermediates. In Hennig's phylogenetic

system, intermediate (or transitional) character

states sometimes provide the only clue for inferring

sister-group relationships (Erwin 1974, fig. 161).

CRITERION OF CHOROLOGICAL PROGRESSION.—Chorology may be used as an index of character state

classification only after the spatial origin of a taxon

has been postulated. When this has been done

structural character states may be correlated with

distributional character states. In general, and in

the absence of contrary evidence, one assumes that

characteristics of relictual taxa are plesiotypic and

that those of taxa from derivative geographical

regions are apotypic. Hennig (1966:95) provided

the terms apochory and plesiochory to express,

respectively, the relative derived and primitive distributional states in chorological transformation

series.

CRITERION OF CORRELATION WITH ADAPTIVE SIGNIFICANCE.—As noted by Munroe (1974:76), "inferred adaptive significance is usually present, but

hidden, in discussions of the distribution of character states in taxa." While it is often difficult to

postulate, much less demonstrate, the adaptive

significance of a substantial number of character

states, most groups of organisms evolved some

character complexes whose adaptive function is

fairly obvious. Character states judged apotypic on

the basis of adaptive significance can be correlated

with characteristics whose adaptive significance is

cryptic, or whose phenotypic expression may not

directly reflect adaptation. The latter kind of

characteristic may represent manifestations of

genetic mechanisms not related to "immediate"

adaptive needs. For example, their expression may

be due to genetic pleiotrophy or based on genetic

recombination generated by drift.

The substantial number of mimetic patterns

(obviously adaptive) present in Cleridae will make

the criterion of correlation with adaptive significance particularly useful for inferring character

phylogeny. Munroe (1974:76) has commented

about the credibility of such inferences involving

characters pertinent to mimicry and to warning

coloration.

CRITERION OF DEVIATION FROM A BASIC PLAN.—

Schlee (1969) and Munroe (1974) discussed this

criterion in considerable detail. According to my

interpretation, the basic tenet involves a search for

plesiotypic states (particularly of transformation

series) of structurally rich characters that in aggregate are considered the basic or ancestral plan. It

is assumed that the relatively apotypic states of

extant descendents ultimately evolved from that

plan. The basic plan (or aggregate of plesiotypic

characteristics) is usually established by the plesiotypic extreme of transformation series or by the

criterion of frequency of occurrence. The taxonomic level at which the ancestral plan is established is, of course, dependent on the taxonomic

level of the group under study and on the categorical level at which such a plan can be credibly

established.

It may be particularly difficult to recognize the

basic plan in recently evolved groups or in those

where the higher classification is not well understood, because in such cases recognition and

interpretation of transformation series may be

particularly difficult. Concordant transformation

series involving complex organ systems are particularly important in this method of establishing

character phylogeny; mainly because it is the

plesiotypic extreme of such series that lends credibility to postulations of what the basic plan is.

ZOOGEOGRAPHIC METHODS

While hypotheses about speciation and phylogeny

have been organized into comprehensive theoretical

models (of species by Mayr [1942], of phylogeny by

Hennig [1966]), those about historical zoogeography

remain unconsolidated. The Darwinian concepts

of center of origin and of dispersal of species

remain the most practical tools for predicting

ancestral distributions. Croizat, et. al. (1974), recently criticized these concepts and provided alternative guidelines for historical biogeography at

biotal level.

According to Croizat, et al. (1974:265), "the general features of modern biotic distributions have

been determined by subdivisions of ancestral biotas

in response to changing geography." I do not reject

the concepts discussed by these authors; indeed the

idea that contemporary macrobiotic distributions

resulted from fragmentations of ancestral biotas is

quite consistant with past geologic and climatic

events, such as continental drift and the less catastrophic events leading to continental refugia. On
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the other hand, I cannot accept their total rejection

of the concept of center of origin and its corollary,

dispersal of species, primarily because faunal fragmentation predicates faunal expansion (expansion

accomplished by various possible mechanisms of

dispersal). It seems to me that a highly vagile species

is capable of at least fortuitous dispersal during its

history, particularly in regions such as Central

America where vicissitudinous climates have

undoubtedly affected dispersal potentials. The

question that remains, therefore, is this: Which of

the aforementioned approaches to historical zoogeography is most useful? I submit that both have

merit, but generally at different biotic and/or

taxonomic levels. Irrespective of biotic or taxonomic

level, it is the nature of the available data that

usually determines the approach to historical

zoogeography.

In general, zoogeographic relationships within

Perilypus are discussed in terms of faunal limits

and vicariance, and because the distributions of

most of the species remain poorly documented the

treatise is brief and fragmentary. Postulations about

historical zoogeography are based on the concept of

center of origin and I assume that vicariance

reflects dispersal. Distributions of the South American species are correlated with Quaternary refugia

postulated by Haffer (1969), and Williams (1970),

and Brown (1975).

Phylogenetics and Zoogeography of Perilypus

INTERGENERIC RELATIONSHIPS

Earlier in this paper (p. 115), I suggested the

possible sister group relationship between the

genera Placopterus and Perilypus, based primarily

on the presence of the lacinia mandibulae in the

larvae. I cannot offer additional statements

regarding intergeneric kinships based on potential

synapotypic similarity. Otherwise, I can comment

that there are at least three Central American

genera (Colyphus, Clerosoma, and Blaxina) whose

phylogenetic placement probably belongs close to

Perilypus and Placopterus. Perhaps it will be necessary to first study the larval stages and adult

internal anatomy before these genera are placed

into credible phylogenetic perspective.

RELATIONSHIPS AMONG THE SPECIES GROUPS

Figure 383 illustrates the proposed phylogeny

of the species groups and Table 1 lists the character states on which the reconstruction is based. The

scheme is provisional, as all proposed phylogenies

must be, and is weakened by several convergences

and reversals. Nevertheless, 1 feel that the analysis

broadly reflects what might have transpired during

the early history of Perilypus; at the very least a

foundation of relationships is presented, one that

is open to the challenge of subsequent discoveries

and resynthesis.

The Placopterus-Perilypus stock probably differentiated in northern Central America perhaps

during mid-Tertiary time when northern Central

America was discontinuous with a series of volcanic

islands (now southern Central America) and isolated by a wide sea gap from South America

(Malfait and Dinkelman, 1972, fig. 3; Ball, 1975,

1,16 l2b,8,15b

4,5^,7,12,24

25,26,27

19,18,31

FICURE 383.—Proposed phylogeny of the Perilypus species

groups (numbers refer to apotypic character states listed in

Table 1).
NUMBER 227 127

TABLE I.—Plesiotypic and apotypic character states used in Figure 383 (references

indicate the criteria by which the character states are judged apotypic)

Number

1

2

2a

21)

3

4

5

6

7

8

9

10

11

12

13

14

15

15b

16

17

18

19

20

21

22

23

24

25

26

27

28

29

Character

$ody depth

Head

antenna

articles 7-8

interocular depression

Thorax

subapical depression

depth

subapical depression

shape

mesoscutellum

Elytra

form

puncture shape

puncture depth

puncture distribution

sutuial cleft

microsculpture

epipleural fold

apical slope

disc color

discal vitta

Legs

front trasal claws

Pygidium

Male genitalia

Tegmen

pigmentation

phallobase width

parameral lateral depression

parameral mesodorsal

margin

phallobasic pigmental

bands

combined length of paramere and phallobasic

apodeme

First connecting membrane

Phallus

phallic plate shape

marginal denticles

Female genitalia

dorsal lamina

Character state

Plesiotypic

shallow (Fig. 260)

clubbed (Fig. 80)

without microsetae (Fig.

60)

not crescemic

strongly impressed

strongly sinuous (Fig. 110)

subquadrate

rectangulate (Fig. 2)

not favous

deep

throughout disc (Fig. 2)

not right angle (Fig. 27)

absent

not convex

gradual (Fig. 30)

predominantly black or

piceous

absent

symmetrical

entire (Fig. 39)

strongly pigmented

normal (Fig. 82)

present

not denticulated

absent

more than 2

serrulated scales absent

normal (Fig. 46)

absent

trilobed (Fig. 71)

Apotypic

deep1

serrate2 (Fig. 96)

moderately serrate

 3

(Fig- 8)

boldy serrate • (Fig. 284)

with microsetae "• *

(Fig. 19)

crescent ic'

feebly impressed *

feebly sinuous

 l (Fig. 342)

transverse

 l

ovatex (Fig. 300)

favous *

shallow'

absent from apical third

or more'

right angle * (Fig. 309)

presentl (Fig. 307)

convex • (Fig. 163)

acute» (Fig. 29)

very gradual '• * (Fig. 31)

predominantly castaneous

or testaceous *

present * (Fig. 2)

asymmetrical ••4

emarginate * (Fig. 94)

feebly pigmented *• *

very broad ' (Fig. 72)

absent *• •

denticulated ••" (Fig. 282)

present •• • (Fig. 344)

less than 2 «• *

serrulated scales present'

uniformly slender *• *

(Fig. 296)

present *• * (Fig. 46)

bilobed •• • (Fig. 49)
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Continued

Number

30

31

32

32a

Character

coxital plates

Male internal reproductive

organs

anterior accessory gland

Distribution

Character state

Plesiotypic

absent (Fig. 198)

not coiled

northern Central America

Apotypic

present • (Fig. 48)

coiled1 (Fig. 51)

southern Central

Americaa

South America *

1

 Frequency of occurrence within Perilypus.

2

 Frequency of occurrence within Clerinae.

1

 Correlation with transformation series.

* Correlation with adaptive significance.

* Deviation from a basic plan.

* Chorological progression.

fig. 119). In that region (north of Nicaragua) occur

the more primitive species of Perilypus, all the

species of Placopterus, and most species of the other

genera closely related to Perilypus. The hypothesis

of a Central American origin of Perilypus requires

the assumption of three independent southern dispersals, and the penetration into South America by

at least one species during each southern movement.

Ancestral Perilypus evolved autapotypic favous

elytral punctations, asymmetrical front tarsal claws,

and coiled male anterior accessory glands. The

second character state was subsequently lost in

several lineages, that is, assuming its genotype is

relatively uncomplicated and easily reversed;

otherwise one would have to make the unlikely

assumption that asymmetry of the front tarsal claws

evolved independently in five out of eight major

lineages. Ancestral Perilypus differentiated into two

lineages, one giving rise to the frontalis stock and

the other to the basic stock of the remaining

species. The stem species of the frontalis group

apparently remained in northern Central America,

evolved an apotypic feebly pigmented tegmen, and

very broad phallobase, and lost the parameral

lateral depression. The first and third of these

characteristics are also present in specimens of

P. levis (viridipennis group) that differ most conspicuously from members of the frontalis group by

having apotypic boldly serrate antennae. In this

scheme, I assume that genital characteristics of

P. levis evolved independently from those of the

frontalis group, and that antennal serration is more

likely to reveal patristic kinships than the aforementioned genital characteristics. Antennal serration undoubtedly developed to maximize surface

area for forebody sensory organs and thus is a

highly adaptive character, one that is probably

genetically complex and less likely lost or evolved

in parallel than genital-phallobasic characteristics

(the reverse would have to be assumed if P. levis

were to be placed in the frontalis group). The

diverging sister lineage of the frontalis stock evolved

apotypic phallic marginal denticles and female

genital bilobed dorsal lamina. The marginal denticles were secondarily lost in P. levis and in the

progenitor of the ornaticollis group.

The ancestor of the criocerides lineage also

remained in northern Central America, retained

an antennal club, and acquired a nearly impunctate

elytron and apotypic pygidium emarginate. Some

reduction of elytron punctation occurred independently in the viridipennis group, although in

that taxon the loss of deeply impressed punctations

involved only the elytral apical third. The complementary stem species evolved apotypic serrate antenna and microsetigerous antennal articles 7 and

8, and differentiated into the progenitor of the

limbatus-viridipennis stock and the one for the

remainder of the species.

An ancestor of the limbatus-viridipennis line

dispersed to southern Central America (exemplifying apochory; page 125) and subsequently penetrated South America. It evolved a convex epipleural fold, a characteristic retained by all

descendants except P. relucens; in that species the

epipleural fold became grooved longitudinally. A
NUMBER 227

129

slightly different convex epipleural fold is also

present in both species of the frontalis group and

in one species of each of the quadrilineatus and

ornaticollis groups. Therefore, convexity of the

epipleural fold may not be a strong indicator of

patristic affinities, but, in the absence of additional

evidence contrary to the proposed grouping, I tentatively consider the presence of that characteristic

in the limbatus-viridipennis assemblage an expression of monophyly. The monophyly of the limbatus

group is based on the presence of autapotypic

female coxital plates; the plates are only feebly

developed in the more primitive Central American

species. From the stock of the limbatus group

evolved the viridipennis group whose members

share apotypic impunctate elytral apical third (or

less). Although I found only one autapotypic characteristic for each of the limbatus and viridipennis

groups, the two lineages probably have been separated for a long time as the considerable habitus

differences between their extant members suggest.

The progenitor of the remaining species groups,

a vicariant with the ancestor of the limbatusviridipennis lineage, acquired apotypic moderately

serrate antenna, the second of two intermediate

states of a transformation series that begins with a

plesiotypic clubbed antenna and ends with an

apotypic boldly serrate antenna. The ancestor of

this lineage also evolved elytral microsculpture

and subsequently differentiated, producing the

reventazon lineage and the stock ancestral to the

chaleloides, quadrilineatus, and ornaticollis groups.

The evidence that the reventazon lineage is

monophyletic is that its members share the apotypic

characteristics of an acute elytral-apical slope and

an elytral discal vitta. Exceptions involving character reversals involve P. orthopleuridus whose

elytral apical slope is gradual, and P. immitis and

P. crassus in which the elytral disc is apparently

avittate (I studied only a few specimens of the last

two species). The lineage complementary to ancestral reventazon evolved shallow elytral punctations

and subsequently differentiated into the basic stock

of the chaletoides lineage and the quadrilineatusornaticollis lineage.

Ancestral chaletoides developed a deep body and

predominantly castaneous elytral color. The progenitor of the complementary stock established two

diverse lines of mimetic species. The most obvious

anatomical reconstructions that materialized during

the evolution of these species were (1) transformation of a moderately deep and rectangular body

form to one that is conspicuously shallow and oval,

and (2) progression of antennal serration from

moderately serrate to boldy serrate. Perfection of a

Batesian mimetic interaction (probably applicable

to most of the mimetic species under consideration)

often requires that the incipient mimic species

evolve toward a model species morphologically and

ecologically. Thus the imitator and the model

usually look alike and occur together in places

where both are maximally exposed to the

selection agent (predator). These requirements

invite the assumption that ancestral quadrilineatusornaticollis negotiated a generalized ecological shift

from a relatively compact vegetation assemblage to

a more open one. Once the shift to a lampyroidtype habitat was initiated genetically pliant species

of Perilypus converged towards the lampyroid

models and concomitantly differentiated. In this

paper, under "Natural History" (p. 4), I provide

some evidence that an ecological shift did indeed

occur during the history of Perilypus.

The monophyletic status of the quadrilineatus

stock is based on apotypic parameral mesodorsal

denticulated margin, a characteristic independently

evolved in one species of the limbatus, viridipennis,

and reventazon groups. From the correlative ancestral species evolved the geographically widespread

ornaticollis group, the most derived species group

in the genus. The species of that group share the

following apotypic characteristics not duplicated

elsewhere in Perilypus: crescentic interocular depression; feebly sinuous pronotal subapical depression; transverse mesoscutellum; right angled

elytral sutural cleft; presence of phallobasic pigmental bands; combined length of paramere and

phallobase over length of phallobasic apodeme less

than 2; first connecting membrane with serrulated

scales; and a uniformly slender phallic-plate shape.

The extent of derivitiveness and the widespread

distribution of the ornaticollis group gives the impression that the group is much older than its

phylogenetic placement suggests. But the derivitiveness of the ornaticollis group may also be

interpreted as a manifestation of accelerated diversification, particularly when considering the positive

influence that ecological shifts might have on

evolutionary rate. The extensive geographic range

of the group is probably a reflection of the general
130

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mimetic interaction between ornaticollis species

and a variety of lampyroid models. A generalized

mimic is not restricted to the range of only one

model species as a more specialized mimic might

be. Therefore, the generalist can overlap the geographical range (become widespread) of several

model species without losing the protective benefits

of the mimetic interaction.

RELATIONSHIPS WITHIN THE SPECIES GROUPS

Patristic and geographic relationships within

each of the species groups are discussed and/or

illustrated to whatever extent the data allow

(Table 2). A fairly complete treatise is possible for

three of the larger species groups, namely the

limbatus, quadrilineatus, and ornaticollis groups.

The reventazon, frontalis, criocerides, viridipennis,

and chaletoides groups are treated briefly and in

discussion form only.

The two species of the frontalis group are very

similar anatomically, and probably are recently

derived: the species are sympatric in Guatemala and

in El Salvador, but P. frontalis extends northward

into southern Mexico and P. sensilis southward

into western Nicaragua; the two species may be

partial latitudinal complements. Perilypus frontalis

evolved an apotypic boldly convex female epipleuron and boldly asymmetrical male front tarsal claws.

Perilypus sensilis lost the asymmetry of the front

TABLE 2.—Plesiotypic and apotypic character states used in Figures 384-386

(references indicate the criteria by which the character states are judged apotypic)

Number

33

34

35

35a

36

37

38

39

39a

40

41

41a

42

43

44

44a

44b

45

46

47

48

Character

Head

gular process

antennal article 11

Pronotum

discal vitta

discal concavities

macrosculpture

transverse wrinkles

Mesoscutellum

Leg color

Elytra

Disc

contour

fascia

Margins

base or posthumeral

apical

Epipleural fold

contour

position

dorsal margin

narrow longitudinal

region proximal to

epipleural fold

Character state

Plesiotypic

rectangulate

not as long as articles 9

and 10 combined

not maculated

not furcate

absent

absent

absent

not stalked

subquadrate

femora concolorous or

bicolorous

plane or feebly convex

not bicristate

absent

avittate

arcuate

not sinuatotruncate

not upcurved

plane

lateral

not explanate

not acutely deflexed

Apotypic

filiform * (Fig. 308)

as long or longer than

articles 9 and 10

combined'

maculatedl (Fig. 293)

furcate' (Fig. 322)

presentJ (Fig. 300)

present*

present •

stalked • (Fig. 310)

triangular *

front and middle femur

concolorous, hind femur

bicolorous *• *

undulated *•4 (Fig. 293)

bicristate * (Fig. 300)

present *

with short vitta •

(Fig. 287)

truncate *• •

sinuatotruncate '•'

upcurved1

obliquely convex •

ventrolateral'

explanate *

acutely deflexed •• *
NUMBER 227

TABLE 2.—Continued

Numbe

49

50

51

52

53

54

55

56

57

58

59

60

61

62

v Character

Vertical seta

Male genitalia

phallic plate

phallic plate marginal

denticles

phallic apex

dorsal sinus

phallobase

Female genitalia

Ventral lamina

length

size

number of lobes

Dorsal lamina

median lobe

number of lobes

Bacculi

dorsal bacculus

contiguous dorsal bacculus

oblique bacculus

Character state

Plesiotypic

normal

not abruptly broadened

medially

far removed from phallic

apex

normal

normal

not explanate

normal (Fig. 131)

normal (Fig. 152)

three

absent

three

not contiguous (Fig. 151)

narrow

not fused with coxital

plate

Apotypic

exceptionally stout'

abruptly broadened

medially! (Fig. 328)

proximal to phallic

apex' (Fig. 130)

acute

reduced '•" (Fig. 338)

explanate1 (Fig. 134)

short'(Fig. 140)

minute2 (Fig. 150)

one1

presentl

seven'

contiguous1 (Fig. 171)

expandedJ (Fig. 180)

fused with coxital

plateJ (Fig. 140)

1

 Frequency of occurrence within Perilypus.

* Frequency of occurrence within Clerinae.

1

 Correlation with transformation series.

4

 Correlation with adaptive significance.

5

 Deviation from a basic plan.

" Chorological progression.

tarsal claws and acquired, in convergence, an

apotypic convex epipleural fold.

In the criocerides group, P. criocerides probably

ranges into southeastern Mexico; no exact locality

in Mexico is available for its sister species, P.

insectus. Except for the autapotypic characteristics

that define the group, the two species do not show

a close relationship to each other. The substantial

habitus differences between them probably reflects

the mimetic interaction between P. criocerides and

an unidentified species of leaf beetle. Perilypus

criocerides evolved apotypic denticulatad outer

margin of phallic plate.

My views of the relationships within the limbatus

group are illustrated in Figure 284. In the group, a

monophyletic group of four Central American species is vicarious with its South American sister

group. The progenitor of the Central American

vicarient evolved apotypic continguous dorsal

baculi, a characteristic convergent in P. reventazon.

130

FIGURE 384.—Proposed phytogeny of the limbatus species

group (numbers refer to apotypic character states listed in

Tables 1 and 2).
132

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Further differentiation of the lineage established

the basic stock of P. decoris characterized by an

apotypic unilobed ventral lamina (also found in

P. reventazon), and the sister stock characterized

by an apotypic acute phallic apex. The latter stock

subsequently gave rise to the latilira-prolixipenis

stock and the testaceicornis stock, which evolved

specializations relevant to leg color. Also in P.

testaceicornis the fused dorsal bacculi of the female

genitalia became expanded in the posterior half. I

did not find a synapotypic characteristic to substantiate the sister group relationship between P. latilira and P. prolixipenis, but their anatomical similarity suggest close relationship; the two species

have probably not been reproductively isolated for

very long.

The South American species of the limbatus

group are known only from the Andes ranging from

Venezuela to Peru; P. apocopatus is vicarious with

P. iris, and P. buga with P. acus. Perilypus limbatus

is particularly widely distributed extending from

northern Venezuela to southern Peru. The known

distributions of these species roughly coincide with

Quaternary Andean forest refugia postulated by

Haffer (1969) and Vanzolini and Williams (1970).

Assuming that the coincidental distributions

between limbatus group species and refugia have

some historical significance, we might assume that

the species in question arose in the Pleistocene

epoch when Andean forest refugia were presumably established. The paucity of external differences among the species, suggesting Recent origin,

lends support to that hypothesis. Ancient refugial

systems, probably also forestal, might also have

influenced the evolution of the Central American

members of the limbatus group. The known geographic occurrence of these species similarly coincides with montane forests, such as those found on

the Panamanian Chiriquf range and Cerro

Campana, which contain more or less coherent

biotas. It seems likely that Pleistocene climatic

changes (Vuilleumier, 1971) which profoundly

influenced South American forests also markedly

affected forest development in southern Central

America.

The South American species of the limbatus

group are more structurally homogeneous externally than their Central American relatives. The

basic stock evolved elytral fascia, an apotypic characteristic independently evolved in P. carbonarius

of the reventazon group and in P. virdipennis of

the viridipennis group. In specimens of these species, however, the fascia are very variable in shape

and may be poorly defined. This lineage subsequently evolved from an ancestral stock, in which

the phallic marginal denticles (of the male genitalia) became more proximal to the phallic apex,

and a basic stock that later differentiated into the

apocopatus-iris and limbatus-columbicus lineages.

I did not find an autapotypic character state for

the latter lineage which subsequently differentiated

into two species. One of these, P. columbicus,

acquired an apotypic explanate phallobase. The

apocopatus-iris lineage is inferred to be monophyletic because the member species share the

following synapotypic characteristics of the female

genitalia: an oblique bacculus fused with coxital

plate and short ventral lamina. The remaining two

taxa, P. buga and P. acus, are probably sister species, but this needs corraboration by synapotypy.

An apotypic characteristic of the female genitalia

present in specimens of P. buga is a dorsal lamina

with median lobe, and in those of P. acus a lateral

lobe of minute ventral lamina.

The sister taxon of the limbatus group is the

viridipennis group; in the group, P. viridipennis

and P. relucens are vicariants. The other known

species of this group, P. levis, ranges widely, occurring in southwestern Colombia, across southern

Venezuela, and eastward into the Guiana Highlands

of Brazil. The progenitor of the group diverged,

producing the levis and viridipennis-relucens lineages. This differentiation probably took place early

in the history of the group; the extant taxa of the

two lineages are very different structurally. The

levis lineage lost the phallic marginal denticles but

evolved in convergence the boldly serrate antenna,

feebly pigmented male genitalia, and the absence of

the lateral depression of the paramere, as well as

the nonconvergent characteristic spatulate tegmen.

I found no unequivocal autapotypic charactristic

for the viridipennis-relucens stock, except perhaps

the predominantly refescent color that is present in

most specimens of the two extant species.

In the reventazon group several species range

widely in northern Central America: P. carbonarius

and P. caliculus occur in Mexico from Sonora to

Oaxaca and apparently are quite abundant in the

Transvolcanic Sierra; P. orthopleuridus is most

prevalent in southeastern Mexico and extends into
NUMBER 227

133

southern Honduras. Most of the other species

appear to be localized in highlands of either central

or southern Mexico, central Guatemala, central

Costa Rica, or northern Panama; P. crassus is the

only species of the group indigenous to South

America.

In aggregate the species of the reventazon group

are very similar externally; consequently, little can

be stated about their patristic relationships. The

species differ substantially in characteristics of the

genitalia, however, but because aedeagal differences

are so pronounced genital character states are generally unsuitable for establishing sister groups; thus

no phylogeny is postulated. Despite these shortcomings some brief statements about relationships

can be made.

The robustness of the male pygidium may be

used to roughly distinguish two clusters of species.

In specimens of P. orthopleuridus, P. calcaris, P.

exilis, and P. rerroazul the pygidium is quite robust (Figure 229) while in the remainder of the

species it ranges from small (Figure 245) to moderately large (Figure 39). I cannot decide to what

extent these groupings reflect phyletic relationships, primarily because the pygidium of some of

the species, such as P. reventazon, is difficult to

group according to size. Also difficult to interpret

are the various degrees of asymmetry of the front

tarsal claws, despite meristic comparisons. The

front tarsal claws are exceptionally asymmetrical in

P. orthopleuridus, P. calcaris, P. exilis, and P. immitis.

There are two particularly discordant species in

the reventazon group, P. orthopleuridus and P.

crassus. Specimens of the first species differ most

conspicuously from members of the other species

of the reventazon group by the following character

state reversals: gradual apical slope of the elytron

and the nonarenose elytral surface. The sister

species of P. orthopleuridus is probably P. calcaris;

specimens of these species share an apotypic mesoventral margin of the emarginate paramere, an

acuminate phallus, inflexed outer margin of the

phallic plate and a very long female genital coxite.

P. crassus, the only South American species of the

group, is notably different because its specimens

have the antenna exceptionally serrate and the

elytron truncate.

The species of the chaletoides group are known

only from Mexico; P. chaletoides are mostly from

the southwest and P. bicolor extends northward

to Durango. The ancestors of the chaletoides group

produced two structurally similar species that are

markedly different in elytral color. Perilypus chaletoides acquired a dark streak along the suture of

an otherwise castaneous elytron while P. bicolor

evolved totally stramineous elytral coloration.

In the quadrilineatus group, P. quadrilineatus

ranges widely in eastern Mexico, but the species

also extends into southern Guatemala; P. telephoroides is vicarious with P. bicristatus. Figure 385

illustrates my views of the phylogenetics of the

quadrilineatus group. Perilypus quardilineatus,

characterized by an apotypic dorsal margin of the

explanate epipleural fold, was apparently the first

offshoot of the ancestral stock. The rest of the

species are derivatives of an ancestor that acquired

a flatter, more oval body and one that evolved the

apotypic characteristic, narrow longitudinal region

proximal to the acutely deflexed elytral posthumeral

margin. This ancestor subsequently differentiated

into the bicristatus-telephoroides and coroniformisfluctus lineages. The progenitor of the first lineage

attained a coarsely macrosculptured pronotum and

subsequently diverged, producing P. telephoroides

and the highly derivative P. bicristatus. Apotypic

character states present in P. bicristatus include a

bicristate elytral disc, triangular scutellum, and

pronotal discal concavities. Perilypus telephoroides

23

FIGURE 385.—Proposed phylogeny of the quadrilineatus species group (numbers refer to apotypic character states listed

in Table 2).
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

acquired the derivatives, the vittate base of the

posthumeral margin, and the transversely wrinkled

pronotum. The elytral surface became undulated

in the complementary progenitor which later diverged into P. fiuctus and P. coroniformis; the latter acquired an apotypic maculated pronotal vitta.

Elytral undulations evolved secondarily in P. ventralis of the ornaticollis group.

In the ornaticollis group two species range widely.

Perilypus ornaticollis occurs in southeastern Mexico, extends northward along the Mexican east

4,5,6,7,12,24,

|25,26,2> '

FIGURE 386.—Proposed phylogeny of the ornaticollis species

group (numbers refer to apotypic character states listed in

Table 2).

coast, and follows a northern track that spans the

Mississippi and Ohio River drainage systems. Perilypus ventralis extends from north of the Isthmus

of Tehuantepec to central Guatemala; P. floralis

is vicarious with the South American P. sinuapicis.

The hypothetical phylogenetic reconstruction of the

ornaticollis group is illustrated in Figure 386. The

most specialized species in the group, and perhaps

in the genus, is P. ventralis, which anatomically

stands far apart from other members of the ornaticollis group. The apotypic character states that

define this early derivative are: a filiform gular

process, stalked mesoscutellum, heptalobed dorsal

lamina, plus various modifications of the elytra including a ventrolateral epipleural fold and the

convergent characteristic undulated elytral surface.

Some of these character states of P. ventralis are

undoubtedly anatomical specializations relevant to

the mimetic existence of that species, particularly

those of the elytra.

The complementary stock of the P. ventralis

lineage acquired a reduction of the dorsal sinus of

the male genitalia and subsequently diverged producing the ornaticollis and the progenitor of the

remaining species. The ornaticollis lineage evolved

at least two apotypic characteristics, furcate pronotal vitta and an abruptly medially broadened

phallic plate. Its sister stem species evolved a truncated elytral apex and subsequently differentiated

into the prolixicornis-galbeus stock and a line in

which the elytral apex became sinuatotruncate.

Article 11 of the antenna became prolonged in the

prolixicornis-galbeus line, which then differentiated

into P. prolixicornis and P. galbeus. The first offshoot of the remaining stock was P. pilatus, which

is characterized by apotypic robust elytral vertical

setae. The remaining three species are derivatives

from a progenitor that acquired an obliquely convex epipleural fold and one that later differentiated

into the antarius and floralis-sinuapicis lineages.

Perilypus antarius acquired an apotypic upcurved

elytral apex. I did not find an autapotypic character

state that may have been present in the ancestor of

P. floralis and P. sinuapicis.
Abstract in Spanish

Se redefine, reclasifica e ilustra el g^nero americano Perilypus. Se sumariza la

historia natural del grupo. Las especies prosperan, por los menos, en tres tipos

de macrohabitat, que son los relativos al roble, a lianas y a plantas herbaceas.

Los escarabajos de cuerpo rectangular habitan en robles y lianas, mientras que

los de cuerpo ovalado viven en hierbas. Se asume que la correlaci6n habitatperfil del cuerpo se debe a interacciones mime'ticas. Los Perilypus mime'ticos se

consideran generalistas que en cuanto a caracterfsticas de aspecto y habitos han

evolucionado simulando mas de un modelo repugnante.

Las especies de Perilypus se encuentran desde el nivel del mar hasta los 3400

metros de altitud, pero la mayoria de ejemplares conocidos se colectaron entre

1000 y 2000 metros. Observaciones de laboratorio prueban que los escarabajos

de Perilypus son extremamente predatores, siendo factores importantes en la

limitation de su agresividad el tamaflo y la rigidez de las presuntas victimas.

Se discuten los criterios usados en el reconocimiento de las especies y de los

grupos infra y supra-gene'ricos; se describen las t£cnicas seguidas para las disecciones, ilustraciones y mediciones; se dan analisis morfologicos de los principales

organos externos e internos, y se ofrece una clave de las especies y de sus grupos

basada en caracteres de adultos. Las fases de larva y ninfa del ge"nero se describen

por vez primera. Se dan los caracteres para cada grupo de especies. Para cada

especie se dan sinonimia, combination diagn6stica, descripci6n, discusi6n de

la variation estructural y cromatica, se dan sinonimia, combinaci6n diagn6stica,

description, discusi6n de la variaci6n estructural y cromatica, discusidn de su

historia natural, distribution geografica, derivation etimol6gica, lot alidades,

notas generales e ilustraciones. Mapas de distribuci6n con simbolos facilitan la

visualizaci6n de las areas geograficas de especies y sus grupos.

En total se reconocen 49 especies en Perilypus y 30 de estas se describen como

nuevas. Se establecen 12 nuevas sinonimias.

Se postula una filoge'nesis de Perilypus usando el m£todo Hennigiano de

analisis filogenetico. Se expone la zoogeografia del g£nero en t&rninos de lfmites

faunisticos y vicarismo. Se discuten sumariamente conceptos de zoogeograffa

historica ("rutas faunisticas", "centro de origen y dispersi6n" y "refugios forestales") y dos de ellos se utilizan para explicar la distribuci6n de los taxa de

Perilypus existentes. Las consideraciones sobre las afinidades entre diversos

groupos varia segun los datos asequibles. Se presenta un estudio bastante completo de los grupos mayores de limbatus, quadrilineatus y ornaticollis, en cambio

el tratamiento del grupo reventazon, el mayor del g^nero, es muy fragmentario.

Es probable que el Perilypus ancestral se desarroll6 en el norte de America

Central durante el Terciario medio evoluci6nado al fin en ocho grupos especificos. Los progenitores de los grupos frontalis, criocerides, chaletoides y quadrilineatus permanecieron en norte de Centro America asi como tambie"n sus

descendientes. Los ancestros de los grupos limbatus y viridipennis se dispersaron

hacia el sur de America Central; algunos miembros de ambos grupos penetraron

ultimamente en Sur America. El grupo ornaticollis, el mas evoluci6nado en el

ge"nero, y el grupo reventazon, especialmente, son de gran dispersi6n geografica.

135
Literature Cited

Ball, G. E.

1975. Pericaline Lebiini: Notes on Classification, a Synopsis of the New World genera, and a Revision of

the Genus Phloeoxena Chaudoir (Coleoptera: Carabidae). Questiones Enlomologicae, 11 (2): 143-242.

Boving, A. G., and A. B. Champlain

1920. Larval of North American Beetles of the Family

Cleridae. Proceedings of the United States National

Museum, 57(2323):575-649.

Brown, K. S., Jr.

1975. Geographical Patterns of Evolution in Neotropical

Lepidoptera: Systematics and Derivation of Known

and New Heliconiini (Nymphalidae: Nymphaline).

Journal of Entomology, series B, 44(3):201-242.

Brundin, L.

1966. Transarctic Relationships and Their Significance as

Evidenced by Chironomid Midges: With a Monograph of the Subfamilies Podonominae and Aphroteniinae and the Austral Heptagyiae. Kongliga

Svenska Vetenskapsakadaniens Handlingar, (4)11:1-

472.

1968. Application of Phylogenetic Principles in Systematics

and Evolutionary Theory. Pages 473—495 in T.

0rvig, editor, Proceedings of the 4th Nobel Symposium: Current Problems of Lower Vertebrate Phytogeny. Stockholm: Almqvist & Wiksell.

1972. Evolution, Causal Biology, and Classification. Zoologica Scripta, 1:107-120.

Campau, E. J.

1940. The Morphology of Chauliognathus pennsylvanicus

(DeGeer) (Coleoptera: Cantharidae). Microentomology, 5(3):57-85.

Chevrolat, M. A.

1843. Coleopteres du Mexique. Magasin Zoologie, series 2

(Insectes), 5:1-26.

1874. Catalogue des Clerides de la collection de M. Chevrolat. Revue et Magasin de Zoologie, series 3,

2(6):252-329.

1876. Mimoire stir la famille des Cle'rites. 51 pages. Paris.

Cholodkovsky, N.

1913. Necrobia ruficollis in St. Petersburg. Zoologischer

Ameiger, 42(12):529-531.

Corporaal, J. B.

1948. Nineteen Notes on Systematics and Synonymy. Entomologische Berichten, 12(281 ):243-246.

Croizat. L.. G. Nelson, and D. Rosen

1974. Center of Origin and Related Concepts. Systematic

Zoology, 23(2): 265-287.

Crowson, R. A.

1944. Further Studies on the Metendosternite in Coleoptera. Transactions of the Royal Entomological Society of London, 94:273-310.

1964. A Review of the Classification of Cleroidea (Coleoptera), with Descriptions of Two New Genera of

Peltidae and of Several New Larval Types. Transactions of the Royal Entomological Society of London, 116(12):275-327.

1970. Classification and Biology. 350 pages. London: Heinemann Educational Books.

Darlington, P. J., Jr.

1970. A Practical Criticism of Hennig-Brundin "Phylogenetic Systematics" and Antarctic Biogeography.

Systematic Zoology, 19(1):1—18.

Desmarest, E.

[I860.] Cleriens. Pages 226-279 of part 2 in J. C. Chenu,

Encyclopedic d'Histoire Naturelle. Paris: Libraire

de Firmin-Didot et C".

Doyen, J. T.

1966. The Skeletal Anatomy of Tenebrio molitor (Coleoptera: Tenebrionidae). Miscellaneous Publications of

the Entomological Society of America, 5(3):102-150.

Dury, C.

1906. Ecological Notes on Some Coleoptera of the Cincinnati Region including Seven New Species. Journal of the Cincinnati Society of Natural History,

20:151-156.

Ehrlich, P. R., R. R. White, M. C. Singer, S. W. McKechnie,

and L. E. Gilbert

1975. Checkerspot Butterflies: A Historical Perspective.

Science, 188(4185):221-228.

Ekis, G.

1975. Taxonomic and Nomenclatural Status of Clerid

Taxa Described by Massimiliano Spinola (1780-

1857) (Coleoptera, Cleridae). Bollettino del Museo

di Zoologia dell'Universita di Torino, 1:1—80.

Ekis, G., and A. P. Gupta

1971. Digestive System of Cleridae (Coleoptera). International Journal of Insect Morphology and Embryology, l(l):51-86.

Erwin, T. L.

1974. Studies of the Subtribe Tachyna (Coleoptera: Carabidae: Bembidiini), Part II: A Revision of the New

World-Australian Genus Pericompsus Le Conte.

Smithsonian Contributions to Zoology, 162:1-96.

Evans, M. E. G.

1961. The Musculature and Reproductive Systems of

Atomaria ruficornis (Marsham) (Coleoptera, Cryptophagidae). Transactions of the Royal Society of Edinburg, 64(14):297-399.

Ferris, G. F.

136
NUMBER 227

137

1940. The Morphology of Plega signata (Hagen) (Neuroptera: Mantispidae). Microentomotogy, 5(2):33-49.

Forbes, W. T. M.

1925. The Second Abdominal Pleurite in the Higher

Coleoptera. Psyche, 32(6):290-292.

Foster, D. E.

1976. Revision of North American Trichodes (Coleoptera: Cleridae). Special Publications of the Museum,

Texas Tech University, 11:1-86.

Gemminger, M., and E. Harold

1869. Familia XL1II, Cleridae. Pages 1722-1759 of volume

6 in Gemminger and Harold, Catalogus Coleopterorum. Monaco: Sumptu E. H. Gummi.

Gorham, H. S.

1878. Descriptions of New Genera and Species of Cleridae,

with Notes on the Genera and Corrections of

Synonymy. Transactions of the Entomological Society of London, 1878(2):153-167.

1882. Fam. Cleridae. Pages 129-168 in part 2 (Coleoptera)

of volume 3 in Godman and Salvin, editors, Biologia

Centrali-A mericana.

1886. Supplement: Colyphus-Poecilochroa. Pages 335-337

in part 2 (Coleoptera) of volume 3 in Godman and

Salvin, editors, Biologia Centrali-Americana.

Haffei, J.

1969. Speciation in Amazonian Forest Birds. Science,

165:131-137.

Hennig, W.

1965. Phylogenetic Systematics. Annual Review of Entomology, 10:97-116.

1966. Phylogenetic Systematics. 263 pages. Urbana: University of Illinois Press.

Hlavac, T. F.

1972. The Prothorax of Coleoptera: Origin, Major Features of Variation. Psyche, 79(3): 123-149.

Holdridge, L. R., W. C. Grenke, W. H. Hathevvay, T. Liang,

and J. A. Tosi, Jr.

1971. Forest Environments in Tropical Life Zones: A

Pilot Study. 747 pages. New York and Toronto:

Pergamon Press, Oxford.

Kavanaugh, D. H.

1972. Hennig's Principles and Methods of Phylogenetic

Systematics. The Biologist, 54(3): 115-127.

Kluge, A. G., and J. S. Faris

1969. Quantitative Phyletics and the Evolution of Anurans. Systematic Zoology, 18(1): 1-32.

Lacordaire, J. T.

1857. Famille XLI: Clerides. Pages 415-496 of volume.4 in

Lacordaire, Histoire Naturelle des Insectes: Genera

des Coleopteres. Paris: Librairie Encyclopedique de

Roret.

LeConte, J. L.

1880. Short Studies of North American Coleoptera. Transactions of the Entomological Society of America,

8:194-195.

Lindroth, C. H.

1957. The Principle Terms Used for Male and Female

Genitalia in Coleoptera. Opuscula Entomologica,

22:241-256.

1969. The Ground-Beetles (Carabidae, excl. Cicindelinae)

of Canada and Alaska, Part I. Opuscula Entomologica (Lund), Supplement 35:I-XIVII.

Malfait, B. T., and M. G. Dinkelman

1972. Circum-Caribbean Tectonic and Igneous Activity,

and the Evolution of the Caribbean Plate. Bulletin

of the Geological Society of America, 83:251-272.

Marx, H., and G. B. Rabb

1970. Character Analysis: An Empirical Approach Applied

to Advanced Snakes. Journal of Zoology (London),

161:525-548.

1972. Phyletic Analysis of Fifty Characters of Advanced

Snakes. Fieldiana Zoology, 63:1-11.

Maslin, T. P.

1952. Morphological Criteria of Phylogenetic Relationships. Systematic Zoology, l(2):49-70.

Matsuda, R.

1970. Morphology and Evolution of the Insect Thorax.

Memoirs of the Entomological Society of Canada,

76:1-431.

Mayr, E.

1942. Systematics and the Origin of Species. 334 pages.

New York: Columbia University Press.

1969. Principles of Systematic Zoology. 328 pages. New

York: McGraw-Hill.

1974. Cladistic Analysis or Cladistic Classification? Zeitschrift fur zoologische Systematik und Evolutionsjorschung, 12:94-128.

Munroe, D. D.

1974. The Systematics, Phylogeny, and Zoogeography of

Symmerus Walker and Australosymmerus Freeman

(Diptera: Mycetophilidae: Ditomyiinae). Memoirs of

the Entomological Society of Canada, 93:1-183.

Pic, M.

1941. Opuscula martialis, II. "L'Echange," Revue Linneene (16 April). [See Cleridae listing on pages 9-13.]

Ross, H. H.

1974. Biological Syste/natics. 345 pages. Reading, Mass.:

Addison-Wesley Publication Co. Inc.

Schaeffer, C.

1904. New Coleoptera. Journal of the New York Entomological Society, 12:215-224.

Schenkling, S.

1898. Zehn neue Cleriden nebst Bemerkungen iiber schon

beschriebene Arten. Deutsche Entomologische Zeitschrift, 1898(2):361-368.

1903. Coleoptera Malacodermata: Fam. Cleridae. Fascicle

13 in P. Wytsman, Genera Insectorum. Bruxelles.

1906. Die Cleriden des Deutschen Entomologischen

National-Museums, nebst Beschreibungen neuer

Arten. Deutsche Entomologische Zeitschrift, 1:241—

320.

Schlee, D.

1969. Hennig's Principle of Phylogenetic Systematics, an

"Intuitive statistico-phenetic Taxonomy"? Systematic

Zoology, 18(1): 127-134.

1971. Die Rekonstruktion der Phylogenese mil Hennig's

Prinzip. 62 pages. Frankfurt.
138

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scotl, H.

1919. Notes on the Biology of Necrobia ruficollis, Fabr.

(Coleoptera, Cleridae.) The Annals of Applied

Biology, 6(2-3): 101-115.

Snodgrass, R. E.

1935. Principles of Insect Morphology. 667 pages. New

York: McGraw Hill.

Sokal, R., and P. Sneath

1963. Principles of Numerical Taxonomy. 359 pages. San

Francisco: Freeman, 359 pp.

Solervicens, A. J.

1973a. El genero Epiclines en Chile. Annales Museo Historia Natural Valparaiso, 6:161-186.

1973. A Revision del genero Natalis Castelnau (Coleoptera-Cleridae-Clerinae). Revista Chilena Entomologia, 7:233-247.

Spinola, M.

1841. Monographie des Terediles. Revue Zoologique par

La Socie'te Cuxnerienne, 4:70-76.

1844a. Essai Monographique sur les Cle'rites: Insectes

Coleopteres. Volume 1, 386 pages. Genes.

1844b. Essai Monographique sur les Cle'rites: Insectes

Coleopteres. Volume 2. 216 pages. Genes.

Tanner, V. M.

1927. A Preliminary Study of the Genitalia of the Female

Coleoptera. Transactions of the American Entomological Society, 53:5-50.

Thomson, M. S.

1860. Materiaux pour servira une monographie nouvelle

de la famille des Cle'rides. Musee Scientifique,

1860:46-67.

Tremblay, E.

1958. Studio morpho-biologico sulla Necrobia rufipes DeG.

Bollettino del R. Laboratorio di entomologia agraria

di Portici, 16:49-140.

Tuomikoski, R.

1967. Notes on Some Principles of Phylogenetic Systematics. Annals Entomologici Fennici, 33(3):137-

147.

Vanzolini, P. E., and E. E. Williams

1970. South American Anoles: The Geographic Differentiation and Evolution of the Anolis chrysolepis

Species Group (Sauria, Iguanidae). Arquivos de

Zoologia Sao Paulo, 19:1-298.

Vuilleumier, B. S.

1971. Pleistocene Changes in the Fauna and Flora of

South America. Science, 173:771-780.

Whitehead, D. R.

1972. Classification, Phylogeny, and Zoogeography of

Schizogenius Putzeys (Coleoptera: Carabidae: Scaratini). Quaestiones Entomologicae, 8:131-348.

Wolcott, A. B.

1910a. Family XLII, Cleridae: The Checkered Beetles.

In W. S. Blatchley, An Illustrated Descriptive Catalogue of the Coleoptera or Beetles (Exclusive of the

Rhynchophora) Known To Occur in Indiana. Bulletin of the Indiana Department of Geology and

Natural Resources, 1:846-862.

1910b. Notes on Some Cleridae of Middle and North

America with Descriptions of New Species. Zoological Series (Field Museum of Natural History Publication 144), 7(10):339-401.

1927a. A Review of the Cleridae of Costa Rica. Coleopterological Contributions, l(l):l-104.

1927b. Descriptions of a New Genus and Four New Species

of American Cleridae. Coleopterological Contributions, 1(1): 105-110.


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