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A revision of the Leptolycini (Coleoptera:Lycidae) with a discussion of paedomorphosis

Miller, Richard Stuart, Ph.D.

The Ohio State University, 1991

Copyright ©1991 by Miller, Richard Stuart. All rights reserved.

UMI 300 N. Zccb Rd. Ann Arbor, MI 48106

A REVISION OF THE LEPTOLYCINI (COLEOPTERA: LYCIDAE)

WITH A DISCUSSION OF PAEDOMORPHOSIS

DISSERTATION

Presented in Partial Fulfillment of the Requirements f

the Degree Doctor of Philosophy in the Graduate

School of The Ohio State University

by

Richard S. Miller, B.A., M.S.

*****

The Ohio State University

1991

Dissertation Committee: Approved By:

Dr. David J. Horn tA Dr. Donald E. Johnston

Dr. Charles A. Triplehorn Advisor

Dr. Barry D. Valentine Department of Entomology Copyright by Richard S. Miller 1991

I To Dr. Roy A. Crowson for his study and his provocative queries

i i ACKNOWLEDGEMENTS

How dull it is to pause, to make an end, To rust unburnished, not to shine in use! As though to breathe were life itself! -A. Tennyson, 1833.

The immensity of biology can either elate or overwhelm the student. There are, essentially, two responses to this conundrum.

One is the quixotic charge here and there at the fog of unexplained phenomena. The other is to succumb to a Faustian despair of ever encompassing the morass of little suspected knowledge. While fully appreciating the absurdity, I chose the former when most of my cohort were increasing their biological fitness (sensu Colinveaux, 1982:394).

This current milestone in my studies has given me pause to reflect on those who have contributed to my formal and informal education. It is impossible to differentiate adequately between them and, thereby, classify them - for they have made varying contributions of infinitely subtle distinction. A "lumper" might thank Western Civilization as probably more than 997. of my thoughts are a result of inculcation from this source. A "splitter" would begin a list of individuals.

Aristotle, a member of Western Civilization, has urged moderation and, so, I will mention a few stellar contributions.

Of course Niki comes to mind first. Besides her readily apparent artistic abilities demonstrated in this dissertation, her support and encouragement have significantly contributed to its conclusion and my

i i i happiness.

Again, my family has been a source of support, even though they kindly find my interests "different." I thank them for everything.

My friends and former office- mates of B & 2 309 have been inspirational in our discussions. These peers include Michael A. Ivie,

William F. Abeles, James B. Stribling, Paul S. Cwikla (really only

309-A), and recently Peter W. Kovarik. I thank Nick Calderone, an honorary member of our clique, who has done much to enrich my appreciation of philosophy and other arcane subjects.

The various members of the faculties of CSUS, UCD, and OSU who have contributed to my education are appreciated. Especially important among these are Harold Wiedmann and Rollo Darby; Martin C. Birch;

Charles A. Triplehorn, Barry D. Valentine, David J. Horn, Donald E.

Johnston, Norman F. Johnson, and J. Bruce Griffing respectively.

The staff of the OSU biology library has been immensely helpful.

I would like to thank them all, and especially Susan Ward, for their patience, understanding, and assistance.

Several of ray colleagues around the world have assisted in various aspects of my work including the loan of specimens. The latter are listed in the materials and methods section, although I would like to acknowledge all here.

This paragraph is reserved for my reading committee should they decide to pass me after the laborious task of reading this manuscript.

Additionally, I would like to thank Dr. David Horn for his question during my general exam on testability by use of hypothesized phylogenies. It was seminal to some of the ideas developed here.

iv Finally, I thank all those natural historians and scientists, whether trained or untrained, who for whatever motives have left their thoughts in print. If I have been remiss in failing to cite their work, I here apologize and can only plead the finite limitations of a human mind.

History beetle evolution and the significance of paedomorphosis in cantharoid evolution. I hope this relationship of ontogeny and phyiogeny can be examined more cogently by providing this arena for further study. If I have failed, the enclosed cladograms must be viewed merely as art to be stored in some attic. Art is the culmination of human imagination, but it was not my purpose.

v VITAE

15 February 1945 ...... Nativity, Los Angeles, California

1970 — 1976 ...... Bachelor of Arts in Biology with a concentration in Zoology, California State University, Sacramento

1975 - 1976...... University of California, Davis

1976 - 1981...... Economic Entomologist I- III, California Department of Food and Agriculture, Sacramento

1981 - 1985 ...... Master of Science in Entomology, The Ohio State University, Columbus

1988 -1989 ...... Insect Identifier, United States Department of Agriculture

1985 - present ...... Ph.D. in Entomology, The Ohio State University, Columbus

PUBLICATIONS

1984. The Buprestidae of the Virgin Islands. Florida Entomologist 67(2 ):288“300. (co-authored with M. A. Ivie)

1988. The behavior of Calooteron reticulatum (F. ) larvae (Coleoptera: Lycidae). Ohio Journal of Science 88(3 ):115— 116-

FIELDS OF STUDY

Major Field: Entomology

vi TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... iii

VITA ...... vi

TABLE OF CONTENTS ...... ix

LIST OF TABLES ...... xi

LIST OF FIGURES ...... xi i

INTRODUCTION ...... 1

MATERIALS AND METHODS ...... 3

Methods ...... 3

Systematic analysis ...... 3

Taxonomic conventions ...... 6

Paedomorphosis ...... 7

Material studied ...... 9

ONTOGENY AND PHYLOGENY ...... 12

Introduction ...... 14

Definition of the problem ...... 18

An example ...... 19

A model ...... 30

Synergism of morphology and systematics ...... 34

Phylogenetic implications ...... 37

Possible taxa for further study ...... 40

PHYLOGENETIC RELATIONSHIPS WITHIN THE CANTHAROIDEA ...... 45

vi i Relations within the Elateriformia ...... 47

Monophyly of the Cantharoidea...... 49

Current hypotheses of Cantharoid relationships ...... 50

Family relations within the Cantharoidea ...... 51

Omalysidae ...... 51

Plastoceridae ...... 53

Cantharidae ...... 54

Omethidae ...... 55

Phengodidae ...... 55

Telegeusidae ...... 57

Dri 1 idae ...... 57

Lampyridae ...... 58

Lycidae ...... 59

THE TRIBE LEPTOLYCINI ...... 62

Key to Genera of Leptolycini ...... 67

Genus Abrolvcus New Genus ...... 69

Genus Anti 1 lolvcus New Genus ..... 109

Genus Ceratoprion Gorham ...... 137

Genus Flabel locaenia Pic ...... 175

Genus Leptol vcus Leng and Mutch ler ...... 215

Genus Pseudacroleptus Pic ...... 259

Genus Pseudoceratoprion New Genus ...... 285

Phylogeny of the Leptolycini ...... 306

Monophyly of the Leptolycini ...... 307

Lycid relationships ofthe Leptolycini ...... 308

Hypothesized Sister Group ...... 312

vi i i Character discussion ...... 313

Relationships within the Leptolycini ...... 342

Phylogenetic Conclusions ...... 343

Biogeographical implications ...... 345

Concluding remarks ...... 349

LIST OF REFERENCES ...... 350

APPENDIX (Illustrations) ...... 361

ix LIST OF TABLES

TABLE

1. Character states found in Thvlodrias ...... 22

2. Possible results of a test as proposed by Lauder .... 35

3. Character states of Leptolycini ...... 314

x LIST OF FIGURES

FIGURE

1. Theoretically available phylogenetic reconstructions for

paedomporphosis CprogenesisD (after Westheide, 1987:850).

2. Models of possible ontogenetic pathways of a character.

3. Hypothesized phylogeny of the genera of the Leptolycini.

4. Abrolvcus bicolor New Species. a-c. Aedeagus - a, dorsal

aspect, b, lateral aspect, c, ventral aspect. d. Right

antenna. e. Right hindwing,

5. Abrolvcus cubensis New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

6. Abrolvcus darlingtoni New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.d.

Lateral aspect head and prothorax.

7. Abrolvcus iviei New Species. a-c. Aedeagus - a, dorsal

aspect, b, lateral aspect, c, ventral aspect.

8. Abrolvcus omalvsiformis New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax.

9. Abrolvcus sandersoni New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect of head and prothorax. e. Internal male

genitalia.

XI 10. Anti1lolvcus auratosuratus New Species. a-c. Aedeagus -

a, dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Last abdominal sternite. e-f. Ultimate and penultimate

abdominal tergites - e. internal view, f. external view.

11. Anti 11olvcus elongatus New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral view of head and thorax.

12. Anti 1lolvcus flavomarginatus New Species. a-c. Aedeagus -

a, dorsal aspect, b, lateral aspect, c, ventral aspect. a.

Lateral aspect head and prothorax.

13. Anti 1lolvcus semiflavus CChevrolat). a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

14. Ceratoprion bordoni New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Right antenna, e. Right hindwing.

15. Ceratoprion chaelae New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

16. Ceratoprion mandibularum New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

right antenna.

17. Ceratoprion nigrum New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Last abdominal sternite. e-f. Ultimate and penultimate

abdominal tergites - e. internal view, f. external view,

g. Internal male genitalia.

IS. Ceratoprion periosus New Species. a-c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect.

19. Ceratoprion serricorne Gorham. a-c. Aedeagus - a, dorsal

aspect, b, lateral aspect, c, ventral aspect. d. Lateral

aspect head and prothorax. e. Left hindwing. f. Left

antenna.

20. Flabellocaenia bourgeoisi Pic. a-c. Aedeagus - a, dorsal

aspect, b, lateral aspect, c, ventral aspect. d. Lateral

aspect head and prothorax. e. Last abdominal sternite.

e-f. Ultimate and penultimate abdominal tergites - e.

internal view, f. external view.

21. Flabel1ocaenia elegantulus New Species. a-c. Aedeagus -

a, dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax.

22. Flabellocaenia leticia New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

23. Flabellocaenia iolei New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax. e. Frontal view of

head.

24. Flabellocaenia pecki New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspecthead and prothorax.

25. Flabellocaenia rimae New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

26. Leptolvcus adiaphorus New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

xi i i Last abdominal sternite. e-f. Ultimate and penultimate

abdominal tergites - e. internal view, f. external view.

27. Leptolvcus brunneus New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax. e. Internal genitalia.

28. Leptolvcus dominicensis New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect.

29. Leptolvcus effeminatus New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax.

30. Leptolvcus flavicol1 is Leng and Mutchler. a-c.

Aedeagus — a, dorsal aspect, b, lateral aspect, c, ventral

aspect. d. Lateral aspect head and prothorax.

31. Leptolycus heterocornis Leng and Mutchler. Habitus of

adult male.

32. Leptolvcus heterocornis Leng and Mutchler. a-c. Aedeagus

- a, dorsal aspect, b, lateral aspect, c, ventral aspect,

d. Lateral aspect head and prothorax. e. Right hindwing.

33. Leptolvcus heterocornis Leng and Mutchler. Habitus of

adult female.

34. Lepto1vcus heterocornis Leng and Mutchler. Habitus of

1arva.

35. Pseudacroleptus caeruleus New Species. a-c. Aedeagus —

a, dorsal aspect, b, lateral aspect, c, ventral aspect.

36. Pseudacroleptus neblinus New Species. a-c. Aedeagus - a,

dorsal aspect, b, lateral aspect, c, ventral aspect. d.

xi v Lateral aspect head and prothorax. e. Rigth wing.

37. Pseudacro1eptus neblinus New Species. a. Thorax - dorsal

aspect on right and ventral aspect on left.

38. Pseudacro1eptus neblinus New Species. a. Head - ventral

aspect. b. Right maxillary palp.

39. Pseudacro1eptus obscuricolor Pic. Habitus of adult male

(Type).

40. Pseudoceratoprion bechvne New Species. a-c. Aedeagus -

a, dorsal aspect, b, lateral aspect, c, ventral aspect. d.

Lateral aspect head and prothorax.

41. Pseudoceratoprion brasi1iensis New Species. a-c.

Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral

aspect. d. Right antenna.

42. Pseudoceratoprion colombiensis New Species. a-c.

Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral

aspect. d. Lateral aspect head and prothorax.

xv INTRODUCTION

Although they have been the object of little recent study, lycids present some interesting questions in both their biology and phylogeny.

Presently, the literature of this group is confined largely to alpha taxonomy. Exceptions occur in the discussion of "trilobite" larvae culminating in the work of Mjorberg (1925), some discussion of mimicry

(Darlington, 1938; Parsons, 1940; Linsley et a l ., 1961; Emmel, 1965; and Burke, 1976), and a few papers on larval feeding (Mjorberg, 1925;

Withycombe, 1926; McCabe and Johnson, 1979 and 1980; Miller, 1988).

Because there has been little synthetic work on a global scale, phylogenetic relationships within the family are obscure. Kleine's catalogue (1933) is the last treatment of the entire family, but it is, perhaps, useful only as a list of described taxa in the New World.

Although he had worked on lycids for 26 years, Kleine’s studies were largely confined to the Old World, especially the Indo-Malaysian region. He published only two papers limited to alpha taxonomy on the

New World fauna (Kleine, 1941 and 1942). As a result, he had to rely heavily on the earlier work of Bourgeois (1905) and vague statements such as "voisin de" of Pic for the relationships of Neotropiaal members of the family.

A further complication in the systematics of the Lycidae is that much of the taxonomic literature is too brief and emphasizes only color characters which are often of little use. This has necessitated the

examination of types before names can confidently be placed on any

Neotropical specimens.

This chaotic state has done little to induce studies in the biology of the Lycidae and, thus, they remain largely unknown. What i known, however, is tantalizing. For example, some species are known t aggregate as adults or larvae, but no species is known to aggregate in both stages. Aggregations are often composed of numerous distantly related species and are thought to serve as models for mimetic complexes. Although lycids are thought to be an evolutionarily recent group (Crowson, 1972), several of the widespread and apparently basal groups have " larviforra" females which are incapable of flight.

The foregoing had prompted me to study one tribe within the

Lycidae, the Leptolycini, but eventually required the examination of ever larger problems that culminated in the following study of some aspects of cantharoid phylogeny and an examination of paedomorphosis. MATERIALS AND METHODS

The methodology discussed here serves two purposes. First, it

serves as a basis for evaluation of my work. Second, it references my

methodology so it need not be repeatedly discussed in fucure work except

in its difference from this approach.

Systematic Analvsis.

The data gathered in this study were analy2ed using the

methodology of phylogentic systematics as proposed by Hennig (1965,

1966) and developed by numerous other workers (see Wiley, 1981 and Ax,

1987 for a review and list of references and, also, recent issues of

Systematic Zoology, Systematic Botany, and Cladistics among others).

This method assumes that 1) evolution occurred, 2) speciation can be

recognized by novel characters, 3) relationships can be recognized by

the recency of these shared novelties (ie. synapomorphies), and 4)

multichotomies are rare. Therefore, in addition to a hypothesis of

character cladogenesis, the resulting tree is also a hypothesis of the

phylogenetic relationships within the taxon. These hypotheses will be

called phylogenetic hypotheses for brevity, but no ancestor - descendant

relationship is implied. A further assumption for transposition of phylogentic hypotheses into the Linnean classification is that evolution

is hierarchical.

3 Species Recognition. The principle method of recognizing species

was the morphological species concept. This implies that some

difference (eg. autapomorphy) was found consistently between populations

in which no intermediates were found to bridge the gap (ie. character

distributions are discrete, not continuous).

Exemplars. When analyzing higher taxa, exemplars are the only

feasible method of study. Their selection was largely a result of

availability of specimens, but attempts were made to find any aberrant

cantharoids for study. Besides the material borrowed from the various

museums and individuals mentioned in the acknowledgements, my study of character states within the Coleoptera was restricted to my personal

collection (RSMC) and the Ohio State University collection (OSUC). No attempt was made to list these specimens studied by repository, but the majority were from the RSMC.

Character Weighting. No character state was arbitrarily weighted. However, implicit weighting must have occurred inadvertently

by choice of characters examined, by availability of material, by human cognition, and by my cultural bias. Additionally, characters were

intentionally weighted a priori if they were the result of loss or reduction, if they were changes of degree, or if they were thought to be affected by paedomorphosis.

Loss was weighted because homology is notoriously difficult to document when it cannot be seen. Lineages supported by reduction or

loss character states were considered suspect and additional support was sought. If increased homoplasy resulted from their inclusion in the analysis as a synapomorphy, then the character states were recoded as unique novelties and the resulting data were reanalyzed. The rationale for such action was not to produce a more parsimonious cladogram, but to draw attention to other homoplasy that might provide more fruitful investigation. Acceptance of hypotheses was subsequently based on parsimony. The homology of recoded character states remains uncertain

(see below) and is so noted.

Presence - absence character states were sought preferentially for analysis. The use of those states resulting in a change of degree, such as shape, was minimized to those with only two states. If a third state is found, it is often not possible to polarize transformation series except by a priori assumptions of parsimony or aesthetics. A case in point is the presence of a square, a transversely rectangular, and a longitudinally rectangular pronotum. Even if the functional out group resolves the basal state, the remaining states have three possible transformation series which can be resolved for use as a synapomorphy only arbitrarily in the absence of studies of allometry during ontogeny.

Such states can, however, be used as autapomorphies.

Characters whose expression were demonstrated to be affected by paedomorphosis were not used in the analysis of paedomorphic species.

This rationale will be further discussed during the treatment of ontogeny and phylogeny (p. 12).

Homology. The homology of characters was hypothesized using the criteria of Remane (1956). These are the positional criterion (Remane,

1956:30), the similarity criterion (Remane, 1956:42), and the criterion of intermediates (Remane, 1956:45). These and the criterion of correlation of transformation series (Hennig, 1966:96) are accepted only as operational rules for pragmatic reasons during hypothesis formation.

The actual test of a hypothesis of homology can be ontogeny which is not

just "complementary to the criterion of out - group comparison" as

stated by Wiley (1981:154). The latter is a phylogenetic criterion

sensu Hennig, but ontogeny can be an actual independent test subject to

falsification sensu Nelson (1978).

Polarization. The outgroup method (Watrous and Wheeler, 1981 )

was used during analysis with consideration of the qualifications

discussed by Madison et a l . (1984). Use of Nelson's ontogenetic rule

(1978) is provisionally rejected because of the difficulty of assessing

its falsifiabi1ity (ie, differentiating between a more general state due

to basal position or one due to paedomorphosis ). Additionally, the

rule assumes a uniform transformation which may or may not be real.

When a sufficiently large data base becomes available, ontogenetic

character precedence may prove useful in analysis (see pp. 28 and 36) if

caenogenesis with polarizable character states can be demonstrated, but

it remains unexamined.

Taxonomic Conventions.

Diagnoses. A combination of both syraplesiomorphies and apomorphies was often used to construct diagnoses. Although the former has information content, it is not sufficient for diagnosis of a

species. Newly discovered taxa are more likely to have such characters

than apomorphies and, hence, may not be separable or even recognized.

Taxa, after all, are composed mostly of symplesiomorphous character

states. Keys. No attempt was made to construct phylogenetic keys.

Although such keys are arguably more stable, it must be remembered that the results of a phylogenetic analysis are only hypotheses. It seems unfair to user the reader to examine apomorphic character states that are perceived only with difficulty if more easily observed plesiomorphic characters are available. Again, additional data may refute the current hypotheses and necessitate an additional key.

However, the cladogram (Figure 3) can be so used.

Classification. The taxonomic hierarchy was constructed using the traditional Linnaean hierarchical method. As there is no known algorithm for deciding the limits of superspecific taxa, their delimitation is at the discretion of the specialist. The merits of combining taxa to show relationships instead of separating them to emphasise uniqueness (Ball 1975:149) were considered, but stability was also a major consideration. In all cases hypothesized monophyly was the final arbiter.

Subgenera are not used. They will only be retained when their names are already available and the lineage is still in need of analysis. Species groups suffice in other cases and do not clutter the literature unnecessarily.

PAEDOMORPHOSIS.

Paedomorphosis was examined to determine its impact on morphology and, hence, on phylogenetic analysis. Its presence was recognized by wingless, "larva-like" females. A species of Dermestidae amenable to culturing, Thvlodrias contractus Motschulsky, was selected for comparative study, because its phylogenetic relationships are well supported.

The methodology followed here is a documentation of character states found in both sexes in a paedomorphic species and subsequent comparison with character states found in other members of the same family. Study of the lineage was primarily restricted to adult material from the RSMC, but was supplemented by OSUC and MCZC material

Genera examined consisted of Dermestes L., Attagenus Latreille,

Novel si s Casey, Trogoderma Dejean, Globicornis Latreille, Megatoma

Herbst, Dearthrus LeConte, Anthrenocerus Arrow, Ctesias Stephans,

Thaumoglossa Redtenbacher, Hemirhopalurn Sharp, Orphinus Motschulsky,

Crvptorhopalum Gu6rin-Menevi1le, Anthrenus Schaeffer, Apsectus LeConte,

Trinodes Dejean Evorinea Beal, and Thvlodrias Motschulsky. This constitutes 18 of 43 genera in 7 of the 9 subfamilies recognized by

Mroczokowski <1968) excluding Orphilinae which has been moved to the

Nosodendridae by Ivie (1985:64). In the latter family, which is hypothesized to be the sister to the Dermestidae, both Orphi1 is

Erickson and Nosodendron Latreille were examined. Additional dermesti genera were not available, and the literature was consulted for their character states. Characters whose states are unknown in the rest of the Dermestidae are not discussed.

The differences in character state trends of the paedomorphic dermestid were then compared to those of other putative paedomorphic species. In a few cases specimens were available, but it was also necessary to consult the literature. Cantharoid * specimens available included Lamovris noctiluca L. male and females, Drilus and Selasia males (Drilidae), Omalvsus males (Omalysidae) and males of numerous genera of Phengodidae and the female of Phengodes. Also, Diphvllostoma fimbrata Fall (Scarabaeoidea ) in the personal collection of Michael A.

Ivie (MAIC) was examined.

Thvlodrias Mots, was not illustrated here as ample illustrations can be found throughout the literature. These include Hinton (1945: figs. 387-390), Franciscolo (1975: figs. 1-7), and Suss and Foggato,

1977-78: figs. 1-22).

As there is an insufficient data base to incorporate ontogenetic data in a phylogenetic analysis using the methodology of Nelson (1978 and 1986), alternative approaches were sought.

Material Studied

The following museums and collections provided material for study.

Examination of the material studied for each species of leptolycine will demonstrate the paucity of that material.

AMNH American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Dr. Lee H. Herman, Jr.

ANSP Academy of Natural Sciences of Philadelphia. 19th and The Parkway, Philadelphia, Pennsylvania 19103. Mr. Donald Azuma and Dr, Daniel Otte.

Even though my studies in progress corroborate Lawrence's (1987) action synonymizing the Cantharoidea with the Elateroidea, I will continue to use the former name for convenience during the discussion of those lineages formerly placed in the superfamily within this paper. 10

BMNH British Museum (Natural History). London, SW7 5BD, England, Great Britain. Dr. C. M. F. von Hayek.

CASC California Academy of Sciences, Goldengate Park, San Francisco, California 94118. Drs. David H. Kavanaugh and Norman Penny

CISC California Insect Survey, University of California, Berkeley, California 94118. Dr. John A. Chemsak

CNCI Canadian National Collection, Agriculture Canada, Biosystematics Research Institute, Ottawa, Ontario K1A 0C6. Mrs. Jean McNamara and Dr. Ales Smetana

FSCA Florida State Collection of Arthropods, Florida Department of Agriculture and Conservation Service. P.O. Box 1269, Gainesville Florida, 32601. Dr. Robert E. Woodruff.

HAHC Personal collection, Drs. Anne and Henry Howden. Carlton University, Ottawa, Ontario K1A 0C6

ICCM Carnegie Museum of Natural History. Pittsburgh, Pennsylvania 15213. Dr. John E. Rawlins

INHS Illinois Natural History Survey. 607 East Peabody Drive, Champaign, IL 61820. Dr. Donald W. Webb.

ISNB Institut Royal des Sciences Naturelles de Belgique, 29 rue Vautier, Brussels, Belgium.

IZAC Instituto de Zoologia, Academia de Ciencias de Cuba, Habana, Cuba.

JARC Personal collection, Dr. J. A. Ramos. Box 1046, Mayaguez, Puerto Rico 00708

MAIC Personal collection, Dr. Michael A. Ivie. Department of Entomology, Montana State University, Bozeman Montana 59717

MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 Dr. Scott R. Shaw

MHMB Naturhistorisches Museum. CH-4001 Basel, Augustinergasse 2, Switzerland. Dr. Michael Brancucci

MIZA Museo Instituto Zoologia Agricola, Universidad Central de 11

Venezuela, Maracay, Venezuela. Dr. J. Luis Joly T.

MNHP Musee National d ’histoire Naturelle, Paris. 45, Rue Buffon, 75005. Dr. Jean J. Menier

NMNH National Museum of Natural History, Smithsonian Institution, Washington, D. C. Drs. Robert Gordon and Paul Spangler.

OSUC The Ohio State University Collection of Insects and Spiders, Department of Entomology, Columbus, Ohio. Dr. Charles A. Triplehorn

RSMC Collection of author.

TAMU Texas A. & M. University Entomological Museum, Department of Ent ‘^ology, College Station, Texas 77843. Dr. Lorace R. Burke.

UCDC University of California, Davis Dr. Robert Schuster.

UPRM University of Puerto Rico, Mayaquez, Puerto Rico 00700. Mr. Rafael Ingles

ZMHB Zoologisches Museum, Museum fur Naturkunde, der Humboldt - Uni versitat zu Berlin, DDR 104, Berlin. Dr. Manfred Uhlig

I would also like to thank the following individuals for material

that was used in this study and that can now be found in the RSMC: Mr.

William F. Abeles, Mr. Patrick Bleuzen, Father Carlo Brivio, Dr. Fortun6

Chalumeau, Dr. Shawn Clark, Dr. Paul S. Cwikla, Dr. Phillip DeWailly,

Dr. Michael A. Ivie, Dr. Norman F. Johnson, Mr. Peter W. Kovarik, Dr.

John F. Lawrence, Drs. Charles and Lois O'Brien, Dr. Luis E. Pena G,,

Dr. Andrew Penniman, Dr. John A. Shuey, Dr. James B. Stribling, Dr.

Barry D. Valentine, and Dr. Charles Withrow.

Material for the study of Thvlodrias contractus Motschulsky was from my private culture supplemented by specimens from a culture maintained by Dr. Barry D. Valentine. ONTOGENY AND PHYLOGENY

"We still do not know the mechanics of evolution in spite of the over-confident claims in some quarters, nor are we likely to make much progress in this by classical methods of paleontology or biology” -White, 1966:8.

"The most incomprehensible thing about nature is that it is comprehensible" —Albert Einstein Or, that we imagine it comprehensible.

While attempting to resolve phylogenetic relationships within the

Leptolycini (Miller, 1991), a paedomorphic lineage within the Lycidae

(Coleoptera ), I encountered unusually high rates of homoplasy and

inexplicable biogeographic patterns that could not be resolved by usual

techniques. Hypothesizing that the anomalous development of the

leptolycines might be causal, I studied another paedomorphic species,

Thvlodrias contractus Motschulsky (Dermestidae), which is more amenable

to laboratory culturing. Additionally, the familial and superfami1ial

relationships of the latter species is more strongly supported than the

relationships within the Cantharoidea. The results revealed unexpected

assumptions about paedomorphosis, why these implicit assumptions might

lead to error, and possible modifications of phylogenetic analysis that could alleviate this problem.

During phylogenetic analysis, systematists attempt to minimize

12 13 assumptions to those that are congruent with evolution as it is

currently understood. Since von Baer's (1Q28 ) pioneering studies of embryology, innumerable attempts have been made to incorporate ontogenetic data while examining evolutionary relationships (Gould,

1977). However, since the repudiation of Haeckel’s views of ontogeny and phylogeny, the study of ontogenetic and evolutionary processes has remained largely separate (Raff and Kauffman, 1983:355) and the relationship of these processes little studied.

The relationship of ontogeny and phylogeny within the Coleoptera is no exception. Ontogeny has not been used extensively in beetle systematics, primarily because so little is known of the immature stages. Newton ( 1990:208 ) has estimated that only 15 7, of the

Staphylinoidea, which comprises about one fifth of all described beetles, are known in the immature stages at the specific level.

Although the extent of our knowledge of other lineages varies, Newton's estimate approximates the level of study throughout the Coleoptera.

Additionally, the embryological literature is at best contradictory

(Crowson, 1981:361) and postembryonic development within the Coleoptera has been examined only in a limited number of species. Even the relatively few descriptions of larvae are often inadequate for recognition. This needs to be rectified, but indubitably will require many years of study. This ignorance in turn may impede understanding of diverse developmental phenomena within the Coleoptera such as paedomorphosis (neoteny of authors, see below).

Paedomorphosis occurs in a number of lineages within the order.

The best known examples are in the fireflies and glowworms of the former 14

Cantharoidea, now Elateroidea (Lawrence, 1987)*. However, the

phenomenon has been documented or is here hypothesized in the

Archeostemata CMicromalthidae, Scott (1938)3, and several superfamilies

within the Polyphaga including Staphylinoidea CStaphy1inidae,

Aleocharinae, Crowson (1981:389)3; Histeroidea CHisteridae, Crowson

(1974, 1981:389)3; Dascilloidea CKarumiidae3, Elateroidea CElateridae,

Cebrionidae3; Cantharoidea sensu strictu CLampyridae, Phengodidae,

Drilidae, Lycidae, and Omalysidae, numerous authors3; Cleroidea

CMelyridae, eg. Apteroma1achius namibensis Wittmer, 19603; Bostrichoidea

CDermestidae, eg. Thvlodrias contractus Motschulsky!; Tenebrionoidea

CRhipiphoridae, eg. Rhioidius Thunb., studied by Besuchet (1956); and

Meloidae, eg. Hornia Riley studied by Linsley (1942)3; and

Curculionoidea CScolytidae, Crowson (1981:390)3. In addition, there

are many other taxa that display paedomorphic tendencies that remain

unstudied. Many of these paedomorphic lineages, if not most, appear to

have had independent origins.

Because paedomorphic, "larviform" females are widespread within

the Cantharoidea and are proposed to be basal in several lineages within

the superfamily (Crowson, 1972:62), the phenomenon of paedomorphosis

must be examined before any meaningful discussion of these lineages is

possible. This is prerequisite because the effect of paedomorphosis on

character states is unknown. Hypothesizing paedomorphic ancestors within the Cantharoidea, however, creates anomalies that require ad hoc explanation (Crowson, 1972).

In addition to possessing hypothesized paedomorphic ancestors, the

Cantharoidea are considered to express basal character states 15

secondarily within the Coleoptera (Crowson, 1967, 1972, 1981;

Geisthardt, 1977; Lawrence, 1982, 1987; Lawrence and Newton, 1982). If

this secondary expression of basal states is related to a paedomorphic ancestor, it may have application to other lineages within the

Coleoptera. Such convergence to apparent basal character states may produce misleading results during phylogenetic analysis if the phenomenon is not addressed. Again, the alternative possibility that these taxa, assumed to be secondarily "primitive," are in fact basal, should not be rejected without reexamination.

Even though knowledge of paedomorphosis is essential to understanding several, if not many, lineages within the Coleoptera, it has remained unknown except from an intuitive morphological perspective. Besides the original descriptions, the literature consists of redescriptions of paedomorphic forms such as the female Omalvsus fontisbellaquei Geoffroy (Geisthardt, 1977 ). Two major exceptions are the excellent and extensive morphological studies of Geisthardt (1974 and 1979 ) of adult Lamprohi za splendidula (L. ) and Lampvris noctiluca

L. However, Geisthardt did not discuss ontogenetic or evolutionary processes in relation to morphology in either paper. Cicero (1988) attempted to study the phenomenon using a restricted data set of only females, but this analysis suffered from a number of problems (Miller, in prep. ) and those discussed below.

The physiological aspects of paedomorphosis have been studied only in Lampvris noctiluca L. Naisse (1966, 1968, and 1969) found that sexual dimorphism was the result of differential titers of juvenile hormone during development. She also demonstrated that the testes, 16 which produce an androgenous hormone, are responsible for male

character states and will masculinize a female when experimentally placed in the abdomen of a female larva. Davydova's (1967) ablation

of the corpora allata and the corpora cardiaca in males of the same

species failed to produce wingless females and her experimental

implantation of 2-6 of these glands in female larvae did not induce wing formation in females. She did not report any effect on the gonads.

A correlation of paedomorphosis with the X-0 sex determination within lineages has been noted (Crowson, 1972), but Smith and Virki

(1907) indicate the loss of the Y - chromosome is widespread throughout the Coleoptera and occurs in many lineages where paedomorphosis is unknown. This karyotype constitutes 14 of the examined species in the order (loc. cit.. p. 90) and their origins often appear independent.

However, the X-0 system may be necessary, but not sufficient for paedomorphosis, and requires further examination. Interestingly, I have found no records of karyotype studies of actual paedomorphic species.

Although little is known about comparative morphology and physiology of paedomorphosis in beetles, we know even less of it as a function of developmental or evolutionary processes. As a result, systematic examination of paedomorphic species has created a number of enigmata. For example, Crowson (1972:49) hypothesized that the character states found in the lycids Platerodrilus Pic, Lvropaeus

Waterhouse, and Puliticola Mjorberg might be basal in the Lycidae. The latter is known to have flightless, paedomorphic females and females are unknown in the other genera, but are probably also paedomorphic. Why, 17

then, do most presumably derived lycid genera have fully flighted males and females? He (loc. cit. . p. 64) implied that the male might carry

the necessary genes that were reincorporated into the females of derived

lineages similar to the tarsomeres of Cryptophagidae. Clearly, he was

troubled with his conclusions and flagged this problem for future cantharoid workers (Crowson, 1972).

Geisthardt (1977) questioned Crowson's concerns, basing his conclusions on the widespread occurrence of paedomorphosis in some

lampyrid genera. Unfortunately, he did not support his argument with a phylogenetic hypothesis demonstrating the non-derived nature of the lampyrids discussed.

More recently, Cicero (19QQ) examined ontogeny and evolution within the Cantharoidea. In so doing, he proposed a discrete classification of developmental stages for female adult beetles, assumed the basal origin of paedomorphosis (his neoteny ) within a restricted

Cantharoidea (Lampyridae, Phengodidae, Drilidae, but not Cantharidae, etc. ) that supports the hypotheses of Crowson (1972) and Geisthardt

(1977). While such an examination of the character states of adult females alone can be fruitful, its implications require further study.

For instance, 1 ) a classification of developmental stages can offer insight into process, but should not be accepted hastily because a discrete classification is arbitrary and, possibly, misleading if the phenomenon is in fact continuous. Current knowledge of ontogenetic processes has not clarified the nature of development over evolutionary time and, hence, it is unknown whether it is discrete or continuous.

2) Ignoring male morphology in favor of female morphology is no more 18

efficacious than studying only males. More importantly, 3) the

homology of developmental character states must be demonstrated and not

assumed. If homology is assumed and, again, the series is assumed to

be a continuum from one extreme to the other, there is an enhanced

possibility of succumbing to the Haeckelian argumentation of "ontogeny

recapitulating phylogeny” - an idea that was rejected four decades ago

(see Gould, 1977 for an extensive review; Miller, in prep. ). Such an

approach can prove disastrous when character states within lineages are

unexamined and assumed to be homologous throughout the entire family

without a phylogentic analysis (See Aiberch, 1985 for a discussion of

the dangers of Haeckelian recapitulation in systematics ).

I here suggest that implicit Haeckelian argumentation remains the

major hurdle to understanding phylogenetic relationships within the

Cantharoidea and that it has yet to be addressed. Because of the

central nature of this problem and because of recurrent problems in

polarization of character states within the Lycidae, a group of particular interest to me, study was initiated to examine this process

and its phylogenetic implications.

Definition of Problem. Gould (1977:484) defined paedomorphosis as "the retention of the ancestral juvenile characters by later ontogenetic

stages of descendants." Aiberch et al. (1979) presented models to

define and explain this and other forms of heterochrony. Because of

the precision of their definitions and because it is not possible to

differentiate paedomorphosis further (ie. neoteny or progenesis) without

supporting developmental data, only their more inclusive term will be 19 used in this discussion.

Another term currently used in discussions of paedomorphosis is

"larviform". It is, however, imprecise. Very few adult beetles are essentially indistinguishable from the larvae (eg. all known female

Phengodidae, some Micromalthus. and Puliticola of the Lycidae). Most are similar to their larvae in only a few characters or in their general facies. Thus, "larviform" is an inappropriate adjective for all but a very few beetles and the other imagoes would more properly be called paedomorphic adults.

A number of character states are commonly associated with paedomorphosis within the Coleoptera. They may be present in any combination, but their expression is normally manifest in extreme sexual dimorphism. Probably the most commonly used character for recognition of the phenomenon is the loss or reduction of wings and elytra. Additionally, the "retention" of more abdominal segments is often noted. Occasionally, the externally undifferentiated meso - and metanotum are discussed. These are comparative characters that require recognition of the opposite sex or sister taxa to evaluate.

However, any one such character is usually not a necessary or sufficient condition to assume paedomorphosis.

Given this imprecise means of identification, how may paedomorphosis be differentiated from other evolutionary phenomena? A putative example of paedomorphosis from the dermestid lineage, Thvlodrias contractus Motschulsky, was examined to clarify possible trends in character states due to the phenomenon.

Adult Thvlodrius contractus was first described (Motschulsky, 1839:75) as a malacoderm near Malachius Fabricius (now in the

Melyridae, Cleroidea). Reitter (1894) moved it to the Drilidae which

is currently placed in the Cantharoidea. Discovery of the larvae

revealed that it was in fact a dermestid (Hinton, 1945:260).

Dermestids are placed currently in the Dermestoidea of Crowson (19Q1)

and Lawrence (1982) and, most recently, Ivie (1985) has placed the

Dermestidae in the Bostrichoidea. Both hypotheses are in agreement

that the sister of the Dermestidae is the Nosodendridae and that other

close relatives include the Bostrichoidea / Bostrichidae. Neither of

these hypotheses place these taxa in the Elateriformia where the

Drilidae (Cantharoidea) is currently placed.

Even though the Dermestidae is probably known better than most families because of its economic status, its higher classification is

in need of review. Currently, Thvlodrias is placed in its own

subfamily (Crowson, 1967:74; Mroczkowski, 1968:158; Suss and Fogato,

1978), although it also has been given monotypic tribal status (Beal

1959:99) as well as superfami1ial status (Franciscolo, 1975). In all

these cases the differences are emphasized, but no relationship is evident. Hinton (1945:258) suggested that Thvlodrias is most closely related to Trinodes based on examination of larvae. Although

I have not seen larvae of the latter genus, the literature seems to support his conclusion.

In the absence of a phylogenetic analysis of the Dermestidae, the sister group relations of Thvlodrias remain obscure. Therefore, general character states within the family and its hypothesized sister group were examined and compared with those in Thvlodrias males and 21 females (Table 1 ). As can be seen, several characters are significantly affected by paedomorphosis.

Sclerotization. The degree of sclerotization is less than other dermestids and nosodendrids examined. This lightly sclerotized cuticle is characteristic of other paedomorphic beetles, although it varies in degree. Historically, this character state was used as a means of grouping taxa. The constituents of the Malacodermata, literally meaning "soft skin" (Gk. ), included most of the lightly sclerotized beetles at some point in the history of beetle classification and all of those currently placed in Cantharoidea.

Compactness. The 'coadaptation' of various sclerites (Crowson,

1981:60) is significantly less than found in other dermestids examined or in the Nosodendridae. It is difficult to ascertain if other paedomorphic species are less compact, as their presumed relatives are also not as compact as many beetles.

Sexual Dimorphism. The sexual dimorphism of Thvlodr ias is much greater than found in other Dermestidae. In fact, unlike the odd beetle, it is often necessary to dissect specimens to determine sex in the other members of the family. Again, the sex of nosodendrids are not as apparent externally, but sexual dimorphism in other studied paedomorphic species is pronounced to extreme.

Antennae. The antennae in both sexes of Thvlodrias are more elongate and the club is less compact than in other dermestids and nosodendrids examined. There are fewer antennomeres (10 in males and 9 in females) and the male club is extremely elongate, but some dermestids 22

Table 1. Character states found in Thvlodrias contractus Motschulsky males and females compared to the general states found in other Dermestidae and Nosodendridae.

Character T. contractus Other Dermestidae females males (and Nosodendridae)

Sclerotization 1 ight 1 ight heavy Coadaptation loose 1 oose compact Retraction of head no no yes Shape of head Globose Elongate Elongate Elongation of 2-6 some more short Antennomeres Number eye facets reduced reduced? not reduced Cryptop1euron unchanged unchanged unchanged Cavities for front no no yes leg retraction Retractibi1ity of no no yes front legs Prosternal carina absent absent present Mesonotal incomplete complete complete differentiation Scutellum caudally fused free free Elytron absent present, present, elongate oval to obovate Metasternal carina absent absent present (absent) Metanotal 1ittle complete complete differentiation Intercoxa1 distance very wide normal normal of metacoxae Metendosterni te strongly weakly normal modified modified Sclerotization of light heavy heavy Metepimeron Hindwing absent present, present +/- reduced Metafemoral cavity absent absent present Ratio Trochanter- normal elongate normal femur length Abdominal Sternites 7 exposed 7 exposed 5 exposed Sexual Dimorphism pronounced pronounced siight 23

(eg, Thorictodes Reitter) have fewer antennomeres also. Examination of the antennae of both sexes of Lampvri s noctiluca L. did not reveal significant size differences, but Phengodes females have larval-type antennae.

Again, the position of antennal placement on the frons is unaffected in both sexes of T. contractus. This placement in the

Coleoptera appears to be restricted to a track between a ventral position over the mandibular bases and a median position found near the vertex in taxa such as Chrysomelidae, Lampyridae and Lycidae. The nosodendrid and dermestid condition, including Thvlodrias. is ventral near the mandibular bases. Other examined paedomorphic species with an adult antennal form are likewise unaffected.

Compound Eyes. The number of eye facets is reduced in the female and appears variable, but a larger sample size is indicated. The male eye facet number may also be slightly reduced, but the taxonomic literature of dermestids does not discuss ommatidial number. The few other dermestids available for study had more facets than Thvlodrias.

Lampvris noctiluca females have significantly fewer ommatidia than males and Phengodes females have no compound eyes.

Prosternal carina. All dermestids examined have a carina that extends the length of the intercoxal process, except it is absent in

Thvlodrias males and females. The carina is also present in

Nosodendron. but not other paedomorphic species examined, nor their probable sister groups.

Cryptopleuron. Only a few genera were examined, but the cryptopleuron appears to be unaffected by the phenomenon in both males 24

and females of L contractus and appears similar to other dermestids and

nosodendrids examined. The cryptopleuron of cantharoid lineages also

appears to be unaffected except in those females indistinguishable from

larvae where it is absent.

Mesonotum. The mesonotum of the male is differentiated

similarly to other dermestids. In the female it is undifferentiated

except the triangular scutellum is visible as a flat plate fused to the

rest of the mesonotum-its apex is not free as in other Dermestidae or

Nosodendridae. Other paedomorphic taxa vary in the extent of

differentiation - those more paedomorphic species having females with

less differentiated mesonota.

Elytra. The elytra are totally absent in the female of

Thvlodrias. but the male has retained less sclerotized and more elongate

version than in other Dermestidae. Although unstudied as yet, the

presence and configuration of trabeculae might be of interest. The epipleura are absent in Thvlodrias. but also are absent in Anthrenus.

Other paedomorphic forms studied retain the epipleura, if they have elytra, and the presence of elytra varies.

Mesosternal carina. This median carina is absent in Thvlodrias.

but present in all other dermestids examined. It is not found in nosodendrids, nor other paedomorphic species studied.

Metanotum. The metanotum is undifferentiated in the female

Thvlodr ias and appears similar to an abdominal tergite, being larger than the mesonotum. The male metanotum is similar to other dermestids examined except that it has a median prescutellary strut over the membranous portion of the notum. This strut has not been noted in 25

Drilidae, Omalysidae, or Phengodidae, but is present in Lycidae and

Lampyridae.

Metafurca. The metafurca (metendosternite of Crowson, 1938,

1944) appears less complex in male Thvlodrias than in the other few

dermestids examined, but differs significantly from the mesofurca. The

female metafurca appears similar to the mesofurca and undifferentiated.

Other paedomorphic taxa are unstudied.

Metepimeron. The metepimeron of female Thvlodrias is less

sclerotized and the dorsal border indistinct. The male appears similar

to other dermestids studied. Again, the extent of expression of the

metepimeron varies between males and females of paedomorphic species

depending on the extent of that paedomorphosis.

Hindwings. The wings are absent in the female, but polymorphic

in the male. According to Mertins (1981:578) they are not apterous and polymorphism is probably genetically controlled. Due to the small

sample size available, I was unable to examine effects on venation, but brachyptery undoubtedly influences the extent of venation. Other flightless Dermestidae are known (eg. some Trogoderma). but

Nosodendridae are not known to be flightless. The condition in the paedomorphic cantharoids studied shows a similar tendency. Venation of

Omalvsus was found to vary considerably among males collected at the same location (Miller, unpub1. data).

Trochantins. The trochantins in Thvlodrias are free and setiferous as in other dermestids examined. In other paedomorphic species character states vary, but do not seem to be correlated with paedomorphos i s. Legs. The limbs are elongate in the male Thvlodrias. The

trochanter to femur ratio is 1:5 instead of 1:3 in other examined dermestids, nosodendrids and Thvlodrias females. The tibiae and

tarsomeres are moreelongate also, but the coxae are not affected. It

is unknown if these relative lengths (trochanter: femur) are a function of allometry or the result of a heterochronic rate change sensu Aiberch et a l . (1979). Another unexamined question is why segments are differentially affected. Lampvri s noctiluca L. does not exhibit sexual dimorphism of limb length, but Phengodes females retain the shorter larval legs.

Abdominal Segmentation. Thvlodrias has seven visible sternites in the male and female. No other dermestid has more than five (Hinton,

1945:258). In other paedomorphic species the males have six, seven, or eight visible sternites.

Intercoxal Process of Third Abdominal Sternite. The third sternite in dermestids and most other beetle families is excavated for the reception of the metacoxae. It is composed of a recessed area and one or two "pegs'* which articulate at the metafurca. One point of articulation is external and ventral between the coxae and is termed the intercoxal process. In addition there may be an internal peg formed by the anterior medial edge of the excavation which does not appear to have a name. The whole abdominal structure allows flexibility in dorsoventral movement, but limits lateral movement. Murray and Tiegs

< 1938 ) have shown how the abdominal segments of the larvae of 5i tophilus orvzae (L. ) (as Calandra ) of the Curculionidae are incorporated within the adult and the formation of the pegs from the ventral portions of 27

the first and second abdominal segments.

This complex structure is undoubtedly homologous throughout the

Coleoptera as it always appears on the third sternite when present. It

has not been found outside the Coleoptera, but is present in all

Coleoptera with five visible sternites examined. It is not found in most taxa with six ventrites and none of those with seven or more.

This is undoubtedly due to the removal of the motacoxae from juxtaposition with the third sternite. The Adephaga with six ventrites do have an excavation, but the second sternite is divided and moved forward so that the metacoxae do articulate with the third sternite.

In all known paedomorphic forms the third sternal excavation is absent in both the males and females because the second segment

intervenes. Thvlodrias is no exception. However, examination of the abdomen indicates that it has the adult configuration and is not

"larval" in both the males and females. Additionally, the male has what appear to be remnants of the intercoxal keel on sternite 3 in the form of a swelling.

Since elements of all suborders possess this complex structure, it must have arisen in an unknown ancestor of the Coleoptera and its loss must be secondary. The null hypothesis of this would be that extant

lineages without such an excavation are basal and the condition plesiomorphic. Efforts to find this plesiomorphic state in the

Coleoptera might be best concentrated in those lineages which have both

5 and 6 abdominal sternites visible as these character states may be indicative of a transition series. The excavation of the third 28

sternite would not be expected in those with 6 visible sternites, but

they should be present in those members of the polymorphic lineage with

5 ventrites if loss of this character is a secondary modification.

Discovery of a lineage with 5 ventrites and no excavation would indicate

an interesting candidate for basal placement within the Coleoptera and

should be investigated if found.

Gonopore. The position of the gonopore in Thvlodrias remains

the same as found in other dermestids. In most beetles it appears

behind the ninth sternite (Snodgrass, 1935:563), but it has been shown

to be in the middle of the ninth sternite in paedomorphic phengodid females (Tiemann, 1967:249) and behind the eighth sternite in the membrane in paedomorphic Puliticola females (Mjorberg, 1925:134).

Many more characters appear to be affected by paedomorphosis, but nonavailability of specimens and lack of character state discussion in the literature prevent further resolution at this time. Recognition of the sister group of Thvlodrias would expedite and enhance the resolution immensely. However, it can be seen that paedomorphosis in

Thvlodrias affects adult beetles in a number of general ways. The cuticle is less sclerotized, the sclerites are less coadapted, limbs are sometimes more elongated, the extent of differentiation of sclerites is often reduced, and the number of abdominal segments is

increased with the concomitant loss of the intercoxal keel of the third sterni te.

More importantly and in summary, examination of these few examples indicates several major trends of paedomorphosis (Table 1). It is apparent that both sexes are affected significantly, contrary to Matsuda (1976:43, 1979:177) and contrary to the implicit assumptions of most systematic coleopterists. A number of character states may not be

found elsewhere within the inclusive lineage or even its sister group.

Because of the lack of differentiation, overall facies tend to converge.

Perhaps more importantly, many characters are not larval, but adult, and their appearance is often presumed to be primitive because of their non-differentiation. Finally, preliminary evidence suggests that the effect on male morphology covaries with that of the female body. Thus, males would be expected to demonstrate more paedomorphic character states in a species with an adult female indistinguishable from its

larvae than one where the female differs little from a non-paedomorphic adult of a sister taxon.

Three major considerations are left unanswered. 1 ) Can character states affected by paedomorphosis be the result of alternative processes? The initial assumption that paedomorphosis is a heterochronic phenomenon (sensu Gould, 1977; Aiberch et al., 1979) requires empirical testing because the phenomenon may be, alternatively, the result of novel developmental pathways. 2) Is the expression of paedomorphosis within various lineages within the Coleoptera the result of the same developmental mechanism (ie. developmentally homologous)?

3) What, if any, is the effect of paedomorphosis on the larval stage?

Cicero (198B) assumed that the major paedomorphic changes occurred during the last stages of development, but earlier developmental stages may also be affected. Certainly, the hypermetamorphic development found in a number of coleopteran lineages is indicative that evolutionary change occurrs throughout the life of the organism and is 30

not limited to terminal phenomena. These unanswered questions provide

many years of productive inquiry. However, empirical elucidation of

the underlying mechanism(s ) and further documention of morphological

variation should be given priority if we hope to more closely

approximate the history of the Coleoptera with phylogenetic hypotheses.

The underlying mechanisms of paedomorphosis in the Coleoptera are

unknown and it is not even certain that the same mechanism operates in

all cases. Although the effect of the phenomenon can be demonstrated,

some priority should be given to elucidating the mechanism(s ). This

will require experimental post embryonic study, but the groundwork for

such effort can be laid prior to its initiation.

A major problem in such a study of paedomorphosis is that growth

in insects appears discontinuous and punctuated with periods of stasis

or equilibrium where little or no growth is apparent. This development may be constrained into a synchronized pattern by the moulting method of

growth. If this be true, how much of the developmental process is not

overtly expressed? This will be examined in the following discussion

and a testable model for development will be proposed.

A Model. As demonstrated above, "malacoderm" character states are expressed during paedomorphosis. These can result from apomorphies of novelty or from apomorphies of reexpression. As many of these character states are adult in nature, they may be merely the expression of an incomplete developmental process which reveals an ancestral character state not necessarily displayed within the immediate sisters of the taxon or even within the Coleoptera. However, this should not 31 be assumed. A model to elucidate character state transformations and test these ideas is proposed in Fig. 1.

The model demonstrates the hypothesized ontogeny of a character state in an insect constrained by the moulting process. The vertical striated bands are periods when the character states are expressed.

The character has been assigned arbitrarily 9 discrete ontogenecic character states in the taxonomic outgroup (Fig. 1, a), but continuous data can be accommodated. In the outgroup the character is only expressed as character state 2 in the first stage, 5 in the second, and

9 in the third (eg. last larval instar, pupa, adult). The other character states are not necessarily visible during development, but still present during ontogeny. An example of this possible masking would be the metanotal cuticle which is only differentiated at end states of development for each instar.

Assuming this model is a close approximation of development in the

Coleoptera, an expressed character state may be the result of several processes. It may be the result of terminal or interstitial addition, deletion or perturbation yielding a new state (Fig. 1, b). Again, character states may be the result of truncation of development (Fig. 1, c). Finally, the expressed character state may be the result of a change in the rate of development such as acceleration (Fig. 1, d).

These later processes, termed heterochrony, is discussed at length in

Alberch et al (1979).

If characters do behave in this manner, the progressive nature of the character transformation may be the expression of an assumption of order. As mentioned earlier, the equivalence of paedomorphosis and 32 heterochrony is untested within the Coleoptera. Also, the implied

transition in Figure 1 may in fact be non-linear. The assumption of ordered directionality might be examined by overlaying a large series of characters for several taxa, isolating novelties of caenogenesis, and determining if the results are ordered. The same method should, parenthetically, reveal a data base that can produce a phylogeny based on the principles of Hennig using outgroup comparison for polarity decisions.

Such a phylogenetic analysis would avoid the problems of using the ontogenetic criterion of Nelson (Nelson, 1978; Nelson and Platnick,

1981) as discussed in Alberch (1985). Foremost among the potential problems is succumbing the Haeckelian recapitulation. His assumption that causality is required for phylogenetic analysis is, however, unjustified. Although a causal sequence may be necessary for heterochrony analysis, a temporal sequence is sufficient for phylogenetic analysis.

Again, Alberch (1985) suggested that the process of determining homology was Haeckelian and not von Baerian in the sense that Haeckel argued that development is recapitulated throughout phylogeny and von

Baer argued for the continuous divergence between species during development. Thus, hypotheses of homology require non-divergence or

"identity cannot be unequivocally established." Alberch seems to imply that there is no middle ground between the arguments whereby homology might be ascertained. However, just as Haeckelian recapitulation is unlikely to provide the sole answer, so is von Baerian recapitulation. Not all characters would be expected to diverge 33

equally and some would not be expected to diverge at all. Thus,

homology decisions based on the criteria of Remane (1956) are adequate

to evaluate at least some ontogenetic characters.

Paedomorphic lineages in the Coleoptera offer a unique

opportunity to examine such development. Because they are sexually

dimorphic they offer two views of development in the same species.

Additionally, a monophyletic lineage may yield a series, although not necessarily directional, of these combinations. A taxon such as the

Phengodidae will, thus, test the proposed presence of unexpressed character states by providing evidence of a transition or none. If the results merit it, the phenomenon can then be examined further by experimental manipulation in a controlled environment. Of course, these procedures presuppose extensive field work in the former approach and an active effort to culture phengodids in the latter.

However, not all characters are hypothesized to develop in this manner. The growth and differentiation of the ventral nerve cord of the beetle CNS appears to be continuous (Miller, unpubl. data). Other organs may behave similarly, but have not been examined. The most likely candidates would be internal organ systems whose continuing function is necessary for survival. These types of characters are amenable to analysis by this model, but because of their continuous nature, the data must be temporally isolated in a similar manner as in heterochrony analysis. These isolated states may be captured for homology decisions by actually measuring time elapsed or maybe comparatively synchronized with other ontogenetic states within the organism. Without such a set of temporal series, the arbitrarily 34 discrete stages imposed on development would be impossible to homologize.

Svnergism of Morphology and Svstematics. Traditionally, the disciplines of comparative morphology and systematics have been mutually dependent. Systematics relies heavily on morphology for its data base and morphology relies on systematic conclusions for deciding which taxa to compare. But there has been recent interest in the testability of both disciplines (Wiley, 1975: Nelson, 1978; Gaffney, 1979; Lauder,

1981; Wiley, 1981; etc.).

Lauder (1981) suggested the use of phylogenetic analysis to test morphological hypotheses. Predictive hypotheses discovered by comparative morphology would be tested by historical patterns of evolution found by cladistic methodology. Falsifiabi1ity results from nonconformity of a phylogeny with expectations. However, this overlooks the nature of phylogenetic hypotheses. Patterns generated by these algorithms are hypotheses of relationship, not observed data.

Because the actual pattern is unknown and will remain unknown even if a true hypothesis is proposed (Xenophanes in Popper, 1965:153 and in

Freeman, 1983:24), there is no means of accessing the validity of the test. Thus, hypotheses cannot be treated as observational data. This can best be seen in the contingency table (Table 2). Thus, Lauder's suggestion may result in discarding a useful generalization when in fact it was valid or in accepting a faulty generalization with no means of assessing error. Although all statistical tests are subject to type I and II errors, those based on data provide a means for estimating such 35

errors. Confidence levels for phylogenetic hypotheses currently only

measure the ability to find the most efficacious tree that meets the

assumptions of the model used such as compatibility, consensus, or

parsimony methods (Felsenstein, 1985). At present there is no means of

knowing whether any algorithm would produce a correct phylogeny, let

alone how a level of confidence can be ascribed to such an algorithm.

The problem with Lauder’s proposal becomes even more complex when we try to discover what is sufficient for rejection of the hypothesis.

A testable hypothesis implies that it can be rejected (Popper, 1965;

Gaffney, 1979; Nelson, 1978). What rate of failure qualifies for rejection? One in ten, one in one thousand, one in a one million? If hypotheses are continuously qualified, they may eventually end in a

series of non-general hypotheses that are little predictive. More

importantly ad hoc qualification is admission of designing an inadequate test. Again, unlike experimental data which is potentially infinite, data based on character state distributions is finite. If the limits of the taxon are known, a hypothesis could conceivably result that

Table 2. Possible results of a test as proposed by Lauder (1981) of a morphological hypothesis using a phylogenetic hypothesis. Type I error is the rejection of a true null hypothesis, a typo II error is the acceptance of a false null hypothesis.

Phylogenetic Morphological Hypothesis

Hypothesi s True False

True Accept Type I Error

Fal se Type 11 Error Reject 36 cannot be tested because doing so would be tautological (all the character states are known and the shortest tree has already been found ).

Interestingly, the phylogenetic and morphological hypotheses will probably not be independent. The phylogenetic hypothesis will probably be constructed from the same morphology that is used to test the morphological hypothesis. Even if caution is taken to use only characters not used for the morphological hypothesis in the phylogenetic analysis, it is not unlikely that some unknown character states will covary with those character states of the morphological hypothesis.

Although this proposed use of phylogenetics presents many problems, testing morphological hypotheses by this means may be useful for isolating the limitations of systematic and morphological analyses.

Nonconformity of results with expectations demonstrate the need to reexamine the model, the phylogeny, or the morphological data base.

Perhaps a more cogent, concurrent use of the methodology of both disciplines is the isolation of heuristic lineages amenable to the study of evolutionary process. Key bifurcations within these lineages can be identified by systematic methodology and the ontogenetic differences at these bifurcations can be demonstrated by comparative morphology. Comparison with other lineages may yield additional insight into process by correlating morphology with process. This data base could then be used to test hypotheses of process by empirical means. This procedure, while not eliminating the possibility of ad hoc falsifiers

just pattern. Otherwise, study of evolutionary process using

systematics is likely to continue to be haphazard, serendipitous,

speculative, or non-existent.

An additional example, but by no means the only other empirical

possibility using this methodology, might be an examination of the post

embryonic development of hypermetamorphosis. Casual examination of the

process suggests an immediate question. Whydo hypermetamorphic

lineages have campodiform primary larvae even though later instar larvae

are vermiform? Clearly, something more than "embryoni2ation" is

occurring here. Resolution of this problem may be facilitated if the

above protocol were followed in monophyletic lineages. Correlations

noted would lend themselves to tests of causality by empirical means.

Until this and other such studies are done routinely we will be

shackled by our lack of knowledge about the process of development and

evolution in the Coleoptera. Nowhere is this lack of knowledge of the

immature stages more apparent than when we attempt to polarize larval

character states. We are left in a sea of inexplicable homoplasy if we

venture afield within the order.

Phylogenetic Implications.

The phylogenetic implications of paedomorphosis have not been examined extensively. Although he wrote a major, informative monograph on the subject, Gould (1977) did not discuss its relevance to systematic methodology. Nelson (1978) proposed a modification of phylogenetic analysis using the ontogenetic criterion to polarize data, but Alberch

(1985) discussed the problems associated with the use of ontogenetic 38

data in phylogenetic and heterochronic analysis. Again, Nelson

(1985:40) has expressed dismay at the possibility of a paedomorphic

outgroup, but did not elaborate. A few systematists have attempted to

consider the phenomenon when looking at their respective taxa, but no

one has proposed possible modifications of systematic technique that

address Alberch's concerns (Miller, in prep.)

Westheide (1987) examined the possible significance of

paedomorphosis (his progenesis) in meiofaunal origins and suggested that

there are three available phylogenetic reconstructions for a

morphological series of paedomorphic organisms. These are 1 ) a

monophyletic lineage from a single ancestor with increasing

paedomorphosis (Fig. 2, a), 2) the independent evolution of paedomorphic

species from non-paedomorphic stem species in a monophyletic lineage

(Figs. 2, a & 2, d) and 3) independent non-monophyletic origins (Fig. 2,

c). The throe reconstructions are shown in Fig. 2. The second case

is shown as originally figured (Fig. 2, b) and refigured in a more

conventional form (Fig. 2, d). It is clear that, while the

possibility for case 2 exists, it is non-parsimonious and requires

numerous hypothesized origins and a series of ancestral stem species

whose existence is unknown except for their assumed non — paedomorphic

character states. In the absence of such evidence, the options are

reduced to 1) monophyletic origin or 2) independent origins.

Additionally, development in case 1 need not be unidirectional as in

Westheide*s a priori assumption although detection of directional

shifts without concomitant caenogenesis remains a problem with current phylogenetic methodology. The examination of double exposures to 39 development in sexually dimorphic species may also alleviate this problem.

There are several apparent implications for phylogenetic analysis that result from the study of Thvlodrias. The presence of presumed paedomorphs requires modification of protocol during cladistic analysis.

These modifications are:

1) recognition that both sexes are likely to be affected

by the phenomenon;

2) because of the possibility of convergent facies, the

monophyly of paedomorphic taxa should be demonstrated

by characters not shown to be affected by

paedomorphosis;

3) although paedomorphic characters should not be used as

synapomorphies, they may be used as autapomorphies with impunity;

4) paedomorphic species might best be placed by synapomorphy after non-paedomorphic members of the lineage have been analyzed to minimize use of an unrecognized paedomorphic character state in the analysis. This is especially true of computer generated phylogenies until such taxa can be accommodated;

5 ) paedomorphic species should be used as an outgroup with extreme caution;

6 ) synapomorphies might best be found in immature stages

although this possibility requires critical examination before implementation; 40

7) erection of higher taxa for paedomorphic forms without

phylogenetic analysis is likely to emphasize

morphological differences rather than indicate

relationships, and should be minimized if phylogenetic

hypotheses are to be useful to anyone other than a

taxonomist

These necessary modifications may suggest apparent potential problems for the cladistic algorithm. If all characters are

independently subject to heterochrony, then it becomes impossible to choose appropriate characters for analysis. Again, how can "minimal" paedomorphosis be detected so that the above procedures may be observed?

Of course, the answer is that neither matter. Phylogenetic systematics attempts to find hypotheses, not Truth, of relationship.

Invalid hypotheses will eventually be disproved, but in the interim the enigmata generated may stimulate study of processes that will resolve these difficulties. Additionally, even a false hypothesis that approximates reality more closely than a former hypothesis is progress as long as it is perceived as only an hypothesis.

Possible Taxa for future study. The introduction to this paper lists a number of lineages within the Coleoptera which should provide fruitful opportunities to study paedomorphosis. Preliminary studies suggest the following comments:

1). Numerous cantharoids such as all known drilids, phengodids, omalysids, and some species of lycids and lampyrids are paedomorphic. 41

Some lineages are probably monophyletic, but none should be assumed so and their relationship to each other must be reinvestigated. Placement of the Cantharoidea in synonymy with the Elateroidea (Lawrence, 1987 ) is corroborated by my studies in progress which suggest that the former taxon is not monophyletic and the maintenance of both superfamilies leaves the Elateroidea paraphyletic. Again, those genera placed basally in the Lampyridae by Crowson (1972) should be reexamined. Most will probably be found to be paedomorphic because females are largely unknown and the male facies are indicative of the phenomenon. Hence, their basal status may be an artifact and they may in fact be derived.

2). The continual debate of the placement of the Strepsiptera may need reexamination. The argument that they are highly derived beetles

(Arnett, 1973; Crowson, 1967) and not the sister group to the beetles

(Kinzelbach, 1966, 1971a, 1971b, and 1978; Lawrence and Newton, 1982;

Kathirithamby, 1989) may have merit. Although strepsipteran females are known to be paedomorphic and males are known to be highly derived, they have not been examined in light of this study. Of course, their hypermetamorphic development presents additional problems in the resolution of homology of character states.

3). Again, the placement of the monotypic Micromalthidae must be examined in the same manner. Micromalthus is paedomorphic and paedogenetic (Scott, 1936, 1938, and 1941; Kuhne, 1972). Its position in the Archeostemata (Lawrence and Newton, 1972:265) is based on a hypothesized transition state (hinge of the cubital vein) and on symplesiomorphies (larval ungues and lack of a cryptopleuron), not shared apomorphies. The difficulty with the pleuron character state 42

{ie. lack of the synapomorphic cryptopleural structure of the Polyphaga)

in M.icromaithidae is that it differs significantly from that of the

Cupedidae also. The propleura of Micromalthus debi1is LeConte is fused and unlike anything else discussed by Hlavac (1972, 1975). As the cryptopleuron of Thvlodrias does not appear to be affected by paedomorphosis, it may be a useful synapomorphy if the studies of Hlavac were completed for the Bostrichoidea-Cucujiformia.

4). Some scolytids (Curculionoidea) and the "probable" histerid

(Histeroidea ) described by Crowson < 1974 ) are paedomorphic. It is interesting to note that the latter paedomorphic histerid males were originally placed in the Scolytidae based on association with paedomorphic scolytid females, on the males being unknown, and on the structure of the proventiculus. They were subsequently moved to the

Histeridae (Crowson, 1974) based on a number of character states. Added to Crowson's argument would be the interesting anomaly of this being the only case of which I am aware where there is sexual reversal of paedomorphic forms if it were in fact a scolytid and not a histerid.

5). Karumiidae is another taxon that has a history of placement within the Cantharoidea (Arnett, 1964; Paulus, 1972). Karumiids are probably paedomorphic as the females are unknown in most cases and males have structural organization indicative of the phenomenon. Karumia estafi1inoides Escalera has been reported with the tenth abdominal tergite visible (Arnett, 1964:65), but the specimen I examined in the

NMNH (probably the one examined by Arnett) has only Q visible tergites.

The mesothoracic and metethoracic terga are undifferentiated and may have been erroneously counted. Crowson (1971:16) moved Anorus. 43 formerly of the Dascillidae, into the Karumiidae. The females of

Anorus p iceus LeConte have been described and are apterous and brachyelytrous and have been captured at the entrances of burrows

similar to the females of Pleocoma (Blaisdell, 1934:322; Howden in

Crowson, 1971:14). The monophyly of the Karumiidae and its relationship within the Dascilloidea should be reassessed.

6). The males of the 3 known species of the monogeneric

Diphy1lostomatidae have 7 visible abdominal sternites (Lawrence,

1982:503). Holloway (1972) removed Diphvllostoma Fall from the

Lucanidae and placed it in its own family. A number of the character states she used are affected by paedomorphoisis and, combined with the reduced compound eyes and the vestigial hindwings of the flightless females, are suggestive of paedomorphosis. I examined 3 males collected in Tulare Co., California (MAIC) with the abdomen removed from one. It does not have an intercoxal process, as predicted.

Of course, all lineages with paedomorphic members should be reexamined, but all these lineages suggest an even larger question.

Why is paedomomorphosis more prevalent in the cantharoid group than in other beetle lineages? The phenomenon is known sporadically throughout the Coleoptera and it sometimes appears to have monotypic origins. The cantharoids, however, contain most of the known paedomorphic species.

Is this an artifact of current methods of classification? If so, there are at least some lineages within the group that appear monophyletic based on the methodological modifications proposed above. These are often fairly large and it would be difficult to justify discarding them as an artifact. Possibly, the high incidence within the cantharoids is 44

a result of a more paedomorphic body plan initially. Any species that

evolves to a more paedomorphic ontogeny from ancestral beetles

exhibiting the malacoderm facies may be more obvious than, say, one that

evolves within the extremely hard-bodied Zopheridae. Thus, the

2opherid may require more change to be recognized as paedomorphic.

Perhaps, the malacoderm facies is merely an intermediate paedomorphic

state?

In summary, paedomorphosis is a widespread phenomenon within the

Coleoptera and its recognition may expedite the study of phylogenetic

relationships. The implicit assumption that its affect on males is negligible, coupled with the lack of recognition of possible convergence

of paedomorphic species to undifferentiated states, may lead to specious conclusions. Therefore, modification of the protocol of phylogenetic analysis is required to accommodate paedomorphosis when its presence has been demonstrated.

All this suggests that adequate analysis of phylogeny must include the total ontogeny of the organism, not just a comparison of selected semiphorants of only one life stage or sex. This is obvious when historical problems of the classification of paedomorphic lineages is contemplated. Therefore, understanding development must be given priority and should be thoroughly documented in at least exemplars of lineages and in those species that are aberrant. This is not to imply that hypotheses of phylogeny without examination of ontogeny should be neglected. Nor does it imply that analysis of selected semiphorants is useless. Both types of study are prerequisite to the efficient initial isolation of heuristic lineages for study of process. PHYLOGENETIC RELATIONS WITHIN THE CANTHAROIDEA.

Iconoclasm is the bane of established order, but the responsibility of science.

Although previously placed basally in the Coleoptera (Crampton,

1928) or Polyphaga (Peyerimhoff, 1933; Lameere, 1938; Jeannel and

Paulian, 1944), the members of the Cantharoidea are now thought to

express basal characters secondarily (Crowson, 1967, 1972, 1981;

Lawrence, 1982, 1988; Lawrence and Newton, 1982). This is partially a

result of Crowson's repeated assertions that the Archeostemata is

basal to the rest of the Coleoptera.

However, the basal nature of the Archeostemata is unconvincing.

It is based on a priori assumptions about evidence of the fossil record

and not polarization by outgroup. Nelson (1978) and Schaeffer et al.

(1972) have discussed the problems associated with correlating origin

with position in strata. Another problem with use of fossils is that

the characters visible are often probable plesiomorphies (eg.

notopleural sutures, transverse metasternal sulcus). Again, the basal

position is supported by characters that appear to have had numerous

origins (ie. reticulate elytra). The resulting aporaorphies used are

absence character states which are difficult to homologize.

Additionally, examination of Ponomarenko (1977) and Arnoldi and

Zherichin (1977) does not show any beetle fossils with clearly

45 identifiable malacoderm facies as would be expected in plesiomorphic character states based on probable outgroups. Lawrence and Newton

(1981:262) have postulated an ancestral ground plan for the Coleoptera resembling the malacoderm facies. Apparently, this was based on outgroup study although not stated. All possible outgroups do suggest a common ancestor with such malacoderm facies. If this hypothesis be true, then a case for an extant, basal, malacoderm beetle cannot be discarded offhand. This implies that the hypothesized secondarily derived malacoderm facies must not be assumed, but must be demonstrated.

Recent studies of tapronomy have examined chi tin decomposition rates in marine environments (Plotnick, 1986) and indicate that decomposition rates are a function of burial depths, exposure, and degree of sclerotization. As yet, no information is available on the effect of the degree of sclerotization in terrestrial environments, but cantharoid absence from the fossil record suggests that the malacoderm facies may be subject to differential preservation and fossi1i2ation by compression. Ashworth (1987, personal communication) noted differential preservation in Chilean peat deposits of Hemiptera, and termites by the presence of only mandibles and other heavily sclerotized parts. This suggests the paucity of the malocoderm facies in the fossil record may be an artifact of taphronomic bias. Several fossil cantharoids have been reported in amber, but compression fossils are rare and placement appears erroneous in the few cases I have been able to evaluate (eg. Chauliognathus pristinus Scudder,

1900:101, pi. 11. fig. 3 is probably a more sclerotized cucujoid and not

46 47

a cantharoid - it is not a member of Chau1iognathus Hentz).

At present the origins of the order cannot be addressed by

phylogenetic methods because there is no agreement on the relations of

the Coleoptera to the rest of the Insecta (Hennig, 1981:302; Kristensen,

1975:29, 1981:150) although the monophyly of the Endopterygota is not

questioned by most workers. Stepsiptera has been suggested as the

sister by Kinzelback (1967, 1971a, 1971b, 1978) and followed by

Boudreaux (1979:237). However, we have seen that the morphology of males as well as females of the latter lineage may be influenced by paedomorphosis and, hence, both are of little use in the polarization of character states within the Coleoptera. Again, Crowson's special relationship of the Megaloptera and Coleoptera is questioned by

Kukalova-Peck (1987, personal communication) because the former lineage

is probably derived within the Neuroptera.

Because of the intractability of the problem and because it is beyond the scope of this study, the superfamily relationships presented by the Crowson school are provisionally accepted.

Relations within the Elateriformia.

Lawrence and Newton (1982) following Crowson (1967, 1972, and

1981 ) suggest that the Artematopoidea, Elateroidea and the Cantharoidea form a monophyletic lineage. This is based on the larval character states: single paired stemmata, bisetose tarsungulus, loss of mandibular mola, coadaptation of maxi 1lolabial complex, and buccal cavity setation. Crowson (1967:61) suggests that the sister group of the Cantharoidea is the Elateroidea based on the wing venation and 48 metendosternite of the adults, and nasale of the larvae.

In addition, the presence of luminous organs is not known outside

the Elateriodea and Cantharoidea in the Coleoptera and is, presumably, a

synapomorphy (Miller, in prep), although Crowson (1972:65) suggests that

it is not. In the Elateroidea it is known in adult and larval

Pvrophorus sensu lat (Elateridae) and has been recently recorded in adult Throscidae (Costa, 1984). Luminescence is reported in the

Lampyridae, Phengodidae, Omalysidae (Bertkau, 1891 ), and Telegeusidae

(Lloyd, 1983:132) within the Cantharoidea. It has been reported in the

Lycidae (Kolbe, 1887 and Shelford, 1901), but is unconfirmed and probably erroneous.

An apparently unnamed structure not currently discussed in relationships is a median prescutellary strut over the membranous portion of the metanotum. I have found it in all Lampyridae and

Lycidae examined, a few Elateridae, and no other families except in

Thvlodrias as mentioned earlier. Its distribution needs further documentation, but may provide further evidence of relationship.

Most recently, Lawrence (1987) has combined the Cantharoidea and

Elateriodea. However, he did not study the relations within these groups and he urged future reexamination of the cantharoid group as a whole and not within families as currently constituted (Lawrence,

1987:43 ).

In the following discussion I will restrict my comments to the

Cantharoidea prior to Lawrence (1987), although the Elateroidea were studied also. This decision was made because in his analysis Lawrence treated the Cantharoidea in the same manner as he treated families of 49 the other Superfami 1ies within the Elateriformia. This lumping has allowed little resolution within the former Cantharoidea. In a like manner, he has not examined these lineages containing many paedomorphic taxa in light of the insight revealed by the study of Thvlodrias Mots.

Monophvlv of the Cantharoidea. This superfamily has a tradition as a

"suitcase" taxon. An unusual life cycle and/or the malacoderm facies

( sensu Crowson, 1967 ) has usually been sufficient for membership during some time of its taxonomic history. Originally, it was named the

Malacodermata (Lacordaire, 1857:289) and consisted of the Cantharidae

(as Telophorides), Drilidae, Lampyridae, Lycidae, Melyridae including

Malachiidae, Phengodidae, and Telegeusidae. Later the Lymexeliidae and Cleridae were added.

Brachvptsectra was elevated by Blair (1939:49) to family status and placed in the Cantharoidea by Boving and Craighead (1933:46), only to be removed later by Crowson (1973:235; 1981:696). Lawrence

(1982:511) replaced Brachypsectridae in the Cantharoidea, but Lawrence

(1987:42) includes it in the Elateroidea sensu lato and mentions that its relationships are unclear.

Crowson (1972:43) elevated Cneoalossa to familial status and placed it in the Cantharoidea. Several years later he moved it to the

Dascilloidea (Crowson, 1978). Then, Crowson (1981:696) moved it back to the Cantharoidea where Lawrence (1982:511) placed it. Lawrence

(1987:43) moved it to his Psephenoidea.

The current composition of the Cantharoidea includes about 11,000 described species in 8 - 11 families. These families are Cantharidae, 50

Drilidae, Lampyridae, Lycidae, Omalysidae, Omethidae, Phengodidae,

Plastoceridae, Teleguesidae. Also, the Brachypsectridae and

Cneoglossidae are often included.

I was not able to demonstrate the monophyly of the Cantharoidea.

Many of the characters used in its support are presumably plesiomorphic states whose secondary derivation cannot be supported or refuted at this time. If all such characters are assumed to be secondarily derived, they are 1) affected by paedomorphosis, 2) the result of reduction or fusion, or 3) losses. A phylogenetic hypothesis was, however, sought and some results are presented here.

Current Hypotheses of Cantharoid Relationships.

The most comprehensive recent treatment of this taxon was by

Crowson (1972) in which he made major rearrangements and discussed many of the remaining problems that even now confront cantharoid workers.

He proposed a basal dichotomy of a Plastoceridae - Cantharidae -

Omethidae lineage and the reminder of the cantharoids in the other lineage. In that latter lineage he again proposed dichotomy of the

Omalysidae - Lycidae lineage and the Drilidae - Lampyridae - Phengodidae

— Telegeusidae, the latter pair being sisters and the most derived

(Crowson, 1972:fig. 35).

Potatskaja (1983) using only the hypostomal cavities and the degree of ventral closure of the gular region has asserted that the

Cantharoidea are composed of two lineages. One lineage is composed of the Lampyridae and the Lycidae. The other is composed of five families with a pair of sister groups being basal - the Brachypsectridae 51 and the Cantharidae. The sister to these are the Phengodidae,

Drilidae, and Omalysidae, the later two having the most recent common ancestry in this branch (Potatskaja, 1983:553, fig.2). She did not mention the Telegeusidae, Omethidae, Plastoceridae nor did she discuss the removal of Brachypsectridae from the Cantharoidea by Crowson (1973).

Based on an outgroup analysis using the Elateroidea, however, only the

Lampyridae and Lycidae are derived for the character states she examined. The remainder would be a taxonomic grade based on symplesiomorphy. Crowson ( 1972:68-69) suggests that the latnpyrid and lycid conditions are convergent and the result of similar ability to retract the larval head within the prothorax.

Fatni lv Relations Wi thin Cantharoidea.

Although I have not been able to satisfactorily resolve the relationships within the superfamily, some interesting ideas have emerged from my attempts to find a functional outgroup for the Lycidae within the Cantharoidea. A brief report of their status is presented below.

Omalysidae

The family name, Homalisidae, proposed by Crowson (1972:44) must be emended to Omalysidae. Silfverberg (1978:117) corrected a number of nomenclatorial errors due to the oversight of Muller's (1764) definition of various genera including Omal.vsus which was subsequently and unjustifiably emended to Homalisus and Omalisus.

The family contains one genus with 10 species and possibly a 52

second genus with 3 species that are distributed in Europe and the Near

East. Crowson removed Thilmanus Redtenbacher and Pseudeuanoma Pic, but

the status of Euanoma Reitter remains uncertain as he did not see it.

Omalvsus fontisbellaouei Fourcroy and 0. nicaeensis Ochs are known to be

paedomorphic (Bertkau, 1891; Ochs, 1949) and it is not unlikely that

other omalysids are also. Our knowledge of the family is based largely

on original descriptions and has not been examined as a whole.

Omalvsus has traditionally been placed in the Lycidae (Kleine,

1933:4), but Bourgeois (1882:26) questioned its placement. Crowson

( 1972:49 ) suggested that the primitive character states of the Lycidae might be found in this family. However, examination of adult males and larvae suggest that it may be best placed in the Elateridae.

The characters uniting the Lycidae and the Omalysidae by Crowson are the lycid long trochanters and the hypothesized transitional mandibles in the larvae. The long trochanters, besides being found in other paedomorphic species are unlike those found in the Lycidae. The metatrochanters are strongly offset as in the elaterids, they are not elongate as in lycids, and they articulate with the coxae in the same manner as in the Elateridae, but not as in the Lycidae.

Again the larval mouthparts of Omalvsus have been suggested as a transitional stage in the evolution of lycid mandibles (Crowson,

1972:47). While they may be, they are no more likely a candidate than those of any predaceous larvae with grooved or tubulate mandibles. In fact the stylet-like mandibles of the Cerylidae (Sen Gupta and Crowson,

1973:422, fig. 237) may be better candidates for a transformation series. Additionally, the more approximate bases of the mandibles in 53

the Lampyridae, as in the Lycidae, suggest a better candidate for a

transition than the distal bases found in Omalvsus larvae.

With this mixture of unclear character states two alternate

scenarios are possible: a paedomorphic eiaterid converging to the cantharoid facies or a paedomorphic lycid that is expressing ancestral

states in the larva as well as adult. The strongest support for the

latter argument is found in the serrate margins of the tergites found in unassociated lycid larvae collected in Ohio (unpubl. pers. observation).

However, the presence of an intercoxa1 process on the elongate prosternum suggests that the former hypothesis is more likely. It was shown in Thvlodrias that the prosterna1 carina was presumably lost, although the relative size of the prosternal bridge appeared unaffected. The intercoxal process of Omalvsus does not articulate with the mesosternal sulcus in the same manner as in elateroids, but this may be a result of the similar incomplete loss observed in

Thvlodrias. There is no evidence that complex ancestral character states are reexpressed due to paedomorphosis and, hence, the mesosternal groove for the reception of the prosternal intercoxal process of

Omalvsus would not be expected if it were a lycid.

Plastoceridae

The Plastoceridae of Crowson (1972:44), not Schwar2 (1907:3) is based on the monotypic genus Plastocerus Schaum (not LeConte), which, according to Crowson, are more closely related to Cebrionidae. The latter nomenclatorial chaos is the result of LeConte's subsequent designation of a type species for the genus after Lacordaire (1857:232) 54 had already done so.

The monotypic genus is known only from Asia minor and Greece.

The status of Plastocerus angulosis Germar requires reexamination as my study of it antedated many of the ideas developed here. Its cantharoid characters are probably the result of paedomorphosis as females are larger than the males, wingless, and have abbreviated elytra

(Crowson, 1972:45). Additionally, males have six visible abdominal sternites and lack an intercoxal process on the third sternite suggestive of paedomorphosis. It will probably be found to be an eiaterid. Lawrence (1987:42) leaves the family in the Cantharoidea, but appears uneasy about this placement.

At least some of the New World Oestodinae (Plastocerus of LeConte and its relatives) of the Elateridae are paedomorphic and the convergent facies explains LeConte's error. Additionally, the cebrionid relationship proposed by Crowson (1971:45) is probably due to convergence during paedomorphic development and the two taxa are unlikely to form a monophyletic lineage.

Cantharidae

The Cantharidae is by far the largest family of the Cantharoidea with over 6000 species. It is cosmopolitan except near the poles and populations are often quite numerous.

The family was examined phylogenetically by Brancucci (1980). He found the reduced wing venation and membranous labrum to be synapomorphies for the family. The paired glands in at least some abdominal segments of the adult and larva and the hydrofuge setation of 55

the larvae may be added to this list.

If the sister of the Cantharidae is the Omethidae as hypothesized

here and by Crowson (1972:62), its status as an outgroup will

necessitate reexamination of the polarities of Brancucci's character

states. But, this action would be premature until larvae of the

Omethidae are known.

Omethidae

Crowson (1972:57) placed three previously disparate lineages in

this family when he defined it. The Matheteinae were formerly placed in the Lampyridae, Dri1ionus in the Drilidae, and the Omethinae proper was a subfamily of the Cantharidae.

The current distribution is North America and the Eastern

Palearctic to include India and Sri Lanka to Japan.

The paired abdominal glands of the Omethinae suggests a possible synapomorphy with the Cantharidae. Their absence in the Driloniinae and Matheteinae may render their relationship equivocal. Whether the apparent polyphyletic nature of this taxon is a result of symplesiomorphies or its basal position is uncertain at this time. No larvae of this family are known and, therefore, resolution of its placement and demonstration of its monophyly is not currently possible.

Phengodidae

The Phengodidae is fairly diverse in adult male morphology, but a well defined lineage by their distinctive larvae and paedomorphic females. The latter are luminescent, elateriform in shape, with 56

exposed heads bearing simple, perforate mandibles whose bases are

distant.

The current distribution is throughout most of the Americas and

the Palearctic from Syria and Iran to China and Southeast Asia.

Wittraer (1972:105) has questioned the placement of Molvchnus

Motschulsky, but it will undoubtedly be demonstrated to be a phengodid.

It appears aberrant from other phengodids only by the presence of a gula. However, this state is undoubtedly plesiomorphic as the only possible outgroup with genal margins closed is the Chauliognathinae of the Cantharidae. Closure of the genal margins over the gula appears to be from the opposite direction than in the cantharids and, thus, is probably not homologous.

All known phengodid larvae are predators of millipedes. Tielman

(1967 ) has discussed the predatory behavior of Zarhipis LeConte in detail. Additionally, I have observed Phengodes Illiger larvae feeding on Narceus Rafinesque and Polvdesmus Latreille as large as themselves in the field. All attacks are not successful though, as a Narceus killed an equal si2ed Phengodes larva by crushing its ventral nerve at the cervix with its mandibles (Miller, unpubl. observ. ). A phenomenon that

I observed in Venezuela and which is not discussed in the literature is the foraging activity of phengodid larvae on the lower vegetation (up to 1 m. high) where millipedes are also found at night. Previous records discuss only ground foraging.

The basal relationship of this family to the Lampyridae has been suggested based on the lack of an organized light organ as seen in the

Lampyridae (Buck, 1948, Cicero, 1988). However, the presence of 57

lanterns in the adult males of Pseudophengodes (Wittmer, 1976;

Lawrence, 1982) similar in gross detail to those of the Lampyridae

suggests that this needs reexamination. Additionally, the light organ

is quite functional Cpers. obs., 19Q6 ). It would be enlightening to know if the fine structure of the modified tracheoles in Pseudophengodes are present and similar with those studied in the adult lampyrids by

Ghiradella (1977, 1973).

Telegeusidae

The Telegeusidae is a small taxon restricted to the Americas between a few kilometers south of the equator in Ecuador to about 35 " north latitude in the deserts of Arizona.

Based on several probable synapomorphies, telegeusids appear to be phengodids with large maxillary palps, but they are, again, probably paedomorphic. If telegeusids and phengodids form a monophyletic lineage, the former are probably basal based on the lack of confluent genal margins (ie. gula present as in Molvchnus Motschuisky) which is synapomorphous for the latter family as currently characterized. As the larvae and females of Telegeusidae are unknown and the known

Phengodidae are paedomorphic, synonymy is not undertaken at this time until more material is available for study.

Dri1idae

Crowson (1972:51) transferred most of the genera formerly thought to be members of the Dri1idae (Wittmer, 1944 and 1948) to the

Lampyridae. Most of the remainder are in the genera Drilus Olivier, 58

Malacogaster Bassi, and Selasia Castelnau which are restricted to the western Paleartic and northern Africa. Their biology has been examined by Crawshay (1903), Cros (1925, 1926, and 1930), and Barker

(1969). No drilids as currently defined (Crowson 1972) are known to be luminescent although elements he moved to the Lampyridae were reportedly so.

Based on several possible synapomorphies, the Drilidae may be

Luciolinae (Lampyridae) with paedomorphic and hypermetamorphic development. Unfortunately we know so little about the latter phenomenon that the placement of hypermetamorphic taxa will remain problematic for some time. Adult characters that suggest relationship may be convergent due to paedomorphosis, and polarization of larval character states would be arbitrary depending on the larval stage selected. It is interesting, however, that the distribution of the

Luciolinae is restricted to the Old World as are the drilids.

Lampyridae

The Lampyridae are a family of about 2000 species which are cosmopolitan except at latitudes higher than 55 degrees.

Crowson (1972:55) redefined the subfamilies of the Lampyridae and changed the composition of the family. He moved Rhagophthalminae and

Mathetinae to the Phengodidae and Omethidae, respectively, and transferred the bulk of the genera of Drilidae into the Otretinae

(Lampyridae). His "higher" (Crowson, 1972:54) and more basal

Lampyridae may be polarized backwards as females of the latter are 59 unknown except in Ototreta Olivier. This absence of females and their morphology is suggestive of paedomorphosis. The phylogenetic relations within this group are still far from resolved and appeal to larvae may take some time as no larvae have yet been associated with adults of the supposed basal taxa. There is some evidence that the Lampyridae may be polyphyletic, but I. have not examined a number of critical genera. At any rate, Crowson's (1972) redefinition of subfamilies has produced a number of significant problems including unplaced taxa.

Lycidae

The Lycidae are a cosmopolitan, well defined family of about 3000 species. Crowson’s claim (1972:53) that no endemic lycids are found in New Zealand must be rejected unless Metriorrhvnchus eraticus Broun is found to be synonymous with an Australian species. Crowson's statement was made in support of his assertion that the Lycidae is a very young group. This hypothesis is difficult to maintain considering the evidence of widespread distribution of a number of lycid genera that can only be explained by their origins antedating the breakup of

Gondwanaland, if not Pangea.

The autapomorphies that define the Lycidae are the morphologically distinctive longitudinally divided styletiform mandibles in the larvae, the longitudinally divided seventh abdominal tergite of the adult male, and the proximal placement of the antennal sockets.

The Lycidae and Lampyridae share a number of derived character states that suggest that they are sister taxa. These states include:

1) reflexed, foliaceous mesepisterna of the adult (found nowhere else in 60

the Coleoptera), 2) onisciform larvae (also in the Silphidae, Cerylidae and a few others), 3) spiracles in upper pleurites instead of the pleural membrane as larvae (widespread, but here hypothesized as a synapomorphy based on outgroup), 4) antennal insertions approximate in the adult (also found in the Chrysomelidae), 5) approximation of bases of larval mandibles, 6) incomplete head closure in the larvae and 7) head retractable within the prothorax of the larvae. The last character state requires additional, comment.

The larvae of the paedomorphic lycid Puliticola paradoxa Mjorberg indicate that Lampyridae and Lycidae are probable sister groups or, alternatively, the Lampyridae is basal to the lineage that gave rise to the Lycidae. This is based on the presence of the completely retractile head in all known lampyrid larvae and the larvae of Lvropaeus

Waterhouse and Puliticola MjBrberg as well as the adult females of the latter. Because the females of Lvropaeus are probably paedomorphic

(Gravely, 1915:361) and Duliticola is known to be paedomorphic

(Mjorberg, 1925), the retractile head may be an expression of paedomorphosis in the larval stage.

Reports of luminescence in trilobite larvae (Kolbe, 1887;

Shelford, 1901) were not confirmed by Mjorberg (1931). If the

Lampyridae is the sister of the Lycidae, luminescence would not be unexpected as a symplesiomorphy. But, as the larvae of lampyrids are often confused with lycids, documentation of the presence of lycid mandibles should accompany any discussion of lycid luminescence.

The abdominal structures found on abdominal segments 2 and 3 of female Thonalmus (Long and Mutchler, 1922; and Miller, 1985) may be 61

evidence of the loss of an intercoxa1 process. They are, however, much

different from the structure found on the abdomen of male Thviodrias

contractus Motschulsky and are not found on Thonalmus males.

Additionally, the genus is not recognizably paedomorphic, but if these

bizarre structures are developmentally a partial reexpression of the

intercoxal piece of the third abdominal segment, it would seem to

indicate that the Lycidae is an unlikely candidate for a basal position

in the Coleoptera.

The relationships within the family need reexamination. It has

been suggested that paedomorphic taxa are basal (Crowson, 1972:49) which would imply higher taxonomic levels (ie. tribal and subfamilial) for

those lineages such as Puliticola. Lvropaeus and the leptolycines.

While initially appealing, such a decision should be approached cautiously in light of the previous discussion of paedomorphosis. Work

in progress suggests a major overhaul of the higher classification within the family is needed and that these taxa are in fact more derived. THE TRIBE LEPTOLYCINI

The Leptolycini was originally conceived for the monotypic

Leptolvcus by Leng and Mutchler (1922:430), Although they did not elaborate their reasons, the decision for tribal status was likely due to the widely divergent morphology of Leptolvcus heterocornis L. & M. from the morphology of most described Lycidae. In fact, the general facies of L. heterocornis might suggest relationship only with Lvropaeus

Waterhouse and Dexoris Waterhouse.

Subsequent discussion of the tribe has since been limited to placement in catalogues (Kleine, 1933:18 and Blackwelder, 1945:343) and a cursory discussion of relationships (Crowson, 1972:48). No new members have been added, even though several species here included have been placed previously in the Calopterini by Pic (1911). He did not note any relationship with Leptolvcus in his brief descriptions, but this may have been the result of several reasons. He probably never saw specimens of L. heterocornis as none were found in his collection or in the NMHP. The lack of diagnostic characters for the tribe except the mention of the included species may have prevented insight into relationship. And, although illustrated, L. heterocornis appears more derived than most other members of the lineage.

Because of Pic's ignorance of the tribe and because of his brief descriptions, it was necessary to examine the types of all the lycid

62 63

genera he described. All except Microlvropaeus notaticol1is Pic

(1929:4) were located and examined. Although not seen, the latter

species is probably a small Lvropaeus based on its description, its distribution, and the large morphological variation observed within that genus (Miller, unpub1. data).

Although several other Neotropical genera not included here have similar facies to the Leptolycini as presently defined, character state distribution suggests that their inclusion in the tribe would result in a polyphyletic classification. The relationships of these disparate elements will be discussed in other work in progress. Additionally, the continued recognition of tribal status of the Leptolycini may render the Calopterini paraphyletic, but more data are required to adequately evaluate this alternative hypothesis. This will be discussed at greater length in the section on phylogeny (p. 306).

The biology of the leptolycines is essentially unknown. The discovery of the paedomorphic females of L. heterocornis and the absence of females resembling adult males in other leptolycines suggests that all may be paedomorphic, but this remains to be ascertained. Even the distributions of the species are poorly known as they usually include only the type locality. The few available collections by Berlese extraction methods suggest that the larvae live on or in the soil and leaf litter. None have been recovered from decomposing wood.

Although the males of 37 species and the females and the larvae of

L. heterocornis are here described or redescribed, the scope of this lineage remains unknown. Because of the small size of the males and their resemblance to gnats while in flight (pers. obs. ) they are not often captured by Coleopterists. The use of malaise and flight intercept traps appears to be ameliorating this artifact, but the number of newly discovered species to previously encountered species throughout this study has remained high. This suggests that many additional species will yet be found. 65

Leptolycini Leng and Mutchler

Leptolycini Leng and Mutchler, 1922:430. Kleine, 1933:18.

Blackwelder, 1945:343.

TYPE: Leptolvcus heterocornis Leng and Mutchler, AMNH.

DIAGNOSIS (Adult males). This tribe may be differentiated from all other lycids by the combination of: the plantar pad of tarsomeres 4 not expanded laterally; the presence of a median longitudinal pronotal carina that is divided from the middle to or near the posterior margin; and the absence of lateral pronotal carinae. However, the pronotum may be only obscurely carinate. The head tends to be more elongate and the antennal prominence more porrect than most lycids except the Oriental

Lvropaeus and the Ethiopian Dexoris. especially in the more derived members of the lineage.

DIAGNOSIS (Adult females). Probably all paedomorphic. The only known female is recognizable by the larval character states of leptolycines except for the absence of mandibles and only one pair of pleurites per abdominal segment.

DIAGNOSIS (Immatures). Larval Leptolycini are distinct from all other lycid larvae examined to date by the membranous gena and the incision on the lateral margin of the antennal prominence. Only Leptolvcus heterocorni s larvae are described herein, but I have seen 6 species that cannot yet be associated with described species. They are, however, undoubtedly leptolycines. The only other known lycids on the Greater 66

Antilles are Thonalmus in which the larvae differ significantly (Miller, in prep). The pupae of Leptolycini remain unknown. 67

KEY TO GENERA OF THE LEPTOLYCINI (ADULT MALES)

1. Pedicel length much smaller than antennomere 3 ...... 2

1'. Pedicel length subequal to antennomere 3 ...... 3

2. (1) Antennae strongly compressed, serrate; elytra 4-costate,

prosternum T- or Y-shaped ...... Pseudoceratoorion (p. 285)

2'. (1) Antennae not or barely compressed, serrate to filiform;

elytra 0-3 costate; prosternum T- or Y- shaped ......

...... Abrolvcus (p. 69)

3. (1*) Antennae serrate or filiform; antennomere 4 longer than 2

and 3 combined ..... 4

3'. (I1 ) Antennae flabellate; length antennomere 4 varies ...... 5

4. (3) Lateral pronotal margins obscure anteriorly; median

longitudinal pronotal carina obscure; prosternum

trianguliform; elytra 2-costate ...... Leptolvcus (p. 215)

4’. (3) Lateral pronotal margins complete; median longitudinal

pronotal carina well defined; prosternum Y- or T-shaped;

elytra 4-costate ...... Ceratoprion (p. 137) Antennomere 4 longer than 2 and 3 combined; prosternum triangular; elytra 2-costate ...... Anti 1lolycus (p. 109)

Antennomere not longer than 2 and 3 combined; prosternum Y— or T-shaped ...... 6

Third antennomere with ramus, modified scaliform setae absent on antennae, elytron 4-costate, ......

...... Pseudacro1eptus (p. 259)

Third antennomere without ramus, modified scaliform setae present on antennae, elytron 3-costate ......

...... F label locaenia (p. 175 ) 69

Abrolvcus NEW GENUS

TYPE. Abrolvcus bicolor new species, INHS.

DIAGNOSIS. This genus can most easily be differentiated from other leptolycines by the noncompressed, filiform antennae with all antennomeres, except 2, subequal in length. Additionally, the prosternum is Y- or T-shaped; the pronotal carinae are obscure and the median fovea is indistinct; the elytra are 0 to 3 costate; the pronotum lacks pronotal stemmata; and there are no scaliform setae on the antennae.

DISTRIBUTION. This genus is known only from the Greater Antillian islands of Cuba, Hispaniola, and Puerto Rico.

DESCRIPTION (Male). Small, slender, elongate, length 1.9-2.9 mm., width 0.3-0.8 mm at humerus. Brown to black; some with golden yellow and white maculation.

Head. Elongate, widest behind eyes, then narrowing to posterior margin; disk shining, sparsely to densely punctate; punctations deep to shallow; setation sparse, semi-decumbent. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally.

Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3-0.7x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect, strongly gibbose; length 0.8-lx width, inserted between middle of eyes; pentagonal to rectangular dorsally,

non-sulcate dorsomedially; ventrally pentagonal to trianguliform,

concave to flat, widest at base, dorsolateral margins straight.

Antennal sockets triangular to oval to subrectangular, separated by 0,1-

0.5x width of epistomal margin and 0.3-2x separation between eye and

antennal socket. Antennifer insertion elevated medially, appearing

basomedial. Interocular distance 0.5-2x width eye dorsally. Vertex

uniformly to irregularly punctate or areolate, transversely convex, almost flat; depression of tentorial maculae obscure to broadly shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, shallow to deep, indistinct to distinct; at or before base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, height 0.5-

0.8x width, with or without triangular gibbosity. Epistomal margin broadly to deeply emarginate to truncate, width 0.4-0.8x frontal distance between eyes. Labrum quadrate, length 0.5-lx width, apically truncate to broadly rounded or broadly emarginate. Maxillary palps 4 segmented, palpomeres 1 and 4 cylindrical, diameter nearly equal to

length; palpomere 2 barrel-shaped, length 2x antennomere 1, diameter equal length; diameter palpomere 3 slightly less than 2; palpomere 4 nearly equal length 3, diameter at base less than 0.5x palpomere 3,

inserted subapically and internally, acuminate. Labial palps 1 segmented, minute and digitiforra or absent. Gula elongate, subrectangular; width slightly greater apically than basally; length 1-

2x width apical margin, inserted anteriorad posterior margin of postgenae. Antennae filiform, length 0.5-0.8x body length, setation dense to moderately dense, uniformly distributed, decumbent to semi-

decumbent on all antennomores, scaliform setae present; scape obconic, width apically 0.5-0.8x length; pedicel cylindrical, short, transverse,

not compressed, length 0.2-0.5x length of antennomere 3; antennomere 3 not adnate to base of antennomere 4, length equal antennomere 4;

antennomeres 3-10 conical, subserrate, expanded apically, weakly compressed; antennomere 11 fusiform, barely compressed, length 1.3x antennomere 3.

Thorax, Pronotum trapezoidal, transverse, length 0.3-0.5x basal width, anterior width 0.4-0.8x basal width; median longitudinal carina absent, median fovea obscure to slightly more distinct and extending up to posterior 0.6x pronotal length; pronotal stemmata absent; all to various margins not swolen; anterior margin straight to slightly convexly arcuate, anterior angles obtuse to acute; lateral margin complete, carinate to foliaceous and erect, straight to slightly concavely arcuate longitudinally; posterior margin weakly to strongly bisinuate; posterior angles acute, expanded laterally 0.8~0,9x width elytral humerus; disk convex, finely rugose, anterior corners deeply

impressed or not, discal setae shorter than elytral setae, decumbent, posterior disk elevated near margin; hypomeron concave to deeply concave, height 0.5~0,6x length, all to various margins swollen, hypomeral stemmata absent; prosternal bridge Y- to T-shaped, may be T- shaped and nearly trianguliform, anterior margin incised to shallowly and broadly emarginate, length 0.3-0.5x width, intercoxal process short and not carinate to carinate. Scutellum pentagonal, distally quadrate, nearly square, medially impressed or not, lateral margins parallel to slightly converging or diverging apically, apex truncate to deeply

emarginate, emargination to 0.5x scutellar length, apices divergent and

acute or not. Mesospiracle prominent to obscure with peritreme not or

barely exposed, orifice not compressed when peritreme exposed.

Mesepisternum elongate, posterior margin straight to convexly arcuate.

Mesepimeron trianguliform, width 0.3-0.5x and height 0.5-0.7x height mesepisternum; posterior margin very little reflexed, straight to arcuate, acutely angulate ventrally. Mesosternum elongate, posterior margin convexly arcuate to truncate. Mesocoxal separation 0.7-lx mesocoxal diameter at base. Elytra ligulate, slightly dehiscent from scutellum to elytral middle, gradually narrowing distally or not, apices rounded; length 7-10x pronotal length; not costate to tricostate, costa

4 distinct when visible 0.1-0.8x elytral length; costa 6 distinct 0.8x elytral length when visible; humeral costa distinct over umbone and 0.8x elytral length when visible; disk irregualrly to regualrly scabrous- punctate to finely punctate to reticulate; setation semi-decumbent to semi-erect, sparse, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight to strongly arcuate. Metasternum completely to

incompletely divided by medial longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent to decumbent, uniformly distributed to distributed along margins; coxae conical, pro- and mesotrochanteral socket 0.3-0.5x coxal length, metacoxae more vertically oriented and directed dorsoanteriorly; trochanters elongate to elongate triangular,

length pro- and mesotrochanters 0.8-lx coxal length, metatrochanters

0.5-0.8x metacoxal length, all femoral insertions oblique and offset to truncate and not offset to only metatrochanter slightly offset; femora strongly compressed, straight to arcuate, fusiform, 2-4x trochanteral length, annulated basally; tibiae weakly to strongly compressed, inner margin slightly sinuate; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi—decumbent to semi-erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing posteriorad uniformly or from segment 4, tergite

8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing posteriad uniformly or from segment 4, sternite 7 deeply emarginate to entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with or without parameres; parameres 0.3x length median lobe when present; basal piece large, dorsoventrally compressed to tubular, length 0.2-3x median lobe length; median lobe tubular, unmodified, membranous apically to ventrally 0.7x length; internal sac not eversible; flagellum absent.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). See the species description of the abberrent leptolycine larva collected on Mt. Britton, Puerto Rico, hypothesized to be Abrolvcus bicolor n. sp. No generic description is attempted at this time, because only the unique specimen has been seen.

DISCUSSION. The interspecific variation in this genus is fairly 74 extensive. For example, the number of costae might suggest numerous genera if the traditional taxonomic methods used in the family were accepted (but, see character discussion). Additionally, the

intraspecific variation of this lineage is unknown, because all species are known from single specimens. These constitute the smallest lycids I have seen to date.

ETYMOLOGY. The combination is from the Greek prefix abro meaning delicate, pretty and the root lvcus. the type genus of the family and

Greek for wolf. 75

KEY TO THE SPECIES OF Abrolvcus (ADULT MALES)

1. Prothorax transverse, width greater than 2x length ...... 2

1 ' . Prothorax width less than 2x length ...... 6

2. (1) Uniformly colored; humeral costa distinct greater than 0.8x

elytral length ...... 3

2'. (1) Not uniformly colored: humeral costa variously expressed ...

...... 4

3. (2) Light brown coloration; elytral surface rugose, not

transversely reticulate; Cuba ..... cubensi s n. sp. (p. 82)

3'. (2) Gray coloration; elytra tranversely reticulate; Cuba ......

...... parda de Zayas (p. 102)

4. <2') Pronotum blackish brown; elytral base yellow; antennomere 11

black; humeral costa greater than 0.8x elytral length;

Puerto Rico ...... bicolor n. sp. (p. 77)

4'. (2' ) Pronotum yellow; antennomere 11 yellow to white ...... 5

5. (41) Humeral costa less than 0.5x elytral length; vertex densely

punctate; Cuba ...... sandersoni n. sp. (p. 104) Humeral costa greater than 0.8x elytral length; vertex sparsely punctate; Hispaniola ...... iviei n. sp. (p. 92)

Brown elytra basally yellow; elytral surface distinctly reticulate, not granulate; humeral costa distinct 0.8x elytral length; femora and tibae barely compressed; Cuba ...

...... omalysiformis n. sp. (p. 97)

Elytra uniformly dark brown colored, except elytral insertions lighter brown,; elytral surface granulate, not reticulate; humeral costa distinct less than 0.5x elytral length; femora and tibiae strongly compressed; Hispaniola ..

...... dar 1 ingtoni n. sp. (p. 87) 77

Abrolvcus bicolor NEW SPECIES

Fig. 4 a-e

TYPE. CNCI, male with the following data: Cwhite printed label] "Vista la Sierra \\ Luquillo Forest \\ P. R. VII 9, 1969 \\ H, & A. Howden".

DIAGNOSIS. Males can be distinguished from other males of Abrolvcus by the combined presence of the bicolored elytra with black antennomere 11; the pronotum is transverse with its length 2x its length; the elytra are bicostate with the humeral costa distinct for 0.8x of the elytral length.

The hypothesized larva of this species can be easily be differentiated from all other known Lycidae by the round head which differs from the normal triangular shape.

DISTRIBUTION. Known only to occur in the mountains of Puerto Rico.

DESCRIPTION (Male). Small, slender, elongate, length 2.1-2.9 mm., width 0.3 mm at humerus. Black; except basal 0.2x elytra, dorsal apex mesepimeron, metepimeron and metepisternum golden yellow; venter of head and scutellum yellowish brown.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, strongly and deeply punctate; setation sparse, semi-decumbent.

Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.7x radius when dorsally viewed.

Antennal prominence distinct from vertex, porrect, strongly gibbose;

length 0.8x width, inserted between middle of eyes; pentagonal dorsally,

narrowly sulcate dorsomedially; ventrally pentagonal, concave, widest at

base, dorsolateral margins straight. Antennal sockets triangular,

separated by O.lx width of epistomal margin and 0.5x distance between

eye and antennal socket. Antennifer insertion elevated medially,

appearing basomedial. Interocular distance 1.5x width eye dorsally.

Vertex uniformly punctate, transversely convex, almost flat; depression

of tentorial maculae broadly shallow, from base of antennal prominence

to posterior eye margin, more distinct anteriorly. Tentorial maculae

separate, shallow, distinct; at base of antennal prominence. Frons

transversely convex, distinct from lower surface of antennal prominence,

height 0.8x width. Epistomal margin broadly emarginate, width nearly

0.7x frontal distance between eyes. Labrum transverse, length 0.5x

width, apical margin broadly rounded. Maxillary palps 4 segmented, palpomeres 1 and 4 cylindrical, diameter nearly equal to length;

palpomere 2 barrel-shaped, length 2x antennomere 1, diameter 2x length; diameter palpomere 3 slightly less than 2; palpomere 4 nearly equal 3,

diameter at base less than 0.5x palpomere 3, inserted subapically and

internally, acuminate. Labial palps 1 segmented, minute and

digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted

anteriorad of posterior margin postgenae. Antennae filiform, length

0.7x body length, setation dense, uniformly distributed, decumbent on all antennomeres, scaliform setae present; scape obconic, width apically 79

0.7x length; pedicel cylindrical, short, transverse, not compressed,

length 0.3x length of antennomere 3; antennomere 3 not adnate to base of

antennomere 4, length equal antennomere 4; antennomeres 3-10 conical,

subserrate, expanded apically, barely compressed; antennomere 11

fusiform, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly 0.8x basal width; median longitudinal

carina absent, median fovea obscure; pronotal stemmata absent; anterior and lateral margins swollen, posterior margin not curbed; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, erect, slightly concavely arcuate longitudinally; posterior margin weakly bisinuate; posterior angles acute, expanded laterally G.8x width elytral humerus; disk convex, finely rugose, anterior corners moderately impressed, discal setae shorter than elytral setae, decumbent, posterior disk slightly elevated near margin; hypomeron deeply concave, height 0.5x length, all margins swollen, hypomerai stemmata absent; prosternal bridge T-shaped, length 0.5x width,

intercoxal process carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex emarginate, emargination nearly O.lx scutellar length, apices not acute and not divergent.

Mesospiracle not prominent, peritreme barely visible, orifice not compressed. Mesepisternum elongate, posterior margin arcuate.

Mesepimeron trianguliform, width 0.3x and height 0.7x mesepisternum; posterior margin not reflexed, straight, acutely angulate ventrally.

Mesosternum elongate, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, slightly ao dehiscent from scutellum, gradually narrowing distally, apices narrowly rounded; length 8x pronotal length; bicostate, costa 4 distinct from base to Q.8x elytral length, humeral costa distinct over umbone and 0.8x elytral length; disk strongly and irregularly punctate; setation semi- decumbent, sparse, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin strongly arcuate. Metasternum divided incompletely by medial

longitudinal sulcus, not reaching mesosternum by mesocoxal diameter at base. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.5x coxal length, metacoxae more elevated and less transversely oriented; trochanters elongate, length pro - and mesotrochanters subequal to coxal length, metatrochanters 0.8x metacoxal length, femoral insertions oblique and slightly offset; femora strongly compressed, fusiform, 3x trochanteral length, annulated basally; tibiae strongly compressed, inner margin slightly sinuate; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads; and covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, but narrowing uniformly posteriad from segment 4, tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, but narrowing uniformly posteriad from segment 4, sternite 7 deeply emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece large, dorsoventraily compressed, length almost

3x median lobe length (Fig. 4 a-c); median lobe tubular, unmodified; 81

internal sac not eversible; flagellum absent.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. A single specimen collected on Mt. Britton is here hypothesized to be the larva of this species. It shares the synapomorphies of other known Leptolycini and ist undoubtedly, not a larva of Thonalmus chevrolati Bourgeois, the only other lycid lineage known from the island. Only three leptolycines are known to inhabit

Puerto Rico. This includes two species of Leptolvcus. which the larva of L- heterocornis L. & M. is known, and A. bicolor. Because this larva is so abberrent, it is presumed to belong to a different genus than L. heterocornis. which narrows the choice to Abrolvcus.

MATERIAL EXAMINED. Puerto Rico, Villalba, Dona Juana Forest, 3 Nov. 1952

(1-JARC); Vista la Sierra, Luquillo Forest, 9 July 1969 (1-CNC, Type);

(1 larva-INHS).

ETYMOLOGY. The specific epithet refers to the black and golden yellow elytra. 82

Abrolvcus cubensis NEW SPECIES

Fig, 5 a-c

TYPE. INHS, male in good condition with the following label data:

Cwhite printed label! "CUBA: Oriente Prov. \\ Loma (Pico) del Gato \\

Sierra Maestra \\ 26-28 May 1959 \\ M. W, Sanderson".

DIAGNOSIS. This species is separable from other leptolycines by the

character states in the key to Abrolvcus and the unique median

triangular gibbosity produced on the frons. It has unicostate elytra

and the anterior corners of the pronotal disk are deeply impressed.

DISTRIBUTION. Known only from the southern coastal range in eastern

Cuba.

DESCRIPTION (Male): Small, slender, elongate, length 2.1 mm., width 0.55

mm. at humerus. Color light brown.

Head. Elongate, widest behind eyes, then narrowing to posterior

margin, shining areolate; setae sparse, semi-decumbent. Genal margin

elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed

dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.5x radius when dorsally viewed. Antennal prominence distinct from vertex, gibbose; length subequal width,

inserted between middle of eyes; pentagonal dorsally, narrowly sulcate dorsomedially; ventrally pentagonal, concave, widest near base, dorsolateral margins straight. Antennal sockets subrectangular, dorsal

and lateral margins evenly rounded, separated by 0.2x width of epistomai

margin and 0.5x distance between eye and antennal socket. Antennifer

insertion elevated medially, appearing basomedial. Interocular

distance nearly 2x width eye dorsally. Vertex areolate, transversely

flat, setae semi-decumbent; depression of tentorial maculae obscure.

Tentorial maculae separate, shallow, distinct; at base of antennal

prominence. Frons transversely convex, distinct from lower surface of

antennal prominence, transversely narrow, height nearly 0.5x width;

produced medially into triangular gibbosity. Epistomai margin

truncate, width 0.8x frontal distance between eyes. Labrum transverse,

rectangular, length 0.5x width, apical margin truncate. Maxillary

palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to

length; length palpomere 4 nearly equal 3, diameter at base less than

0.5x palpomere 3, inserted, acuminate. Labial palps 1 segmented, digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length lx width apical margin, inserted anteriorad posterior margin of postgena. Antennae filiform, 0.8x body

length; setation moderately dense, semi-decumbent, uniformly distributed, scaliform setae present; scape obconic, width apically 0.6x

length; pedicel cylindrical, short, transverse, not compressed, length

0.3x length of antennomere 3; antennomere 3 not adnate with base antennomere 4, length equal antennomere 4; antennomeres 3— 10 conical,

subserrate, expanded apically, barely compressed; antennomere 11 fusiform, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly G.7x basal width; median longitudinal

carina obscure, median fovea slightly more distinct; pronotal stemmata

absent; all discal margins swollen; anterior margin straight, anterior

angles obtuse; lateral margin complete, foliaceous, explanate, slightly

concavely arcuate; posterior margin weakly bisinuate; posterior angles

acute, expanded laterally 0.8x width elytral humerus; disk convex,

anterior corners deeply impressed, discal setae shorter than elytral

setae, decumbent, posterior disk elevated near margin; hyporaeron

shallowly concave, height 0.5x width, margins not swollen, hypomerai

stemmata absent; prosternal bridge T-shaped, length 0.3x width,

intercoxal process carinate. Scutellum pentagonal, distally quadrate,

nearly square, lateral margins parallel, apex deeply emarginate,

emargination nearly 0.3x scutellar length, apices acute and divaricate.

Mesospiracle obscure, peritreme not visible. Mesepisternum elongate,

posterior margin straight. Mesepimeron trianguliform, width 0.5x and

height 0.7x height mesepisternum; posterior margin very little reflexed,

nearly straight, acutely angulate ventrally. Mesosternum elongate,

posterior margin truncate. Mesocoxal separation 0.8x mesocoxal

diameter at base. Elytra ligulate, slightly dehiscent from elytral

middle, barely narrowing distally, apices broadly rounded; length 9x

pronotal length; unicostate, humeral costa distinct 0.8x elytral length;

disk irregularly scabrous-punctate; setation semi-erect, sparse, uniformly distributed. Metepisternum with ventral margin increasingly

arcuate posteriorly. Metepiraeron with ventral margin arcuate.

Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent, uniformly distributed; coxae 85 conical, trochanteral socket less than 0.5x coxal length, metacoxa more vertically oriented and directed dorsoanteriorly; trochanters short, length pro- and mesotrochanters subequal to coxa! length, metatrochanters 0.7x metacoxal length, femoral insertions truncate and not offset; femora strongly compressed, straight, length moderate, 4x trochanteral length, annulated basally; tibiae strongly compressed, internal margin strongly sinuate; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length equal length of median lobe (Fig. 5 a-c); median lobe tubular, unmodified, membranous ventrally at apex; internal sac not eversible; flagellum absent.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. CUBA, Oriente Provence, Loma (Pico) del Gato, Sierra

Maestra, 26-28 May 1959, M. W. Sanderson (1 INHS, holotype). 86

ETYMOLOGY. The specific epithet refers to the island where the type and only specimen was collected. 87

Abrolvcus dar1ingtoni NEW SPECIES

Fig. 6 a-d

TYPE: MCZC, male with the following label data: Cwhite printed label]

"fthills. Cord. Cent. \\ S. of Santiago \\ June '38. Dom. Rep. \\

Dariington."

DIAGNOSIS: This species is easily distinguished from other known

Abrolvcus by the non-costate elytra, the stout, non-compressed antennae with short bristling setae, the stout compressed legs with offset trochanters, and the glabrous, shining, and areolate head with a rectangular antennal prominence.

DISTRIBUTION. Only known from northern Hispaniola.

DESCRIPTION (Male): Small, slender, elongate, length 2.7 mm., width

0.4 mm at humerus. Blackish brown, except elytral base briefly testaceous.

Head, Elongate, widest behind eyes, then narrowing to posterior margin, shining areolate; setae sparse, semi-decumbent. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.5x radius when viewed dorsally. Antennal prominence distinct from vertex, gibbose; length O.Qx width, inserted between middle of eyes; rectangular dorsally, broadly sulcate dorsomedially, sulcus anteriorly narrow; ventrally trianguliform,

concave, widest near base, sides uniformly and convexly arcuate.

Antennal sockets oval, separated by 0.5x width of epistomai margin and

0.3x distance between eye and antennal socket. Antennifer insertion

elevated medially, appearing basomedial. Interocular distance 2x width

eye dorsally. Vertex areolate, transversely flat, setae semi-erect;

depression of tentorial maculae broadly shallow, widened posteriorly to

near posterior eye margin. Tentorial maculae separate, deep, distinct;

slightly anterior to base of antennal prominence. Frons transversely

convex, distinct from lower surface of antennal prominence, transversely

narrow, height nearly 0.5x width. Epistomai margin broadly and deeply

emarginate, width 0.4x frontal distance between eyes. Labrum

transverse, rectangular, length 0.5x width, apical margin broadly and

distinctly emarginate. Maxillary palps 4 segmented, palpomeres 1-3

cylindrical, diameter nearly equal to length; length palpomere 4 nearly

equal 3, diameter at base less than 0.5x palpomere 3, acuminate.

Labial palps 1 segmented, digitiform. Gula elongate, subrectangular;

width slightly greater apically than basally; length 2x width apical

margin; inserted anteriorad posterior margin of postgenae. Antennae

filiform, 0.5x body length; setation short, moderately dense, erect and bristling, uniformly distributed, scaliform setae present; scape

obconic, width apically 0.8x length; pedicel cylindrical, short,

transverse, not compressed, length 0.5x length of antennomere 3; antennomere 3 not adnate to base of 4, length lx antennomere 4; antennomeres 3-10 conical, subserrate, expanded apically, barely

compressed; antennomere 11 fusiform, length 1.3x antennomere 3. Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly 0.8x basal width; median longitudinal carina obscure, median fovea obscure; pronotal stemmata absent; all discal margins swollen; anterior margin broadly emarginate, anterior angles obtuse; lateral margin complete, not foliaceous, not reflexed, concavely arcuate longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk arcuate, surface granulate, discal setae sparse, semi-erect; hypomeron shallowly concave, height 0.6x width; all margins not swollen; hypomeral stemmata absent; prosternal bridge T-shaped, slightly incised medially, length 0.5x width, intercoxal process carinate. Scutellum pentagonal, distally trapezoidal, lateral margins divergent posteriorly, apex deeply emarginate, emargination nearly 0.5x scutellar length, apices acute and divergent. Mesospiracle obscure, peritreme not visible. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.5x and height 0.5x mesepisternum; posterior margin very little reflexed, nearly straight, acutely angulate ventrally. Mesosternum elongate, posterior margin convexly arcuate.

Mesocoxal separation 0.8x mesocoxal diameter at base. Elytra ligulate, slightly dehiscent behind scutellum, not narrowing distally, apices broadly rounded; length lOx pronotal length; not costate; disk irregularly scabrous; setation semi-erect, sparse, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight.

Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent, uniformly distributed; coxae 90

conical, pro- and mesotrochanteral socket less than 0.5x coxal length,

metacoxa more vertically oriented and directed dorsoanteriorly;

trochanters elongate triangular, length pro- and mesotrochanters

subequal to coxal length, metatrochanters 0.7x metacoxal length, all

femoral insertions oblique and slightly offset; femora stout, slightly

compressed, straight, length moderate, 4x trochanteral length, annulated

basally; tibiae slightly compressed, internal margin slightly arcuate;

tibial spines small and similar; tarsi elongate, tarsomeres without

laterally expanded plantar pads; covered with semi-erect bristling

setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9

medially divided longitudinally; tergite 10 shield shaped. Sternites

rectangular, wider than long, length 1-8 subequal, but narrowing

uniformly posteriad; sternite 7 entire. Spiracles in pleural membrane,

located subraedially from apical margin of tagma. Aedeagus with

parameres; paramere length 0.3x median lobe; basal piece compressed

dorsoventrally, length 0,2x aedeagal length (Fig, 6a-c); median lobe

tubular, unmodified, membranous ventrally at apex; internal sac not

eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. In the description the anterior margin of the unique type 91 is described as broadly emarginate, but his may be an artifact of preservation.

MATERIAL EXAMINED. DOMINICAN REPUBLIC, Cordillera Central, South of

Santiago, June 1938, P. Darlington (1-MCZC, holotype).

ETYMOLOGY. This species is named in honor of the late Phillip

Darlington who advanced our knowledge of lycids in the Greater Antilles by his collecting efforts and by his study of mimicry (Darlington,

1938 ). 92

Abrolvcus i viei NEW SPECIES

Fig. 7a-c

TYPE. NMNH, male with the following data: Cwhite printed label]

"DOMINIC. REP.: Prov. La Vega \\ 12 km NE Jarabacos, 550 m, \\ 01-07 Sep

19BB, flight inter. \\ trap. M. A. Ivie, T. K. Phillips \\ & K. A.

Johnson coirs."

DIAGNOSIS. Males can be distinguished from other males of Abrolvcus by the black coloration with golden yellow prothorax and frons and the unicostate elytra. Additionally, the anterior corners of the pronotal disk are deeply impressed which also occurs in the unicolorous A. cubensis. Again, unlike other Abrolvcus the ventral surface of the antennal prominence is flat.

DISTRIBUTION. Known only to occur in the central mountains of the

Dominican Republic on the island of Hispaniola.

DESCRIPTION

0.5 mm at humerus. Black; except pronotum and frons golden yellow and ultimate antennomere white.

Head. Elongate, widest behind eyes, then narrowing to posterior margin; disk shining, sparsely punctate, punctations strong and deep; setation sparse, semi-decumbent. Genal margin elongate, length greater than 2x eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect, strongly gibbose; length equal width, inserted between middle of eyes; pentagonal dorsally, not sulcate dorsomedially; ventrally pentagonal, flat, widest at base, dorsolateral margins straight. Antennal sockets oval, separated by 0.3x width of epistomai margin and equidistant separation between eye and antennal socket.

Antennifer insertion elevated medially, appearing basomedial.

Interocular distance 2x width eye dorsally. Vertex irregularly punctate, transversely convex, almost flat; depression of tentorial maculae broadly shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, height 0.7x width. Epistomai margin broadly emarginate, width nearly 0.7x frontal distance between eyes. Labrum quadrate, length equal width, broadly rounded apically. Maxillary palps 4 segmented, palpomeres 1 and 3 cylindrical, diameter nearly equal to length; palpomere 2 barrel-shaped, length 2x antennomere 1, diameter equal length; diameter palpomere 3 slightly less than 2; palpomere 4 nearly equal length 3, diameter at base less than 0.5x palpomere 3, inserted subapically and internally, acuminate. Labial palps absent. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted anteriorad posterior margin of postgenae. Antennae filiform, length 0.8x body length, setation dense, uniformly distributed, semi- decumbent on all antennomeres, seal iform setae present; scape obconic, 94

width apically 0.5x length; pedicel cylindrical, short, transverse, not

compressed, length 0.3x length of antennomere 3; antennomere 3 not

adnate to base of antennomere 4, length equal antennomere 4;

antennomeres 3-10 conical, subserrate, expanded apically, weakly

compressed; antennomere 11 fusiform, barely compressed, length 1.3x

antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.3x basal

width, anterior width 0.7x basal width; median longitudinal carina

absent, median fovea obscure; pronotal stemmata absent; anterior and

lateral margins swollen, posterior margin not curbed; anterior margin

straight, anterior angles obtuse; lateral margin complete, foliaceous,

erect, straight longitudinally; posterior margin strongly bisinuate;

posterior angles acute, expanded laterally 0,9x width elytral humerus;

disk convex, finely rugose, anterior corners deeply impressed, discal

setae shorter than elytral setae, decumbent; hypomeron deeply concave

anteriorly, height 0.6x length, lateral hypomeral margins weakly

swollen, ventral margin not swollen, hypomeral stemmata absent;

prosternal bridge T-shaped, almost trianguliform, anterior margin

shallowly and broadly emarginate, length less than 0.5x width,

intercoxal process short and not carinate. Scute11urn pentagonal,

distally quadrate, nearly square, lateral margins slightly converging

apically, apex truncate, emargination nearly 0.lx scutellar length,

apices not divergent or acute. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.3x and height 0.5x height mesepisternum; posterior margin very little reflexed, nearly straight, acutely angulate ventrally. Mesosternum elongate, posterior margin

convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base.

Elytra ligulate, slightly dehiscent from middle, narrowing slightly

distally, apex broadly rounded; length 7x pronotal length; unicostate,

humeral costa distinct over umbone and 0.8x elytral length; disk

irregularly scabrous-punctate, not reticulate; setation semi-decumbent,

sparse, uniformly distributed. Metepisternum with ventral margin

increasingly arcuate posteriorly. Metepimeron with ventral margin

arcuate. Metasternum divided completely by medial longitudinal sulcus.

Legs elongate, setation sparse, semi-decumbent, uniformly distributed;

Coxae conical, pro- and mesotrochanteral socket 0.5x coxal length,

metacoxae more vertically oriented and directed dorsoanteriorly;

trochanters elongate, length pro- and mesotrochanters subequal to coxal

length, metatrochanters 0.7x metacoxal length, all femoral insertions

truncate and not offset except metatrochanter slightly offset; femora

strongly compressed, fusiform, 2.5x trochanteral length, annulated

basally; tibiae strongly compressed, inner margin slightly sinuate;

tibial spines small and similar; tarsi elongate, tarsomeres without

laterally expanded plantar pads; covered with semi-decumbent bristling

setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing posteriorad segment 4, tergite 8 entire; tergite

9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing

uniformly posteriad from segment 4, sternite 7 deeply emarginate.

Spiracles in pleural membrane, located submedially from apical margin of 96

tagma. Aedeagus without parameres; basal piece large, dorsoventrally

compressed, length almost equal median lobe length (Fig. 7a-c>; median

lobe tubular, unmodified, ventrally membranous 0.2x length at middle;

internal sac not eversible; flagellum absent.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. Although a considerable number of leptolycines were captured on the same trip, this is the only Abrolvcus and is represented by only one specimen. Either members of this genus are much more rare than other leptolycines or their behavior differs significantly.

MATERIAL EXAMINED. DOMINICAN REPUBLIC, Province of La Vega, 12 km NE

Jarabacos, 550 m, 01-07 Sep 1988, M. A. Ivie, T. K. Phillips, & K. A.

Johnson coirs (1-NMNH, holotype).

ETYMOLOGY. The specific epithet is to acknowledge the efforts Michael

A. Ivie in his efforts to aquaint the world with the beetle fauna of the

Virgin Islands and more recently the Greater Antilles, as well as his specific efforts to provide me with additional material for this study. 97

Abro1vcus omalvsiformi s NEW SPECIES

Fig. 8 a-d

TYPE. INHS, male in good condition with the following label data: Cwhite

printed label] "CUBA: Oriente Prov. \\ Loma (Pico) del Gato \\ Sierra

Maestra \\ 26-28 May 1959 \\ M. W. Sanderson".

DIAGNOSIS. This species can be easily separated form other Abrolvcus by its weakly compressed legs versus other Abrolvcus strongly compressed, the elytra are bicostate, and the head is punctate dorsally and areolate ventrally. In addition the elongate, tubular, and medially hinged male genitalia is very distinctive within the Lycidae.

DISTRIBUTION. Known only from Cuba.

DESCRIPTION (Male): Small, slender, elongate, length 2.9 mm., width 0.8 mm at humerus. Dark brown; except apical 0.3x elytra yellow, maxillary palps testaceous.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, strongly and deeply punctate dorsally and areolate ventrally; setation sparse, semi-decumbent. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect, strongly gibbose; length 0.8x width, inserted between middle of eyes; pentagonal dorsally, narrowly and deeply sulcate dorsomedially, nearly bigibbous; ventrally pentagonal, concave, widest near base, dorsolateral margins straight. Antennal sockets subquadrate, separated by O.lx width of epistomai margin and 2x distance between eye and antennal socket. Antennifer insertion elevated medially, appearing basomedial. Interocular distance 2x width eye dorsally. Vertex densely punctate, transversely convex, almost flat; depression of tentorial maculae indistinct. Tentorial maculae indistinct. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width. Epistomai margin broadly truncate, width nearly 0.7x frontal distance between eyes. Labrum transverse, length 0.5x width, apical margin truncate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; palpomere 2 barrel-shaped, length 2x antennomere 1, diameter 2x length; diameter palpomere 3 equal 2; length palpomere 4 nearly equal 2, diameter at base less than Q.5x palpomere 3, inserted apically, acuminate. Labial palps absent. Gula elongate, subrectangular; width slightly greater basally than apically; length 2x width apical margin, inserted anteriorad posterior margin postgena.

Antennae filiform, 0.6x body length, setation dense, uniformly distributed, serai-decumbent on all antennomeres, scaliform setae present; scape obconic, width apically 0.8x length; pedicel cylindrical, short, transverse, not compressed, length 0.2x length of antennomere 3; antennomere 3 not adnate to base of antennomere 4, length equal length antennomere 4; antennomeres 3-10 conical, subserrate, expanded apically, barely compressed; antennomere 11 fusiform, weakly compressed, length 99

1,3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly 0.6x basal width; median longitudinal carina obscure, median fovea slightly more distinct posterior 0.6x pronotal length, closed before posterior margin; pronotal stemmata absent; anterior margin swollen; anterior margin straight, anterior angles acute; lateral margin complete, weakly carinate, slightly concavely arcuate longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk convex, finely rugose, discal setae shorter than elytral setae, semi-decumbent, posterior disk elevated near margin; hypomeron shallowly concave, height 0.5x length, all margins not swollen, hypomeral stemmata absent; prosternal bridge T—shaped, barely emarginate, length 0.5x width, intercoxal process carinate and short. Scuteilum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex slightly emarginate, emargination nearly O.lx scutellar length, apices not divergent or acute. Mesospiracle prominent, peritreme exposed, orifice compressed. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.3x and height 0.5x mesepisternum; < posterior margin very little reflexed, straight, acutely angulate at base. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separation 0,7x mesocoxal diameter at base.

Elytra ligulate, slightly dehiscent from scuteilum, narrowing uniformly distally, apices narrowly rounded; length 7x pronotal length; bicostate, costa 4 distinct basal 0.3x elytral length, humeral costa distinct over umbone and 0.8x elytral length; disk strongly and irregularly punctate- 100

reticulate basally, scabrous punctate near apex; setation semi-

decumbent, sparse, uniformly distributed. Metepisternum with ventral

margin increasingly arcuate posteriorly. Metepimeron with ventral

margin arcuate. Metasternum divided completely by median longitudinal

sulcus. Legs elongate, shining, setation sparse, semi-decumbent,

distributed along margins; coxae conical, pro- and mesotrochanterai

sockets 0.3x coxal length, metacoxa more vertically oriented and

directed dorsoanteriorly; trochanters short, length pro- and

mesotrochanters O.Sx coxal length, metatrochanters 0.5x metacoxal

length, all femoral insertions oblique and offset; femora weakly

compressed, fusiform, 3x trochanteral length, annulated basally; tibiae weakly compressed, internal margin slightly sinuate, tibial spines small and similar; tarsi elongate, tarsoraeres without laterally expanded plantar pads; covered with semi-erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-6

subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece large, tubular, length 1.3x median lobe

(Fig, 8 a-c); median lobe tubular, unmodified, ventraliy membranous near apex; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown. 101

DESCRIPTION (Immatures ). Unknown.

MATERIAL EXAMINED. CUBA, Oriente Province, Loma (Pico) del Gato, Sierra

Maestra, 26-28 May 1959 , M. W. Sanderson (1-INHS, holotype)

ETYMOLOGY. The specific epithet refers to the superficial resemblance to the European Omalvsi s fontisbellaguei Foucroy. 102

Abrolvcus parda (de Zayas >

Leptolvcus parda de Zayas, 1988:47, fig. 38.

Abrolvcus parda (de Zayas). NEW COMBINATION.

TYPE. A male with the following locality data: Cuba, Provincia

Oriente, Loma del Gato, Julio 1959 [Not Seen],

DIAGNOSIS. From the original description, this species differs from all other Abrolvcus by its uniformly dark grey coloration without maculation and its three elytral costae.

DISTRIBUTION. Known only from the type locality at Loma del Gato,

Oriente, Cuba.

DESCRIPTION (Male): Small, slender, elongate, length 2.0 mm., width at humerus unknown. Color dark grey, except elytral margins translucent.

Head. Elongate. Genal margin elongate, length greater than eye length. Eyes hemispherical, projecting anteriolaterally when viewed dorsally. Antennal prominence distinct from vertex, gibbose.

Antennae 0.7x body length; antennomeres 3-10 conical, subserrate, expanded apically, barely compressed.

Thorax. Pronotum trapezoidal, transverse, nearly rectangular, length less than O.Sx basal width. Scuteilum pentagonal, distally quadrate, nearly square, lateral margins parallel. Elytra ligulate, not dehiscent, apices broadly rounded; length lOx pronotal length; tricostate, costa 4 distinct at least basal 0.7x elytral length, costa 6 distinct at least basal 0.8x humeral costa distinct at least basal 0.8x 103 elytral length; reticulation regularly arranged.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immature Stages). Unknown.

DISCUSSION. This species was described by de Zayas sometime after

1959, but the description was not published until after his death. He tentatively placed this species in Leptolvcus L. & M. , although he questioned that placement. I have not yet seen the type or any specimens referable to it, but I am confident in its placement here based on the figure and description. Many characters used here are not known at this time and, thus, the description is incomplete. The presence of three distinct elytral costae is anomalous in the Abrolvcus

I have examined, but not entirely unexpected. I have modified the generic description to accomodate that state. However, the antennal states of 11 (left) and 12 (right) antennomeres (de Zayas, fig. 38) is undoubtedly rendered incorrectly, as there should only be 11 and, perhaps, only 10 should be visible at the scale drawn, because of the small size of the pedicel.

MATERIAL EXAMINED. None seen.

ETYMOLOGY. The specific epithet refers to the ground color of the species - a Spanish adjective for dark grey. 104

Abrolvcus sandersoni NEW OPECIES

Fig. 9 a-e

TYPE. INHS, male in good condition with the following label data:

Cwhite printed label! "CUBA: Gran Piedra \\ nr. Santiago, Oriente \\

Prov. 0-31 May 1959 \\ M. W. Sanderson."

DIAGNOSIS. Unlike all other Abrolycus seen, the median pronotal fovea is distinct, although still not bounded by definable carinae. Besides the couplets in the key, the bicostate elytra are dehiscent from the scuteilum, the scuteilum is medially impressed, and, then, more strongly dehiscent beyond the elytral middle.

DISTRIBUTION. Known only from Cuba.

DESCRIPTION (Male): Small, slender, elongate, length 1.9 mm., width 0.5 mm at humerus. Brown; except pronotum yellow; ultimate antennomere yellowish white.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, strongly and deeply punctate; setation sparse, semi-decumbent.

Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3x radius when dorsally viewed.

Antennal prominence distinct from vertex, porrect, strongly gibbose; length 0.8x width, inserted between middle of eyes; pentagonal dorsally, broadly sulcate dorsomedially, sulcus visible only posteriorly; ventrally pentagonal, convex, widest at base, dorsolateral margins straight. Antennal sockets triangularly oval, separated by O.lx width of epistomai margin and 0.5x distance between eye and antennal socket.

Antennifer insertion elevated medially, appearing basomedial.

Interocular distance 2x width eye dorsally. Vertex uniformly punctate, transversely convex; depression of tentorial maculae shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width.

Epistomai margin broadly emarginate, almost straight, width 0.5x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly rounded. Maxillary palps 4 segmented, palpomeres 1 and 3 cylindrical, diameter nearly equal to length; palpomere 2 barrel-shaped, length 2x antennomre 1, diameter 2x length; length palpomere 4 nearly equal 2, diameter at base less than 0.5x palpomere 3, inserted apically, acuminate. Labial palps absent. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted anteriorad posterior margin postgena. Antennae filiform, length 0.8x body length, setation moderately dense, uniformly distributed, semi-decumbent on all antennomeres, seal iform setae present; scape obconic, width apically

0.8x length; pedicel cylindrical, short, transverse, not compressed, length 0.3x length of antennomere 3; antennomere 3 not adnate to base of antennomere 4, length equal antennomere 4; antennomeres 3-10 conical, 106

subserrate, expanded apically, barely compressed; antennomere 11

fusiform, not compressed, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length nearly 0.5x basal width, anterior width 2x length and 0.4x basal width; median

longitudinal carina obscure, median fovea slightly more distinct; pronotal stemmata absent; anterior and lateral discai margins swollen, posterior margin not swollen; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, erect, straight

longitudinally; posterior margin weakly bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk convex, finely rugose, anterior corners deeply impressed, discai setae shorter than elytral setae, semi-decumbent; hypomeron deeply concave anteriorly, all margins not swollen, height 0.5x length, hypomeral stemmata absent; prosternal bridge T-shaped, slightly incised medially, length 0„5x width, intercoxal process carinate. Scutellum pentagonal, distally quadrate and nearly square, medially impressed, lateral margins slightly converging apically, apex emarginate, emargination nearly 0.2x scutellar length, apices not acute or divergent. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, 0.5x width and less than 0.7x height mesepisternum; posterior margin very little reflexed, nearly straight, acutely angulate ventrally. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, slightly dehiscent from scutellum, more strongly so from elytral middle, narrowing distally, apices broadly rounded; length lOx pronotal length; bicostate, costa 4 distinct from base to 0.5x elytral length, humeral costa distinct over umbone and 0.5x elytral length; disk irregularly and strongly punctate-reticulate; setation decumbent, sparse, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin arcuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket

0.3x coxal length, metacoxae more vertically oriented and directed dorsoanteriorly; trochanters elongate, length pro- and mesotrochanters subequal to coxal length, metatrochanters 0.5x metacoxal length, all femoral insertions oblique and offset; femora strongly compressed, fusiform, 2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin sinuate; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; sternite 7 deeply emarginate.

Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length subequal length of median lobe (Fig. 9 a-c); median lobe tubular, unmodified, ventrally membranous 0.7x length; internal sac not eversible; flagellum absent. 108

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. CUBA: Oriente Province, Gran Piedra, near

Santiago, May 1959, M. W. Sanderson (1-INHS, holotype).

ETYMOLOGY. Named for the collector of the type and over half the known specimens of this genus, Milton W. Sanderson. 109

Anti11olveus NEW GENUS

TYPE. Anti 1lolvcus elongatus NEW SPECIES, Basel.

DIAGNOSIS. This genus can be separated from other leptolycine genera by the combination of: a triangular prosternal bridge; the antenna is flabel late beginning with the fourth antennomere with their rami inserted apically; the pedicel is adnate to antennomere 3; the prothorax has 2 pair of pronotal stemmata on the pronotal disk and the 2 pair on the hypomeron; and the terminal sternite is apically cleft.

Additionally, the subantennal sulcus meets the eye instead of the epistomal margin; the elytra is ligulate, dehiscent, and 2 costate; and the antennae have modified scale-like setae on their antennomeres.

DISTRIBUTION. This genus is known from both Hispaniola and Cuba.

Given this distribution and the paucity of material available, it is not unlikely that it will be found also on Puerto Rico.

DESCRIPTION (Male). Small, slender, elongate, length 2.Q-4.9 mm., width 0.7-1.1 mm. at humerus. Black to brownish black; except yellow maculation.

Head. Elongate, widest at or behind eyes, then narrowing to posterior margin, punctate; setae sparse, short, semi-decumbent. Genal margin elongate, length greater than eye length, convexly arcuate to parallel when viewed dorsally. Eyes subhemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate; radial origin at: or inside head margin 0.3x radius when dorsally viewed.

Antennal prominence distinct from vertex, porrect and gibbose; length

0.7-lx width, inserted between middle to posterior margin of eyes; pentagonal dorsally, slightly wider or narrower distally, broadly and deeply to indistinctly sulcate dorsomedially; ventrally trianguliform,

strongly convex, sides nearly straight to uniformly and feebly convexly arcuate, widest near base. Antennal sockets oval to subrectangular, separated by 0.1-0.5x width of epistomal margin and 0.3-lx distance between eye and antennal socket, Antennifer insertion elevated medially, appearing basomedial. Interocular distance 2-4x dorsal eye width. Vertex punctate, transversely flat, setae semi-decumbent; depression of tentorial maculae broadly shallow, from base or posterior

0.3x of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae shallow to deep, distinct to indistinct, separate or not; at base of antennal prominence. Frons transversely convex, slightly distinct from lower surface of antennal prominence, height 0.5-*1.5x width. Epistomal margin truncate to emarginate, width

0.4-0.8x frontal distance between eyes. Labrum transverse to elongate,

length 0.5x width, apical margin truncate to broadly rounded.

Maxillary palps 3-4 segmented, palpomeres 1~2 or 3 cylindrical, diameter nearly equal to length; length apical palpomere nearly equal penultimate, diameter at base less than 0.5x penultimate palpomere, acuminate. Labial palps 1 segmented and digitiform or absent. Gula transverse to elongate, subrectangular to trapezoidal; width slightly greater apically than basally; length 0.5-2x width apical margin;

inserted anteriorad of postgenal posterior margin. Antennae Ill flabellate, length 0.7-lx body length, rami between 1-3.5x axial length and their compression perpendicular to axial compression; setation sparse to dense, decumbent to semi-decumbent, uniformly distributed, scaliform setae present or absent; scape obconic, width apically 0.6x-

0.8x length; pedicel cylindrical, short, transverse, not compressed, length 0.5x-0.7x length of antennomere 3, adnate to antennomere 3; antennomere 3 slightly compressed and filiform, not flabel late, not adnate to base of antennomere 4, length 0,l-0.5x antennomere 4; antennomeres 4-10 elongate, strongly compressed; extent of compression uniform to apex, flabel late with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, transverse, length 0.6x basal width, anterior width nearly 0.6-0.7x basal width; median longitudinal carina obscure, median fovea more distinct, Q.5-0,7x pronotal length and not reaching posterior margin; pronotal stemmata present, anterior pair on or behind anterior margin 1 stemmatal diameter and 1-2 stemmatal diameter from lateral margin, posterior pair in front of posterior margin 1.5-3 stemmatal diameters and 2-3 stemmatal diameters from lateral margin; discai margins variously swollen; anterior margin straight and slightly reflexed; anterior angles obtuse; lateral margin complete to incomplete anteriorly and posteriorly, foliaceous or not, reflexed or not, slightly concavely arcuate; posterior margin bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk flat to sLightly convex, barely elevated at posterior margin, setation decumbent; hypomeron concave, height 0.4-0.7x length; margins variously swollen, hypomeral stemmata 1 stemmatal diameter below dorsal margin, anterior pair behind anterior margin 3-9 stemmatal diameters and separated from posterior pair by 1-3 stemmatal diameters, posterior pair

5-8 stemmatal diameters before posterior margin; prosternal bridge trianguliform, length 0.5-lx width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, nearly square, disk longitudinally impressed or not, lateral margins parallel, apex rounded to truncate and entire to emarginate, emargination to 0.1-0.3x scutellar length. Mesospiracle prominent, peritreme exposed, orifice not to slightly compressed. Mesepisternum elongate, posterior margin straight to convexly arcuate. Mesepimeron trianguliform to subrectangular to lobate, width 0.5x and height 0.5-0.8x mesepisterum; posterior margin not refiexed, nearly straight to weakly arcuate, angulate at ventral

0.3x height or ventrally. Mesosternum transverse to elongate, posterior margin truncate to convexly arcuate. Mesocoxal separation

0.8-lx mesocoxal diameter at base. Elytra ligulate, dehiscent from basal 0.5x elytral length, uniformly narrowing distally, slightly or not reconvergent apically, apices narrowly rounded to acuminate; length 7- lOx pronotal length; bicostate, costa 4 distinct basal 0.3-0.6x elytral length, costa 8 distinct 0.7x-0.8x elytral length; disk irregularly scabrous-punctate to irregularly reticulate; setation decumbent to erect, sparse to dense, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin sinuate to arcuate. Metasternum completely to incompletely divided medially by longitudinal sulcus and sulcus not reaching mesosternum by lx mesocoxal diameter. Legs elongate, setation sparse, decumbent to semi-decumbent, uniformly distributed; coxae 113 conical, trochanteral socket 0.2-0.5x coxal length, metacoxae strongly

elevated and more obliquely oriented; trochanters elongate, length pro- and mesotrochanters lx coxal length, metatrochanters 0.8-1.lx metacoxal

length, femoral insertions truncate and not offset; femora strongly compressed, straight to clavate, short, 2-4x trochanteral lengch, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tibial spines small and similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; tarsomere 4 extended by membrane beyond insertion of tarsomere 5 or not; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped and apically cleft. Sternites rectangular, wider than long, length 1-Q subequal, but width narrowing uniformly posteriorad; sternite 7 entire.

Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with incompletely differentiated or without parameres; basal piece large, dorsoventrally compressed, length 0.5-3x length of median lobe; median lobe tubular, unmodified, ventrally membranous; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. Chevrolat (1870:78) described Calopteron semiflavum from 114

Cuba. I have been unable to see the type in National Collection of

Cuba (Gundlach and Poey collections), but due to the efforts of Michael

A. Ivie, I have seen two specimens identified as that species from the

National collection of Cuba. The description does not completely agree with the specimens, because Chevrolat did not mention the presence of flabel late antennae - only that they were compressed, long, and wide.

Consequently, this may be an incorrect attribution and may have to be changed later. However, I have placed it here because material was at hand, I did not want to create another name and possible synonym, and did not want to leave Chevrolat's species incerata sedis. His description indicates that it is undoubtedly a leptolycine.

Although the membranous extensions of tarsomeres 4 extending apically below tarsomere 5 are slightly expanded and might be interpreted as plantar pads, their structure differs significantly from that of other lycids, including the state found in the outgroup. It is therefore, hypothesized to be non-homologous and independently derived within Anti 1lolvcus.

ETYMOLOGY. The prefix refers to the geographical location of all known species within this genus and the Greek root means wolf and is the type genus of the family. 115

KEY TO SPECIES OF Antillolvcus (ADULT MALES)

1. At least basal 0.3x elytral length golden yellow; legs

entirely black ...... 2

1'. Basal 0.3x elytral length at most only yellow in part; legs

with yellow annulations ...... 3

2. (1) Scutellum broadly emarginate, Hispaniola ......

...... elongatus n. sp. (p. 121)

2'. (1) Scutellum truncate, Cuba ... semiflavus (Chevrolat) (p. 132)

3. (I1) Humeral angles yellow, elytral suture black at base; apical

antennomeres yellow; Hispaniola ......

...... auratosuratus n . sp. (p . 116)

3. (I1) Elytra irregular yellow, giving a mottled appearance,

elytral suture blackish brown; all antennomeres black;

Hispaniola ...... flavimarginatus n. sp. (p. 127) 116

Anti 1lolvcus auratesuratus NEW SPECIES

Fig. 10 a-f

TYPE: MHMB, a male in good condition with the following label data:

[white printed label 1 "Cazabita 31.10 \\ 1250 m. 1971;" [white printed

label! "Rep. Dominic. \\ J. & S. Klapperich."

DIAGNOSIS. This species is separable from congenerics by the tibiae

being golden yellow basally, the yellow apical antennomeres, and the

genitalia (Fig. 10).

DISTRIBUTION. Known only from the Dominican Republic on Hispaniola.

DESCRIPTION (Males). Small, slender, elongate, length 3.4-4.3 mm.,

width 0.7-0.9 mm. at humerus. Black; except basal 0.3x elytra along

margins, apices of last 3 antennomeres in part, bases of trochanters,

femora, and tarsi golden yellow.

Head. Elongate, widest behind eyes, then narrowing to posterior

margin, punctate; setae sparse, short, serai-decumbent. Genal margin elongate, length greater than 2x eye length, convexly arcuate when

viewed dorsally. Eyes subhemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate; radial origin

inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect and gibbose; length 0.7x width,

inserted between middle of eyes; pentagonal, slightly wider distally, broadly and deeply sulcate dorsomedially, posterior sulcus distinct 0.5x length; ventrally trianguliform, strongly convex, sides nearly straight, widest near base. Antennal sockets oval, separated by 0.2x width of epistomal margin and 0.5x distance between eye and antennal socket.

Antennifer insertion elevated medially, appearing basomedial.

Interocular distance 2x dorsal eye width. Vertex punctate, transversely flat, setae semi-decumbent; depression of tentorial maculae broadly shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae deep, distinct, separate; at base of antennal prominence. Frons transversely convex, slightly distinct from lower surface of antennal prominence, height l.Sx width. Epistomal margin broadly truncate, width 0.5x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly rounded. Maxillary palps 3 segmented, palpomeres cylindrical, diameter nearly equal to length; length palpomere 3 nearly equal 2, diameter at base less than 0.5x palpomere 2, acuminate.

Labial palps absent. Gula transverse, subrectangular; width slightly greater apically than basally; length 0.5x width apical margin; inserted anteriorad of postgenal posterior margin. Antennae flabel late, length

0.7x body length, rami between 1.6-3x axial length and their compression perpendicular to axial compression; setation sparse, semi-decumbent, uniformly distributed, scaliform setae present; scape obconic, width apically 0.6x length; pedicel cylindrical, short, transverse, not compressed, length 0.5x length of antennomere 3; antennomere 3 slightly compressed and filiform, not flabellate, not adnate to base of antennomere 4, length 0.4x antennomere 4; antennomeres 4-10 elongate, 118

strongly compressed; extent of compression uniform to apex, flabel late with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, transverse, length 0.6x basal width, anterior width nearly 0.6x basal width; median longitudinal carina obscure, median fovea more distinct, 0.7x pronotal length; pronotal stemmata present, anterior pair behind anterior margin 1

stemmatal diameter and 1 stemmatal diameter from lateral margin, posterior pair in front of posterior margin 3 stemmatal diameters and 2 stemmatal diameters from lateral margin; anterior and lateral discai margins swollen, posterior margin not swollen; anterior margin straight and slightly reflexed; anterior angles obtuse; lateral margin incomplete anteriorly and posteriorly, not foliaceous, not reflexed, slightly concavely arcuate; posterior margin bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk flat, setation decumbent; hypomeron concave, height 0.4x length; all margins not swollen, hypomeral stemmata present 1 stemmatal diameter below dorsal margin, anterior pair behind anterior margin 3 stemmatal diameters and separated from posterior pair by 3 stemmatal diameters, posterior pair 5 stemmatal diameters before posterior margin; prosternal bridge trianguliform, length 0.5x width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex incised to slightly emarginate, emargination to 0.lx scutellar length. Mesospiracle prominent, peritreme exposed, orifice not compressed. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.5x and height 0.7x mesepisterum; posterior margin not reflexed, nearly straight, angulate at ventral 0.3x height. Mesosternum transverse, posterior margin truncate. Mesocoxal separation subequal to coxal diameter at base. Elytra ligulate, dehiscent from basal 0.5x elytral length, uniformly narrowing distally, slightly reconvergent apically, apices narrowly rounded; length 8x pronotal length; bicostate, costa 4 distinct basal 0,5x elytral length, costa 8 distinct Q.8x elytral length; disk irregularly scabrous- punctate; setation decumbent, moderately dense, uniformly distributed.

Metepisternum with ventral margin increasingly arcuate posteriorly.

Metepimeron with ventral margin sinuate. Metasternum incompletely divided medially by longitudinal sulcus, sulcus lx mesocoxal diameter from mesosternum. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5x coxal length, raetacoxae strongly elevated and more obliquely oriented; trochanters elongate, length pro- and mesotrochanters equal to coxal length, metatrochanters equal to metacoxal length, femoral insertions truncate and not offset; femora strongly compressed, clavate, short, 2.5x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tibial spines short and similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae, tarsomere 4 not extended by membrane beyond insertion of tarsomere 5; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped and apically cleft. Sternites rectangular, wider than long, length 1-8 120

subequal, but width narrowing uniformly posteriorad; sternite 7 entire.

Spiracles in pleural membrane, located submedially from apical margin of

tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length equal length of median lobe (Fig. 10 a-c); median

lobe tubular, unmodified, ventral.ly membranous; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures ). Unknown.

MATERIAL EXAMINED. DOMINICAN REPUBLIC, Cazabita, 1250 m., 24 September and 31 October 1971, J, & S. Klapperich (2—MHMB, including holotype).

ETYMOLOGY. The Latin feminine prefix of the specific epithet, aurate, and the Latin root, suratus. refers to the golden yellow bases of the tibiae, which is more prominent, although not unique, in the known

Anti 11o 1vcus. 121

Anti 1lolvcus elongatus NEW SPECIES

Fig. 11 a-d

TYPE. MHMB, a male with the following label data: [white printed

label!) "Colonia, 1000 m. , 18, III, 1972;" Cwhite printed label!) "Rep.

Dominic. \\ J. & S. Klapperich."

DIAGNOSIS. This species can be separated from congenerics by the complete, basal yellow fascia of the elytra, the scutellum broadly emarginate, and the genitalia (Fig. 11 a-c). A. elongatus and A. semiflavus look very similar, but can most easily be separated without dissection examining the ratio of the pronotal to the elytra lengths and

the relative width at the elytral humerus to overall body si2e. A. elongatus is more elongate and less robust.

DISTRIBUTION. Known only from Hispaniola.

DESCRIPTION (Males): Small, slender, elongate, length 2.8-4.9 mm., width

0.8-1.1 mm at humerus. Color black, except basal 0.3-0.4x elytra and apices antennal rami 9-11 golden yellow, scutellum varies from all, partially, to not yellow.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, shining punctate; setae sparse, directed anteriorly, seroi- decumbant. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes subhemispherica1, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3x

radius when dorsally viewed. Antennal prominence distinct from vertex,

gibbose; length 0.8x width, inserted between posterior margin of eyes;

pentagonal dorsally, broadly sulcate dorsomedially, sulcus more narrowly distinct posteriorly; ventrally trianguliform, convex, sides uniformly

and convexly arcuate, widest near base. Antennal sockets

subrectangular, separated by 0.5x width of epistomal margin and 0.5x distance between eye and antennal socket. Antennifer insertion elevated medially, appearing basomedial. Interocular distance 4x width eye dorsally. Vertex shallowly punctate, transversely flat, setae

semi-decumbent; depression of tentorial maculae well defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae not separate, shallow, indistinct; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width. Epistomal margin broadly emarginate, almost straight, width

0.4x frontal distance between eyes. Labrum transverse, rectangular, width 0.5x length, apical margin broadly and distinctly emarginate.

Maxillary palps 3 segmented, palpomere 1 cylindrical, diameter nearly equal to length; length palpomere 2 nearly equal 3, diameter at base

less than 0.5x palpomere 2, acuminate. Labial palps 1 segmented and digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted anteriorad of posterior margin of postgena. Antennae flabellate, length lx body length; rami between l,4-3.5x axial length and their compression perpendicular to axial compression; setation sparse, 123 decumbent on all antennomeres, uniformly distributed, scaliform setae present; scape obconic, width apically O.Qx length; pedicel cylindrical, short, transverse, not compressed, length 0.5x length of antennomere 3; length antennomere 3 nearly 0.5x antennomere 4; antennomeres 4-10 elongate, compressed, flabel late with ramal insertions apical; extent of compression uniform to apex; antennomere 11 without axis indistinguishable from ramus.

Thorax. Pronotum trapezoidal, transverse, length 0.6x basal width, anterior width 0.6x basal width; median longitudinal carina obscurely present, median fovea broadly distinct, not reaching posterior margin; pronotal stemmata present, anterior pair separated from anterior margin by 1 stemmatal diameter and separated from lateral margin by 2 stemmatal diameters, posterior pair separated from lateral margin by 2 stemmatal diameters, from posterior margin by 2 stemmatal diameters; anterior discai margin swollen; anterior margin straight, slightly reflexed; anterior angles obtuse; lateral margin incomplete anteriorly, not foliaceous, not reflexed, slightly concavely arcuate; posterior margin weakly bisinuate; posterior angles acute; disk strongly convexly arcuate, strongly elevated, setation shorter than elytral setae, decumbent; hypomeron concave, height 0.5x length; ventral and lateral margins swollen, hypomeral stemmata hypomeral stemmata 1 stemmatal diameter below dorsal margin, anterior pair behind anterior margin 9 stemmatal diameters and separated from posterior pair by 3 stemmatal diameters, posterior pair 8 stemmatal diameters before posterior margin; prosternal bridge trianguliform, length 0.5x width, intercoxal process carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly

0,3x scutellar length, apices truncate, not divaricate. Mesospiracle prominent, peritreme exposed, oriface slightly compressed.

Mesepisternum elongate, posterior margin convexly arcuate. Mesepimeron lobate, 0.5x width and 0.5x height mesepisternum; posterior margin very little reflexed, weakly arcuate. Mesosternum elongate, posterior margin convexly arcuate. Mesocoxal separation 0.8x mesocoxal diameter at base. Elytra ligulate, slightly dehiscent from anterior margin of elytral fascia, uniformly narrowing distally, not reconvergent apically, apices acuminate; length 9x pronotal length; bicostate, costa 4 distinct

0.5x basal elytral length, humeral costa distinct 0.7x elytral length; reticulation irregular at elytral middle and rugose apically and basally; setation semi-erect, sparse, uniformly distributed.

Metepisternum with ventral margin inreasingly arcuate posteriorly.

Metepimeron with ventral margin sinuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent, uniformly distributed; coxae conical, trochanteral socket Q.5x coxal length; trochanters elongate, length pro- and mesotrochanters lx coxal length, metatrochanters l.lx metacoxal length, femoral insertions truncate, not offset; femora strongly compressed, straight, short, 4x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly bisinuate; tibial spurs small and similar, nearly 0.3x length tarsomere 1; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered dorsally with erect bristling setae; tarsomere 4 extended by membrane beyond insertion 5; ungues simple. 125

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing uniformly posteriorad; tergite 0 entire; tergite

9 medially divided longitudinally; tergite 10 shield shaped and apically cleft. Sternites 1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedialiy from apical margin of tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length almost 3x length of median lobe (Fig.

11 a-c); median lobe tubular, unmodified, internal sac not eversible, flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The extent of the basal yellow fascia varies binomially in size for the small sample available (n=16).

MATERIAL EXAMINED. DOMINICAN REPUBLIC: Provincia Trujillo. Coionia

Ranfis, 26 December 1955, J. Maldanado-Capriles (1-NMNH). The following have same data, except the dates differ as follows: Coionia,

1000 m., 30 January 1972, J. & S. Klapperich (l-MHMB). 26 November 1970

(1-MHMB ). 12 February 1972 (l-MHMB). 18 March 1972 (2-MHMB ). 21

April 1972 (l-MHMB). 19 May 1972 (l-MHMB). La Vega, 24 km. SE.

Constanza, 4 August 1979, C. W. O'Brien (1-RSMC). La Vega, 18 km. E.

El Rio, 11 August 1979, crest in cloud forest, L. B. O'Brien (1-RSMC).

La Vega, 20 km. SE Constanza, 26 May 1978, C. W. and L. B. O'Brien (1- 126

RSMC). El Siebo, 93 km. N. Pedro Sanchez, 2 August 1979, C. W. O ’Brien

(1-RSMC). Valle Nuevo, SE. Constanza, August 1938, ca. 2100m,

Darlington (1-MCZC). Loma Rucilla & mts. N . , June 1938, 1,500-2,300 m,

Darlington (1-MCZC). Provincia . Cazabita, 1250 m . , 26 June 1974, J. £

S. Klapperich (l-MHMB).

ETYMOLOGY. The specific epithet refers to the elongate body form. 127

Anti 1lolvcus flavimarginatus NEW SPECIES

Fig. 12 a-d

TYPE. MCZC, a male in good condition with the following label data:

Cwhite printed label] "fthills Cord. Cent. \\ S. of Santiago W June

’38 Dom. Rep. \\ Darlington."

DIAGNOSIS. This species can be easily recognized by its mottled yellow and brown elytral coloration with the yellow coloration primarily along the elytral margins. The irregular border between the ground color and the lateral yellow maculation of the elytra is unique within the

Leptolycines as they are currently understood. Also, the yellow basal annulations of all the leg segments, the median impressed scutellum, and the ramal length nearly equal to the length of the axis of the following antennomere will separate this species from other Leptolycini.

DISTRIBUTION. Known from Hispaniola only from the type specimen which was collected in 1938 by Phillip Darlington. Presumably, the indefinite locality indicated is on the north-facing slope of the island, which is more mesic and part of the Northern Hispaniolan Plate.

DESCRIPTION (Male). Small, slender, elongate, length 4.2 mm., width

1.0 mm at humerus. Brownish black; except bases of all leg segments, lateral margin of pronotum and lateral elytral margins excluding apex pale yellow.

Head. Elongate, widest at eyes, then narrowing to posterior margin; shallowly punctate; setae dense, short, semi“decumbent. Genal

margin elongate, length greater than eye length, parallel when viewed

dorsally. Eyes subhemispherica1, projecting anteriolaterally when

viewed dorsally; hind margin shallowly emarginate, nearly straight;

radial origin inside head margin 0.3x radius when dorsally viewed.

Antennal prominence distinct from vertex, gibbose; length 0.8x width,

inserted between middle of eyes; pentagonal dorsally, narrowly sulcate

dorsomedially, sulcus indistinct posteriorly; ventrally trianguliform,

strongly convex, sides uniformly and feebly convex-arcuate, widest near base. Antennal sockets oval, separated by 0.3x width of epistomal margin and lx distance between eye and antennal socket. Antennifer

insertion elevated medially, appearing basomedial. Interocular distance 2x dorsal eye width. Vertex transversely arcuate, densely pubescent, setae semi-decumbent; depression of tentorial maculae shallow, from posterior 0.3x of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae deep, distinct, separate; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width. Epistomal margin nearly truncate, slightly emarginate, width 0.8x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin trucate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter lx

length; length palpomere 4 elongate, 2x antennomere 3, diameter at base

0.5x palpomere 3, digitiform, apex acuminate. Labial palps 1 segmented, digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted 129 anteriorad posterior margin of postgena. Antennae flabel late, length

0.7x body length, rami between 1-1.3x axial length and their compression perpendicular to axial compression; setation dense, decumbent, uniformly distributed, scaliform setae absent; scape obconic, width apically 0.7x

length; pedicel cylindrical, short, transverse, not compressed, length

0.5x length of antennomere 3; antennomere 3 slightly compressed and filiform, not flabel late, not adnate to base of antennomere 4, length

0.5x antennomere 4; antennomeres 4-10 elongate, strongly compressed, extent of compression uniform to apex, flabel late with raroal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, transverse, length equal anterior width, anterior width nearly 0.7x basal width; median longitudinal carina obscure, median fovea a linear impression distinct 0.5x pronotal length and not reaching posterior pronotal margin; pronotal stemmata present, anterior pair adjacent to anterior margin and separated from lateral margin by 2 stemmatal diameters, posterior pair separated from lateral margin by 3 stemmatal diameters and from posterior margin by 2 stemmatal diameters; all discai margins swollen, posterior margin weakly so; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, reflexed, slightly concavely arcuate; posterior margin weakly bisinuate; posterior angles acute; disk slightly convex, setation decumbent, barely elevated at posterior margin; hypomeron concave, height 0.5x width, lateral and ventral margins not swollen, hypomeral stemmata present 1 stemmatal diameter below dorsal margin, anterior pair behind anterior margin 7 stemmatal diameters and separated from posterior pair by 3 stemmatal diameters, posterior pair 6 stemmatal diameters before posterior margin; prosternal bridge trianguliform, length lx width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, nearly square; disk with deep median longitudinal sulcus; lateral margins parallel; apex convexly arcuate and shallowly emarginate, emargination nearly 0.lx scutellar length.

Mesospiracle prominent, peritreme exposed, orifice slightly compressed.

Mesepisternum elongate, posterior margin convexly arcuate. Mesepimeron subrectangular, width 0.5x and height 0.7x mesepisternum; posterior margin very little reflexed, nearly straight, obtusely angulate ventrally. Mesosternum elongate, posterior margin convexly arcuate.

Mesocoxal separation 0.8x mesocoxal diameter at base. Elytra ligulate, slightly dehiscent at elytral middle; uniformly narrowing distally, not reconvergent apically, apices accuminate; length lOx pronotal length; bicostate, costa 4 distinct basal 0.3x elytral length, humeral costa distinct basal 0.7x elytral length; disk irregularly reticulate, obscured by setation; setation erect, uniformly dense, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin sinuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5x coxal length, metacoxae strongly elevated and more obliquely oriented; trochanters elongate, length lx coxal length, all femoral insertions truncate and not offset; femora strongly compressed, straight, short, 2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly bisinuate; tibial spines short and similar; tarsi elongate, tarsomeres without laterally 131 expanded planar pads; covered with erect bristling setae; tarsomere 4 extended by membrane beyond insertion of tarsomere 5; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped, apically cleft. Sternites 1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without completely differentiated parameres; basal piece, dorsoventrally compressed, length almost 0.5x length of median lobe (Fig. 12 a-c); median lobe tubular, membranous ventrally, internal sac not eversible; flagellum absent.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immafcures). Unknown.

MATERIAL EXAMINED. DOMINICAN REPUBLIC: Prov., S. of Santiago, June

1938, Darlington (1-MCZC, holotype).

ETYMOLOGY. Flava (Latin) meaning yellow and marsinatus (Latin) meaning margined refers to the coloration of the elytra. 132

Anti 1lolvcus semiflavus (Chevrolat)

Fig. 13 a-c

Calopteron semiflavum Chevrolat, 1870:78. Gemminger and Harold,

1869:1632. Gundlach, 1891:256. Leng and Mutchler, 1922:429.

Blackwelder, 1945:345.

TYPE: CUBA, a male. Not Seen.

DIAGNOSIS. This species is separable from congenerics by the combination of the yellow basal facia of the elytra, the entirely black legs, the truncate scutellum, and the distinctive genitalic configuration. A. elongatus and A. semiflavus look very similar, but can most easily be separated without dissection by examining the ratio of the pronotal to the elytra lengths and the relative width at the elytral humerus to overall body size. semiflavus is less elongate and more robust.

DISTRIBUTION. Known only from Oriente Province in Cuba.

DESCRIPTION (Males). Small, slender, elongate, length 3,2-4.4 mm., width 0.7-0.8 mm. at humerus. Black; except basal 0.3x elytra, apices of last 3 antennomeres in part golden yellow.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, strongly punctate; setae sparse, short, semi-decumbent. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes subhemispherical, projecting anteriolaterally when viewed dorsally; hind margin emarginate; radial origin at lateral head margin when dorsally viewed. Antennal prominence distinct from vertex, porrect and gibbose; length Ix width, inserted between middle of eyes; pentagonal, broadly and deeply sulcate dorsomedially, sulcus distinct entire length; ventrally trianguiiform, convex, sides nearly straight, widest near base. Antennal sockets lenticular, separated by

O.lx width of epistomal margin and 0.3x distance between eye and antennal socket. Antennifer insertion elevated medially, appearing basomedial. Interocular distance 2x dorsal eye width. Vertex strongly punctate, transversely flat, setae semi-decumbent; depression of tentorial maculae broadly shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae deeply distinct, separate; at base of antennal prominence. Frons transversely convex, little to not distinct from lower surface of antennal prominence, height 0.8x width. Epistomal margin broadly emarginate, width 0.8x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin emarginate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps absent. Gula transverse, trapezoidal; width slightly greater apically than basally; length 0.3x width apical margin; inserted anteriorad of postgenal posterior margin. Antennae flabel late, length lx body length, rami between l-3x axial length and their compression perpendicular to axial compression; setation sparse, decumbent, uniformly distributed, 134 scaliforni setae present; scape obconic, width apically 0.7x length; pedicel cylindrical, short, transverse, not compressed, length 0.7x length of antennomere 3; antennomere 3 compressed and filiform, not flabellate, not adnate to base of antennomere 4, length O.lx antennomere

4; antennomeres 4-10 elongate, strongly compressed; extent of compression uniform to apex, flabellate with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, transverse, length 0.6x basal width, anterior width 0.6x basal width; median longitudinal carina obscure, median fovea more distinct, O.Sx pronotal length; pronotal stemmata present, anterior pair behind anterior margin 1 stemmatal diameter and 1 stemmatal diameter from lateral margin, posterior pair in front of posterior margin 1.5 stemmatal diameters and 3 stemmatal diameters from lateral margin; anterior and lateral discal margins swollen, posterior margin not swollen; anterior margin straight and slightly reflexed; anterior angles obtuse; lateral margin incomplete anteriorly and posteriorly, not foliaceous, not reflexed, slightly concavely arcuate; posterior margin bisinuate; posterior angles acute, expanded laterally 0.7x width elytral humerus; disk flat, slightly elevated at posterior margin, setation dense, semi-erect; hypomeron shallowly concave, height 0.7x length; all margins not swollen, hypomeral stemmata present 1 stemmatal diameter below dorsal margin, anterior pair behind anterior margin 3 stemmatal diameter and separated from posterior pair by 1 stemmatal diameter, posterior pair 5 stemmatal diameters before posterior margin; prosternal bridge trianguliform, length 0.6x width, intercoxal process not carinate. Scutellum 135 pentagonal, distally quadrate, nearly square, lateral margins parallel, apex truncate. Mesospiracle prominent, peritreme exposed, orifice not compressed. Hesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.5x and height 0.8x mesepisterum; posterior margin not reflexed, nearly straight, angulate at ventral 0.3x height. Wesosternum transverse, posterior margin truncate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, dehiscent from scutellum, uniformly narrowing distally, slightly reconvergent apically, apices acuminate; length 7x pronotal length; bicostate, costa

4 distinct basal 0.6x elytral length, costa 8 distinct 0.7x elytral length; disk irregularly scabrous-punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin arcuate. Metasternum incompletely divided medially by longitudinal sulcus, sulcus lx mesocoxal diameter from mesosternum.

Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.2x coxal length, metacoxae strongly elevated and more obliquely oriented; trochanters elongate, length pro- and mesotrochanters lx coxal length, metatrochanters 0.8x metacoxal length, femoral insertions truncate and not offset; femora strongly compressed, clavate, short, 2.5x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tibial spines short and similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; tarsomere 4 extended by membrane beyond insertion of tarsomere 5; ungues simple. 136

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but width narrowing uniformly posteriad; tergite Q entire;

tergite 9 medially divided longitudinally; tergite 10 shield shaped and

apically cleft. Sternites rectangular, wider than long, length 1-8

subequal, but width narrowing uniformly posteriorad; sternite 7 entire.

Spiracles in pleural membrane, located submedially from apical margin of

tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length equal length of median lobe (Fig. 13 a-c); median

lobe tubular, unmodified, ventrally membranous; internal sac not eversible; flageilum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. CUBA, Prov. Oriente, La Gran Piedra, July 1970, P.

Alayo coir. (1-IESH). Cobre Range, Loma del Gato, 3-8 July 1936, P.

Darlington coir, (1-MCZC). Pico Turquino, south side 900-1200 m, June

1936, P. Darlington coir. (1-MCZC).

ETYMOLOGY. The Latin prefix of the specific epithet, semi. meaning half and the Latin root, flavum. meaning yellow refers to the basal 0.3x of elytron's golden yellow coloration. 137

Ceratoprion Gorham

Ceratoprion Gorham, 1884:348. Kleine, 1933:33. Blackwelder,

1945:348.

TYPE: Ceratoprion serricorne Gorham, BMNH, by original monotypy.

DIAGNOSIS. Ceratoprion can be distinguished from other leptolycines by combination of the serrate antennae longer than 0.5X length of the body, antennomere 4 more than 3X the combined length of antennomeres 2 and 3, antennomere 2 subequal to 3, antennomere 3 and 4 adnate, the apical width of antennomere 3 being equal to the basal width of antennomere 4, and the 3 - 4 costate elytra. Like Anti 11olvcus it has discal pronotal stemmata, but differs by the absence of hypomeral stemmata and the presence of well developed and often internally dentate parameres. The appearance of Ceratoprion is most similar to Pseudoceratoprion. but is easily differentiated by the antennal states. The pedicel to antennomere 3 ratio in the latter genus is much smaller, antennomere 3 is not adnate to the base of 4, and the length of antennomere 4 is sunequal to that of antennomere 3.

DISTRIBUTION. This genus appears to be restricted to the highlands of

Central America and the Andes south to Ecuador.

DESCRIPTION (Males). Slender, elongate; length 3.6 - 7.2 mm., widthat humerus 0.6 - 1.3 mm. Brown to black; except various yellow maculation. Head. Elongate, widest at eyes, then narrowing to posterior margin, shining to dull; setae sparse, decumbant to semi - decumbent.

Genal margin elongate, length greater to slightly less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly emarginate; dorsally radial origin at head margin to inside head margin

0.3X radius. Antennal prominence indistinct to slightly distinct from vertex, shelf ~ like and almost gibbose; length 0.4X - IX width, inserted between middle of eyes; pentagonal to nearly quadrate dorsally, anterior margin transversely arcuate to barely produced medially, broadly and obscurely sulcate dorsomedially, sulcus variously distinct; ventrally trainguliform to trapezoidal or nearly quadrate, shallowly to deeply concave, widest near base. Antennal sockets subrectangular, separated by 0.IX - 0.2X width of epistomal margin and 0.1X - 2X distance between eye and antennal socket. Antinnifer elevated medially, appearing basomedial. Interocular distance 2X width eye dorsally. Vertex reticulate, transversely flat to slightly arcuate, setae decumbent to serai - decumbent; depression of tentorial maculae shallow to deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, distinct to deeply distinct; at base of antennal prominence.

Frons transversely flat, distinct from lower surface of antennal prominence, transversely narrow, height 0.3X - 0.5X width. Epistomal margin entire and almost straight to broadly and deeply emarginate, width 0.7X - 0.8X frontal distance between eyes. Labrum transverse, rectangular, length 0.5X width, apical margin narrowly and distinctly emarginate to broadly rounded. Maxillary palps 3 or 4 segmented, basal

palpomeres cylindrical, diameter nearly equal to length; length ultimate

palpomereO.3X - IX penultimate, diameter at base Q.5X pneunultimate palpomere at base, acuminate. Labial palps absent to 2 segmented, digitiform. Gula transverse, pentagonal; width slightly greater apicaliy than basally; length approximately 0.5X - 2X width apical margin. Antennae filiform - serrate to serrate, 0.6X ~ 0.7X body

length, setation dense to sparse, uniformly distributed, decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically

0.7X - 0.8X length; pedicel cylindrical, short, transverse, not compressed, length subequal length of antennomere 3; antennomere 3 adnate with base antennomere 4; length antennomere 3 0.1X - 0.2X antennomere 4; antennomeres 4-10 filiform - serrate to serrate, expanded apically, strongly compressed; antennomere 11 fusiform, length

1.3X antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5X basal width, anterior width slightly more than 0.5X - 0.6X basal width; median longitudinal carina distinct to obscure, median fovea distinct, slightly more than 0.3X — 0.5X pronotal length, fovea closes at or before posterior margin; pronotal stemmata absent to present, anterior lateral pair when present on anterior margin by 1 - 2 stemmatal diameters and separated from lateral margins by 1 stemmatal diameter, anterior internal pair when present on or separated from anterior margin by 1 - 2 stemmatal diameters and separated from lateral margins by 1 - 8 stemmatal diameters, posterior lateral pair on or separated from posterior margin by 1 - 2 stemmatal diameters and separated from lateral margins by 2 stemmatal diameters, posterior internal pair on or

separated from posterior margin by 3 stemmatal diameters and separated

from lateral margins by 2 - 12 stemmatal diameters; all discal margins

curbed; anterior margin nearly straight, anterior angles obtuse; lateral margin complete, foliaceous, not to slightly reflexed, slightly

concavely arcuate to nearly straight; posterior margin bisinuate to

strongly bisinuate; posterior angles acute, expanded laterally 0.8X width elytral humerus; disk flat, discal setae shorter than elytral

setae, setae serai - decumbent; hypomeron shallowly concave, height 0.3X

- 0.6X length, all margins curbed, hypostomal stemmata absent; prosternal bridge Y - shaped, anterior margin broadly emarginate, length

0.5X width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly 0.3X scutellar length, apices truncate to acute and divaricate. Mesospiracle obscure, peritreme not exposed.

Mesepisternum elongate, posterior margin straight. Mesepimeron 0.3X -

IX width and O.SX - IX height mesepisternum width and height; posterior margin very little to strongly reflexed, nearly straight to convexly arcuate, angulate ventrally or not. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separation O.SX - 1.5X mesocoxal diameter at base. Elytra ligulate, slightly dehiscent from

0.3X - O.SX elytral length, reconverging near apices; length 9X - 10X pronotal length; tricostate to quadricostate, costa 2 distinct basal

0.3X - 0.7X elytral length and distally absent or evansecent to apex, costa 4 distinct O.SX - IX elytral length and distally evanencent or absent, costa 6 absent or distinct 0.5X - 0.8X elytral length and distally absent or evanescent, humeral costa distinct over umbone 0.5X to elytral apex; reticulation irregular anteriorly to scabrous punctate, at times more distinct posteriorly, aerolets present or absent and quadrate to irregular; setation semi - erect, sparse, uniformly arranged. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight to strongly arcuate. Metasternum anteriorly carainate or not, completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxa conical, trochanteral socket 0.4X - 0.5X coxal length; trochanters elongate, length pro - and mesotrochanters 0.8X - IX coxal length, raetatrochanters 0,8X - IX metacoxal length, all femoral insertions truncate and not or slightly offset; femora strongly compressed, straight, short, 1.7X - 3X trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tarsi elongate, tarsomeres without laterally expanded planar pads; covered with semi - decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1 - 8 subequal, but narrowing uniformly posteriad; tergite 6 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1 - 8 subequal,but narrowing uniformly posteriorad; sternite 7 entire to emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma.

Aedeagus with parameres; parameres 0.5X - 2X larger than median lobe, serrate or not internally; basal piece large, arcuate, not or slightly compressed, length nearly 0.4X - 0.7X length of median lobe; median lobe 142

tubular, unmodified to expanded apically with apices acute and

divergent, apex membranous to ventrally membranous 0.5X medina lobe

length; internal sac not eversible; flagellum present or absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The biology of this genus is basically unknown except for the few comments on the biology of C. nigrum n. sp. 143

KEY TO THE SPECIES OF Ceratoprion (ADULT MALES)

1. Bicolored species with yellow pronotum ...... 2

1*. Unicolorous or bicolorous species with brown or black

pronotum ...... 3

2. (1) Elytra tricostate; antennomeres trapezoidal and nearly

filiform; Venezuela ...... bordoni n. sp. (p. 145)

2'. (1) Elytra quadricostate; antennomeres trianguliform and nearly

pectinate; Panama to Ecuador .... serricorne Gorham (p. 170)

3. (I1) Elytra tricostate; elytral disk granulate, not reticulate;

elytral setation moderately dense and uniformly distributed;

Venezuela ...... nigrum n. sp (p. 160)

3'. (!') Elytra quadricostate

4. (3*) Elytral humerus yellow; reticulation densely and uniformly

quadrate; elytral setation sparse, inserted only on costae;

Costa Rica ...... chaelarum n. sp. (p. 150)

4'. (31) Elytra unicolorous ...... 5

5. (4' ) Elytral reticulation regularly quadrate, ocassionally 144

interneures missing to yield elongate reticulation; elytral

setation sparse, inserted only on costae; Colombia ......

...... periosus n. sp. (p. 165)

5'. (4') Elytral reticulation irregularly quadrate; elytral setation

moderately dense, inserted on costae and floors of areolae;

Venezuela ...... mandibularum n. sp. (p. 155) 145

Ceratoprion bordoni NEW SPECIES

Fig. 14 a-e

TYPE. MIZA, male with the following label data: [Printed white label] "VENEZUELA Aragua \\ Rancho Grande \\ 1500 m. 19 XI 86 \\

10'21'N, 67'41'0." [Handwritten white label] "Col. \\ C. Bordon."

DIAGNOSIS. This species may be differentiated from other known

Ceratoprion by its 3 costate elytra with a scabrous-punctate disk. It also may be separated by the combined character states of: a yellow pronotum, the trapezoidal antennomeres, and the presence and location of

1 pair of anterior pronotal stemmata.

DISTRIBUTION. Known only from the Henri Petier National Park,

Venezuela, above the Rancho Grande Biological Station.

DESCRIPTION (Males): Small, slender, elongate, length 3.6-5.5 mm., width

0.6-1.1 mm at humerus. Black; except pronotum, basal process of elytra, and antennomere 11, penultimate and ultimate abdominal segments yellow; coxae and trochanters testaceous.

Head. Elongate, widest at eyes, then narrowing to posterior margin, shining; setae sparse, decumbent. Genal margin elongate, length greater than eye length, convexly narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly emarginate; dorsally radial origin inside head margin 0.3x radius. Antennal prominence indistinct from vertex, shelf-like; length 0,7x width, inserted between middle of eyes; pentagonal dorsally, barely produced medially, broadly and shallowly sulcate dorsomedially, sulcus indistinct posteriorly, poorly defined

0.5x anterior length; ventrally trianguliform, deeply concave, sides uniformly and convexly arcuate, widest near base. Antennal sockets subrectangular, separated by 0.lx width of epistomal margin and O.lx distance between eye and antennal socket. Antennifer insertion elevated medially, appearing basomedial. Interocular distance 2x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transversely narrow, height 0.3x width.

Epistomal margin broadly emarginate, almost straight, width 0.7x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly and distinctly rounded. Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps absent, Gula transverse, short, pentagonal; width slightly greater apically than basally; length 0.5x width apical margin. Antennae filiform-serrate, 0.6x body length; setation dense, uniformly distributed, decumbent on all antennomeres, seal iform setae absent; scape obconic, width apically 0.7x length; pedicel cylindrical, short, transverse, not compressed, length subequal antennomere 3; antennomere 3 adnate with base antennomere 4; 147

antennomeres 4-10 filiform-serrate, barely expanded apically, strongly

compressed; length antennomere 3 0.2x antennomre 4; antennomere 11

fusiform, compressed, length 8.8x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly 0.6x basal width; median longitudinal

carina obscure, median fovea slightly more distinct 0.3x pronotal

length, fovea closes at posterior margin; pronotal stemmata present, anterior pair separated from anterior margin by 2 stemmatal diameters and separated from lateral margins by 8 stemmatal diameters, posterior

lateral pair separated from posterior margin by 2 stemmatal diameters and separated from lateral margins by 2 stemmatal diameters, posterior

internal pair separated from posterior margin by 3 stemmatal diameters and separated from lateral margins by 12 stemmatal diameters; all discal margins swollen; anterior margin nearly straight, anterior angles obtuse; lateral margin complete, foliaceous, slightly reflexed, nearly straight; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, discal setae seroi-decumbent; hypomeron shallowly concave, height 0.6x length; all margins swollen, hypomeral stemmata absent; prosternal bridge Y- shaped, anterior margin broadly emarginate, length less than 0.5x width, intercoxal process not carinate. Scute11urn pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly 0.3x scutellar length, apices truncate, slightly divergent. Mesospiracle obscure, peritreme not exposed.

Mesepisternum elongate, posterior margin convexly arcuate. Mesepimeron width and height subequal mesepisternum; posterior margin strongly 148

reflexed, evenly arcuate. Mesosternum length equal width, posterior

margin convexly arcuate. Mesocoxae separated by 0.8x coxal diameter at

base. Elytra ligulate, slightly dehiscent at middle, reconverging at

apices; length lOx pronotal length; tricostate, costa 2 distinct basal

0.3x elytral length, costa 4 distinct most elytral length, humeral costa

distinct most elytral length, just short of costa 4; disk irregularly $

scabrous-punctate, occasionally obscure and irregular transverse costae

between the apices of costae 4 and 8; setation semi“decumbent, sparse, uniformly arranged. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin strongly arcuate.

Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5x coxal length; trochanters elongate,

length pro- and mesotrochanters subequal to coxal length, metatrochanters subequal metacoxal length, femoral insertions truncate, not offset; femora strongly compressed, straight, short, 1.7x trochanteral length, annulated basally; tibiae strongly compressed,

internal margin slightly sinuate; tarsi elongate, tarsomeres without

laterally expanded plantar pads; covered with semi-decumbent bristling

setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, 1-8 narrowing uniformly posteriorad; sternite 7 emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; 149 paramere length 0.5x median lobe, serrate internally; basal piece large, arcuate, not compressed, length about 0.4x length of median lobe (Fig.

14 a-c>; median lobe tubular, unmodified, ventrally membranous 0.5x

length, internal sac not eversible, fiagellum present.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures ). Unknown.

DISCUSSION. All specimens were caught in a flight intercept trap

(Joly, personal communication, 1987).

MATERIAL EXAMINED. VENEZUELA, Aragua, Rancho Grande, 1500 m, 19

November 1986, C. Bordon (13-MIZA, including holotype; 2-NMNH; 1-BMNH;

1-MNHP; 3-RSMC; l-MAIC).

ETYMOLOGY. This species is named after Carlos Bordon who collected all known specimens. 150

Ceratoprion chelarum NEW SPECIES

Fig. 15 a-c

TYPE. HAHC, male with the following data - [white printed label]

"COSTA RICA, Limon P. \\ Valle de la Estrella \\ Pandora. 17-20 Feb.

1984 \\ H. & A. Howden."

DIAGNOSIS. C. chelarum may be distinguished by the combined presence

of the yellow humeral angles on quadracostate elytra with reticulation

quadrate, and strongly serrate antennae with trianguliform antennomeres.

It is the only known species of Ceratopri on without pronotal stemmata.

DISTRIBUTION. Known only from the type locality in Costa Rica.

DESCRIPTION (Male). Slender, elongate; length 4.0 mm., width at

humerus 1.0 mm. Black; except hypomera and posterior angles of

pronotum, humeral angles extending 0,5x elytral length, trochanters,

penultimate and ultimate tarsoroeres yellow to testaceous.

Head. Elongate, widest behind eyes, then narrowing to posterior

margin, shining; setae sparse, decumbant. Genal margin elongate,

length greater than eye length, uniformly narrowing when viewed

dorsally. Eyes hemispherical, projecting anterior dorsally when viewed dorsally; hind margin broadly emarginate; dorsally radial origin inside

head margin 0.3x radius. Antennal prominence slightly distinct from vertex, shelf-like, almost gibbose; length 0.8x width, inserted between middle of eyes; nearly quadrate dorsally, anterior margin transversely arcuate, broadly and obscurely sulcate dorsoraedially, sulcus more distinct posteriorly, indistinct 0.5x anterior length; ventrally nearly quadrate, slightly wider near base. Antennal sockets subrectangular, separated by 0.2x width of epistomal margin and 2x distance between eye and antennal socket, Antennifer elevated medially, appearing basomedial. Interocular distance nearly 2x width eye dorsally.

Vertex reticulate, transversely flat, setae semi-decumbent; depression of tentorial maculae shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transversely narrow, height less than 0.5x width. Epistomal margin broadly and deeply emarginate, width 0.8x frontal distance between eyes.

Labrum transverse, rectangular, length O.Sx width, apical margin narrowly and distinctly emarginate. Maxillary palps 3 segmented, palpomeres 1-2 cylindrical, diameter nearly equal to length; length palpomere 3 nearly equal 2, diameter at base less than O.Sx palpomere 2, acuminate. Labial palps 1 segmented, digitiform. Gula transverse, pentagonal; width slightly greater apically than basally; length approximately 2x width apical margin. Antennae serrate, Q.7x body length, setation dense, uniformly distributed, decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically 0.8x length; pedicel cylindrical, short, transverse, not compressed, length subequal length of antennomere 3; antennomere 3 adnate with base antennomere 4, antennomeres 4-10 serrate, expanded apically, strongly compressed; length antennomere 3 O.lx antennomere 4; antennomere 11 152

fusiform, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal

width, anterior width slightly more than 0.5x basal width; median

longitudinal carina distinct, median fovea distinct, slightly more than

0.5x pronotal length, fovea closes at posterior margin; pronotal

stemmata absent; all discal margins swollen; anterior margin nearly

straight, anterior angles obtuse; lateral margin complete, foliaceous, not reflexed, slightly concavely arcuate; posterior margin bisinuate; posterior angles acute, expanded laterally 0.8x width elytral humerus; disk flat, setae semi-decumbent; hypomeron shallowly concave, height

0.5x length, all margins swollen, hypostomal stemmata absent; prosternal bridge Y-shaped, length O.Sx width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly 0.3x scutellar length, apices acute and divaricate. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, 0.5x width and 0.7x height mesepisternum width and height; posterior margin very little reflexed, nearly straight, angulate ventrally. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separated by coxal diameter at base. Elytra ligulate, slightly dehiscent from elytral middle; length 9x pronotal length; quadricostate, costa 2 distinct 0.7x elytral length, costa 4 and 6 distinct O.Sx elytral length, humeral costa distinct over umbone to elytral apex; reticulation irregular anteriorly, more distinct posteriorly, aereolets quadrate in posterior

0.3x elytra; setation semi-erect, sparse, uniformly arranged. 153

Metepisternum with ventral margin inreasingly arcuate posteriorly.

Metepimeron with ventral margin nearly straight. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket O.Sx coxal length; trochanters elongate, length pro- and mesotrochanters Q.8x coxal length, metatrochanters subequal metacoxal length, all femoral insertions truncate and slightly offset; femora strongly compressed, straight, short, 2.3x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal,but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; parameres slightly larger than median lobe, not serrate internally; basal piece large, arcuate, not compressed, length nearly

0.7x length of median lobe (Fig. 15 a-c); median lobe tubular, expanded apically, apices acute and divergent, apex membranous; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown. 154

MATERIAL EXAMINED. COSTA RICA, Limon, Valle de la Estrella Pandora,

17-20 February 1984, H. & A. Howden (1-HAHC, holotype).

ETYMOLOGY. Name refers to the chelate - like appearance of the parameres in the male. 155

xxxxxxxxxxxxxxxxxxxxx

Maxilla with 4 segmented palp; 1 - 3 cylindrical, diameter subequal to length; 4 small, diameter less than O.Sx segment 3, pointed distally.

Costa 1 obscure posterior 0.6x, anterior 0.3x incomplete, angularly serpentine; costae 2 and 4 complete, linear, approaching apex; costa 3 obscure basal 0.5x, complete only apical 0.3x; pubescence dimorphic, costal and interneural setae long and erect, floors with decumbent dense setae 0.lx length of erect setae; fenestrae irregular, mostly trapeziform. 155

Ceratoprion mandibularum NEW SPECIES

Figs. 16 a-d

TYPE. MIZA, a male with the following data: CPrinted white labelD

"VENEZUELA, Meri- \\ da. Carbonera. \\ 2600 m. 28 - VI - 68;" CPrinted

white label3 "J. & B. Bechyne \\ leg."

DIAGNOSIS. This species differs from others in the genus by the

following combined character states: the coloration is uniformly brown

except for the pronotal stemmata and the last antennomer which are

yellowish white, the elytra is quadricostae with irregular quadrate

reticulation, and the elytral setae are found in the areolae as well as

the on the costae.

DESCRIPTION

mm. at humerus. Brown; except ultimate antennomere and pronotal

stemmata yellowish white.

Head. Transverse, widest at eyes, then narrowing to posterior

margin, dull; setae sparse, semi-decumbant. Genal margin elongate,

length greater than eye length, narrowing posteriorly when viewed

dorsally. Eyes hemispherical, projecting anteriolaterally when viewed

dorsally; hind margin broadly and shallowly emarginate; dorsally radial

origin inside head margin 0.3x radius. Antennal prominence distinct from vertex, gibbose; length equal width, inserted between middle of eyes; pentagonal dorsally, narrowing at base, shallowly sulcate dorsomedially, sulcus distinct posteriorly, poorly defined 0.5x anterior length; ventrally trianguliform, shallowly concave, sides uniformly and

convexly arcuate, widest near base. Antennal sockets subrectangular,

separated by less than O.lx width of epistomal margin and subequal to

distance between eye and antennal socket. Antennifer elevated

medially, appearing basomedial. Interocular distance 2x width eye

dorsally. Vertex transversely flat, setae semi-decumbent; depression

of tentorial maculae shallow and broadly defined, from base of antennal

prominence to behind posterior eye margin, somewhat more distinct

anteriorly. Tentorial maculae separate, shallow, distinct; at base of

antennal prominence. Frons transversely flat, distinct from lower

surface of antennal prominence, transversely narrow, height 0.3x width.

Epistomal margin broadly emarginate, almost straight, width 0.7x frontal

distance between eyes. Labrum transverse, rectangular, length 0.3x

width, apical margin truncate. Maxillary palps 4 segmented, palpomeres

1-3 cylindrical, diameter nearly equal to length palpomere 2; length

palpomeres, except palpomere 2 0.5x length 2; diameter palpomere 4 less

than 0.5x diameter palpomere 3 basally, acuminate. Labial palps 1

segmented, digitiform. Gula elongate, subrectangular, width slightly

greater apically than basally, length 2x width anterior margin, inserted anteriorad posterior margins of postgena. Antennae serrate, 0.7x body

length; setation sparse, uniformly distributed, decumbent on all antennomeres, seal iform setae absent; scape elongate obconic, width apically 0.4x length; pedicel cylindrical, short, transverse, not compressed, length subequal antennomere 3; antennomere 3 adnate with base of antennomere 4; antennomeres 4-10 serrate, expanded apically,

strongly compressed; length antennomere 3 0.2x antennomere 4; 157

antennomere 11 fusiform, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width 0.5x basal width; median longitudinal carina

indistinct, median fovea distinct 0.5x pronotal length, fovea closed before posterior margin; pronotal stemmata present, anterior pair on anterior margin and separated from lateral margins by 4 stemmatal diameters, posterior lateral pair on posterior margin at lateral margin, posterior internal pair on posterior margin and separated from lateral margins by 2 stemmatal diameters; all discal margins swollen; anterior margin convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, slightly reflexed, nearly straight; posterior margin strongly bisinuate; posterior angles acute, expanded laterally

0.8x width at elytral humerus; disk flat, discal setae sparse, semi- decumbent; hypomeron shallowly concave, height 0.3x length, all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, anterior margin broadly emarginate, length less than 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex incised, apices truncate.

Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin convexly arcuate. Mesepimeron trianguliform, 0.5x width and 0,7x height mesepisternum; posterior margin slightly reflexed, strongly angulate at ventral 0.7x. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separation by coxal diameter at base. Elytra ligulate, dehiscent at 0.3x elytral length from base, reconverging at apices; length lOx pronotal length; quadricostate, costa 2 distinct 0.5x basal elytral length, costa 4 distinct most elytral length, costa 6 distinct apical 0.5x elytral

length, humeral costa distinct over umbone to near elytral apex and

sinuate; disk irregularly scabrous-punctate, setation semi-decumbent, dense, uniformly distributed. Metepisternum with ventral margin

increasingly arcuate posteriorly. Metepimeron with ventral margin

straight. Metasternum anteriorly carinate, completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.4x coxal length; trochanters elongate, length pro- and mesotrochanters subequal to coxal length, metatrochanters aubequal to metacoxal length, all femoral insertions truncate and offset; femora strongly compressed, straight, short, 3x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi- decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 deeply emarginate, emargination 0.5x segmental length. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; paramere length

2x median lobe, extends beyond apex of median lobe, crossing at apices; basal piece large, slightly compressed, length 2.5x length of median lobe (Fig. 16 a-c); median lobe tubular, unmodified apically, ventrally membranous 0.5x length; internal sac not eversible; flagellum absent. 159

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISTRIBUTION. Known only from the type.

MATERIAL EXAMINED. VENEZUELA, Merida, Carbonera, 2600 m, 28 June

1968, J. & B. Bechyne (1-MIZA, holotype).

ETYMOLOGY. The specific epithet refers to the resemblance of the crossed parameres to predacious mandibles. 160

DESCRIPTION. Slender, elongate; length 7.0 mm., width at humerus 1.4 mm. Brown except antennomeres 10 - 11; pronotal gibosities on anterior and posterior margin yellow.

Eyes hemispherical, hind margin slightly emarginate, nearly straight

Antennal prominence shallowly sulcate dorsomedially, sulcus widens

posteriorly into broad deep pretentorial depression.

Interocular distance 0.66x eye width dorsally. posterior margin rounded; deepest posteriorly, shallow and narrow at

sulcus of antennal prominance.

Pretentorial pits separate, distinct.

Epistomal margin broadly emarginate; defexed posteriad.

Labrum square; width subequal to length.

Maxilla with 4 segmented palp; 1 - 3 cylindrical, diameter subequal to

length; 4 small, diameter less than 0.5x segment 3, pointed

di stally.

Gula transverse, width ZZx length.

Antennae elongate, ZZx body length; scape pyriform; pedicel

cylindrical, transverse, length subequal to antennomere 3;

antennal segment 3 serrate, compressed; antennomeres 4-10

expanded laterally from base.

Pronotum trapezoidal, transverse, length 0.5x basal width; anterior

width nearly 0.5x basal width; median longitudinal carina divided posterior 0.33; anterior margin nearly straight, incised medially, one pair pronotal stemmata; posterior margin strongly bisinuate; posterior angles acutely projecting laterally, not beyond humerus discal setae long, decumbent, oriented mesad posterior margin sinuate disk with 1 pair pronotal stemmata in front of elytral insertions

Scutellum exposed, longer than wide, apex broadly emarginate/excavate.

Elytra 4 costate; costa 1 obscure posterior 0.25; costae 2 and 4 to

near apex; costa 3 evanescent basally, more defined posterior

0.25; costa 4 pronounced at umbone; interneures poorly defined;

reticulation [obscure!, longitudinally elongate, rectangular;

cell floor vestiture short, dense; costal and interneural setae

long, decumbent. PLEASE NOTE

Page(s) missing in number only; text follows. Filmed as received.

162-164

University Microfilms International 165

Ceratoprion periosus NEW SPECIES

Fig. 18 a-c

TYPE. ZMHB, a male with the following label data; CPrinted green

label! "Columbien \\ Paso del Quindiu \\ 12000' 0. Thiermo S."

[Handwritten green label! "Palo del Quin. \\ die."

DIAGNOSIS. This brown species is immediately recognized by its relatively large size combined with the quadricostate, fenestrate elytra whichyhave regular quadrate reticulation and sparse setation restricted to the costae,

DISTRIBUTION. Known only from the type specimen from Colombia.

DESCRIPTION (Male): Si 2e moderate, slender, elongate, length 6.3 mm., width 1.2 mm at humerus. Brown; except ultimate and penultimate antennomeres, apical tergite and sternite yellow.

Head. Elongate, widest at eyes, then narrowing to posterior margin, shining; setae sparse, semi-decumbent. Genal margin short, length less than eye length, narrowing posteriorly when viewed dorsally.

Eyes hemispherical, projecting anterior laterally when viewed dorsally; hind margin broadly and deeply emarginate; dorsally radial origin inside head margin 0.3x radius. Antennal prominence barely distinct from vertex, shelf-like; length equal width, inserted between middle of eyes; quadrate dorsally, barely produced medially, broadly and shallowly sulcate dorsomedially; ventrally pentagonal, concave, sides uniformly and convexly arcuate, widest near base. Antennal sockets rectangular,

separated by 0.lx width of epistomal margin and 2.Ox distance between eye and antennal socket. Antennifer elevated medially, not appearing basomedial. Interocular distance subequal width eye dorsally. Vertex

slightly arcuate, nearly transversely flat, setae decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly.

Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transverse, height 0.3x width. Epistomal margin broadly emarginate, width 0.7x frontal distance between eyes. Labrum transverse, rectangular, length approximately 0.5x width, apical margin broadly and distinctly rounded. Maxillary palps 4 segmented, palpomere length 1 equal length 3, palpomere 2 2x palpomere 1, diameter 1 and 2 nearly equal to length 2; length palpomere 4 nearly 0,3x 3, diameter palpomeres 3 and 4 at base less than Q.5x palpomere 2, palpomere 4 acuminate. Labial palps absent. Gula transverse, short, pentagonal; width slightly greater apically than basally; length 0.5x width apical margin. Antennae serrate, 0,6x body length; setation sparse, uniformly distributed, decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically 0.6x length; pedicel cylindrical, short, transverse, not compressed, length subequal antennomere 3; antennomere 3 adnate with base antennomere 4; antennomeres 4-10 serrate, expanded apically, strongly compressed; length antennomere 3 0.2x antennomere 4; antennomere 11 fusiform, compressed.

Thorax. Pronotum trapezoidal, transverse, length Q.5x basal width, anterior width nearly 0.5x basal width; median longitudinal

carina well defined, median fovea slightly more distinct 0.7x pronotal

length, carinae narrowly separated, fovea closed before posterior

margin; pronotal stemmata present, anterior pair separated from anterior margin by 1 stemmatal diameters and separated from lateral margins by 8

stemmatal diameters, posterior lateral pair separated from posterior margin by 1 stemmatal diameters and separated from lateral margins by 2 stemmatal diameters, posterior internal pair separated from posterior margin by 3 stemmatal diameters and separated from lateral margins by 12 stemmatal diameters; all discal margins swollen; anterior margin convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, concavely arcuate; posterior margin very strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, discal setae semi-decumbent; hypomeron shallowly concave, height 0.5x length; all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, anterior margin broadly emarginate, length less than 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly

0.2x scutellar length, apices divergent and rounded. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight, Mesepimeron trianguliform, width 0.3x and height 0.7x mesepisternum; posterior margin not reflexed, straight; angulate at venter. Mesosternum transverse, posterior margin convexly arcuate.

Mesocoxal separation 2x mesocoxal diameter at base. Elytra ligulate, slightly dehiscent at middle, reconverging at apices; length 18x pronotal length; quadricostate, costa 2 distinct basal 0.7x elytral length, costa 4 distinct most elytral length, costa 6 distinct 0.8x elytral length, humeral costa distinct O.flx elytral length; disk irregularly scabrous to reticulate, basally obscure and irregularly transverse to square areolets distally; setation semi-decumbent, sparse, uniformly arranged. Metepisternum with ventral margin increasingly arcuate anteriorly. Metepimeron with ventral margin strongly straight.

Metasternum incompletely divided medially by longitudinal sulcus, not reaching mesosternium by 0.5x mesocoxal diameter. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5x coxal length; trochanters elongate, length pro- and mesotrochanters 1.5x coxal length, metatrochanters subequal metacoxal length, all femoral insertions oblique and offset; femora strongly compressed, straight, short, 1.4x trochanteral length, annulated basally; tibiae strongly compressed, internal margin nearly straight; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-0 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

1-8 narrowing uniformly posteriorad; sternite 7 broadly emarginate.

Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; paramere length 0.5x median lobe, serrate internally; basal piece large, arcuate, not compressed, length about 0.4x length of median lobe (Fig. 18 a-c); median lobe tubular, unmodified, ventrally membranous 0.5x length, internal sac not 169 eversible, flagellutn present.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. COLOMBIA, Pao del Quindiu, 4175 m, December

(1-ZMHB, holotype).

ETYMOLOGY. Although this species is not large by comparison to other lycid genera, it is immense for Leptolycini as here defined. Hence, the Greek specific epithet meaning immense. 170

Ceratoprion serricorne Gorham

Figs, 19 a-f

Ceratopri on serricorne Gorham, 1884:248. Kleine, 1933:34.

Blackwelder, 1945:348.

TYPE. BMNH, a male with the following labels: CPrinted circular red labell "Type." CPrinted white label! "Bugaba \\ 800-1500 ft. \\

Champion." CHandwritten label! "Ceratoprion \\ serricorne \\ Gorham."

CPrinted label! "B. C. A. Col. III(2J Ceratoprion \\ serricorne W

Gorham."

DIAGNOSIS. This brown species is distinguished by the combination of its quadricostate elytra with regular reticulation, the distal antennomeres are trianguliform, the presence of two pair of anterior pronotal stemmata as well as two pair of posterior pronotal stemmata, and the yellow pronotum,

DISTRIBUTION. Known from central Panama south to Ecuador. Although I have seen few specimens of this species, its distribution is more widespread than any other known leptolycine.

DESCRIPTION (Male): Small, slender, elongate, length 3.8-4.5 mm., width

0.8-0.9 ram, at humerus. Brown; except prothorax, pedicel and antennomere 11 yellow; occassionally antennomere 3, apex of 10, pro- and mesocoxae, trochanters, bases of femora and tibiae yellow and pronotum ultimate abdominal segment testaceous; pronotal stemmata yellow. Head. Elongate, widest at eyes, then narrowing to posterior

margin, dull; setae sparse, decumbent. Genal margin short, length

slightly shorter than eye length, narrowing posteriorly when viewed

dorsally. Eyes hemispherical, projecting anteriolaterally when viewed

dorsally; hind margin broadly and shallowly emarginate; dorsally radial

origin at head margin. Antennal prominence slightly distinct from

vertex, shelf-like; length 0.4x width, inserted between middle of eyes;

quadrate dorsally, not produced medially, broadly and shallowly sulcate dorsomedially, sulcus distinct; ventrally trapezoidal, shallowly concave, antenniferae inserted at dorsolateral angles. Antennal

sockets subrectangular, separated by O.lx width of epistomal margin and equidistant separation between eye and antennal socket. Antennifer elevated medially, not appearing basomedial. Interocular distance 2x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct, at base of antennal prominence. Frons transversely flat, distinct from

lower surface of antennal prominence, transversely narrow, height 0,3x width. Epistomal margin broadly and deeply emarginate, width 0.7x frontal distance between eyes. Labrum transverse, rectangular, length

0.5x width, apical margin broadly rounded. Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length;

length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps 2 segmented, digitiform. Gula transverse, short, trapezoidal; width slightly greater basally than 172

apically; length 0.5x width apical margin. Antennae serrate, O.Qx body

length; setation sparse, uniformly distributed, decumbent on all

antennomeres, scaliform setae absent; scape obconic, width apically Q.7x

length; pedicel cylindrical, short, transverse, not compressed, length

subequal antennomere 3; antennomere 3 adnate with base antennomere 4,

length 0.2x antennomere 4;; antennomeres 4-10 filiform-serrate, strongly

expanded apically, strongly compressed antennomere 11 fusiform,

compressed, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal

width, anterior width 0.5x basal width; median longitudinal carina

distinct, median fovea slightly more distinct 0.5x pronotal length,

fovea closes before posterior margin; pronotal stemmata present,

anterior lateral pair on anterior margin at lateral margin, anterior

internal pair on anterior margin and separted from lateral margin by 1

stemmatal diameter, posterior lateral pair on posterior margin at

lateral margin, posterior internal pair on posterior margin and

separated from lateral margins by 1 stemmatal diameter; all discal

margins swollen; anterior margin nearly straight, anterior angles

obtuse; lateral margin complete, foliaceous, erect, nearly straight; posterior margin bisinuate; posterior angles acute, expanded laterally

Q.9x width at elytral humerus; disk flat, discal setae semi-decumbent;

hypomeron shallowly concave, height 0.3x length; all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, anterior margin

broadly emarginate, length 0.5x width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex shallowly emarginate, emargination 0.3x scutellar length, 173 apices truncate, slightly divergent. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin convexly arcuate. Mesepimeron 0.3x width and 0.5x height mesepisternum; posterior margin slightly reflexed, strongly angulate at ventral 0.8x.

Mesosternum width lx length, posterior margin convexly arcuate.

Mesocoxal separation 1.5x mesocoxal diameter at base. Elytra ligulate, strongly dehiscent at middle, reconverging at apices; length 9x pronotal

length; quadricostate, costa 2 distinct basal 0.5x elytral length and evanescent distally to apex, costae 4 and 6 distinct basal 0.8x elytral length and evanescent distally and convergent near apex, humeral costa distinct to middle elytral length and evanescent to 0.7x elytral length; disk reticulate to irregularly scabrous-punctate; setation semi-erect, sparse, uniformly arranged. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin straight. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket Q.5x coxal length; trochanters elongate, length pro- and mesotrochanters O.Qx coxal length, metatrochanters 0.8x metacoxal length, femoral insertions truncate, offset; femora strongly compressed, straight, short, 2.7x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly sinuate; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 174

medially divided longitudinally; tergite 10 shield shaped. Sternites

rectangular, wider than long, length 1-8 subequal, nut narrowing

uniformly posteriorad; sternite 7 broadly emarginate. Spiracles in

pleural membrane, located submedially from apical margin of tagma.

Aedeagus with parameres; paramere length 0.5x median lobe, dentate

internally; basal piece large, arcuate, slightly compressed, length

about 0.6x length of median lobe (Fig. 19 a-c); median lobe tubular,

apically spatulate, ventrally membranous more than 0.5x length, internal

sac not eversible, flagellum not present.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Imraatures). Unknown.

MATERIAL EXAMINED, PANAMA, Bugaba, 245-460 m, Champion (1-BMNH, holotype; 1-MNHP); Darien Province, Santa Fe, April to May, D. M.

DeLong and C. A. Triplehorn, malaise trap (1-0SUC; 1-RSMC). ECUADOR,

Napo Province, Limoncocha, 250 m., 15-28 June 1976, S. & J. Peck, malaise trap (1-CNCI).

ETYMOLOGY. The specific epithet refers to the serrate antennae. 175

Flabellocaenia Pic

Flabellocaenia Pic, 1929. Kleine, 1933:31. Blackwelder, 1945:347.

TYPE: Flabellocaenia bourgeoisi Pic, MNHP, by original monotypy.

DIAGNOSIS. Flabellocaenia can be separated from other leptolycines by

the combined presence of: the flabellate antennae with pedicel subequal

to antennnomere 3 and rami on antennomeres 4-11; by the 3 costate, slightly explanate-1igulate elytra; by the absence of a short rostrate epistoma; by the small mouthparts; and by the absence of pronotal stemmata. The venter of the head is horizontally flat and the mouthparts narrow which gives the impression that the mouthparts are highly reduced or absent.

DISTRIBUTION. This genus is known only from the Amazon basin.

DESCRIPTION (Males). Small, slender, elongate, but elytra expanded distally 0.7x their length and then narrowing to apex; length 4.1-5.0 ram, width at humerus 0.9-1.1 mm. Blackish brown, sometimes testaceous; except yellow maculation.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining to dull; setae sparse, decumbent to serai-decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin at or 0.3x inside head margin when dorsally viewed. Antennal prominence slightly distinct from vertex, shelf-like to slightly

gibbose; length 0.3-0.6x width, inserted between middle of eyes; pentagonal dorsally to rectangular and slightly produced apically, not

to shallowly impressed dorsomedially; ventrally trianguliform, flat to

slightly or deeply concave, often longitudinally impressed, widest near base, sides nearly straight. Antennal sockets subrectangular, separated by O.lx width of epistomal margin and l-3x distance between eye and antennal socket. Antennifer insertion elevated medially, not appearing basomedial. Interocular distance 2-4x width eye dorsally.

Vertex transversely flat, setae semi-decumbent; depression of tentorial maculae broad, deep to shallow and poorly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly.

Tentorial maculae separate, shallow to deep, distinct; at base of antennal prominence. Frons transversely flat or concave, distinct from lower surface of antennal prominence, transversely narrow, height 0.3-

Q.4x width. Epistomal margin truncate to broadly or narrowly and shallowly to strongly emarginate, width 0.3-0.6x frontal distance between eyes. Labrum transverse, rectangular, length 0.3-Q.5x width, apical margin truncate. Maxillary palps minute, 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps absent to 1 segmented and digitiform. Gula transverse, linear; length 0.1-0.3x width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabellate, length

0.6-0.7x body length; rami between 3-1lx axial length and their compression perpendicular to axial compression; setation sparse, 177 uniformly distributed to distributed along margins, decumbent to semi- decumbent on all antennomeres, seal iform setae present; scape obconic, width apically subequal length; pedicel cylindrical, short, transverse, not compressed, length l-2x length of antennomere 3; antennomere 3 slightly compressed, adnate to antennomere 4 basally, length 0.8x antennomere 4; antennomere 4 flabellate with raraal insertions basal to medial, more strongly compressed; antennomeres 5-10 compressed, extent of compression uniform or decremental to apex, flabellate with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5-Q.7x basal width, anterior width 0.6-Q.7x basal width; median longitudinal carina present, divided in middle 0.3-0.7x pronotal length before posterior margin, fovea present and 0.5-lx pronotal length, closed or not at posterior margin; pronotal stenunata absent; all discal margins swollen or variously not; anterior margin straight to slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect to slightly explanate, slightly concavely arcuate longitudinally; posterior margin moderately to strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat to slightly transversely arcuate, but anterior 0.3-0.5x inflated allowing reception of head, lateral margins deeply impressed or not, setation sparse, decumbent to semi-decumbent; hypomeron concave, height 0,5-0.7x width, all to various margins swollen, hypomeral stemmata absent; prosternal bridge T- or Y-shaped, lateral arms broadly emarginate or obtusely to acutely separated, anterior emargination shallow, length 0.5-0.6x width, intercoxal process not carinate. Scutellum pentagonal, distally

trianguliform to quadrate and nearly square, longitudinally impressed or not, lateral margins parallel to divergent apically, apex shallowly to deeply emarginate, emargination 0.1-0.5x scutellar length, apices acute or rounded and divergent. Mesospiracle prominent, peritreme exposed,

orifice slightly or not compressed. Mesepisternum elongate, posterior margin straight to slightly convexly arcuate. Mesepimeron lobate; width

0.5-lx and height O.Bx mesepisternum; posterior margin moderately to strongly reflexed, evenly rounded. Mesosternum transverse to almost quadrate, length 0.4-lx width, posterior margin convexly arcuate.

Mesocoxal separation 1-1.3x mesocoxal diameter at base. Elytra slightly explanate-1igulate, slightly dehiscent from 0.3-0.5x elytral length, expanded to 0.7x and narrowed distally to apex, slightly to strongly reconvergent near apex, apices acute; length 7-10x pronotal length; tricostate, costa 2 distinct basal 0.3-0.6x elytral length, costae 4 and 8 variously distinct to elytral length, costa 8 nearly arcuate or sinuate and convergent apically to near 4 or not; disk irregularly scabrous-punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly to increasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight to slighlty arcuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, decumbent to semi-decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.5-0.8x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters l.l-1.3x coxal 179

length, metatrochanters 0.9-1,4-x metacoxal length, all femoral insertions truncate and not offset or metatrochanters slightly oblique and offset; femora compressed, clavate, short, 1.8-2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight to arcuate to sinuate; tibial spines small and similar; tarsi short to very short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire to produced and bilobate; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire to emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; paramere length 0.9-lx median lobe, differentiated from median lobe apical 0.3-lx length of lobe, sinuate apically to laterally divergent apically or deflexed dorsally; basal piece dorsoventrally compressed, length almost 0.3~0.7x length of median lobe; median lobe tubular, unmodified to deflected apically, membranous ventrally at apex to most its length; internal sac not eversible; fiagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown. ETYMOLOGY. The prefix f1abe11o (Latin) meaning a fan and the root caenia (Greek) meaning new refers to this genus' superficial similarity in appearance to the calopterine genus Caenia Newman. However, the two genera are only distantly related and reference to the flabellate antennae does not distinguish it from the latter, as the males of that genus also have flabellate antennae. In fact the resemblance is mostly a result of that antennal state. 181

KEY TO SPECIES OF Flabellocaenia

1. Elytra unicolorous ...... 2

1'. Elytra bicolorous ...... 3

2. (1) Epistomal margin deeply and narrowly emarginate; distance

between eye and antennal socket 3x interantennal socket

distance; genitalia as in fig. 23 a-c; Venezuela ......

...... Flabellocaenia iolvi n. sp. (p. 194)

2'. (1) Epistomal margin truncate; distance between eye and antennal

socket lx interantennal socket distance; genitalia as in

fig. 25 a-c; Venezuela ......

...... Flabellocaenia rimae n. sp. (p. 209 )

3. (1*) Yellow elytra with blackish brown narrow post median fascia

and along sutural margin ...... 4

3'. (1* ) Elytral blackish brown with apex and base briefly yellow ...

...... 5

4. (3) Epistomal margin broadly emarginate; elytral costae reach

apical margin; genitalia as in fig. 21 a-c; Ecuador ......

...... Flabel locaenia elegantulus n. sp. (p. 189) Epistomal margin narrowly emarginate; elytral costae do not reach apical margin; genitalia as in fig. 24 a-c; Ecuador

...... Flabellocaenia pecki n. sp. (p. 204)

Venter entirely yellow, except tibiae distally and tarsi brownish black; yellow of costae at base extends further posteriorad than on elytral floors; Bra2 il ......

...... Flabellocaenia bourgeoisi Pic (p. 183 )

Venter mostly brown, except leg segments briefly yellow at insertions; base of elytra yellow at insertions, not extending along costae further than on elytral floors; elytral apex briefly yellow; Colombia ......

...... Flabel 1 ocaenia leticia n. sp. (p. 199 ) 183

Flabellocaenia bourgeoisi Pic

Fig. 20 a-f

Flabellocaenia bourgeois! Pic, 1929:76. Pic, 1931:104. Kleine,

1933:31. Blackwelder, 1945:347.

TYPE: MNHP, a card mounted male in poor condition with the following

label data: white handwritten label [Amaz.]; white handwritten label Cex

collection \\ Gorham]; white handwritten label CGorhamia \\ flabellifera

Bourg. \\ genus voisins de \\ Acroleptis]; white handwritten label

CFlabellocaenia \\ n. gen W Bourgeoi si]: blue printed label [Museum

Paris \\ coll. M. PicD.

DIAGNOSIS. The yellow venter combined with the basal yellow along the

costae extending further posteriorad than on the floors is distinctive

within the genus.

DISTRIBUTION. Known only from the Amazon Basin, but the exact locality

of any specimen is unknown.

DESCRIPTION (Males). Elongate, but elytra expanded distally 0.6x their

length and then narrowing to apex; length 4.4-5.6 mm, width at humerus

0.9-1.3 mm. Blackish brown; except yellow maculation; elytra at humeral angles, extending along lateral margin to 0.5x elytral length,

apical 0.3x elytral length yellow; antennal axis except distal 3

antennoroeres, apices of rami antennoneres 6-11 yellow; mouthparts, 184 coxae, femora and basal 0.3x tibia yellow except femora testaceous band encircling middle.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin at head margin when dorsally viewed. Antennal prominence slightly distinct from vertex, shelf-like and slightly gibbose; length 0.4x width, inserted between middle of eyes; quadrate dorsally, not impressed dorsomedially; ventrally trianguliform, concave, impressed longitudinally, sides nearly straight, widest near base. Antennal sockets subrectangular, separated by 0.lx width of epistomal margin and 0.5x to distance between eye and antennal socket. Antennifer insertion elevated medially. Interocular distance 2.2x width eye dorsally. Vertex transversely arcuate, setae semi-decumbent; depression of tentorial maculae broad and shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely concave, distinct from lower surface of antennal prominence, transversely narrow, height 0.3x width.

Epistomal margin shallowly emarginate, width 0.4x frontal distance between eyes. Labrum unseen. Maxillary palps unseen. Labial palps absent. Gula unseen. Antennae flabellate, length 0.6x body length; rami between 3-4x axial length and their compression perpendicular to axial compression; setation sparse, distributed along margins, semi- decumbent on all antennomeres, scaliform setae present; scape obconic, 185 width apically subequal length; pedicel cylindrical, short, transverse, not compressed, length 2x length of antennomere 3; antennomere 3 adnate to antennomere 4 basally, slightly compressed; antennomere 4 flabellate with ramal insertions medial, more strongly compressed; antennomeres 5-

10 compressed, extent of compression decremental to apex, flabellate with ramal insertions apical; length antennomere 3 0.8x antennomere 4; antennomere 11 without axis.

Thorax. Pronotum trapezoidal, length 0.6x basal width, anterior width 0.6x basal width; median longitudinal Carina present, divided in middle D.3x pronotal length before posterior margin, median fovea 2x pronotal length, closing at posterior margin; pronotal stemmata absent; all discal margins swollen; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, slightly concavely arcuate; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk fiat, but anterior 0.3x inflated allowing reception of head, setation decumbent; hypomeron concave, height 0.6x width, anterior and posterior margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms strongly V-shaped, length 0.5x width, intercoxal process not carinate. ScuteHum pentagonal, distally quadrate and nearly square, lateral margins parallel, apex deeply emarginate, emargination 0.3x scutellar length, apices acute and divergent. Mesospiracle prominent, peritreme exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin slightly convexly arcuate. Mesepimeron lobate; width equal and height 0.8x mesepisternum; posterior margin strongly reflexed, evenly rounded. Mesosternum length equal width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, slightly dehiscent from middle, slightly reconvergent at apex, apices acute; length lOx pronotal length; tricostate, costa 2 distinct basal 0.3x elytral length, costa 4 and 8 distinct 0.8x elytral length; disk irregularly scabrous- punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight.

Metasternuo completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, semi-decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.6x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 1.lx coxal length, metatrochanters 1.2x metacoxal length, all femoral insertions truncate and not offset; femora compressed, clavate, short, 2.Ox trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight; tibial spines small, similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniforml posteriad; tergite 8 entire; tergite 9 not medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma.

Aedeagus with parameres; paramere length 0.4x median lobe, 187 differentiated from median lobe apical 0.3x length of lobe, dentate bassally, sinuate apically; basal piece dorsoventrally compressed, length almost 0.6x length of median lobe (Fig, 20 a-c); median lobe tubular, unmodified, membranous ventrally O.Sx its length; internal sac not eversible; flagelium not present.

DESCRIPTION (Female). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. As the type and other material available are card mounted and appear fragile, the venter was not visible without the possibility of destruction. Therefore, many characters were not included in the description.

Pic evidently named a specimen from the Gorham collection he purchased after examining the unique specimen in the Bourgeois collection. Bourgeois' specimen has a label with the manuscript name

Gorhamia flabellatis. but that generic name was not available as Pic had used it earlier (Pic, 1911:186) for a genus he placed in the Drilidae.

Pic's type was examined and it is not congeneric with the latter species. Crowson (1972) later moved Gorhamia from the Driliidae to the Larapyridae.

MATERIAL EXAMINED. AMAZON. (2-MNHP, holotype and Bourgeois’ specimen). ETYMOLOGY. Named for Jules Bourgeois, a French lycid taxonomist, who was apparently respected by Pic, as he did not begin describing lycids until a month after Bourgeois' death. In that paper Pic advised the world that he was planning to continue in the tradition of his good friend. Bourgeois' work was excellent. 189

F 1abe11ocaenja elegantulus NEW SPECIES

Fig. 21 a-d

TYPE. CNC1, a male with the following label data: [white printed

label] "ECU. Napo Prov. \\ Limoncocha, 250 m. \\ 15-28.VI.1976 \\ S. &

J. Peck W Malaise trap."

DIAGNOSIS. It is most easily differentiated from F. pecki. which it

ressembles in coloration by the rami being 5x versus 4x axial length

basally, the elytral costae 4 and 8 reaching apex of the elytra versus only near it, the epistomal margin broadly versus narrowly emarginate, and the median lobe of the aedeagus is ventrally membranous at apex only

instead of along entire venter.

DISTRIBUTION. This species is known only from the type specimen which was collected near Limoncocha, Equador.

DESCRIPTION (Male). Elongate, but elytra expanded distally 0.7x their

length and then narrowing to apex; length 5.0 mm, width at humerus 0.9 mm. Yellow; except elytra briefly at base, fascia at apical 0.7x elytra, pronotum, head, pleurites, middle of all rami, apices of tibiae and tarsi blackish brown.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, semi-decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate; radial origin 0.3x inside head margin when dorsaliy viewed. Antennal prominence slightly distinct from vertex, shelf-like; length 0.6x width, inserted between middle of eyes; pentagonal dorsaliy, not impressed dorsomedially; ventrally trianguliform, deeply concave, longitudinally impressed, impression well defined, sides nearly straight, widest near base. Antennal sockets subrectangular, separated by O.lx width of epistomal margin and equidistant to distance between eye and antennal socket. Antennifer insertions elevated medially, not appearing basomedial. Interocular distance 2,7x width eye dorsaliy. Vertex transversely flat, setae semi-decumbent; depression of tentorial maculae broad and deep, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transversely narrow, height 0.4x width.

Epistomal margin broadly emarginate, width 0.6x frontal distance between eyes. Labrum not seen. Maxillary palps minute, 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps absent. Gula transversely rectangular; length 0,3x width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabellate, length 0.6x body length; rami between 3-

5x axial length and their compression perpendicular to axial compression; setation sparse, distributed along margins, semi-decumbent on all antennomeres, seal iform setae present; scape obconic, width apically subequal length; pedicel cylindrical, short, transverse, not 191

compressed, length 2x length of antennomere 3; antennomere 3 slightly

compressed, adnate to antennomere 4 basally, length 0.7x antennomere 4;

antennomere 4 flabellate with ramal insertions medial, more strongly

compressed; antennomeres 5-10 compressed, extent of compression

decremental to apex, flabellate with ramal insertions apical;

antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.7x basal width, anterior

width 0,6x basal width; median longitudinal carina present, divided in

middle 0.5x pronotal length before posterior margin, median fovea

present and 0.5x pronotal length, closing at posterior margin; pronotal

stemmata absent; anterior and lateral discal margins swollen; anterior

margin convexly arcuate, anterior angles obtuse; lateral margin

complete, foliaceous, erect, straight; posterior margin strongly

bisinuate; posterior angles acute, expanded laterally O.fix width at

elytral humerus; disk flat, but anterior 0.3x inflated allowing

reception of head, setation semi-decumbent; hypomeron deeply concave,

height 0.7x width, margins not swollen, hypomeral stemmata absent;

prosternal bridge Y-shaped, lateral arms obtusely separated, length 0.5x

width, intercoxal process not carinate. Scutellum pentagonal, distally

quadrate and nearly square, lateral margins parallel, apex deeply

emarginate, emargination 0.3x scutellar length, apices acute and

divergent. Mesospiracle prominent, peritreme exposed, orifice slightly

compressed. Mesepisternum elongate, posterior margin convexly arcuate,

nearly straight. Mesepimeron lobate; width lx and height 0.8x mesepisternum; posterior margin strongly reflexed, evenly rounded.

Mesosternum transverse, length 0.4x width, posterior margin convexly 192 arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra slightly explanate-1igulate, slightly dehiscent from middle, slightly reconvergent near apex, expanded Q.7x and narrowed distally to apex, apices acute; length 9x pronotal length; tricostate, costa 2 distinct basal 0.6x elytral length, costae 4 and 8 distinct to elytral apex; disk irregularly scabrous-punctate; setation semi“decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly arcuate posteriorly. Metepimeron with ventral margin slightly arcuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.5x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 1.3x coxal length, metatrochanters 1.2x metacoxal length, femoral insertions truncate and not offset except metatrochanters slightly oblique and slightly offset; femora compressed, clavate, short, 2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly arcuate; tibial spines small and similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues s i mp 1e .

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural 193

membrane, located submedially from apical margin of tagma. Aedeagus

with parameres; paramere length Q.9x median lobe, differentiated from

median lobe apical 0.5x length of lobe, sinuate apically; basal piece

dorsoventrally compressed, length almost 0.3x length of median lobe

(Fig. 21 a-c>; median lobe tubular, unmodified, membranous ventrally at

apex; internal sac not eversible; flagellum absent.

DESCRIPTION Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The similarity of coloration of this species and F. pecki demonstrates a major problem in lycid taxonomy. A significant portion of lycid descriptions in the primary literature is largely a discussion of coloration. Often, these are insufficient to determine what species

is at hand, because of putative mimicry within the family. This is especially true in the Neotropical fauna.

MATERIAL EXAMINED. ECUADOR, Provincia Napo, Limoncocha, 250 m . , 15-28

June 1976, S, & J. Peck (1— CNCI, holotype).

ETYMOLOGY. The species is named in reference to its appearance. 194

Flabellocaenia iolvi NEW SPECIES

Fig. 23 a-e

TYPE: MIZA, a card mounted male in good condition with the following

data: Cwhite printed labels] "VENEZUELA T. F. \\ Amazonas Dept. \\ Rio

Negro;" "Rio Baria \\ 140 m \\ 0 ’55’N, 66'10'W" "L. J. Joly W A.

Chacon \\ 4-11 II 84;" Cgreen label] "Venezuela Inst. \\ Zool. Agricola

\\ Fac. Agronomia \\ U. C. V. Maracay."

DIAGNOSIS. This species can be separated from other known members of

the genus by a combination of characters: its unicolorous, dark coloration; the epistomal margin is deeply and narrowly emarginate; the distance between the eye and antennal socket is 3X the interantennal

socket distance; and the genitalia, which may be most reliable until a

larger series is seen.

DISTRIBUTION. Known only from the type locality near Cerro de la

Neblina, Venezuela.

DESCRIPTION (Male). Elongate, but elytra expanded distally 0.7X their

length and then narrowing to apex; length 4.4 mm, width at humerus 0.9 mm. Blackish brown except trochanters, antennomeres 2 and 3 and tips of rami yellow brown.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsaliy. Eyes hemispherical, projecting laterally when viewed dorsaliy; hind margin shallowly emarginate; radial origin at head margin when dorsaliy viewed.

Antennal prominence distinct from vertex, shelf-like, slightly gibbose;

length 0.3X width, inserted between middle of eyes; rectangular

dorsaliy, slightly produced apically, impressed dorsoraedially; ventrally

trianguliform, slightly concave, sides nearly straight, widest near

base. Antennal sockets subrectangular, separated by 0.1X width of

epistomal margin and 3X distance between eye and antennal socket.

Antennifer insertion elevated medially, not appearing basomedial.

Interocular distance 3.2X width eye dorsaliy. Vertex transversely

flat, setae decumbent; depression of tentorial maculae broad and

shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely flat, distinct from

lower surface of antennal prominence by strong transverse impression below antennal sockets, transversely narrow, height 0.3X width.

Epistomal margin deeply and narrowly emarginate, width 0.6X frontal distance between eyes. Labrum transverse, rectangular, length 0.5X width, apical margin truncate. Maxillary palps 4 segmented, palpomeres

1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5X palpomere 3, acuminate.

Labial palps absent. Gula transverse, linear; length 0.1X width apical margin; inserted slightly anteriorad postgenal posterior margin.

Antennae flabellate, length 0.7X body length; rami between 5-10X axial

length and their compression perpendicular to axial compression; setation sparse, distributed along margins, semi-decumbent on all antennomeres, sealiform setae present; scape obconic, width apically 196

subequal length; pedicel cylindrical, short, transverse, not compressed,

length IX length of antennomere 3; antennomere 3 slightly compressed,

adnate to antennomere 4 basally, length 0.5X antennomere 4; antennomere

4 flabellate with ramal insertions medial, more strongly compressed;

antennomeres 5-10 compressed, extent of compression decremental to apex,

flabellate with ramal insertions apical; antennomere 11 without axis

distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.6X basal width, anterior

width 0.7X basal width; median longitudinal carina present, divided in

middle 0.8X pronotal length before posterior margin, median fovea

present and 0.8X pronotal length, not closed at posterior margin;

pronotal stemmata absent; all discal margins swollen; anterior margin

slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, slightly concavely arcuate; posterior margin strongly bisinuate; posterior angles acute, expanded laterally

0.8X width at elytral humerus; disk transversely arcuate slightly with anterior 0.3X slightly inflated allowing reception of head, lateral margin deeply impressed, setation sparse and decumbent; hypomeron concave, height 0.6X width, anterior margins swollen, hypomeral stemmata absent; prosternal bridge T-shaped, broadly emarginate, length 0.5X width, intercoxal process not carinate. Scutellum pentagonal, distally trianguliform, longitudinally impressed, lateral margins divergent, apex emarginate, emargination 0.3X scutellar length, apices rounded and divergent. Mesospiracle prominent, peritreme exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin slightly convexly arcuate. Mesepimeron lobate; width equal and height 0.8X mesepisternum; posterior margin strongly reflexed, evenly rounded. Mesosternum

transverse, length 0.5X width, posterior margin convexly arcuate.

Mesocoxal separation 1.3X mesocoxal diameter at base. Elytra slightly explanate-1igulate, slightly dehiscent from middle, slightly reconvergent near apex, expanded to 0.7X and narrowed distally to apex, apices acute; length 8.6X pronotal length; tricostate, costa 2 distinct basal 0.6X elytral length, costae 4 and 8 distinct to near elytral apex, humeral costa convergent to near 4 apically; disk irregularly scabrous- punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly arcuate posteriorly. Metepimeron with ventral margin slightly arcuate.

Metasternum completely divided medially by longitudinal sulcus. Legs elongate; setation moderately dense, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.5X coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 1.IX coxal length, metatrochanters 0.9X metacoxal length, all femoral insertions truncate and not offset; femora compressed, clavate, short, 2X trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly arcuate; tibial spines small and similar; tarsi very short, tarsomeres without laterally expanded plantar pads; covered with semi- decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing 198

uniformly posteriorad; sternite 7 entire. Spiracles in pleural

membrane, located submedially from apical margin of tagma. Aedeagus

with parameres; paramere length subequal median lobe, differentiated

from median lobe entire length of lobe, apices deflexed dorsaliy,

acuminate; basal piece dorsoventrally compressed, length almost 0.7X

length of median lobe (Fig. 23 a-c); median lobe broadly tubular,

deflexed apically, membranous ventrally at apex; internal sac not

eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The distinctive frons of the type and only specimen may be an artifact of crushing in preservation. Therefore, this character

state is not used in the diagnosis.

MATERIAL EXAMINED. VENEZUELA; Territoria Federal Amazonas, Departmento

Rio Negro, Rio Baria, 140 m . , 0 * 5 5 ^ 60”10*W, 4-11 February 1984, L. J.

Joly and A. Chacon (1-MIZA, holotype).

ETYMOLOGY. This species is named in honor of Dr. Luis J. Joly-Tinoco, the curator of the beetle collection at Maracay, who is among the most knowledgable general coleopterists I have met. This species was discovered on his memorable collecting trip via boat to Cerro de la

Neblina. 199

Flabellocaenia leticia NEW SPECIES

Fig. 22 a-c

TYPE: HAHC, a male with the following label data: [white printed

label] "LETICIA, Araa2onas \\ Colombia 700 ft. \\ Feb. 24-28" [white

printed label] "S. Peck pan \\ trans for."

DIAGNOSIS. This species may be separated from congenerics by the

combination of: elytra blackish brown with apex and base briefly yellow

and the venter is mostly brown; the basal yellow coloration does not

extend caudally along the costae beyond the elytral floor; the brown

legs are yellow at their segmental insertions.

DISTRIBUTION. This species is known only from the type specimen which

was collected near Leticia, Colombia.

DESCRIPTION (male). Elongate, but elytra expanded distally 0.7x their

length and then narrowing to apex; length 4.7 mm, width at humerus 0.9 mm. Blackish brown; except apices of rami briefly and elytra briefly, venter of pro- and mesothorax, and annulations briefly at femoral and tibial insertions yellow.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk dull; setae sparse, semi-decumbent. Genal margin short, length less

than eye length, uniformly narrowing when viewed dorsaliy. Eyes hemispherical, projecting laterally when viewed dorsaliy; hind margin shallowly emarginate, nearly straight; radial origin at head margin when dorsaliy viewed. Antennal prominence slightly distinct from vertex,

shelf-like; length 0.5x width, inserted between middle of eyes; pentagonal dorsaliy, shallowly impressed dorsomedially; ventrally

trianguliform, concave, sides straight, widest near base. Antennal

sockets subrectangular, separated by 0.lx width of epistomal margin and equidistant to distance between eye and antennal socket. Antennifer

insertion elevated medially, not appearing basomedial. Interocular distance 2x width eye dorsaliy. Vertex transversely flat, setae semi- decumbent; depression of tentorial maculae broad and shallow, from base of antennal prominence to posterior eye margin. Tentorial maculae separate, shallow, distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transversely narrow, height 0.3x width. Epistomal margin deeply and narrowly emarginate, width Q.3x frontal distance between eyes. Labrum not seen. Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, inserted, acuminate.

Labial palps absent. Gula transversely rectangular; length approximately 0.2x width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabellate, length 0.6x body length; rami between 6-8x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, decumbent on all antennomeres, sealiform setae present; scape obconic, width apically subequal length; pedicel cylindrical, short, transverse, not compressed, length 2x length of antennomere 3; antennomere 3 slightly compressed, adnate to antennomere 4 basally, length subequal antennomere 4; 201 antennomere U flabellate with ramal insertions medial, more strongly compressed; antennomeres 5-10 compressed, extent of compression decremental to apex, flabellate with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.6x basal width, anterior width 0,6x basal width; median longitudinal carina present and divided

in middle 0.5x pronotal length before posterior margin, median fovea

0.5x pronotal length, closing at posterior margin; pronotal stemmata absent; all discal margins swollen except posterior incomplete; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, slightly explanate, slightly concavely arcuate; posterior margin strongly bisinuate; posterior angles acute, expanded laterally Q.8x width at elytral humerus; disk slightly transversely arcuate with anterior 0.3x inflated allowing reception of head, setation decumbent; hypomeron concave, height 0.5x width, all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms broadly and obtusely separated, length 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate and nearly square, lateral margins divergent, apex deeply emarginate, emargination

0.5x scutellar length, apices acute and divergent. Mesospiracle prominent, peritreme exposed, orifice not compressed. Mesepisternum elongate, posterior margin slightly convexly arcuate. Mesepioeron lobate; width 0.5x and height 0.8x mesepisternum; posterior margin moderately reflexed, evenly rounded. Mesosternum length lx width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra slightly explanate-ligulate, slightly dehiscent 0.4x elytral length, strongly reconvergent at apex, expanded

to 0.7x elytral length and narrowed distally to apex, apices narrowly

acute; length 9x pronotal length; tricostate, costa 2 distinct basal

0.4x elytral length, costae 4 and 8 distinct to near elytral apex, costa

8 sinuate and converging to near 4 at apex; disk irregularly scabrous-

punctate; setation semi-decumbent, moderately dense, uniformly

distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight.

Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.8x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 1.3x coxal length, metatrochanters lx to metacoxal length, all femoral insertions truncate and not offset; femora compressed, clavate, short, 1.8x trochanteral

length, annulated basally; tibiae strongly compressed, internal margin sinuate; tibial spines small and similar; tarsi very short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 produced and bilobate; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 emarginate. Spiracles in pleural membrane, located subroedially from apical margin of tagma.

Aedeagus with parameres; paramere length 0.9x median lobe, 203

differentiated from median lobe apical 0.3x length of lobe, laterally

divergent laterally apically; basal piece dorsoventrally compressed,

length almost 0.7x length of median lobe (Fig. 22 a-c); median lobe

tubular, unmodified and not deflexed ventrally; membranous apically;

internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The type was captured in a pan trap. These traps have captured a number of specimens of other Leptolycini.

MATERIAL EXAMINED. COLUMBIA, Amazonas, Leticia, ca. 200 m, 24-28 Feb.,

S. Peck (l-HAHC, holotype).

ETYMOLOGY. This species is named for the town in which it was captured. 204

F 1abe11ocaenia pecki NEW SPECIES

Fig. 24 a-d

TYPE: CNCI, a male with the following label data: Cwhite printed label! "ECU. Napo Prov. \\ Limoncocha. 250 \\ m. 15-28-VI— 78 \\ S. & J.

Peck."

DIAGNOSIS. Separation from the simlarly marked F. eleeantulus is most easily seen in the rami being 4x versus 5x axial length basally, the elytral costae 4 and 8 not reaching apex of the elytra versus reaching it, the epistomal margin is narrowly versus broadly emarginate, and the median lobe of the aedeagus is membranous along the entire venter instead of only at the apex.

DISTRIBUTION. This species is known only from the type specimen which was collected near Limoncocha, Equador.

DESCRIPTION (Male). Small, slender, elongate, but elytra expanded distally 0.7x their length and then narrowing to apex; length 5.0 mm, width at humerus 0.9 mm. Blackish brown except yellow maculation; elytra at humeral angles, extending along lateral margin to 0.5x elytral length, apical 0.3x elytral length yellow; antennal axis except distal 3 antennomeres, apices of rami antennomeres 6-11 yellow; mouthparts, coxae, femora and basal 0.3x tibia yellow except femora testaceous band encircling middle.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk dull; setae sparse, semi-decumbent. Genal margin short, length less

than eye length, uniformly narrowing when viewed dorsaliy. Eyes

hemispherical, projecting laterally when viewed dorsaliy; hind margin

shallowly emarginate, nearly straight; radial origin at head margin when

dorsaliy viewed. Antennal prominence slightly distinct from vertex,

shelf-like; length 0.3x width, inserted between middle of eyes;

pentagonal dorsaliy, not impressed dorsomedially; ventrally

trianguliform, concave, impressed longitudinally, widest near base,

sides nearly straight. Antennal sockets subrectanguiar, separated by

0.lx width of epistomal margin and equidistant to distance between eye

and antennal socket. Antennifer insertion elevated medially, not

appearing basomedial. Interocular distance 2.3x width eye dorsaliy.

Vertex transversely flat, setae semi-decumbent; depression of tentorial

maculae broad and shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate,

shallow, distinct; at base of antennal prominence. Frons transversely concave, distinct from lower surface of antennal prominence,

transversely narrow, height 0.3x width. Epistomal margin deeply and narrowly emarginate, width 0.4x frontal distance between eyes. Labrum

transverse, rectangular, length 0.5x width, apical margin truncate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base

less than 0.5x palpomere 3, inserted, acuminate. Labial palps absent.

Gula transverse, linear; length O.lx width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabellate, length

0.6x body length; rami between 3-4x axial length and their compression 206

perpendicular to axial compression; setation sparse, distributed along

margins, semi-decumbent on all antennomeres, sealiform setae present;

scape obconic, width apically subequal length; pedicel cylindrical,

short, transverse, not compressed, length 2x length of antennomere 3;

antennomere 3 slightly compressed, adnate to antennomere 4 basally,

length 0.8x antennomere 4; antennomere 4 flabellate with ramal

insertions medial, more strongly compressed; antennomeres 5-10

compressed, extent of compression decremental to apex, flabellate with

ramal insertions apical; antennomere 11 without axis distinguishable

from ramus.

Thorax. Pronotum trapezoidal, length 0.6x basal width, anterior

width 0.6x basal width; median longitudinal carina present and divided

in middle 0.3x pronotal length before posterior margin, median fovea

0.5x pronotal length, closing at posterior margin; pronotal stemmata

absent; all discal margins swollen; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, slightly concavely arcuate longitudinally; posterior margin

strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, but anterior 0.3x inflated allowing reception of head, setation decumbent; hypomeron concave, height 0.6x width, anterior and posterior margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms narrowly and acutely separated, length 0.5x width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate and nearly square, lateral margins parallel, apex deeply emarginate, emargination 0.3x scutellar

length, apices acute and divergent. Mesospiracle prominent, peritreme 207 exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin slightly convexly arcuate. Mesepimeron lobate; width equal and height 0.8x mesepisternum; posterior margin strongly reflexed, evenly rounded. Mesosternum length lx width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra slightly explanate-1igulate, slightly dehiscent from middle, slightly reconvergent at apex, expanded to 0.7x elytral length and narrowed distaliy to apex, apices acute; length lOx pronotal length; tricostate, costa 2 distinct basal 0.3x elytral length, costae 4 and 8 distinct 0.8 elytral length; disk irregularly scabrous-punctate; setation semi- decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight, Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, semi-decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.6x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro - and mesotrochanters l.lx coxal length, metatrochanters 1.2x metacoxal length, all femoral insertions truncate and not offset; femora compressed, clavate, short, 2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight; tibial spines small and similar; tarsi short, tarsoraeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 208 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; paramere length 0.9x median lobe, differentiated from median lobe apical 0.3x length of lobe, sinuate apically; basal piece dorsoventrally compressed, length almost 0.4x length of median lobe

(Fig. 24 a-c); median lobe tubular, unmodified, membranous ventrally most its length; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Iramatures). Unknown,

DISCUSSION. The type was captured in a flight intercept trap.

MATERIAL EXAMINED. ECUADOR, Napo Province, Limoncocha, 15 - 28 June

1978, S. and J. Peck (1 - CNCI, holotype).

ETYMOLOGY. This species is named in honor of Drs. Stewart Peck and

Ludmilla Kukalova-Peck who have substantially aided my study of the

South American Lycidae by providing material for study. 209

Flabellocaenia rimae NEW SPECIES

Fig. 25 a~c

TYPE: NMNH, a male with the following label data: Cwhite printed label] "VENEZUELA, T. F. Amaz. \\ Cerro de la Neblina \\ 1 km. SE

Basecamp \\ O ’SG'N, 66',10'W \\ 140 m. 3 Feb. 1985;" Cwhite printed label] "On low foliage \\ rainforest trail \\ W. E. Steiner \\ col lector."

DIAGNOSIS. F. riroae can be separated from other known congenerics by its unicolorous elytra, the truncate epistomal margin, the equidistance between the antenna1 fovea and the eye and antenna1 fovea, and the genitalia. Its appearance is very similar to F. iolvi. but the genitalia will easily separate the species.

DISTRIBUTION. This species is known only from the type locality on the flanks of Cerro Neblina, Venezuela. As collecting effort has been concentrated over January and February over a two year period, little is known about its temporal distribution.

DESCRIPTION (Males). Small, slender, elongate, but elytra expanded distally 0.7x their length and then narrowing to apex; length 4.1-4.7 mm, width at humerus 0.9-1.1 mm. Blackish brown; except elytral insertions, trochanters, apical abdominal segment and apices of all antennal rami briefly testaceous.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, semi-decumbent. Genal margin short, length less

than eye length, uniformly narrowing when viewed dorsally. Eyes

hemispherical, projecting laterally when viewed dorsally; hind margin

shallowly eraarginate, nearly straight; radial origin at head margin when

dorsally viewed. Antennal prominence slightly distinct from vertex,

shelf-like; length 0.3x width, inserted between middle of eyes;

pentagonal dorsally, narrowly impressed dorsomedially; ventrally

trianguliform, flat to slightly concave, widest near base, sides nearly

straight. Antennal sockets subrectangular, separated by O.lx width of

epistomal margin and equidistant to distance between eye and antennal

socket. Antennifer insertion elevated medially, not appearing

basomedial. Interocular distance 4x width eye dorsally. Vertex

transversely flat, setae semi-decumbent; depression of tentorial maculae

broad and shallow, poorly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae

separate, distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transversely

narrow, height Q.3x width. Epistomal margin truncate, width 0.5x

frontal distance between eyes. Labrum transverse, rectangular, length

0.3x width, apical margin truncate. Maxillary palps minute, 4

segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length;

length palpomere 4 nearly equal 3, diameter at base less than 0.5x

palpomere 3, acuminate. Labial palps 1 segmented, digitiform. Gula

transverse, linear; length 0.2x width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabel late, 0.6x body length; rami between 7-1lx axial length and their compression perpendicular to 211

axial compression; setation sparse, distributed along margins, semi-

decumbent on all antennomeres, scaliform setae present; scape obconic,

width apically lx length; pedicel cylindrical, short, transverse, not

compressed, length lx length of antennomere 3; antennomere 3 slightly

compressed, adnate to antennomere 4 basally, length lx antennomere 4;

antennomere 4 flabel late with ramal insertions basal, more strongly

compressed; antennomeres 5-10 compressed, extent of compression uniform

to apex, flabel late with ramal insertions apical; antennomere 11 without

axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior

width 0.7x basal width; median longitudinal carina present, divided in

middle 0.7x pronotal length before posterior margin, median fovea

present and lx pronotal length, closing at posterior margin; pronotal

stemmata absent; all discal margins swollen; anterior margin straight,

anterior angles obtuse; lateral margin complete, foliaceous, erect,

slightly concavely arcuate longitudinally; posterior margin bisinuate;

posterior angles acute, expanded laterally 0.8x width at elytral

humerus; disk flat, but anterior 0.5x inflated allowing reception of

head, setation semi-decumbent; hypomeron concave, height 0.7x width,

anterior and ventral margins swollen, hypomeral stemmata absent;

prosterna1 bridge Y—shaped, anterior emargination shallow, length 0.6x

width, intercoxal process not carinate. Scutellum pentagonal, distally

trianguliform, lateral margins expanded apically, apex shallowly

emarginate, emargination O.lx scutellar length, apices rounded and divergent. Mesospiracle prominent, peritreme exposed, orifice slightly

compressed. Mesepisternum elongate, posterior margin straight. Mesepimeron lobate; width lx and height 0.8x mesepisternum; posterior margin strongly reflexed, evenly rounded. Mesosternum length 0.5x width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra slightly expanded-1igulate, slightly dehiscent from 0.3x elytral length, slightly reconvergent near apex, expanded to 0.7x elytral length and narrowed distally to apex, apices acute; length 7-8x pronotal length; tricostate, costa 2 distinct basal 0.5x elytral length, costae 4 and 8 distinct 0.9x elytral length; disk irregularly scabrous-punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin decreasingly arcuate posteriorly. Metepimeron with ventral margin nearly straight. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.6x coxal length, roetacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters l.lx coxal length, metatrochanters 1.4x metacoxal length, all femoral insertions truncate and not offset; femora compressed, clavate, short, 1.9x trochanteral length, annulated basally; tibiae strongly compressed, internal margin sinuate; tibial spines small and similar; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing 213 uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus with parameres; pararaere length 0.9x median lobe, differentiated from median lobe apical 0,3x length of lobe, sinuate apically; basal piece dorsoventrally compressed, length almost 0.4x length of median lobe

(Fig. 25 a-c); median lobe tubular, unmodified, membranous ventrally most its length; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The type series were captured in flight intercept traps

(10), sweeping (1), and at blacklight (1).

MATERIAL EXAMINED. VENEZUELA, Territorio Federal Ama2onica, Cerro de la Neblina O'SO'N, 66°9'44"W, basecamp at 140 m, 13-20 February 1984, D.

Davis and T. McCabe (1-MIZA, 1-NMNH); Same locality, 66'10’W, basecamp at 140 m, 24-31 January 1985, P. J. and P. M. Spangler, P. Faitoute, W.

Steiner (1-RSMC); Same locality, 26 January 1985, P. J. and P. M.

Spangler, P. Faitoute, W. Steiner (1-RSMC); Same locality, 10-20

February 1985, P. J. and P. M. Spangler, P. Faitoute, W. E. Steiner (2-

MIZA, 2-NMNH); Same locality, 21-24 February 1985, P. J. and P. M.

Spangler, P. Faitoute, W, Steiner (1-NMNH, 1-RSMC); Same locality, 21-28

February 1985, P. J. and P. M. Spangler, P. Faitoute, W. Steiner (1-

MIZA); Same locality, 1 km, SE Basecamp at 140 m, 3 February 1985, W, E. 214

Steiner <1-NMNH, holotype).

ETYMOLOGY. This species is named for the allegorical Rima of W. H.

Hudson's novel, Green Mansions. 215

Leptolvcus Leng and Mutchler

Lepfcolvcus Leng and Mutchler, 1922:430. Kleine, 1933:18.

Blackwelder, 1945:343. Crowson, 1972:48.

TYPE. Leptolvcus heterocornis Leng and Mutchler, by original designation, AMNH.

DIAGNOSIS. These very small and elongate lycids can be recognized in the male by their porrect tubular heads with antennal insertions on a gibbous prominence, the subantennal suclus meeting the eye margin instead of directly meeting the epistomal margin, the elongate and bristling antennal setae, and the dehiscent elytra. Known females are paedomorphic and can be differentiated from larvae by their lack of mouthparts which are represented only by the maxillae as protuberant undifferentiated lobes. Larvae may be distinguished from other known lycid larvae by the presence of a subantennal sulcus in addition to the genal sulcus (ecdysial line).

DISTRIBUTION, This genus is known from Hispaniola, Puerto Rico and St.

John Is. of the U. S. Virgin Islands. Although it has not been found in Cuba, it is expected to occur there.

DESCRIPTION (Male). Small, slender, elongate, length 2.1-4.2 mm., width 0.5-1.0 mm. at humerus. Black; except yellow and white maculation. Head. Elongate, widest at eyes, then narrowing to posterior

margin, punctate to punctate-areolate; setae sparse to dense and

occluding the disk, semi-decumbent. Genal margin elongate, length

greater than eye length, narrowing posteriorly when viewed dorsally.

Eyes hemispherical, projecting anteriolaterally when viewed dorsally;

hind margin shallowly emarginate, to nearly straight; radial origin

inside head margin 0.3-05x radius when dorsally viewed. Antennal

prominence distinct from vertex, porrect to shelf-like and gibbose dorsally; length 0.8-1.3x width, inserted between middle of eyes;

dorsally pentagonal to quadrate with anterior margin slightly produced, variously sulcate dorsomedially or not, sometimes appearing bigibbose;

ventrally trianguliform or a thin transverse ridge, concave or not.

Antennal sockets oval to lenticular to rectangular, separated by 0.1-

0.2x width of epistomal margin and 0.3-lx distance between eye and antennal socket. Antennifer elevated medially, appearing basomedial,

inserted at apex or below. Interocular distance 1.7-3x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae semi-decumbent; depression of tentorial maculae shallow to deep, broadly defined, from base of antennal prominence to eye margin or behind eye at midvertex, more distinct anteriorly. Tentorial maculae separate, indistinct to deeply distinct, at base of antennal prominence. Frons transversely convex, distinct to indistinct from lower surface of antennal prominence, height 0.3-lx width. Epistomal margin truncate to narrowly emarginate, width 0.6-0,8x frontal distance between eyes.

Labrum transverse, length 0.5-lx width, apical margin rounded to truncate. Maxillary palps 4 segmented, palpomeres 1-4 cylindrical, length 1-3 equal, diameters decrements1; palpomere 4 acuminate. Labial

palps 1 segmented and digit iform to absent. Gula elongate to

transverse, pentagonal to trapezoidal to almost linear; width slightly

greater basally than apically; length 2-5x width apical margin; inserted

anteriorad posterior margin of postgenae. Antennae filiform, 0.7-1.3x

body length; basal setation sparse, uniformly distributed, decumbent on

antennomeres 1-3; distal setation on antennomeres 4-11 distributed along

margins, elongate, semi-erect and bristling; scaliform setae present along margins; scape obconic, width apically 0.5-0.7x length; pedicel cylindrical, short, transverse, not compressed, length l-2x antennomere

3; antennomere 3 adnate with base antennomere 4 Cor not]; antennomeres

4-10 elongate filiform to subserrate, strongly compressed, lateral margins irregular to regular; length antennomere 3 7-20x antennomere 4; antennomeres 9-11 not expanded laterally beyond width antennomere 4; antennomere 11 fusiform, compressed, length 1.3x antennomere 3.

Thorax. Pronotum trapezoidal, elongate to transverse, length

0.4-2x basal width, anterior width nearly 0.6-0.7x basal width; median

longitudinal carina not visible, median fovea not visible to obscure anteriorly and well defined posteriorly, closed at posterior margin; pronotal stemmata absent; all discal margins variously swollen; anterior margin convexly arcuate to nearly straight, anterior angles obtuse;

lateral margin complete to incomplete anteriorly and carinate posteriorly, foliaceous or not, erect or reflexed, straight to concavely arcuate longitudinally; posterior margin weakly to strongly bisinuate; posterior angles acute, expanded laterally 0.7-0.9x width at elytral humerus; disk transversely arcuate, ocassionally rugose, deeply impressed laterally or not, elevated near posterior margin Cor notD,

discal setae decumbent to semi-decumbent, sparse to dense and occluding

disk; hypomeron shallowly to deeply concave, height 0.5-0.6x length;

anterior, ventral and lateral margins swollen, hypomeral stemmata

absent; prosternal bridge trianguliform, anterior margin truncate to

slightly emarginate, length 0.5-O.Qx width, intercoxal process nearly

defined or not, not carinate. Scutellum pentagonal, apically quadrate

and nearly square to trapezoidal; lateral margins parallel or divergent

apically; apex slightly to deeply emarginate, emargination to 0.5x

scutellar length; apices divergent or not, acute; setation sparse to

dense and occluding disk, decumbent. Mesospiracle obscure to

prominent, peritreme exposed or not, oriface compressed or not.

Mesepisternum elongate, posterior margin straight. Mesepimeron lobate

to trianguliform, width 0.3-05x and height 0.8-lx mesepisternum;

posterior not to little reflexed, straight to evenly arcuate.

Mesosternum transverse to elongate, posterior margin convexly arcuate to

truncate. Mesocoxal separation lx mesocoxal diameter at base. Elytra

ligulate, strongly dehiscent variously from scutellum to elytral middle,

divergent to apices, uniformly narrowing most elytral length, rate of

lateral convergence increasing near apex or not, slightly expanded and

spatulate apically or not, apex acuminate; length 5-9x pronotal length;

bicostate, costa 4 distinct basal 0.3-0.5x elytral length, humeral costa more distinct 0.5-lx elytral length; disk scabrous-punctate to to

strongly punctate-reticulate to irregularly reticulate; setation sparse, uniformly distributed, semi-decumbent. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral 219

margin arcuate or straight. Metasternum completely divided medially by

longitudinal sulcus. Legs thin and elongate, setation sparse,

decumbent, uniformly distributed; coxae conical, trochanteral socket

0.3-0.4x coxal length; trochanters elongate, length pro- and mesotrochanters 1-1.2x coxal length, metatrochanters 0.8-1.3x metacoxal

length, all femoral insertions oblique to truncate and offset to

slightly offset; femora strongly compressed, straight, 2-5x trochanteral

length, annulated basally; tibiae strongly compressed, internal margin

sinuate to straight; tibial spines similar, short; tarsi elongate,

tarsomeres without laterally expanded plantar pads; covered with semi- decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorad; sternite 7 broadly emarginate to entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece absent to large, dorsoventrally compressed, expanded posteriorly or not, length about 0.4-lx length of median lobe; median lobe tubular, acuminate, punctate or not, ventrally and medially to subapically membranous 0.3-

0.5x length; internal sac not eversible; flagellum absent.

DESCRIPTION (Female). See description of L, heterocornis L. & M.

DESCRIPTION (Immatures). See description of L. heterocornis L. & M, DISCUSSION. Like other Leptolycini, the biology of these species is

largely unknown. The best known apecies, is only known from label

data accompanying preserved material. From locality data, it appears

that larvae are litter dwellers and can withstand considerable xeric

conditions much as in Lvcus sensu lato. but uncharacteristic for most

Lycidae. The females of at least L. heterocornis L. & M. are

paedomorphic and the females of the other species are undoubtedly

somewhat effected by the phenomenon and are probably all flightless.

ETYMOLOGY. Lepto (Greek), slender and 1vcus (Greek), wolf and the name

of the type genus of the family. 221

KEY TO SPECIES OF Leptolvcus (ADULT MALES >

1. Brown species with slightly indicated yellow humeralangles;

antennal prominence expanded apically; pronotum strongly

transverse and trapezoidal; Hispaniola ......

...... Leptolvcus brunneus n. sp. (p. 227)

1'. Black and yellow species, other characters vary ...... 2

2. (I*) Pronotum yellow, not black ...... 3

2*. (lr) Pronotum black, not yellow ...... 5

3. (2) Lateral pronotal margins distinct, foliaceous and erect;

length pronotum 0.4 basal width; prothorax pale yellow;

erect and bristling antennal setae moderately dense;

Hispaniola ...... Leptolvcus adiaphorus n. sp. (p. 223)

3’.(2) Lateral pronotal margins indistinct anteriorly ; prothorax

golden yellow; erect and bristling antennal setae sparse ...

...... 4

4. (3' ) Length pronotum O.Ox basal width; venter of head and

metathorax black; Puerto Rico ......

...... Leptolvcus flavicollis L. & M. (p. 242) Length pronotum 0.6x basal width; venter of head and

metathorax golden yellow; Hispaniola ......

...... Leptolvcus dominicensis n. sp. (p. 232)

Antennae serrate with antennomeral margins regular; erect

and bristling antennal setae absent; lateral margins of

pronotum distinct, but not foliaceous; Puerto Rico ......

...... Leptolvcus effeminatus n. sp. (p. 237)

Antennae irregularly serrate with margins irregular; erect and bristling antennal setae present; lateral margins of pronotum indistinct anteriorly; Puerto Rico and Virgin

Islands ...... Leptolvcus heterocornis L. & M. (p. 247) 223

Leptolvcus adiaphorus NEW SPECIES

Fig, 26 a-f

TYPE: NMNH, a male with the following data: Cwhite printed labelD

"DOMINIC, REP.: Prov. La Vega \\ 12 km NE Jarabacoa, 550 m . W 01-07 SEP

1988, flight inter.\\ trap. M. A. Ivie, T. E. Phillips, and K. A.

Johnson coirs.”

DIAGNOSIS. This species canm be recognized by the combination of: black coloration except the pale yellow pronotum and last abdominal segment; the lateral pronotal margins are distinct, foliaceous and erect; the pronotal length is 0.4x its basal width; and the erect and bristling antennal setae are moderately dense.

DISTRIBUTION. Known only from the type locality in the Dominican

Republic on the island of Hispaniola.

DESCRIPTION (Males). Small, slender, elongate, length 2.8-4,1 mm., width

0.7-1.0 mm. at humerus. Black; except prothorax and apical abdominal segment pale yellow; meso- and metathorax testaceous; antennomere 11 white.

Head. Elongate, widest at eyes, then narrowing to posterior margin, densely punctate-aereolate; setae sparse, semi-decumbent.

Genal margin elongate, length greater than eye length, narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin 0.3 radius when dorsally viewed. Antennal prominence distinct from vertex, shelf-like, gibbose dorsally; length equal to width, inserted between middle of eyes; quadrate dorsally, anterior margin slightly produced, not sulcate dorsomedially; ventrally a thin transverse ridge. Antennal sockets rectangular, separated by O.lx width of epistomal margin and equidistant to distance between eye and antennal socket. Antennifer elevated medially, appearing basomedial. Interocular distance 1.7x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae semi-decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to behind eye at midvertex, much more distinct anteriorly. Tentorial maculae separate, deep, at base of antennal prominence. Frons transversely convex, indistinct from lower surface of antennal prominence, height equal to width.

Epistomal margin truncate, width 0.8x frontal distance between eyes.

Labrum transverse, length 0.5x width, apical margin rounded. Maxillary palps 4 segmented, palpomeres 1-4 cylindrical, length 1-3 equal, diameters decremental; palpomere 4 acuminate. Labial palps absent.

Gula elongate, almost linear; length 0.2x width apical margin; inserted slightly anteriorad posterior margin of postgenae. Antennae filiform,

0.8x body length; basal setation sparse, uniformly distributed, decumbent on antennomeres 1-3; distal setation on antennomeres 4-11 distributed along margins, elongate, semi-erect and bristling; scaliform setae present along margins; scape obconic, width apically 0.5x length; pedicel cylindrical, short, transverse, not compressed, length equal to antennomere 3; antennomere 3 adnate with base antennomere 4; 225 antennomeres 4-10 elongate filiform, strongly compressed, lateral margins irregular; length antennomere 3 8x antennomere 4; antennomeres

9-11 not expanded laterally beyond width antennomere 4; antennomere 11 fusiform, compressed.

Thorax. Pronotum trapezoidal, transverse, length 0.4x basal width, anterior width nearly 0.6x basal width; median longitudinal carina not visible, median fovea obscure anteriorly, well defined posteriorly; pronotal stemmata absent; all discal margins swollen; anterior margin convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, straight longitudinally; posterior margin weakly bisinuate; posterior angles acute, expanded laterally 0.7x width at elytral humerus; disk transversely arcuate, rugose, deeply impressed laterally, discal setae semi-decumbent, sparse; hypomeron shallowly concave, almost flat, height 0.6x length; anterior, ventral and lateral margins swollen, hypomeral stemmata absent; prosternai bridge trianguliform, anterior margin truncate to slightly emarginate, length

0.5x width, intercoxal process nearly defined, not carinate. Scutellum pentagonal, apically quadrate and nearly square; lateral margins parallel; apex deeply emarginate, emargination 0.5x scutellar length; apices not divergent, acute; setation sparse, decumbent. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight. Mesepimeron lobate, width 0.3x and height 0.8x mesepisternum; posterior little reflexed, evenly arcuate. Mesosternum transverse, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, strongly dehiscent behind elytral middle, divergent to apices; length 9x pronotal length; 226

bicostate, costa 4 distinct basal 0.3x elytral length, humeral costa

more distinct 0.7x elytral length; disk scabrous-punctate to irregularly

reticulate; setation sparse, uniformly distributed, semi-decumbent.

Metepisternum with ventral margin increasingly arcuate posteriorly.

Metepimeron with ventral margin arcuate. Metasternum completely

divided medially by longitudinal sulcus. Legs elongate, setation

sparse, decumbent, uniformly distributed; coxae conical, trochanteral

socket 0.4x coxal length; trochanters elongate, length pro- and

mesotrochanters equal to coxal length, metatrochanters O.Bx metacoxal

length, all femoral insertions oblique and offset; femora strongly

compressed, straight, 4x trochanteral length, annulated basally; tibiae

strongly compressed, internal margin, sinuate; tibial spines similar,

short; tarsi elongate, tarsomeres without laterally expanded plantar

pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but width narrowing uniformly posteriorly; tergite 8 entire;

tergite 9 not medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; sternite 7 broadly emarginate.

Spiracles in pleural membrane, located submedially from apical margin of tagrna. Aedeagus without parameres; basal piece undifferentiated (Fig.

26 a-c); median lobe tubular, acuminate, ventrally membranous 0.9x

length; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown. 227

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. Dominican Republic, Provincia de La Vega, 12 km NE

Jarabacoa at 550 m,, in flight intercept trap, 01-07 September 1988, M.

A. Ivie, T. E. Phillips, and K. A. Johnson coirs. (1-NMNH, holotype;

3-MAIC; 1-RSMC).

ETYMOLOGY. The Greek specific epithet, adiaophorus. refers to the intermediate states of this species within the genus. 228

Leptolvcus brunneus NEW SPECIES

Fig. 27 a-e

TYPE: INHS, a male with the following data: Cwhite printed label]

"HAITI, Furcy \\ Dept, de 1'ouest \\ May 16, 1959 \\ M. W. Sanderson \\

H 59-4."

DIAGNOSIS. This species may be differentiated from other Leptolvcus

spp. by its light brown coloration with slightly indicated yellow

humeral angles and its strongly transverse and trapezoidal pronotum.

Unlike other known Leptolvcus. the apical antennomere is not white, but

the same brown as the other antennomeres.

DISTRIBUTION. Known only from the unique type collected in Furcy,

Haiti on the island of Hispaniola.

DESCRIPTION (Male). Small, slender, elongate, length 3.2 mm., width 0.7

ram. at humerus. Yellow brown; except at humeral urabone briefly yellow.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, densely punctate; setae semi-decumbent. Genal margin elongate,

length greater than eye length, narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate; radial origin inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect, strongly gibbose; length 1.3x width, inserted between middle of eyes; pentagonal dorsally, deeply and broadly sulcate dorsomedially, appearing bigibbose; ventrally trianguliform, sides nearly straight, widest near base. Antennal sockets oval, separated by

0.2x width of epistomal margin and equidistant to distance between eye and antennal socket. Antennifer elevated medially, appearing basomedial. Interocular distance 2x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae semi-decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly.

Tentorial maculae separate, deeply distinct, at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width. Epistomal margin narrowly emarginate, width 0.6x frontal distance between eyes.

Labrum rectangular, length Q.5x width, apical margin truncate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than G.5x palpomere 3, inserted subapically and internally, acuminate. Labial palps absent. Gula elongate, pentagonal; width slightly greater basally than apically; length 2x width apical margin; inserted anteriorad posterior margin of postgenae. Antennae filiform,

0.7x body length; basal setation sparse, uniformly distributed, decumbent on antennomeres 1-3 and basally on 4; distal setation on antennomeres 4-11 distributed along margins, elongate, semi-erect and bristling; scaliform setae present along margins; scape obconic, width apically 0.6x length; pedicel cylindrical, short, transverse, not compressed, length subequal to antennomere 3; antennomere 3 adnate with base antennomere 4, length 0.2x antennomere 4; antennomeres 4-10 230 elongate filiform, strongly compressed, lateral margins irregular; antennomeres 9-11 expanded laterally 1.5x width antennomere 4; antennomere 11 fusiform, compressed, length 4x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 2x basal width, anterior width nearly 0.6x basal width; median longitudinal carina absent, median fovea obscure; pronotal stemmata absent; anterior and lateral discal margins swollen, posterior margin not; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, slightly convexly arcuate; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.9x width at elytral humerus; disk flat, anterior corners deely impressed, posterior disk elevated near posterior margin, discal setae semi-decumbent, pronotal stemmata absent; hypomeron shallowly concave, almost flat, height 0.5x length; anterior and posterior margins swollen, hypomeral stemmata absent; prosternal bridge trianguliform, anterior margin shallowly emarginate, length 0.8x width, not carinate. Scutellum pentagonal, apically quadrate and nearly square, lateral margins parallel, apex shallowly emarginate, dense setation occluding disk. Mesospiracle prominent, peritreme exposed, orifice slightly compressed.

Mesepisternum elongate, posterior margin straight. Mesepimeron triangulate, width 0.5x and height 0.8x mesepisternum; posterior not reflexed, nearly straight. Mesosternum transverse, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base.

Elytra ligulate, strongly dehiscent at scutellum, divergent to apices, uniformly narrowing most elytral length, rate lateral convergence increases near apex, apex acuminate; length 0x pronotal length; bicostate, costa 4 distinct basal 0.5x elytral length, humeral costa

distinct to near apex; setation semi-decumbent, sparse, uniformly

arranged. Metepisternum with ventral margin increasingly arcuate

posteriorly. Metepimeron with ventral margin strongly arcuate.

Metasternum completely divided medially by longitudinal sulcus. Legs

thin and elongate, setation sparse, decumbent, uniformly distributed;

coxae conical, trochanteral socket 0.4x coxal length; trochanters

elongate, length pro- and mesotrochanters subequal to coxal length,

metatrochanters 0.9x metacoxal length, femoral insertions slightly

oblique, metatrochanter slightly offset, others not; femora strongly

compressed, straight, short, 3x trochanteral length, annulated basally;

tibiae strongly compressed, internal margin straight; tibial spines

similar, short; tarsi elongate, tarsomeres without laterally expanded

plantar pads; covered with semi-decumbent bristling setae; ungues

simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but width narrowing uniformly posteriorly; tergite 8 entire;

tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; sternite 7 broadly emarginate.

Spiracles in pleural membrane, located submedially from apical margin of

tagma. Aedeagus without parameres; basal piece absent

length; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown. 232

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The only available individual of this brown species apppears to be teneral. Thus, its faint yellow humeral angles may be more distinct, if it were captured later in life. Nevertheless, the yellow indications may be of diagnostic assistance, because they are at the humeral angles and do not meet the median suture as the basal fascia of other Leptolvcus species.

MATERIAL EXAMINED. HAITI, Dept, de l'Ouest, Furcy, 16 May 1959, M. W.

Sanderson coir. (1-INHS, holotype).

ETYMOLOGY. This species is named for its brown coloration. 233

Leptolvcus dominicensis NEW SPECIES

Fig. 28 a-c

TYPE: NMNH, a male with the following data: Cwhite printed

label] "DOM. HEP. La Vega \\ 18 km. E. El Rio. Aug. \\ 4, 1979 crest,

cloud \\ forest. G.B. Marshall."

DIAGNOSIS. This black species with the prothorax, the venter of the

head and the metathorax golden yellow, can be distinguished by the lack

of complete and well-defined lateral pronotal margins and by the sparse

erect and bristling antennal setae. This species strongly resembles L.

f lavicol1i s from Puerto Rico, but differs by the coloration of the

venter of the head and metathorax and the more transverse prothorax.

The later is most easily seen in comparative study of both species, but

the ratios should be of use.

DISTRIBUTION. Known only from the type locality in the Dominican

Republic on Hispaniola.

DESCRIPTION (Male). Small, slender, elongate, length 3.5 mm., width 0.8

mm. at humerus. Black; except thorax, basal 0.2x elytra, coxae,

trochanters basally, venter of head golden yellow; and antennomeres 10

distally and 11 white.

Head. Elongate, widest behind eyes, then narrowing to posterior margin; setae dense, semi-decumbent, occluding disk. Genal margin

elongate, length greater than eye length, narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when

viewed dorsally; hind margin shallowly emarginate; radial origin inside

head margin 0,5x radius when dorsally viewed. Antennal prominence

distinct from vertex, porrect, strongly gibbose; length equal width,

inserted between middle of eyes; pentagonal dorsally, sulcate

dorsomedially, sulcus broad anteriorly and indistinct posteriorly;

ventrally trianguliform, shallowly concave, sides nearly straight,

widest near base. Antennal sockets lenticular, separated by O.lx width

of epistomal margin and equidistant to distance between eye and antennal

socket. Antennifer elevated medially, appearing basomedial, inserted near apex. Interocular distance 3x width eye dorsally. Vertex

slightly arcuate, nearly transversely flat, setae semi-decumbent; depression of tentorial maculae shallow and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct, at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.7x width.

Epistomal margin truncate, width 0.6x frontal distance between eyes.

Labrum transverse, length 0.5x width, apical margin truncate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical and lengths equal; diameter 1 equal to length 3, diameter 2 l.Sx palpomere 1;

length palpomere 4 2.Ox palpomere 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps absent. Gula transverse, trapezoidal; width slightly greater basally than apically; length 2x width apical margin; inserted anteriorad posterior margin of postgenae.

Antennae filiform, 1.3x body length; basal setation sparse, uniformly 235 distributed, sparse, semi-decumbent on scape; distal setation on antennomeres 2-11 distributed along margins, elongate, semi-erect and bristling; scaliform setae present along margins; scape obconic, width apically 0.7x length; pedicel cylindrical, short, transverse, not compressed, length subequal antennomere 3; antennomere 3 not adnate with base antennomere 4, length O.lx antennomere 4; antennomeres 4-10 elongate filiform, strongly compressed, lateral margins irregular;; antennomeres 9-11 not expanded laterally greater than width antennomere

4; antennomere 11 fusiform, compressed, length 9x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width nearly 0.6x basal width; median longitudinal carina not visible, median fovea obscure, 0.5x pronotal length, closed at posterior margin; pronotal stemmata absent; apical and lateral discal margins swollen; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, concavely arcuate longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally Q,9x width at elytral humerus; disk transversely arcuate, elevated near posterior margin, discal setae semi-decumbent, dense, obscuring disk; hypomeron deeply concave, height 0.5x length; anterior, ventral and posterior margins swollen, hypomeral stemmata absent; prosternal bridge trianguliform, anterior margin broadly emarginate, length 0.5x width, not carinate. Scutellum pentagonal, apically trapezoidal, lateral margins divergent, apex strongly emarginate, emargination 0.3x scutellar length.; apices divergent, acute.

Mesospiracle prominent, peritreme exposed, orifice slightly compressed.

Mesepisternum elongate, posterior margin straight. Mesepimeron lobate, width 0.3x and height 0.8x mesepisternum; posterior not reflexed, evenly arcuate. Mesosternum transverse, posterior margin convexly arcuate,

Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, strongly dehiscent near base in fascial middle, divergent to apices, uniformly narrowing most elytral length, slightly expanded and spatulate near apex, apex acuminate; length 9x pronotal length; bicostate, costa 4 distinct basal 0.5x elytral length, humeral costa, distinct 0.7x elytral length; disk irregularly and strongly punctate-reticulate; setation sparse, semi-decumbent, uniformly arranged. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin slightly arcuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.4x coxal length; trochanters elongate, length pro- and mesotrochanters lx coxal length, metatrochanters lx metacoxal length, femoral insertions truncate, metafemora slightly offset, others not; femora strongly compressed, straight, short, 2x trochanteral length, annulated basally; tibiae strongly compressed, internal margin sinuate; tibial spines similar and short; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorly; sternite 7 broadly emarginate. 237

Spiracles in pleural membrane, located submedially from apical margin of

tagma. Aedeagus without parameres; basal piece small, compressed,

length about 0.4x length of median lobe (Fig. 28 a-c); median lobe

tubular, acuminate, ventrally and subapically membranous O.Sx length;

internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. An additional structure that warrants further study is the scutellum which is strongly emarginate with the apices divaricate in L. dominiecensis versus the non-emarginate scutellum with apex narrow and truncate in L. flavicol 1 is. However, a larger series of both species

is necessary to determine the limits of variation.

MATERIAL EXAMINED. DOMINICAN REPUBLIC: La Vega, 18 km. E. El Rio. 4

August 1979, crest in cloud forest, G.B. Marshall coir, (1-NMNH, holotype >

ETYMOLOGY. The specific epithet refers to the country where the type locality is found. 238

Leptolvcus effeminatus NEW SPECIES

Fig. 29 a-d

Leptolvcus heterocornis Leng and Mutchler, 1922:430, fig. 22a (pars,

"female" ).

TYPE: AMNH, the allotype of Leptolvcus heterocornis L. & M. , a male with the following data: [white printed label] "Aibonito, P. R. \\

July 14 - 17, * 14;" [red prnted labelD "ALLOTYPE;" [printed white

label] "Am. Mus. Nat. Hist. \\ Dept. Invert. Zool. W No. 24526;"

[handwritten white label] "Leptolycus \\ heterocornis \\ type L & M."

DIAGNOSIS. This species is separable from congenerics by the combination of the strongly serrate antennae with regular margins and the V-shaped impression on the lower surface of the antennal prominence.

It is most easily identified by the ultimate white antennomere with blackish apex which is unknown in other Leptolycini.

DISTRIBUTION. Known only from Puerto Rico.

DESCRIPTION (Males). Small, slender, elongate, length 2.1-2.8 mm., width 0.5-0.6 mm. at humerus. Color black, except basal 0.2-0.3x elytra golden yellow and ultimate antennomere white excluding apex.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, shining punctate; setae sparse, short, semi-decumbant. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when

viewed dorsally; hind margin shallowly emarginate, nearly straight;

radial origin inside head margin 0.3x radius when dorsally viewed.

Antennal prominence distinct from vertex, gibbose; length 0.8x width,

inserted between middle of eyes; quadrate, anterior margin slightly

produced dorsally, broadly and deeply sulcate dorsomedially, sulcus

distinct entire length; ventrally trianguliform, deeply concave, with V-

shaped impression, sides uniformly and convexly arcuate, widest near base. Antennal sockets subrectangular, separated by 0.2x width of epistomal margin and 0.3x distance between eye and antennal socket.

Antennifer elevated medially, appearing basomedial. Interocular distance 2x dorsal eye width. Vertex shallowly punctate, transversely flat, setae semi-decumbent; depression of tentorial maculae broadly shallow, from base of antennal prominence to median eye length, more distinct anteriorly. Tentorial maculae indistinct; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transverse, height 0,5x width. Epistomal margin truncate, width 0.7x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly rounded.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps 1 segmented and digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin, inserted anteriorad posterior margin of postgena. Antennae 0.7x body length; setation sparse, bristling, distributed along margins; scape obconic, 240 width apically 0.5 length; pedicel cylindrical, short, transverse, not compressed, length lx antennomere 3 length; antennomere 3 length O.lx antennomere 4; antennomeres 4-10 elongate, subserrate, strongly compressed, margins not irregular; antennomere 11 fusiform, compressed, length 1.3x antennomere 4.

Thorax. Pronotum trapezoidal, transverse, length O.Qx basal width, anterior width nearly O.Qx basal width; median longitudinal carina obscure, median fovea obscure; pronotal stemmata absent; anterior and lateral discal margins swollen; anterior margin slightly produced and reflexed; anterior angles obtuse; lateral margin complete, not foliaceous, not reflexed, strongly concavely arcuate; posterior margin bisinuate; posterior angles acute, expanded laterally O.Qx width elytral humerus; disk flat, more finely reticulate than head or elytra, margin elevated near posterior margin, setation shorter than elytral setae, decumbent; hypomeron convex, medial height 0.5x length; posterior and lateral margins swollen, hypomeral stemmata absent; prosternal bridge trianguliform, length 0.5x width, not carinate. Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex truncate, slightly incised. Mesospiracle prominent, peritreme exposed, orifice not compressed. Mesepisternum elongate, posterior margin straight. Mesepimeron trianguliform, 0.5x width and subequal mesepisternal height; posterior margin very little reflexed, nearly straight. Mesosternum elongate, posterior margin truncate. Nesocoxal separation lx coxal diameter at base. Elytra ligulate, dehiscent from scutellum; length 6x pronotal length; bicostate, costa 2 distinct basal

0.3x elytral length, humeral costa distinct 0.7x elytral length; reticulation irregular; setation decumbent, sparse, uniformly arranged.

Metepisternum with ventral margin increasingly arcuate posteriorly.

Metepimeron with ventral margin straight. Metasternum completely

divided medially by longitudinal sulcus. Legs elongate, setation

sparse, semi-decumbent, uniformly distributed; coxae conical,

trochanteral socket nearly 0.3x coxal length; trochanters elongate,

length pro- and mesotrochanters subequal to coxal length,

metatrochanters subequal to metacoxal length, femoral insertions

truncate, not offset; femora strongly compressed, straight, short, 5x

trochanteral length, annulated basally; tibiae strongly compressed,

internal margin slightly sinuate; tibial spines similar and short; tars elongate, tarsomeres without laterally expanded plantar pads; covered with erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma.

Aedeagus without parameres; basal piece large, dorsoventrally compressed, length equal length of median lobe (Fig. 29 a-c); median

lobe tubular, unmodified, ventrally membranous 0.5x length, membrane distal from apex O.lx length median lobe; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown. 242

DESCRIPTION (Immatures ). Unknown.

DISCUSSION. Leng and Mutchler (1922) mistook the sex of this species and sonsidered it the female of L heterocornis.

MATERIAL EXAMINED. Puerto Rico, Aibonito, 14 July 1917 (3-AMNH, including holotype); Cambalalache Forest Res., 28 July 1979, C. W.

O'Brien (1-RSMC); Guajutaca For. Res., 27 July 1979, C. W. O'Brien

(1-RSMC); Guilarte For. Res., Hwy 131 and 158, 23 July 1979, L. B.

O'Brien (1-RSMC, 1-MAIC); Cthere is supposedly a specimen in the NMNHD.

ETYMOLOGY. The specific epithet refers to the type's erroneus former status as a female allotype. 243

Leptolvcus flavicol1 is NEW SPECIES

Fig. 30 a-d

Leptolvcus heterocornis var. flavicollis Leng and Mutchler,

1922:431. Blackwelder, 1945:343.

TYPE: AMNH, a male with the following data: Cwhite printed labels!

"Aibonito, P. R. \\ July 1914;’’ "H. G. Barber;” ”Am. Mus. Nat, Hist. \\

Dept. Invert. Zool. No 24529."

DIAGNOSIS. This species is characterized by the combination of: pronotum golden yellow; the venter of the head and metathorax are black;

the lateral pronotal margins are not well-defined anteriorly; the sparse

setation on the antennae is erect and bristling; and the pronotal length

is O.Qx its basal width. It differs from L. dominicensis. which it most closely resembles, by ventral coloration.

DISTRIBUTION. Known only from Puerto Rico,

DESCRIPTION (Males). Small, slender, elongate, length 3.1-4.2 mm., width

0.8-1.0 mm. at humerus. Black; except basal 0.3x elytra golden pro— and mesothorax and base trochanters golden yellow; base metatrochanters and antennomere 11 white.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, shining; setae sparse, decumbent. Genal margin elongate, length greater than eye length, narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin shallowly emarginate; radial origin inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, porrect, strongly gibbose; length 1.lx width, inserted between middle of eyes; pentagonal dorsally, not sulcate dorsomedially; ventrally trianguliform, convex, sides nearly straight, widest near base. Antennal sockets lenticular, separated by O.lx width of epistomal margin and 0.3x distance between eye and antennal socket.

Antennifer elevated medially, appearing basomedial. Interocular distance 2x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae decumbent; depression of tentorial maculae deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae not separate, deep, at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.3x width. Epistomal margin truncate, width 0.5x frontal distance between eyes. Labrum quadrate, length equal width, apical margin truncate. Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, inserted subapically and internally, acuminate. Labial palps absent. Gula elongate, pentagonal; width slightly greater basally than apically; length 2x width apical margin; inserted anteriorad posterior margin of postgenae. Antennae filiform, 1.3x body length; basal setation sparse, uniformly distributed, decumbent on antennomeres 1-3 and basally on 4; distal setation on antennomeres 4-11 distributed along margins, elongate, semi-erect and bristling; sealiform setae absent; scape 245

obconic, width apically 0.6x length; pedicel cylindrical, short,

transverse, not compressed, length 2x antennomere 3; antennomere 3

adnate with base antennomere 4, length O.lx antennomere 4; antennomeres

4-10 elongate filiform, strongly compressed, lateral margins irregular;;

antennomeres 9— 11 expanded laterally 1.5x width antennomere 4;

antennomere 11 fusiform, compressed, length 0.9x antennomere 3.

Thorax. Pronotum trapezoidal, elongate, length 0.8x basal width,

anterior width nearly 0.7x basal width; median longitudinal carina

barely visible anteriorly, median fovea obscure; pronotal stemmata

absent; apical discal margins swollen, others not; anterior margin

strongly convexly arcuate, anterior angles acute; lateral margin

incomplete anteriorly and carinate posteriorly, strongly concavely arcuate longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk

transversely arcuate, posterior disk elevated near posterior margin, discal setae semi-decumbent, dense, obscuring disk; hypomeron shallowly concave, almost flat, height 0,5x length; anterior, ventral and lateral margins swollen, hypomeral stemmata absent; prosternal bridge trianguliform, anterior margin truncate to slightly emarginate, length

0.8x width, not carinate. Scutellum pentagonal, apically quadrate and nearly square, lateral margins parallel, apex truncate, dense setation occluding disk. Mesospiracle prominent, peritreme exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin straight. Mesepimeron lobate, width 0.5x and height 0.8x mesepisternum; posterior not reflexed, evenly arcuate. Mesosternum transverse, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, strongly dehiscent near

base in fascial middle, divergent to apices, uniformly narrowing most

elytral length, slightly expanded and spatulate near apex, apex

acuminate; length 5x pronotal length; bicostate, costa 4 distinct basal

0.3x elytral length, humeral costa distinct 0.5x elytral length; disk

occluded by dense, semi“decumbent setae. Metepisternum with ventral

margin increasingly arcuate posteriorly. Metepimeron with ventral

margin strongly arcuate. Metasternum completely divided medially by

longitudinal sulcus. Legs thin and elongate, setation sparse,

decumbent, uniformly distributed; coxae conical, trochanteral socket

0.4x coxal length; trochanters elongate, length pro-and mesotrochanters

1.2x to coxal length, metatrochanters 1.3x metacoxal length, femoral

insertions truncate, none offset; femora strongly compressed, straight,

short, 2.3x trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight; tibial spines similar, short;

tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but width narrowing uniformly posteriorad; tergite 8 entire;

tergite 9 medially divided longitudinally; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but width narrowing uniformly posteriorad; sternite 7 broadly emarginate.

Spiracles in pleural membrane, located submedially from apical margin of

tagma. Aedeagus without parameres; basal piece large, compressed, expanded posteriorly, length about 0.7x length of median lobe (Fig. zz ); median lobe tubular, acuminate, punctate, ventrally and medially 247 membranous 0.6x length, membrane distal from apex 0.2x length median lobe; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. This species was considered by Leng and Mutchler as to be a variety of L. heterocornis based on a single specimen which they thought was a female. The International Code of Nomenclature does not recognize varietal names, so the name flavico11is dates from the present work.

MATERIAL EXAMINED. PUERTO RICO: Aibonito, July 1914, H. G. Barber

(1-AMNH, holotype); Caribbean Nat. For., Mt. Britton Trail, 19 July

1979, G. B. Marshall (1-RSMC); Guilarte For. Res., Hwys. 131 & 158, 23

July 1979, L. B. O ’Brien (1-RSMC).

ETYMOLOGY. Flavis (Latin), yellow and c o 1lum (Latin), neck. 248

Leptolvcus heterocorn is Leng and Mutchler

Figs. 31, 32 a-c, 33, 34

Leptolvcus heterocornis Leng and Mutchler, 1922:430, fig. 12. Kleine,

1933:18. Beatty, 1944:136. Blackwelder, 1945:343. MLskimen and

Bond, 1970:85

TYPE: AMNH, a male with the following data: [white printed label]

"Aibonito, Porto Rico, \\ July 14-17, 1914, \\ H. G. Barber;" [red

printed label] "Holotype;" Cwhite printed label] "Am. Mus. Nat. Hist.

\\ Dept. Zool. \\ No. 24525."

DIAGNOSIS, Males can be separated form other male leptolycines by the following characters in combination: pronotum black, lateral margins of pronotum indistinct anteriroly, antennomeres irregularly serrate with margins irregularly, and erect and bristling antennal setae present.

Females appear similar to conspecific larvae, but differ by the presence of non-fenestrated cuticle, the non-differentiated maxilla and labium which only consists of a pair lobes without palps, the presence of one pair of free abdominal pleurites, the abdominal sternites reach the

lateral margin of the abdomen instead of being restricted to small central plates, and the presence of a gonopore. Larvae can be

separated from other known lycids by the characters given in the generic treatment.

DISTRIBUTION. Known from throughout Puerto Rico, St. John and St. 249

Thomas of the U. S. Virgin Islands and Guana Is. of the British Virgin

Islands. Beatty (1944:136) and Miskimen and Bond (1970:85) reported a species of Leptolvcus on St. Croix which is presumably L. heterocornis.

I have yet to find any specimens verifying this distribution, although a search was specifically conducted for the reported specimens.

DESCRIPTION (Males): Small, slender, elongate, length 3.2-3.8 mm., humeral width 0.6-0.9 mm. Black; except basal 0.2-0.3x elytra golden yellow, trochanters pale yellow, antennomere 11 white.

Head. Elongate, widest behind eyes, then narrowing to posterior margin, shining areolate; setae sparse, directed anteriorly, semi- decumbant. Genal margin elongate, length greater than eye length, convexly arcuate when viewed dorsally. Eyes small, hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly and shallowly emarginate, nearly straight; radial origin inside head margin 0.3x radius when dorsally viewed. Antennal prominence distinct from vertex, gibbose; length 0.8x width, inserted between middle of eyes; pentagonal dorsally, broadly sulcate dorsomedially, sulcus distinct posteriorly, indistinct 0.5x anterior length; ventrally pentagonal, concave, sides uniformly and convexly arcuate, widest near base. Antennal sockets lenticular, ventral margin almost straight separated by O.lx width of epistomal margin and 0.5x distance between eye and antennal socket. Antennifer elevated medially, appearing inserted basomediaily. Interocular distance 2x width eye dorsally.

Vertex transversely arcuate, nearly flat, setae decumbent; depression of tentorial maculae shallow, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate,

shallow, distinct; at base of antennal prominence. Frons transversely convex, distinct from lower surface of antennal prominence, transversely narrow, height 0.5x width. Epistomal margin broadly emarginate, almost straight, width 0.7x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly rounded.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.5x palpomere 3, acuminate. Labial palps 1 segmented and digitiform. Gula elongate, subrectangular; width slightly greater apically than basally; length 2x width apical margin. Antennae lx body length, setation sparse, uniformly distributed, erect and bristling on antennomeres 3-11; scape pyriform, width apically 0.7x length; pedicel cylindrical, short, transverse, not compressed, length lx length of antennomere 3; antennomere 3 not adnate to antennomere 4, length O.lx antennomere 4; antennomeres 4-10 conical, irregularly filiform, strongly compressed; antennomere 11 fusiform, compressed, length 0.2x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length O.Qx basal width, anterior width nearly 0.7x basal width; median longitudinal carina obscure, median fovea indistinct; pronotal stemmata absent; apical discal margin swollen; anterior margin convexly arcuate and angulate medially, anterior angles obtuse; lateral margin incomplete, weakly carinate posteriorad, strongly concavely arcuate longitudinally; posterior margin weakly bisinuate; posterior angles acute, expanded laterally 0.7x elytral humerus; disk slightly arcuate, nearly flat, posterior disk strongly elevated at margin discal setae shorter and sparser than elytral setae, semi-decumbent; hypomeron convex, height

0,5x width, posterior and lateral margins not swollen; prosternal bridge trianguliform, length lx width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, nearly square, lateral margins parallel, apex deeply emarginate, emargination nearly 0.5x scutellar length, apices truncate. Mesospiracle prominent, peritreme exposed, oriface slightly compressed. Mesepisternum elongate, posterior margin straight. Mesepimeron lobate, 0.6x width and 0.7x height mesepisternum; posterior margin very little reflexed, uniformly arcuate. Mesosternum elongate, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, dehiscent from scutellum, divergent to apex, uniformly narrowing distally except apex spatulate; length 5x pronotal length; bicostate, costa 4 distinct basal 0.5x elytral length, costa 8 distinct 0.8x elytral length; disk irregularly punctate; setation semi-decumbent, moderately dense, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron nearly straight. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, semi-decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5x coxal length; trochanters elongate, length pro- and mesotrochanters 0.8x coxal length, metatrochanters lx metacoxal length, femoral insertions truncate and not offset; femora strongly compressed, straight, short, 2.5x trochanteral length, annulated basally; tibiae strongly compressed, internal margin slightly bisinuate; tibial spines small and similar; tarsi elongate, tarsomeres without 252

plantar pads; covered with erect bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9

medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing

uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece large, dorsoventrally compressed, length almost lx length of median lobe (Fig. 32 a-c); median lobe tubular, unmodified, ventrally membranous 0.6x length and subapically 0.2x length of median lobe, internal sac not eversible, flagellum absent.

DESCRIPTION (Females). Small, onisciform; length 2.6 - 3.2 mm; widest width 0.6 - 0.7 mm. No sclerites fenestrate. Pale yellow.

Head. Trianguliform, widest apically, width 2x length; partially recessed within prothorax; all sclerites reticulate-fenestrate. Eyes a single pair of stemmata above ecdysial line at base of antennal prominence, redessed, dark black. Antennal prominences porrect anteriolaterally; apical orifices large, incised laterally along lateral genal margin to near eye. Antennae recessed, 2 segmented, apical segemnt bulbous. Vertex depressed between two arms from base to antennal oriface. Epistomal margin nearly straight, not incised medially. Mandibles absent. Maxillary palps a undifferentiated lobe protudeced apically.

Thorax. Elongate, length 0.4x body length; all sclerites reticulate-fenestrate. Tergites transversely arcuate, nearly flat, 253 margins reflexed ventrally at sides. Pronotal length 1.3x mesonotal

length. Prosternum a triangular sclerite, widest apically, length 0.5 width. Mesonotal length lx metanotal length. Mesosternum not

sclerotized. Metasternum a square sclerite. Pleurites scelrotized above all legs, pleural sutures distinct. Mesospiracle central in circular, protruding pleural sclerite. Legs 3 segmented; tarsunguius

long, attenuated distal 0.3x, diameter at apical 0.3x less than 0.5x base; terminated by single claw.

Abdomen. Ten segmented, last segment modified to as a pygopodium; pygopodium an undifferentiated disk. Tergites wider than

long, not explanate at posteriolateral corners; tergite 9 without urogomphi. One differentiated pair of pleurites per each abdominal segment; dorsal pair rectangular rectangular, length 0.5x height, biforous spiracle inserted medially; ventral pair absent. Sternites transverse, length approximately 0.5x width, reaching lateral margins.

DESCRIPTION (Immatures>. Small, onisciform; length 2.5 - 3.2 mm; widest width 0.6 - 0.7 mm. Most sclerites fenestrate. Yellow, except pro - and mesosegmental and 1 - 3 abdominal sclerites brown entirely and occassionally other abdominal tergites medially.

Head. Trianguliform, widest apically, width 2x length; partially recessed within prothorax; all sclerites reticulate-fenestrate. Eyes a single pair of stemmata above ecdysial line at base of antennal prominence, redessed, dark black. Antennal prominences porrect anteriolaterally; apical orifices large, incised laterally along lateral genal margin to near eye. Antennae recessed, 2 segmented, apical 254

segemnt bulbous. Vertex depressed between two arms from base to

antennal oriface. Epistomal margin nearly straight, not incised

medially. Mandibles divided, short, less than O.Sx epistomal margin;

inserted medially, directed ventrally.

Thorax. Elongate, length 0.4x body length; all sclerites

reticulate-fenestrate. Tergites transversely arcuate, nearly flat,

margins reflexed ventrally at sides. Pronotal length 1.3x mesonotal

length. Prosternum a triangular sclerite, widest apically, length

O.Sx width. Mesonotal length lx metanotal length. Mesosternum not

sclerotized. Metasternum a square sclerite. Pleurites scelrotized

above all legs, pleural sutures distinct. Mesospiracle central in

circular, protruding pleural sclerite. Legs 3 segmented; tarsungulus

long, attenuated distal 0.5x, diameter at apical 0.3x less than 0.3x

base; terminated by single claw.

Abdomen. Ten segmented, last segment modified to as a

pygopodium; pygopodium an undifferentiated disk. Tergites wider than

long, not explanate at posteriolateral corners; tergite 9 without

urogomphi. Two differentiated pair of pleurites per each abdominal

segment; dorsal pair rectangular rectangular, length 0.5x height,

biforous spiracle inserted medially; ventral pair trianguliform, widest

caudally. Sternites transverse, length approximately O.Sx width, not

reaching lateral margins.

DISCUSSION. This species was originally described from a series

containing two species. The holotype is a male of one species and the

allotype is, in fact, a male of the other, although it was mistaken for 255 a female. The later is redescribed here as J.. effeminatus.

In addition to this error, a number of character states in the original description of Leng and Mutchler (1922) are not as in the series before me. It is unlikely that this deviation is the result of the combination of character states of the two species, but probably results from insufficiently powerful optics.

A single male adult was found associated with larvae from St.

John, U. S. Virgin Islands. Numerous larvae were collected throughout the island, primarily by William Muchmore in his studies of pseudoscorpions of that island and by Michael A. Ivie in his studies of the beetle fauna of the Virgin Islands. A second color morph wasfound which were often mixed in the same sample. One is here hypothesized to be the last instar larva and the other the paedomorphic female. Only one species of lycid is known as adults from St. John and the only other likely non-leptolycine would be Thonalmus spp. I have seen the larvae of that genus and these larvae differ significantly. No other species of adult leptolycines have been collected on St. John and, hence, the hypothesis of conspecifity in both thelarval morphs and female is thought to be well supported.

MATERIAL EXAMINED (Adult Males). PUERTO RICO: Aibonito, 14-17 July

1914, H. G. Barber (1-AMNH, Type No. 24525); Mayaguez, 19 Aug. 1945,

J. A. Ramos (1-JARC); Insular Forest, Maricao, 29 Mar. 1947 (1-JARC);

Maricao For. Res., Hwy. 120, K134H8, 25 July 1979, L.B. O'Brien and G.

B. Marshall (3-RSMC); Maricao Forest, 600-900 m., 30 May-2 June,

Darlington (2- MCZC); Guilarte For. Res., Hwys. 131 & 158, 23-27 July 256

1979, L. B. O'Brien & G. B, Marshall (11-RSMC); Caribbean Nat. For., El

Toro Negro, Hwy. 143, DK19H9, 22 July 1979, G. B. Marshall & C. B.

O'Brien (3-RSMC); Toro Negro, 19 Nov. 1947 (1- UPRC); Luquillo Forest,

El Yunque Sta., 10-16 July, 1969, H. & A. Howden (1-CNCI); El Yunque, c. 900 m. , Darlington (4-MCZC); El Yunque, 17 Feb. 1926, H. L. Do2 ier

(1-NMNH); Vilalba, 26 Oct. 1933, R. G. Oakley (1-NMNH); Cayey, 30-31 May

1915 (1- NMNH, Paratype 61029 = Paratype 24527 of AMNH).

U. S. VIRGIN ISLANDS: St. John, 29 May 1940, D. DeLeon, on grass

(1-NMNH). Estate Carolina, NW Coral Bay, 8 m, 18 May 1984, ex ground litter, W. B. Muchmore (1-MAIC).

BRITISH VIRGIN ISLANDS: Guana Is., 13 July 1988. (1-MAIC). Guana

Is., 1 October 1989, V. 0. Becker coir. (1-MAIC).

(Adult Females). U. S. VIRGIN ISLANDS: St. John, Lameshur Bay,

12 June 1980, at base of large tamarind (2-MAIC); Gray Gut, 12 June

1980, among rocks and pieces of fallen termite nest (2-MAIC). Francis

Bay Pond, 29 February 1984, at base of large tree on shore of pond, W.

B. Muchmore (2-MAIC). Windberg Ruins, 8 October 1979, ex litter, W.

B. Muchmore (1-MAIC). Estate Hermitage, Ruins, 20 June 1980, at base of well, W. B. Muchmore (3-MAIC; 1-RSMC). Cinnamon Bay Nature Trail, 6

June 1980, ex kapok buttresses, W. B. Muchmore (1-MAIC).

Catherineburg, 25 May 1982, in litter under old log, W. B. Muchmore

(1-MAIC). Trunk Bay, 9 June 1980, at base of tree and Antherium. W. B.

Muchmore (1-MAIC). Brown Bay, 4 May 1984, litter under calabash tree,

W. B. Muchmore, (3-MAIC).

St. Thomas. Magens Bay, 30 July 1980, ex rotten stump, M. A. 257

Ivie (1-MAIC).

(Immatures). U. S, VIRGIN ISLANDS: St. John. Coral Bay,

butler's house, 10 October 1979, ex ground litter, W. B. Muchmore

(1-MAIC). Estate Hermitage ruins, 20 June 1980, at base of well, W.

B. Muchmore (2-MAIC; 2-RSMC). Estate Carolina, NW Coral Bay, 8 in, 18

May 1984, ex ground litter, W. B. Muchmore (1-MAIC, adult association).

Salt Pond Bay, 10 October 1979, ex litter, W. B. Muchmore (2-MAIC).

St. Thomas. Magens Bay Arboretum, 30 July 1980, ex litter in kapok buttresses, M. A. Ivie (1-MAIC).

ETYMOLOGY. The prefix, Hetero (Greek), meaning different and root, cornu

(Latin), meaning horn refers to the supposed sexual dimorphism of the

antennae in the mixed type species. PLEASE NOTE

Page(s) missing in number only; text follows. Filmed as received.

258

University Microfilms International 259

Pseudacroleptus Pic

Pseudacroleptus Pic, 1911:165. Kleine, 1933:34. Blackwelder,

1945:348.

TYPE: Pseudacroleptus obscuricolor Pic, MNHP.

DIAGNOSIS. This genus can be differentiated from other leptolycines by the presence of a short, rostrate epistomal area; by the flabellate antennae with long rami on antennomeres 3-11; and by the pedicel length nearly, but less than length of antennomere 3.

DISTRIBUTION. Psedacroleptus appears to be restricted to central South

America east of the Andes; two of the four species are known from the

Tepui highland region.

DESCRIPTION (Males). Elongate, but elytra expanded distally 0.6x their length and then narrowing to apex; length 5.6-7.4 mm, width at humerus

1.2-1.6 mm. Black to blackish brown; with no or various maculation of reddish brown or yellow.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin at or inside head margin 0.5x when dorsally viewed. Antennal prominence distinct from vertex, gibbose; length 0.8x width, inserted between middle of eyes;

rectangular dorsally, convex, narrowly impressed dorsomedially, anterior

margin bisinuate to truncate; ventrally pentagonal, deeply concave,

broadly impressed longitudinally, dorsolateral sides nearly carinate

along antennal socket, widest near base. Antennal sockets rectangular,

separated by O.lx width of epistomal margin and 0.3-5x distance between eye and antennal socket. Antennifer insertion elevated medially, not appearing basomedial. Interocular distance 2.8-3x width eye dorsally.

Vertex transversely flat, setae decumbent; depression of tentorial maculae indistinct. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely convex, indistinct from lower surface of antennal prominence, transverse, height 0.6-lx width, shortly rostrate apically, height rostrum 0.3-0.4x width epistoma. Epistomal margin broadly emarginate, width 0.5x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin truncate to broadly emarginate. Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length;

length palpomere 4 nearly equal 3, diameter at base less than 0.3x palpomere 3, directed internally, acuminate. Labial palps 1 segmented, digitiform. Gula transversely rectangular to trapezoidal; length 0.5- lx width apical margin; inserted anteriorad postgenal posterior margin.

Antennae flabel late, length Q.6-0.7x body length; rami between 6-24x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, decumbent to semi-decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically

0.6-lx length; pedicel cylindrical, short, transverse, not compressed, 261

length 0.7-0.8x length of antennomere 3; antennomere 3 flabel late with

ramal insertions basal, slightly compressed, not adnate to base of

antennomere 4, length 0.4-lx antennomere 4; antennomere 4 flabel late

with ramal insertions medial to apical, more strongly compressed;

antennomeres 5-10 compressed, extent of compression decremental to apex,

flabellate with ramal insertions apical; antennomere 11 without axis

distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior

width 0.6x basal width; median longitudinal carina absent anteriorly,

poorly defined and divided posteriorly, median fovea 0.4-0.9x pronotal

length, closing or not at posterior margin; pronotal stemmata present, anterior pair on anterior margin and at lateral margin to 10 stemmatal diameters from lateral margins, posterior pair on posterior margin or cephalad 0.5x stemmatal diameter and 2-4 stemmatal diameters from

lateral margin; anterior and lateral to all discal margins swollen; anterior margin sraight to slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect to erect-explanate, straight to concavely arcuate longitudinally; posterior margin strongly bisinuate, incised or not; posterior angles acute, expanded laterally

0.7-0.Qx width at elytral humerus; disk flat, but anterior 0.3-0.5x

inflated allowing reception of head, setation decumbent; hypomeron shallowly to moderately concave, height 0.4-0.6x width, anterior and posterior to all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms strongly acutely separated to uniformly rounded, length 0.5-0.8x width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrately transverse, lateral margins parallel, apex deeply to shallowly emarginate, emargination 0.1-0.3x

scutellar length, apices acute and divergent or not. Mesospiracle

prominent, peritreme exposed, orifice slightly compressed.

Mesepisternum elongate, posterior margin slightly convexly arcuate.

Mesepimeron lobate to trianguliform; width 0.3-0.5x and height 0.8x

mesepisternum; posterior margin straight to nearly straight, acutely to

obtusely angulate ventrally and dorsally, reflexed to slightly reflexed,

evenly rounded. Mesosternum transverse, length 0.3-0.5x width,

posterior margin convexly arcuate. Mesocoxal separation 1-1,6x

mesocoxal diameter at base. Elytra ligulate, strongly dehiscent from

apical 0.2-0.3x elytral length, slightly reconvergent near apex, apices

acute; length 9-10x pronotal length; bicostate to quadricostate, costa 2

distinct basal 0.3x elytral length or absent, costa 4 and Q distinct

0.8x to near elytral apex, costa 6 faintly distinct middle 0.5-0.8x

elytral length; disk irregularly scabrous-punctate to irregularly

reticulate, cells absent to rare and mostly quadrate; setation semi-

decumbent, sparse, uniformly distributed. Metepisternum with ventral

margin increasingly to decreasingly arcuate posteriorly. Metepimeron with ventral margin arcuate. Metasternum completely to incompletely divided medially by longitudinal sulcus to near mesosternum. Legs elongate, setation moderately dense, decumbent to semi-decumbent, uniformly distributed; coxae conical, pro— and mesotrochanteral socket

0.5-0.8x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 0.7-0.8x coxal length, metatrochanters 0.9-lx metacoxal length, femoral

insertions slightly oblique and not offset except metafemoral insertion 263 slightly to moderately offset; femora compressed, short, 1.8-2.8X trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight to sinuate; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads, covered with decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire to produced and bilobed on postei'ior margin; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire to emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece dorsoventrally compressed or not, broad or narrower, length

0.5-0.6x length of median lobe; median lobe awl-shaped, membranous ventrally 0.1-0.6x its length; internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

ETYMOLOGY. The Latin pseudo. meaning false, refers to superficial similarity of appearance to Acrolepbus Bourgeois, which is left in the

Calopterini and not here placed in the Leptolycini. 264

KEY TO THE SPECIES OF Pseudacroleptus (ADULT MALES)

1. Elytra bicostate, unicolorus, Venezuela ......

...... Pseudacro1eptus obscuricolor Pic (p. 277)

1 * More than 2 visible costae on elytron ...... 2

2. (I1) Elytra tricostate ...... 3

2'. (1 * ) Elytra quadricostate, disk scabrous puncatate, not

reticulate; Venezuela ......

...... Pseudacro1eptus caeruleus New Species (p. 265)

3. (2) Black, Black, with with rustrust brownbrown pronotum; pronotum; French French Guyana Guyana......

...... Pseudacroleptus sinuatis Pic (p. 281)

3. (2*) Mostly black, pronotum rust brown, and antennal axis, rami

basally and distally, venter of metathorax yellow; Venezuela

...... Pseudacroleptus neblinus New Species (p. 270) 265

Pseudacroleptus caeruleus NEW SPECIES

Fig. 35 a-c

TYPE: MIZA, a male in fair condition (the front legs partially missing) with the following locality data: [white printed label!

’’Venezuela. Meri- \\ da, La Azulita \\ 2000 m. 2-X-69;” [white printed label! "J. & B. Bechyne \\ leg;’’ [green printed label! "Venezuela-Inst.

\\ Zool. Agricola \\ Fac. Agromia \\ U.C.V. Maracay."

DIAGNOSIS. This species is easily separable from other Pseudacropletus by the couplets in the key. Unlike the other known species it is quadricostate.

DISTRIBUTION. This species is known only from the type specimen. It was collected in the mountains of western Venezuela near the Colombian border.

DESCRIPTION (Male). Elongate, but elytra expanded distally 0.6x their length and then narrowing to apex; length 7.4 ram, width at humerus 1.6 mm. Blackish brown; except basal O.Qx elytral margin at humerus, elytral apices briefly, pronotal magins anteriorly and last 3 antennomeres apically golden yellow.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin at head margin when dorsally viewed. Antennal prominence distinct from vertex, gibbose; length 0.8x width, inserted between middle of eyes; quadrate dorsally, nearly flat, narrowly impressed dorsomedially, posterior 0.5x divided and divergent to tentorial maculae, anterior margin nearly straight; ventrally pentagonal, deeply concave, broadly impressed longitudinally, dorsolateral sides nearly carinate along antennal socket, widest near base. Antennal sockets rectangular, separated by O.lx width of epistomal margin and 0.3x distance between eye and antennal socket.

Antennifer insertion elevated medially, not appearing basomedial.

Interocular distance 3x width eye dorsally. Vertex transversely flat, setae decumbent; depression of tentorial maculae indistinct. Tentorial ffla&Uiae deebi dietiflEfj dt fea&e bf' 3ft£enfial

Frons transversely convex, indistinct from lower surface of antennal prominence, height approximately equal width, shortly rostrate apically, height rostrum 0.3x width epistoma, Epistomal margin broadly emarginate, width 0.5x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin broadly emarginate. Maxillary palps 4 segmented, paipomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.3x palpomere 3, directed internally, acuminate. Labial palps 1 segmented, digitiform. Gula trapezoidal; length equal width apical margin; inserted anteriorad postgenal posterior margin. Antennae flabel late, O.&x body length; rami between

6-10x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, decumbent on all 267

antennomeres, scaliform setae absent; scape obconic, width apically 0.6x

length; pedicel cylindrical, short, transverse, not compressed, length

0.7x length of antennomere 3; antennomere 3 flabel late with ramal

insertions basal, slightly compressed, not adnate with base of antennomere 4, length 0.4x antennomere 4; antennomere 4 flabel late with ramal insertions apical, equally compressed; antennomeres 5-10 compressed, extent of compression uniform to apex, flabel late with ramal

insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior width 0.6x basal width; median longitudinal carina absent anteriorly, poorly defined and divided posteriorly, median fovea 0.4x pronotal length, not closed at posterior margin; pronotal stemmata present, anterior pair on anterior margin and at lateral margins, posterior pair before posterior margin 0.5x stemmatal diameter and 2 stemmatal diameters from lateral margin; anterior and lateral discal margins swollen; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, erect, concave longitudinally; posterior margin strongly bisinuate, incised medially; posterior angles acute, expanded laterally 0.7x width at elytral humerus; disk flat, but anterior 0.5x inflated allowing reception of head, setation decumbent; hypomeron shallowly concave, height 0.4x width, all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms uniformly arcuate, length O.flx width, intercoxal process not carinate.

Scutellum pentagonal, distally quadrate, lateral margins parallel, apex deeply emarginate, emargination 0.3x scute!lar length, apices acute and divergent. Mesospiracle prominent, peritreroe exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin straight.

Mesepimeron iobate and almost rectangular; width 0.5x and height 0.8x mesepisternum; posterior margin nearly straight, obtusely angulate ventrally and dorsally, reflexed. Mesosternum transverse, length 0.3x width, posterior margin convexly arcuate. Mesocoxal separation 1.6x coxal diameter at base. Elytra ligulate, strongly dehiscent from apical 0.2x elytral length, slightly reconvergent at apex, apices acute; length lOx pronotal length; tricostate, costa 4 distinct to near elytral apex, costa 6 faintly distinct middle Q.5x elytral length, costa 8 distinct 0.9x elytral length and strongly sinuate; disk irregularly scabrous-punctate to irregular reticulate, cells rare and mostly quadrate; setation decumbent, sparse, uniformly distributed.

Metepisternum with ventral margin decreasingly arcuate posteriorly.

Metepimeron with ventral margin arcuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation moderately dense, decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.5x coxal length, metacoxa more elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 0.8x coxal length, metatrochanters 0.9x metacoxal length, femoral insertions slightly oblique and not offset except metafemoral insertion slightly offset; femora compressed, elongate, 2.8x trochanteral length, annulated basally; tibiae strongly compressed, internal margin straight; tibial spines small and similar; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple. Abdomen. Tergites rectangular, wider than long, length 1—8

subequal, but narrowing uniformly posteriad; tergite 8 produced on

posterior margin, bilobed; tergite 9 medially divided longitudinally;

tergite 10 shield shaped. Sternites rectangular, wider than long,

length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 emarginate. Spiracles in pleural membrane, located submedially from apical margin of tagma. Aedeagus without parameres; basal piece not

compressed, length almost 0.6x length of median lobe (Fig. zz); median

lobe awl-shaped, membranous subapically and ventrally 0.4x its length;

internal sac not eversible; flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The incision of the posterior margin of the pronotum and the posterior divaricate median sulcus of the antennal prominenece may be developmental abberations.

MATERIAL EXAMINED. VENEZUELA: Merida, La Azulita at 2000 m., 2

October 1969, J. & B. Bechyne (1-MIZA, holotype).

ETYMOLOGY. The specific epithet, caeruleus. is Latin for sky blue and refers to the type locality, La Azulita. 270

Pseudacropletus neblinus NEW SPECIES

Figs. 36 a-e, 37 a, 38 a-b

TYPE: NMNH, a male in good condition with the following data: Cwhite printed label3 "VENEZUELA, T. F. Amaz. \\ Cerro de la Neblina \\

Basecamp, 140 m. \\ CT50’N, 68'10’W \\ 10-20 February 1985;" Cwhite printed labe3 "Malaise trap \\ in rain forest \\ P. J. and P. M.

Spangler, \\ R. A. Faitoute & \\ W. E. Steiner coirs."

DIAGNOSIS. This species can be differentiated from P. obscuricolor Pic and P. caeruleus n. sp. by the presence of three costae on each elytron, instead of 2 and 4. It is most easily differentiated from P. sinuatis. which also has a reddish brown pronotum, by its distinctive yellow coloration as discussed in the key.

DISTRIBUTION. Spatial and temporal distribution known only from the

Cerro de la Neblina joint tepui study by the Smithsonian and the

Venezuelan scientists at MIZA. Specimens were collected at the base camp by malaise trap from 13-29 February 1984 and 27 January-25

February. An additional specimen was collected at black light at the same locality on 27 January. Whether adults live throughout the rest of the year is unknown.

DESCRIPTION (Males). Elongate, but elytra expanded distally 0.6x their length and then narrowing to apex; length 5.6-6.5 mm, width at humerus

1.2-1.6 mm. Blackish brown; except pronotum reddish brown; antennal 271 axis, l-ami basally and apically, last two abdominal segments, venter of metathorax, and legs along margins yellow.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight; radial origin inside head margin

0.5x when dorsally viewed. Antennal prominence distinct from vertex, gibbose; length 0.8x width, inserted between middle of eyes; rectangular dorsally, convex, narrowly impressed dorsomedially, anterior margin bisinuate; ventrally pentagonal, deeply concave, broadly impressed longitudinally, dorsolateral sides nearly carinate along antennal socket, widest near base. Antennal sockets rectangular, separated by

O.lx width of epistomal margin and 0.5x distance between eye and antennal socket. Antennifer insertion elevated medially. Interocular distance 2.8x width eye dorsally. Vertex transversely flat, setae decumbent; depression of tentorial maculae indistinct. Tentorial maculae separate, deep, distinct; at base of antennal prominence.

Frons transversely convex, indistinct from lower surface of antennal prominence, transverse, height 0.6x width, shortly rostrate apically, height rostrum 0.4x width epistoma. Epistomal margin broadly emarginate, width 0.5x frontal distance between eyes. Labrum transverse, rectangular, length 0.5x width, apical margin truncate.

Maxillary palps 4 segmented, palpomeres 1-3 cylindrical, diameter nearly equal to length; length palpomere 4 nearly equal 3, diameter at base less than 0.3x palpomere 3, directed internally, acuminate. Labial 272 palps 1 segmented, digitiform. Gula transversely rectangular; length

0.5x width apical margin; inserted anteriorad posterior margin of postgena. Antennae flabel late, length 0.7x body length; rami between

20-24x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, semi-decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically subequal length; pedicel cylindrical, short, transverse, not compressed, length 0.8x length of antennomere 3; antennomere 3 flabel late with ramal insertions basal, slightly compressed, not adnate to base of antennomere

4, length subequal to antennomere 4 ; antennomere 4 flabel late with ramal insertions medial, more strongly compressed; antennomeres 5-10 compressed, extent of compression decremental to apex, flabeliate with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior width 0.6x basal width; median longitudinal carina absent anteriorly, poorly defined and divided posteriorly, median fovea Q.5-0.9x pronotal length, closing at posterior margin; pronotal stemmata present, anterior pair on anterior margin and 10 stemmatal diameters from lateral margins, posterior pair on posterior margin and 4 stemmatal diameters from lateral margin; all discal margins swollen; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect-explanate, straight longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, but anterior 0.3x inflated allowing reception of head, setation decumbent; hypomeron concave, height 0.6x width, anterior and posterior margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, lateral arms acutely separated, length 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrately transverse, lateral margins parallel, apex deeply to shallowly emarginate, emargination 0.1-0.3x scutellar length, apices acute and not divergent. Mesospiracle prominent, peritreme exposed, orifice slightly compressed. Mesepisternum elongate, posterior margin slightly convexly arcuate. Mesepimeron trianguliform; width 0.3x and height O.flx mesepisternum; posterior margin straight, acutely angulate ventrally, slightly reflexed, evenly rounded. Mesosternum transverse, length 0.5x width, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base.

Elytra ligulate, strongly dehiscent from apical 0.3x elytral length, slightly reconvergent at apex, apices acute; length 9x pronotal length; quadricostate, costa 2 distinct basal 0.3x elytral length, costa 4 and 8 distinct 0.8x to near elytral apex, costa 6 faintly distinct middle 0.8x elytral length; disk irregularly scabrous-punctate; setation semi- decumbent, sparse, uniformly distributed. Metepisternum with ventral margin increasingly arcuate posteriorly. Metepimeron with ventral margin arcuate. Metasternum incompletely divided medially by longitudinal sulcus to near mesosternum. Legs elongate, setation moderately dense, semi-decumbent, uniformly distributed; coxae conical, pro- and mesotrochanteral socket 0.8x coxal length, metacoxa less elevated and more transversely oriented; trochanters elongate, length pro- and mesotrochanters 0.7x coxal length, metatrochanters equal to metacoxal length, femoral insertions slightly oblique and not offset 274

except metafemoral insertion moderately offset; femora compressed,

short, l.Qx trochanteral length, annulated basally; tibiae strongly

compressed, internal margin sinuate; tibial spines small and similar;

tarsi elongate, tarsomeres without laterally expanded plantar pads,

covered with decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9

medially divided longitudinally; tergite 10 shield shaped. Sternites

rectangular, wider than long, length 1-8 subequal, but narrowing

uniformly posteriorad; sternite 7 entire. Spiracles in pleural

membrane, located submedially from apical margin of tagma. Aedeagus

without parameres; basal piece dorsoventrally compressed, broad, length almost 0.5x length of median lobe (Fig. 36 a-c); median lobe awl-shaped,

membranous ventrally D.6x its length; internal sac not eversible; flagellum not present.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The intraspecific variation in this species is more

extreme than found in any North American temperate species. Whether

this is the result of heterochrony or some other phenomenon is unknown,

but many of the character states that vary appear to be those affected

by paedomorphosis as demonstrated in the Thvlodrias study. For example, the median cell of the pronotum is nearly absent in some 275 specimens and varies in length from half to most of the pronotal length.

The scutellum is variously emarginate from almost truncate to 0.3x its length. Again, the elytral costae 1,5, and 7 are suggested by a broken zigzag reticulation in some specimens, but not all. Future studies should investigate this variation with reference to its ontogeny. Possibly, this species varies more because some characters have developmental cusps which are more sensitive to paedomorphic shifts than others.

In addition to this extreme morphological variation, coloration in this species varies as in other lycids. Thus, the extent of yellow brown coloration of the legs varies from individual to individual.

MATERIAL EXAMINED. VENEZUELA: Territory Federal Amazonica, Cerro de la

Neblina, 0”50'N, 68'9'W, at base camp, 13-20 February 1984, D. Davis and

T. McCabe (1-MIZA, 1-NMNH); Same locality, 21-29 February 1984, D. Davis and T. McCabe (1-NMNH); Same locality, 0 ’50'N, at base camp,

27 January 1985, at blacklight, P. J. and P. M. Spangler, R. A.

Faitoute, W. E. Steiner (1-MIZA); Same locality, 26-31 January 1985, P.

J. and P. M. Spangler, P. Faitoute, W. E. Steiner (1-NMNH, 1 MIZA); Same locality, 1-9 February 1985, P. J. and P. M. Spangler, P. Faitoute, W.

E. Steiner (2-NMNH, 2 MIZA, 3—RSMC); Same locality, 10-20 February 1985,

P. J. and P. M. Spangler, P. Faitoute, W. E. Steiner (2-NMNH including type, 2-MIZA, 1 BMNH, 1-MNHP, 1-ZMHB); Same locality, 21-24 February

1985, P. J. and P. M. Spangler, P. Faitoute, W. Steiner (1-NMNH, 1-

MIZA); Same locality, 21-25 February 1985, P. J. and P. M. Spangler, P.

Faitoute, W. Steiner (2 MIZA, 2-NMNH). 276

ETYMOLOGY. The specific epithet refers to the type location. The base camp where most specimens were captured was in a rain forest on the side of the tepui, Cerro de la Neblina. 277

Pseudacroleptus obscuricolor Pic

Fig. 39

Pseudacro1eptus obscuricolor Pic, 1911:165. Kleine, 1933:165.

Blackwelder, 1945:348.

TYPE: MNHP, a card mounted male without left antenna, but in good condition otherwise, with the following label data: Cwhite printed

label with lined margins] "S. Antonio da Barra \\ Pr. de Bahia \\

Gounelle 11-12.88;" Chandwritten white labels] "Acroleptus ou \\ genus voisin sp. \\ nov. o;11 "29;" "Acroleptus \\ n. sp.;" "type;" Cred printed label! "TYPE;" [blue printed label] "Museum Paris \\ Coll. M.

Pic.;" Chandwritten white label] "Pseuacroleptus \\ obscuricolor \\

Pic. "

DIAGNOSIS. This species can be separated from other known members of the genus by its unicolorous dark coloration, the bicostate elytra, and

the weakly reconvergent elytral apices.

DISTRIBUTION. Known only from the type locality near Antonio de Barra

in the state of Bahia, Brasil.

DESCRIPTION (Male): Small, slender, elongate, length 5.0 mm., width 1.3 mm at humerus. Color blackish brown.

Head. Transverse, widest at eyes, narrowing slightly caudad; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical,

projecting laterally when viewed dorsally; hind margin shallowly

emarginate, nearly straight. Antennal prominence distinct from vertex,

gibbose; length unknown width, inserted between middle of eyes;

rectangular dorsally, convex, narrowly impressed dorsomedially;

ventrally pentagonal, deeply concave, broadly impressed longitudinally,

doroslateral sides nearly carinate along antennal socket, widest near

base. Antennal sockets rectangular. Antennifer insertion elevated

medially. Vertex transversely flat, setae decumbent; depression of

tentorial maculae indistinct. Tentorial maculae separate, deep,

distinct; at base of antennal prominence. Frons transversely convex,

indistinct from lower surface of antennal prominence, transverse,

shortly rostrate apically. Epistomal margin broadly emarginate.

Labrum transverse, rectangular, length 0.5x width, apical margin

truncate. Maxillary palps 4 segmented, palpomere 4 acuminate. Labial

palps 1 segmented, digitiform. Antennae flabel late, 0.7x body length; rami between 20-24x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, semi- decumbent on all antennomeres, sealiform setae absent; scape obconic;

pedicel cylindrical, short, transverse, not compressed, length nearly

equal length of antennomere 3; antennomere 3 flabel late with ramal

insertions basal, slightly compressed; antennomere 4 flabellate with

ramal insertions medial, more strongly compressed; antennomeres 5-10

compressed, extent of compression decremental to apex, flabellate with ramal insertions apical; antennomere 11 without distinguishable axis.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior width 0.6x basal width; median longitudinal carina absent anteriorly, poorly defined and divided posteriorly, median fovea 0.4x pronotal length, closing at posterior margin; margins swollen; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect-explanate, straight longitudinally; posterior margin bisinuate; posterior angles acute, expanded laterally; disk flat, but anteriorly inflated allowing reception of head, setation decumbent; hypomeron concave; prosternal bridge Y-shaped, length 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, lateral margins parallel, apex shallowly emarginate.

Mesospiracle prominent, peritreme exposed, orifice slightly compressed.

Mesepimeron reflexed. Mesosternum transverse. Mesocoxal separation lx mesocoxal diameter at base. Elytra iigulate, strongly dehiscent from apical O.lx elytral length, slightly reconvergent at apex, apices acute; length lOx pronotal length; bicostate, costa 4 distinct basal

0.5x elytral length, costa 8 distinct 0.9x elytral length; disk irregularly scabrous-punctate; setation semi-decumbent, sparse, uniformly distributed. Metasternum incompletely divided medially by longitudinal sulcus to near mesosternum. Legs elongate, setation moderately dense, semi-decumbent, uniformly distributed; coxae conical; trochanters elongate, femoral insertions slightly oblique and offset; femora compressed, short; tibiae strongly compressed, internal margin sinuate; tarsi elongate, tarsomeres without laterally expanded plantar pads; covered with decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad. Sternites rectangular, 280 wider than long, length 1-8 subequal, but narrowing uniformly posteriorad. Aedeagus unknown.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

DISCUSSION. The type was examined, but the genitalia were not removed and many of the characters used here were developed afterwards. Thus, the description is incomplete and, although the type must be reexamined, the characters in the key are sufficient to distinguish it from known congenerics.

MATERIAL EXAMINED. Brazil, Prov. de Bahia, S. Antonio da Barra,

November-December 1808 (1-NMHP, holotype).

ETYMOLOGY. The specific epithet undoubtedly refers to the type specimen's brownish black color. 281

Pseudacropletus sinuatis Pic

Pseudacroleptus sinuatis Pic, 1911:166. Kleine, 1933:34.

Blackwelder, 1945:348.

TYPE: MNHP, a card mounted male without hind legs and right antenna and with the following label data: Cwhite printed label! "Julliet;"

"Guyane francaise \\ Charven;" Cwhite printed label! "Bas Maroni \\

Coll. Le Moult;" Chandwritten white label! "n m;" Chandwritten white label! "type;" Cred printed label! "TYPE;" Cgreen-blue printed label!

"Museum Paris \\ Coll. M. Pic;" Chandwritten white label!

CPseudacroleptus \\ sinuatis Pic."

DIAGNOSIS. This species can most easily be differentiated by the characters in the key couplets. The sinuate margins of the elytra are not diagnostic.

DESCRIPTION (Male). Small, slender, elongate, but expanded distally

0.6x elytra! length; length 6.0 mm, width at humerus 1.3 mm. Blackish brown; except pronotum reddish brown.

Head. Transverse, widest at eyes, narrowing slightly caudad, disk shining; setae sparse, decumbent. Genal margin short, length less than eye length, uniformly narrowing when viewed dorsally. Eyes hemispherical, projecting laterally when viewed dorsally; hind margin shallowly emarginate, nearly straight. Antennal prominence distinct from vertex, gibbose, inserted between middle of eyes; rectangular dorsally, convex, narrowly impressed dorsomedially; ventrally pentagonal, deeply concave, broadly impressed longitudinally, dorsolateral sides nearly carinate along antennal socket, widest near base. Antennal sockets rectangular. Antennifer insertion elevated medially. Vertex transversely flat, setae decumbent; depression of tentorial maculae indistinct. Tentorial maculae separate, deep, distinct; at base of antennal prominence. Frons transversely convex, indistinct from lower surface of antennal prominence, shortly rostrate apically. Epistomal margin broadly emarginate. Labrum transverse, rectangular, length 0.5x width, apical margin truncate. Maxillary palps 4 segmented. Labial palps 1 segmented, digitiform. Antennae flabel late, length 0.7x body length; rami between 15~20x axial length and their compression perpendicular to axial compression; setation sparse, uniformly distributed, semi-decumbent on all antennomeres, scaliform setae absent; scape obconic, width apically subequal length; pedicel cylindrical, short, transverse, not compressed, length less than length of antennomere 3; antennomere 3 flabellate with ramal insertions medial, slightly compressed, not adnate to base of antennomere 4, length subequal to antennomere 4 ; antennomere 4 flabellate with ramal insertions apical, more strongly compressed; antennomeres 5-10 compressed, extent of compression decremental to apex, flabellate with ramal insertions apical; antennomere 11 without axis distinguishable from ramus.

Thorax. Pronotum trapezoidal, length 0.5x basal width, anterior width 0.6x basal width; median longitudinal carina absent anteriorly, poorly defined and divided posteriorly, median fovea Q.5x pronotal length, closing at posterior margin; margins swollen; anterior margin slightly convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect-explanate, straight longitudinally; posterior margin strongly bisinuate; posterior angles acute, expanded laterally; disk flat, but anteriorly inflated allowing reception of head, setation decumbent; hypomeron concave; prosternal bridge Y-shaped, lateral arms V-shaped, length 0.5x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, lateral margins parallel, apex shallowly emarginate, apices acute and not divergent.

Mesospiracie prominent, peritreme exposed, orifice slightly compressed.

Mesosternum transverse. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, strongly dehiscent, slightly reconvergent at apex, apices acute; tricostate, costa 2 distinct basal 0.3x elytral length, costa 4- and 8 distinct to near elytral apex; disk irregularly scabrous-punctate; setation semi-decumbent, sparse, uniformly distributed. Metasternum incompletely divided medially by longitudinal sulcus to near mesosternum. Legs elongate, setation moderately dense, semi-decumbent, uniformly distributed; coxa conical; trochanters elongate; femora compressed, short; tibiae compressed, internal margin sinuate; tarsi elongate, tarsomeres without laterally expanded plantar pads, covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 10 shield shaped.

Sternites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriorad; sternite 7 entire. Aedeagus unknown. 284

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures ). Unknown.

DISTRIBUTION. Known from the type locality.

DISCUSSION. The type was examined, but the genitalia were not removed.

Many of the characters used in this study were developed after

examination of the type and, therefore, the type will have to be

reexamined.

MATERIAL EXAMINED. GUAYANE FRANCAISE, Bas Moroni, Charven, July 19, Le

Moult (1-MNHP, holotype).

ETYMOLOGY. Apparently named for the sinuate internal margins of the elytra, which is characteristic of known species of this genus and not unique to the species. 285

Pseudoceratoprion NEW GENUS

TYPE. Pseudoceratoprion colombiensis n. sp., ANSP.

DIAGNOSIS. The combination of strongly compressed and serrate antennomeres 3 - 10, antennomere 3 at least 3x the length of 2, and the flanged median genal margin which is vertically oriented will separate this genus from all other leptolycines.

DISTRIBUTION. This genus is known only from northern South America, but it is likely that it will be found further south and, perhaps, in

Central America.

DESCRIPTION (Male): Size moderate, slender, elongate, length 2.9-5.6 mm., width 0.7-1.1 mm at humerus. Brown to brownish black, with or without whitish-yellow maculation.

Head. Elongate, widest at eyes, then narrowing to posterior margin, shining; setae sparse, semi-decumbent. Genal margin short, length less than eye length, narrowing posteriorly when viewed dorsally.

Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly and deeply emarginate; dorsally radial origin inside head margin 0.5-0.7x radius. Antennal prominence distinct from vertex, shelf-like; length 0.6-0.8x width, inserted between middle of eyes; dorsally pentagonal to quadrate and barely produced medially, deeply and distinctly sulcate dorsomedially; ventrally pentagonal, concave, sides uniformly and convexly arcuate, widest near base. Antennal sockets oval to rectangular, separated by O.lx width of epistomai margin and lx distance between eye and antennal socket. Antennifer elevated medially, not appearing basomedial. Interocular distance 2-2.5x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae decumbent to semi-decumbent; depression of tentorial maculae shallow to deep and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transverse, height 0.3-0.4x width. Epistomai margin broadly and shallowly to deeply emarginate, width 0.4-0.6x frontal distance between eyes.

Labrum transverse, rectangular, length approximately 0.5x width, apical margin truncate. Maxillary palps 4 segmented, palpomere length 1 0.5x length 3, palpomere 2 2x palpomere 1, diameter 1-3 lx length 3; length palpomere 4 lx palpomere 3, diameter palpomere 4 at base 0.5x palpomere

3, palpomere 4 acuminate. Labial palps absent. Gula transverse, short, linear to trapezoidal; length 0.2-0,3x width apical margin.

Antennae serrate, 0.7-lx body length; setation sparse to moderately dense, uniformly distributed, decumbent on all antennomeres, scaliform setae present; scape obconic, width apically 0.6-lx length; pedicel cylindrical, short, transverse, not compressed, length 0.2-0.3x antennomere 3; antennomere 3 not adnate with base antennomere 4, length

0.9-lx antennomere 4; antennomeres 4-10 elongate-serrate to filiform- serrate, uniformly expanded apically to immediately expanded and parrallel to apex, strongly compressed, trapezoidal to trianguliform with apical margin excavate or not; antennomere 11 fusiform, compressed, 287

length 1.4-2x antennomere 3.

Thorax. Pronotum trape2oidal, transverse, length 0.4-0.5x basal

width, anterior width 0.5-0.6x basal width; median longitudinal carina

well defined, median fovea slightly more distinct and 0.7x pronotal

length, carinae narrowly separated, fovea closed at posterior margin;

pronotal stemmata absent; all discal margins variously swollen; anterior

margin straight to slightly convexly arcuate, anterior angles obtuse;

lateral margin complete, foliaceous, erect, concavely arcuate; posterior

margin very strongly bisinuate; posterior angles acute, expanded

laterally 0.8-0.9x width at elytral humerus; disk flat to slightly

transversely arcuate, discal setae semi-decumbent; hypomeron shallowly

concave, height 0.5x length; all margins swollen, hypomeral stemmata

absent; prosternal bridge T- or Y-shaped, anterior margin broadly to

slightly and narrowly emarginate, length 0.3x width, intercoxal process

not carinate. Scutellum pentagonal, distally quadrate to

trianguliform, lateral margins parallel to divergent, apex deeply

emarginate, emargination 0.3-0.5x scutellar length, apices acute and not

divergent. Mesospiracle obscure to prominent, peritreme exposed and

compressed or not. Mesepisternum elongate, posterior margin straight.

Mesepimeron trianguliform, width 0.3-0.5x and height 0.7-0.8x

mesepisternum; posterior margin not reflexed, straight; angulate at

venter. Mesosternum transverse, posterior margin convexly arcuate.

Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate,

slightly dehiscent variously at scutellum to 0.3x elytral length, reconverging or not at apices; length 9-lOx pronotal length; bicostate,

costa 4 distinct to strongly elevated basal 0.9x elytral length, humeral costa distinct 0.9x elytral length; disk scabrous-punctate to regularly

reticulate; setation semi-erect, sparse to moderately dense, uniformly

arranged. Metepisternum with ventral margin increasingly arcuate

anteriorly. Metepimeron with ventral margin sinuate to nearly

straight. Metasternum completely divided medially by longitudinal

sulcus. Legs elongate, setation sparse, decumbent to semi-decumbent,

uniformly distributed; coxae conical, trochanteral socket 0.4-0.6x coxal

length; trochanters elongate, length pro- and mesotrochanters 0.9-lx

coxal length, metatrochanters lx metacoxal length, all femoral

insertions oblique and slightly offset; femora strongly compressed,

straight, short, 3-3.5x trochanteral length, annulated basally; tibiae

strongly compressed, internal margin sinuate, external margin foliaceous

or not; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi—decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8

subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located at caudal 1/3 at edge sternite. Aedeagus with parameres; paramere length 0.3~0.8x median lobe, serrate or notched apicolaterally; basal piece arcuate, slightly or not dorsoventrally compressed, length 0.3-0.7x length of median lobe; median lobe tubular, unmodified or apically expanded, ventrally membranous 0.1-0.8x length, membrane apical or subapical nearly 0.2x from apex, internal sac not eversible, flage11urn absent. 289

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures ). Unknown.

DISCUSSION.

ETYMOLOGY. Psuedo. Latin, means false and refers to the superficial resemblance of this genus to Ceratoprion Gorham. KEY TO THE SPECIES OF Pseudoceratoprion (ADULT MALES)

Lateral margins of antennomeres parallel, not expanded

apically; elytra regularly reticulate; Brazil ......

...... Pseudoceratoprion brasi1iensis n. sp. (p. 296 )

Lateral margins of antennomeres divergent, antennomeres

trianguliform; elytra not regularly reticulate; ...... 2

Apical margin of antennomeres emarginate; brownish black

species with apices of ultimate antennomeres whitish yellow;

Equador to Venezuela ......

...... Pseudoceratoprion colombiensis n . sp. (p. 301)

Apical margin of antennomeres entire, not emarginate;

uniformly brown species; Venezuela ......

...... Pseudoceratoprion bechvnae n. sp. (p. 291) For DESCRIPTION or DIAGNOSIS:

Mouthparts ventral and reduced, mandibles not evident.

Antennae large, at least 0.5x length of body

Length pedicel less than 0.3x 3rd antennomere

Antennomeres 3-11 subequal, strongly serrate;

Prosternal bridge Y-shaped

Spiracles in pleural membrane caudal 0.3x at edge sternite.

Posterior sternite reflexed dorsally posterior Q.5x.

Apical tergite rounded distally.

Parameres present. 291

Pseudoceratoprion bechvnae NEW SPECIES

Fig. 40 a-d

TYPE. MIZA, a male in good condition with the following label data:

[white printed labels! "Venezuela Her- \\ ida, Carbonera \\ 2600 m \\

28-VI-68;" "J. & B. Bechyne \\ leg."

DIAGNOSIS. This species is most easily separated from congenerics by

the trianguliform antennomeres with apical margins emarginate, and the

absence of maculation on the ultimate antennomeres. Additionally, the

bicostate elytra are irregularly punctate-reticulate.

DISTRIBUTION. Known only from the type locality in the mountains of western Venezuela.

DESCRIPTION (Male): Size moderate, slender, elongate, length 4.4-4.6 mm., width 1.0-1.1 mm at humerus. Brownish black.

Head. Elongate, widest at eyes, then narrowing to posterior margin, shining; setae sparse, semi-decumbent. Genal margin short,

length less than eye length, narrowing posteriorly when viewed dorsally.

Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly and deeply emarginate; dorsally radial origin inside head margin 0.7x radius. Antennal prominence distinct from vertex,

shelf-like; length 0.6x width, inserted between middle of eyes; quadrate dorsally, barely produced medially, deeply and distinctly sulcate dorsomedially; ventrally pentagonal, concave, sides uniformly and 292 convexly arcuate, widest near base. Antennal sockets rectangular,

separated by O.lx width of epistomai margin and lx distance between eye and antennal socket. Antennifer elevated medially, not appearing basomedial. Interocular distance 2x width eye dorsally. Vertex slightly arcuate, nearly transversely flat, setae semi-decumbent; depression of tentorial maculae shallow and broadly defined, from base of antennal prominence to posterior eye margin, more distinct anteriorly. Tentorial maculae separate, deeply distinct; at base of antennal prominence. Frons transversely flat, distinct from lower surface of antennal prominence, transverse, height 0.4x width.

Epistomai margin broadly and deeply emarginate, width 0.6x frontal distance between eyes. Labrum transverse, rectangular, length approximately 0.5x width, apical margin truncate. Maxillary palps 4 segmented, palpomere length 1 0.5x length 3, palpomere 2 2x palpomere

1, diameter 1-3 lx length 3; length palpomere 4 lx palpomere 3, diameter palpomere 4 at base 0.5x palpomere 3, palpomere 4 acuminate.

Labial palps absent. Gula transverse, short, linear; length 0.3x width apical margin. Antennae serrate, 0.7x body length; setation sparse, uniformly distributed, decumbent on all antennomeres, scaliform setae present; scape obconic, width apically 0.6x length; pedicel cylindrical, short, transverse, not compressed, length 0.2x antennomere 3; antennomere 3 not adnate with base antennomere 4, length 0.9x antennomere 4; antennomeres 4-10 elongate-serrate, slightly expanded apically, strongly compressed, trianguliform with apical margin excavate; antennomere 11 fusiform, compressed, length 1.4x antennomere

3. Thorax. Pronotum trapezoidal, transverse, length 0.4x basal width, anterior width 0.5x basal width; median longitudinal carina well defined, median fovea slightly more distinct and 0.7x pronotal length, carinae narrowly separated, fovea closed at posterior margin; pronotal stemmata absent; all discal margins swollen; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, erect, concavely arcuate; posterior margin very strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, discal setae semi-decumbent; hypomeron shallowly concave, height

0.5x length; all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, anterior margin broadly emarginate, length 0.3x width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, lateral margins parallel, apex deeply emarginate, emargination

0.5x scutellar length, apices acute and not divergent. Mesospiracle obscure, peritreme not exposed. Mesepisternum elongate, posterior margin straight. Mesepimeron trianguliform, width 0.3x and height 0.8x mesepisternum; posterior margin not reflexed, straight; angulate at venter. Mesosternum transverse, posterior margin convexly arcuate.

Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, slightly dehiscent at 0.3x elytral length, reconverging at apices; length lOx pronotal length; bicostate, costa 4 distinct basal 0.9x elytral length, humeral costa distinct 0.9x elytral length; disk irregularly punctate-reticulate, basally obscure and distally more distinct; setation semi-erect, sparse, uniformly arranged.

Metepisternum with ventral margin increasingly arcuate anteriorly.

Metepimeron with ventral margin sinuate. Metasternum completely 294 divided medially by longitudinal sulcus. Legs elongate, setation

sparse, semi-decumbent, uniformly distributed; coxae conical, trochanteral socket 0.6x coxal length; trochanters elongate, length pro- and mesotrochanters 0.9x coxal length, metatrochanters lx metacoxal

length, all femoral insertions oblique and slightly offset; femora strongly compressed, straight, short, 3x trochanteral length, annulated basally; tibiae strongly compressed, internal margin sinuate; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located at caudal 1/3 at edge sternite. Aedeagus with parameres; paramere length 0.4x median lobe, serrate laterally; basal piece arcuate, dorsoventrally compressed, length 0.5x length of median lobe (Fig. 40 a-c); median lobe tubular, apically expanded, ventrally membranous O.lx length and subapical nearly 0.2x from apex, internal sac not eversible, flageilum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown.

MATERIAL EXAMINED. VENEZUELA, Merida, Carbonera, 2600 m., 28 June

1968 (1-MIZA); Merida, Culata, 21 February 1969, J. & B. Bechyne / leg. 295

{1-RSMC >.

ETYMOLOGY. Named in honor of Mrs. Bechyne who helped collect half the type series and was most helpful during my visits to Venezuela. 296

Pseudoceratoprion brasj1iensis NEW SPECIES

Fig. 41 a-d

TYPE. CNCI, a male in good condition with the following label data:

Cwhite printed labelD "BRASIL: Rio de \\ Janiero, Munudu \\ Campos VIII-

1978 \\ M. Alvarenga."

DIAGNOSIS. This species is most easily separated from congenerics by the non-trianguliform antennomeres which appear to be compressed and fi1irform-serrate. Additionally, the bicostate elytra are regularly reticulate.

DISTRIBUTION. Known only from the type locality in Rio de Janiero,

Brazi1.

DESCRIPTION (Male): Size small, slender, elongate, length 2.9 mm., width

0.7 mm at humerus. Brownish black.

Head. Elongate, widest at eyes, then narrowing to posterior margin; setae sparse, decumbent. Genal margin short, length less than eye length, narrowing posteriorly when viewed dorsally. Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly and shallowly emarginate; dorsally radial origin inside head margin 0.5X radius. Antennal prominence distinct from vertex, shelf-like; length 0.8X width, inserted between middle of eyes; pentagonal dorsally, broadly and deeply sulcate dorsomedially; ventrally pentagonal, concave, sides uniformly and convexly arcuate, widest near base. Antennal sockets oval, separated by 0.1X width of epistomai

margin and IX distance between eye and antennal socket. Antennifer

elevated medially, not appearing basomedial. Interocular distance 3X

width eye dorsally. Vertex slightly arcuate, nearly transversely flat,

setae semi-decumbent; depression of tentorial maculae shallow and

broadly defined, from base of antennal prominence to posterior eye

margin, more distinct anteriorly. Tentorial maculae separate, deeply

distinct; at base of antennal prominence. Frons transversely flat,

distinct from lower surface of antennal prominence, transverse, height

0.4X width. Epistomai margin broadly and shallowly emarginate, width

0.8X frontal distance between eyes. Labrum transverse, rectangular,

length approximately 0.5X width, apical margin truncate. Maxillary

palps 4 segmented, palpomere length 1 0.5X length 3, palpomere 2 2X

palpomere 1, diameter 1-3 IX length 2; length palpomere 4 IX palpomere

3, diameter palpomere 4 at base less than 0.5X palpomere 3, palpomere 4

acuminate. Labial palps absent. Gula transverse, short, linear;

length 0.2X width apical margin. Antennae filiform-serrate, 0.6X body

length; setation moderately dense, uniformly distributed, decumbent on all antennomeres, scaliform setae present; scape obconic, width apically

IX length; pedicel cylindrical, short, transverse, not compressed,

length 0.3X antennomere 3; antennomere 3 not adnate with base antennomere 4, length IX antennomere 4; antennomeres 4-10 filiform-

serrate, expanded immediately, lateral margins parallel to apices,

strongly compressed, width 2X depth; antennomere 11 fusiform, compressed, length 2X antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.4X basal width, anterior width 0.5X basal width; median longitudinal carina

indistinctly defined, median fovea slightly more distinct and 0.7X pronotal length, carinae narrowly separated, fovea closed at posterior margin; pronotal stemmata absent; anterior and lateral discal margins strongly swollen, posterior margin weakly swollen; anterior margin straight, anterior angles obtuse; lateral margin complete, foliaceous, erect, concavely arcuate; posterior margin very strongly bisinuate; posterior angles acute, expanded laterally 0.9X width at elytral humerus; disk slightly transversely arcuate, discal setae semi- decumbent; hypomeron shallowly concave, height 0.5X length; all margins swollen, hypomeral stemmata absent; prosternal bridge T-shaped, anterior margin slightly and narrowly emarginate, length 0.3X width, intercoxal process not carinate. Scutellum pentagonal, distally quadrate, lateral margins parallel, apex deeply emarginate, emargination 0.3X scutellar length, apices acute and not divergent. Mesospiracle prominent, peritreme exposed, slightly compressed. Mesepisternum elongate, posterior margin straight. Mesepimeron trianguliform, width 0.5X and height 0.7X mesepisternum; posterior margin not reflexed, straight; angulate at venter. Mesosternum transverse, posterior margin convexly arcuate. Mesocoxal separation IX mesocoxal diameter at base. Elytra ligulate, slightly dehiscent at scutellum, not reconverging at apices; length 9X pronotal length; bicostate, costa 4 strongly elevated basal

0.9X elytral length, humeral costa distinct 0.9X elytral length; disk regularly reticulate, basally obscure to distinctly transverse or square areolets distally; setation semi-erect, moderate, uniformly arranged.

Metepisternum with ventral margin increasingly arcuate anteriorly. 299

Metepimeron with ventral margin sinuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0.5X coxal length; trochanters elongate, length pro-and mesotrochanters 0.9X coxal length, metatrochanters IX? metacoxal length, all femoral insertions oblique and slightly offset; femora strongly compressed, straight, short, 3.5X trochanteral length, annulated basally; tibiae strongly compressed, internal margin sinuate, external margin foliaceous; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues s i mp 1e .

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located at caudal 1/3 at edge sternite. Aedeagus with parameres; paramere length 0.3X median lobe, notched apically; basal piece small, arcuate, slightly dorsoventrally compressed, length

0.3X length of median lobe (Fig. 41 a-c); median lobe tubular, unmodified, ventrally membranous apical 0.IX length, internal sac not eversible, flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown. DISCUSSION. A distinctive character state of the type is the produced posterior angles of the pronotum, but this may be an artifact of preservation or development and should be used with caution until more material is available. Again, costa 2 and costa 6 are faintly indicated by occassional swellings along the probable location of those costae, but are seen only with difficulty (or imagination?) and were not used. A larger series may present individuals with costae more distinct which might, thus, not be recognized as conspecific.

MATERIAL EXAMINED. BRASIL: Rio de Janiero, Munudu Campos VII1—1978, M.

Alvarenga (1- CNCI, holotype).

ETYMOLOGY. Named after the country in which it was found. 301

Pseudoceratoprion colombiensis NEW SPECIES

Fig. 42 a-d

TYPE. ANSP, a male in good condition except a broken left antenna with

the following label data: [white printed labels] "Cincinati \\ Trail

\\ VI11-19-2;" "Sierra de \\ S. Lorenzo \\ 4-6000 ft.;" "Magdalena, \\

Colombia \\ F. R. Mason."

DIAGNOSIS. This species is most easily separated from congenericsby the combination of the trianguliform antennomeres with apical margins straight and not emarginate, the absence of maculation on the ultimate antennomeres, and the bicostate elytra are irregularly scabrous- punctate.

DISTRIBUTION. Known from Ecuador, Colombia, and Venezuela.

DESCRIPTION (Males): Si2e moderate, slender, elongate, length 4.7-5.6 mm., width 0.6-0.7 mm at humerus. Brown, except apical antennomere whitish yellow; occasionally posterior angles of prothorax, elytral humerus or penultimate antennomere whitish-yellow.

Head. Elongate, widest at eyes, then narrowing to posterior margin, shining; setae sparse, semi-decumbent. Genal margin short, length less than eye length, narrowing posteriorly when viewed dorsally.

Eyes hemispherical, projecting anteriolaterally when viewed dorsally; hind margin broadly and shallowly emarginate; dorsally radial origin inside head margin 0.5x radius. Antennal prominence distinct from vertex, shelf-like; length 0.6x width, inserted between middle of eyes;

quadrate dorsally, not produced medially, distinctly sulcate

dorsomedially; ventrally pentagonal, concave, sides uniformly and

convexly arcuate, widest near base. Antennal sockets rectangular,

separated by O.lx width of epistomai margin and lx distance between eye

and antennal socket. Antennifer elevated medially, not appearing

basomedial. Interocular distance 2.5x width eye dorsally. Vertex

slightly arcuate, nearly transversely flat, setae semi-decumbent;

depression of tentorial maculae deep and broadly defined, from base of

antennal prominence to posterior eye margin, more distinct anteriorly.

Tentorial maculae separate, deeply distinct; at base of antennal

prominence. Frons transversely flat, distinct from lower surface of

antennal prominence, transverse, height 0.3x width. Epistomai margin

broadly and shallowly emarginate, width 0.6x frontal distance between

eyes. Labrum transverse, rectangular, length 0.5x width, apical margin

truncate. Maxillary palps 4 segmented, palpomere length 1 0.5x length

3, palpomere 2 2x palpomere 1, diameter 1-3 lx length 2; length palpomere 4 lx palpomere 3, diameter palpomere 4 at base less than 0.5x

palpomere 3, palpomere 4 acuminate. Labial palps absent. Gula

transverse, short, trapezoidal; length lx width apical margin.

Antennae serrate, lx body length; setation sparse, uniformly distributed, decumbent on all antennomeres, scaliform setae present;

scape obconic, width apically 0.7x length; pedicel cylindrical, short,

transverse, not compressed, length 0.2x antennomere 3; antennomere 3 not adnate with base antennomere 4, length lx antennomere 4; antennomeres 4-

10 serrate, slightly expanded apically, strongly compressed, 303 trianguliform with apical margin straight and not excavate; antennomere

11 fusiform, compressed, length 1.8x antennomere 3.

Thorax. Pronotum trapezoidal, transverse, length 0.5x basal width, anterior width 0.6x basal width; median longitudinal carina well defined, median fovea slightly more distinct and 0.5x pronotal length, carinae narrowly separated, fovea closed at posterior margin; pronotal stemmata absent; all discal margins swollen; anterior margin convexly arcuate, anterior angles obtuse; lateral margin complete, foliaceous, erect, concavely arcuate; posterior margin very strongly bisinuate; posterior angles acute, expanded laterally 0.8x width at elytral humerus; disk flat, discal setae semi-decumbent; hypomeron shallowly concave, height 0.5x length; all margins swollen, hypomeral stemmata absent; prosternal bridge Y-shaped, anterior margin broadly and deeply emarginate, length 0.5x width, intercoxal process not carinate.

Scutellum pentagonal, distally trianguliform, lateral margins divergent, apex deeply emarginate, emargination 0.3x scuteliar length, apices acute and not divergent. Mesospiracle obscure, peritreme not exposed.

Mesepisternum elongate, posterior margin straight. Mesepimeron trianguliform, width 0.5x and height 0.7x mesepisternum; posterior margin not reflexed, straight; angulate at venter. Mesosternum transverse, posterior margin convexly arcuate. Mesocoxal separation lx mesocoxal diameter at base. Elytra ligulate, slightly dehiscent at

0.3x elytral length, reconverging at apices; length 9x pronotal length; bicostate, costa 4 distinct basal 0.9x elytral length, humeral costa distinct 0.9x elytral length; disk irregularly scabrous-punctate basally to obscurely and irregularly reticulate apically; setation semi- 304 decumbent, sparse, uniformly arranged. Metepisternum with ventral margin increasingly arcuate anteriorly. Metepimeron with ventral margin sinuate. Metasternum completely divided medially by longitudinal sulcus. Legs elongate, setation sparse, decumbent, uniformly distributed; coxae conical, trochanteral socket 0,4x coxal length; trochanters elongate, length pro- and mesotrochanters lx coxal length, metatrochanters 0.9x metacoxal length, all femoral insertions oblique and slightly offset; femora strongly compressed, straight, short, 2.3x trochanteral length, annulated basally; tibiae strongly compressed, internal margin nearly straight; tarsi short, tarsomeres without laterally expanded plantar pads; covered with semi-decumbent bristling setae; ungues simple.

Abdomen. Tergites rectangular, wider than long, length 1-8 subequal, but narrowing uniformly posteriad; tergite 8 entire; tergite 9 medially divided longitudinally; tergite 10 shield shaped. Sternites

1-8 narrowing uniformly posteriorad; sternite 7 entire. Spiracles in pleural membrane, located at caudal 1/3 at edge sternite. Aedeagus with parameres; paramere length O.Qx median lobe, serrate apicolaterally; basal piece arcuate, not dorsoventrally compressed, length 0.7x length of median lobe (Fig. 42 a-c); median lobe tubular, unmodified, ventrally membranous 0.8x length, internal sac not eversible, flagellum absent.

DESCRIPTION (Females). Unknown.

DESCRIPTION (Immatures). Unknown. 305

MATERIAL EXAMINED. COLOMBIA: Magdalena, Sierra de San Lorenzo,

Cincinnati Trail, 1200-1600 m, 19 July 1920, F. R. Mason (1-ANSP, hoiotype).

ECUADOR: Provincia Napo, 12 km. SW Tena, 500 m., 8-11 July 1976, malaise trap, S. & J. Peck (1-CNCI).

VENEZUELA: Aragua, 24 km. N. Maracay, Henri Petier National Park, 1000 m., 24 December 1985, P. Kovarick and R. Jones (1-TAMU); Aragua, Rancho

Grande, 1100 m. 21 April 1953, J. A. Gonzales (1-RSMC); Aragua, Rancho

Grande, Portachuelo, 1100 m . , 28 May 1983 (1-MIZA).

ETYMOLOGY. Named for the country in which the first specimen of this species was seen. PHYLOGENY OF THE LEPTOLYCINI

The members of the Leptolycini are here hypothesized to be paedomorphic. Such a hypothesis does not imply, however, that all leptolycine females are "larviform." They may exhibit any degree of paedomorphic states from almost unaffected to strongly affected, but all are probably flightless. Because of the constraints of paedomorphic development on phylogenetic analysis as demonstrated by the Thvlodrias study, the resulting proposed changes in methodology were employed in the analysis of the Leptolycini and its relationships within the

Lycidae. This resulted in the elimination of a number of traditional character states from the analysis and indicated a number of others that should be scrutinized in the future.

Elimination of those character states which are presumably affected by paedomorphosis, suffers from a similar limitation of clique analysis. By removing non-congruent characters during clique analysis, all similarities of the removed character states are assumed to be homoplasious and not synapomorphic. Thus, potential information may be discarded, if these states are in fact not homoplasious (Farris, 1982).

In a like manner information may be lost by discarding characters thought to be affected by paedomorphosis. The two methods, however, differ by their approaches. Clique analysis is ad hoc in the sense that it removes those characters found to be statistically inconsistent with the rest of the data. It attempts to find the most parsimonious

306 307

distribution of the data given the assumptions of the evolutionary model

found in phylogenetic analysis (Hennig, 1966; Wiley, 1981; etc). These

assumptions may or may not reflect natural parsimony, assuming such

parsimony exists and is not a merely an expression of human reasoning

processes. On the other hand, the proposed removal of paedomorphic

states is an a priori assumption about the effect of development on certain characters and it attempts to recapture developmental

information as currently understood. Although it is unlikely that heterochrony is the only phenomenon currently not successfully

incorporated in the methodology of phylogenetic analysis, it has been demonstrated to be one such phenomenon. If the proper intent of phylogenetic analysis is the study of evolution, these discordant phenomena should not be ignored. After all, nothing in biology makes

sense except in the light of ontogeny.

Monophvlv of the Leptolycini. A hypothesis of the monophyly of the

Leptolycini is based on the loss of the laterally expanded plantar pads on all tarsomeres, the presence of pronotal stemmata

in Leptolvcus and Abrolvcus). and the dorsoventral oblique position of the metacoxae. Additionally, there is a trend in displacement of the mouth parts to a more posterior position behind the apex of the frons in the more derived members of the lineage, but this may result from it3 anomalous development.

Confounding this analysis, there are indications that the

Leptolycini may not be monophyletic. If these taxa have convergent character states due to heterochronic development, detection of this 308

trend with current methodology is very difficult. The proposed

methodological modifications suggest finding synapomorphies, but the

means of recognizing them within a lineage is not yet formalized. As a

consequence, the use of characters in the analysis may tend to appear ad

hoc. An attempt was made to minimize such arguments and, therefore,

some states that might have been discarded have been incorporated into

the analysis. In all cases, the discussion of characters that follows

mentions any reasons for exclusion in the analysis.

Lycid Relationships of the Leptolycini. The sister group of the

Leptolycini is here hypothesized to be members of the Calopterini.

This hypothesis is supported by few characters which are at times found

in other genera within the Lycidae, but the latter are thought to be

homoplasous by reduction or loss and not synapomorphic (Miller, in

prep. ). Because this hypothesis varies significantly from that

suggested by Crowson (1972) and apparently followed by Bocak and

Bocacova (1988:203, 1989), a discussion of that alternative hypothesis

is required.

Crowson (1972:49) suggested that the basal Lycidae might be found

in the assemblage including Lvropaeus Waterhouse, Pristolvcus Pic, and

Puliticola Mjorberg. In fact Kieine (1941) attempted to exclude the

latter genus from the Lycidae by arguments based on presumed

plesiomorphic states. Initially, my analyses of the relationships within the family indicated that these taxa did not share the

synapomorphies of the "more derived" Lycidae and, hence, were basal.

However, I continually encountered problems of conflicting character 309

states in all lineages, which produced high rates of homoplasy.

Additionally, Crowson's (1972:64) implication of the reexpression of the

lost wings of paedomorphic ancestors in an X-0 sex-determining system is

unsatisfying. Such re-expression would have to be dependent on some

sex-linked, concentration threshold mechanism, because most males do not have the development of their wings impaired, but females do. Although

Naisse's (1966, 1968, and 1969) work does support differential titers of

JH as predicted by this hypothesis, the karyotypes of the alate Lycidae apparently do not differ from the X-0 system (Smith and Virkki, 1978) that would be expected in support of it. Crowson (1972:69) closed his paper suggesting that future cantharoid specialists should study the problem of heterochrony (his neoteny). Therefore, the study of

Thvlodrias was indicated.

Also, during their ongoing studies of lycid relationships, Bocak and Bocakova (1989) elevated the monogeneric African Dexorini Kieine

(1933) to subfamilial status and elevated Lvropaeus Waterhouse, an endemic genus in the Oriental region, as a second tribe within the subfamily. This action is predicated on the the reduction in mouthparts and the extent of morphological divergence of the two lineages from other lycid subfamilies and each other (Bocak and

Bocacova, 1989:718). There is no doubt that both genera differ considerably from most other Lycidae and are easily characterized.

However, the decision was probably the result of not recognizing the implications of their probable paedomorphic nature.

Current studies in progress, using the proposed modifications in phylogenetic analysis for heterochrony, suggest that they are not in 310 fact closely related to each other, nor are either closely related to the Leptolycini. Although they share reduction and loss character states that might be thought synapomorphous, they also share synapomorphies of novelty with various differing taxa within the

Lycidae. Thus, the most parsimonious solution is that their paedomorphic development is independent and the apparent convergence of general facies.

In support of a hypothesis of independent origins, no females are known in any of these except the paedomorphic Puliticola paradoxa studied by Mjorberg and, now Leptolvcus heterocornis. There are numerous reports and some speculation that the remaining taxa are all paedomorphic, but no paedomorphic females have been described For example, Gravely (1915:361) suggested that Lvropaeus Waterhouse is

"larviform" and Bocak and Bocakova (1988:195; 1989:718) suggested that

Dexoris Waterhouse has flightless females. If these taxa are paedomorphic and the higher classification of the Lycidae is examined in light of the Thvlodrias study, the Dexorinae is found to be polyphyletic and each included genus has its closest common ancester in genera endemic to the same continental region. Clearly, more work needs to be done on this problem.

In the interim, I have provisionally resolved this issue by a preliminary analysis of the family using the immature stages as well as adults. The results to date support such independent origins because more homoplasy is explicable. Thus, the Leptolycini share the most recent common ancestry within the Calopterini. Whether continued recognition of the Leptolycini creates a paraphyletic Calopterini is not 311 yet known. Unfortunately, the state of our knowledge of lycid immatures is very poor and most of the calopterine immatures, like those of most of the family as a whole, are unknown. The above hypothesis might best be examined by a phylogenetic analysis of the Calopterini and eventually the entire family, but field work including collecting and life history studies would augment the database immeasurably.

In addition to the genera already discussed, there are a number of other disparate taxa within the Lycidae that may be paedomorphic.

Several genera currently placed within the Calopterini sensu lato were initially included in the Leptolycini, but sufficient evidence exists using the proposed modifications that their origins are independent, that they were excluded. These include the Brazilian Aporrhipis flexi1is Pascoe, which was originally described as a rhipiphorid. Also attributed to the South American Calopterini are an undescribed monotypic "genus" of uncertain afffinities, the monotypic

Flabellolycinella Kieine, and two species of Acroleptus Bourgeois - A. chevrolati Bourgeois and another undescribed species,. Again, the

Australian Metriorrhvnchus apterus Lea, which occurs in Queensland, is clearly only a paedomorphic member of that genus.

Parenthetically, geographic distribution supports either hypothesis. Two ad hoc arguments can be convincingly proposed to explain the data. One argument is that the paedomorphic lineages discussed here are all basal and their distribution is a relict basal fauna. Thus, the distribution of these taxa seems to suggest that the tropics might provide a refugium for basal taxa. The alternative argument supports a phylogenetic hypothesis of the independent origin of 312 paedomorphic taxa. Thus, the distribution of these taxa would seem to suggest that the tropics may readily allow the expression of paedomorphos i s.

Hvpothesized Sister Group. The sister group relationships of the

Leptolycini remain obscure, but were here hypothesised to be various members of the Calopterini (Calocladon Gorham, Cvrtopteron Bourgeois),

Emplectus Erichson, Eurrhacus Waterhouse, and Metapteron Bourgeois.

These genera were used as sister taxa in the analysis, but the relationships of those genera within the latter tribe are presently unresolved. The Calopterini is undoubtedly polyphyletic and current studies in progress are addressing this problem. This condition, in turn, is a reflection of the hypothesized relationships within the family as a whole which are in need of reanalysis. In addition there is substantial evidence that the continued recognition of the

Leptolycini may yield a paraphyletic Calopterini if all the Leptolycini are indeed paedomorphic and derived as here proposed. Further resolution of these problems is not possible at this time because it will require examination of more adult material and, more importantly, it will require associated larvae. As these larvae are rare in collections, it is not anticipated that the problem will be resolved immediately. Because work in progress at the family level may refute current hypotheses of relationships within the Leptolycini, unique specific epithets were chosen throughout the entire lineage for nomenclatural stability.

Synapomorphies supporting the outgroup hypothesis presented here 313

are the presence of a median longitudinal carina divided medially with

lateral prenatal carinae and anteriolateral longitudinal carina absent.

A hypothesis of monophyly based on a single character state may seem

imprudent, but the extreme homoplasy found throughout the Leptolycini

and Calopterini

the present time. The reassessment of the relationships indicated by

the current classification of the family is required because characters

often used to support it, while more easily seen, are perhaps too

variable within lineages to be of use.

Characters examined. The characters examined here for taxonomic

treatment and phylogentic analysis are often the same as traditional characters, but the consideration of paedomorphosis and the possibility

of convergent evolution required their re-examination and a change in perspective. Consequently, it has been necessary to discuss new characters and to implement a non - traditional and, hopefully, more

descriptive terminology for these states. Again, many of the characters that have been used to support higher taxa are quite variable within this tribe and sometimes within a single genus. The

implications of this variation are discussed in reference to future

studies of the higher relationships within the family. The characters examined, the rationale for their inclusion or exclusion, and their

states as used in the phylogenetic analysis is discussed. The character numbers following the character states are 314

Table 3. Character states used in the phylogenetic analysis of the Leptolycini and their hypothesi2ed outgroup.

Character Plesiomorphic Apomorphic

01 . Prosterna1 shape T- or Y- shaped Trianguliform 02. Antenna1 compression Compressed Not compressed 03. Pronotal stemmata Absent Present 04. Genal margin length Short Elongate 05. Mouthparts Anteriad Retracted 06. Frons length Elongate Short 07 Epistomal width Hide Narrow 08. Plumose antennal setae Absent Present 09. Scaliform antennal setae Absent Present 10. Length antennomere 2:3 2 < 3 Subequal 11. Length antennomere 3:4 Subequal 4 > 2 + 3 12. Shape antennomere 3 Serrate~filiform Flabellate 13. Shape antennomere 4 Serrate-fi1 iform Flabellate 14. Lateral margins of Elevated above disk Subdorsal above pronotum disk 15. Posterior margins of Subequal lateral Elevated above pronotum margins lateral margins 16. Frons shape Concave Convex 17. Median genal margins Unmodified Flanged 18. Plantar pads Latera11y expanded Not expanded 19. Femoral base Not annulated Annulated basally 20. Rostrate epistoma Not rostrate Rostrum short 21. Metacoxae Transverse Vertically oblique 22. Subantennal sulcus Meets epistoma Meets eye 23. Hypomeral stemmata Absent Present 24. Antennomere 2 and 3 Free Adnate 25. Antennomere 3 and 4 Free Adnate 26. Apical abdominal Entire Apically cleft tergite 27. Antennifer insertion Appears internally Appears elevated basomedial 315 those used in the phylogenetic analysis and can be found in Table 3.

Problems of the use of reduction and shape change are not extensively discussed here, but in methodology. Although great care was taken in the measurement data, all ratios based on those measurements are rounded to tenths and more precision should not be assumed (eg. 2/3 vs. 0.7).

These rounded ratios usually subsumed the intraspecific variation encountered for most states. The use of such ratios has limitations in morphometric analysis (Bookstein et al, 1986), but I believe they may be of use in specific delimitation. Finally, it can be assumed that any character not discussed here was not examined for this study.

Possibly, because of paedomorphosis, character states in this tribe varied more than is usual within lycid genera, with the exception of Lvropaeus Waterhouse (Miller, unpubl. data). This extensive variation and the concomitant rampant homoplasy would indicate numerous monotypic genera if traditional methodology and characters were used.

Alternatively, a single heterogenous genus might be recognized. The hypotheses of numerous monophyletic genera here proposed are supported by hypothesized synapomorphies, thought not to be affected by paedomorphosis. Further support is found in a number of unassociated and, hence, as yet undescribed ieptolycine larvae from Hispaniola.

However, the extensive variation in larvae may also be the result of anomalous development.

In addition to the extent of interspecific variation, intraspecific variation was also extensive. This often made identification of specific limits troublesome. These decisions were, again, exacerbated by the small sample sizes of most taxa. It is very 316 impossible to determine the limits of intraspecific variation in a species represented by a unique specimen, although it makes the process of description much easier. In this study specific limits were hypothesized from extrapolations based on the extent of variationfound in other taxa within the lineage with the largest sample si2es.

Although this implicit procedure in taxonomy is standard, it mayerr in either splitting or lumping taxa. Larger sample sizes would undoubtedly help, but this problem remains a major problem for all biologists, no matter how large the sample sizes.

Size. Overall length and the width at the humerus were discussed solely for diagnostic reasons and were not used in the analysis.

Interestingly, the Leptolycini contain some of the smallest lycids known and this may be a result of their development. However, there are other lycids nearly as small in which non-paedomophic females are known.

Length was measured from the anterior apex of the antennal prominence to the posterior apex of the elytra at its longest point. Width was measured at the widest point at the humeral angles. Although no formal allometric analysis was conducted, si2e was considered by examination of other small taxa within the family. For example, loss of laterally expanded plantar pads does not appear to be soley an allometric phenomenon because Haplobothris Bourgeois of similar size still has the expanded state.

Setation. The general setal condition was discussed, but setae may be rubbed and difficult to access. States varied from dense to sparse, but were not used in the analysis because of the difficulty in assigning discrete states. States of setal orientation measured from 317 the perpendicular are: decumbent (0o-30 ">, semi-decumbent (SI*—45"), semi-erect (46"-75°) to erect (76"-90*) respectively. These angles were determined near the basal insertions of the setae, because the setae may be straight, arcuate, to apically hooked.

Punctation. The surface may be slightly to deeply punctate.

Additionally, some lycids have punctation appearing areolate.

Discussion of punctation refers to the widespread state, but regional variation is not discussed if localized. Again, no discrete states could be assigned.

HEAD. The head shape, orientation, and length offer some useful character states, but were used with reservation because of the constraints of shape change definition and polarization. Shape is indicated by various measurements. Head length is measured by the ratio of the longest eye length to length of the genal margin when examined dorsally. This measurement (Character 04) was made with the head inserted normally on pinned specimens. As there is considerable intraspecific variation in the extent the head is retracted in death, all measurements were not as accurate as desired. Therefore, the data were lumped into two classes - greater than and less than length of eye.

Unique material, including types, often had to be estimated. The interoccular distance varies interspecifically and was used to further indicate separation of the eyes relative to change in head shape. This measurement is the relative distance between the eyes to the greatest width of the eyes when viewed dorsally.

Although trends related to shape change and orientation of the head are used in the analysis, there is some evidence that they might be 318

related to degree of paedomorphic expression. They present a

transition series similar to that found in Lvropaeus. Dexoris. and

several other Neotropical genera here excluded from the Leptolycini.

Similar head structure and orientation has been observed also in

Cantharidae and Lampyridae; in all cases the taxa are thought to be paedomorphic. Generally, there is a transition from a dorsally transverse appearance in the basal lineages to the more derived elongate condition where the genal margins are relatively long. This apparent correlation requires further study.

Genal Margin. See discussion of head shape.

Eyes. The eyes in Lycidae are usually directed laterally which produces a head with a transverse appearance when viewed dorsally. The more derived members of the Leptolycini and a few other genera thought to be paedomorphic have eye placement more anteriolateral and yield a more elongate appearance. This is partly the result of the head shape

(ie., the extension of the genal margins as discussed above). The size of the eyes is smaller in the more derived members of the lineage and appear to be correlated with the shift. The reduction of ommatidia number is not unexpected in paedomorphic forms. In addition to the eyes varying in size, they also vary in the convexity of their appearance, which I attempted to convey (not entirely successfully) as the ratio of the estimated radial length to the difference between the estimated center of the eye from the lateral margin of the head when viewed dorsally. Lycid eyes vary from heraispherically prominent to more convex and vary within the Leptolycini in such a manner from basal to derived, respectively. Whether this is an artifact of paedomorphic 319 ontogeny, or not, is unknown. Again the hind margin of the eye may be

slightly emarginate or straight in leptolycinesias well as other lycids.

Antennal Prominence. In most Lycidae the antennal insertions are on variably developed antennal prominence. Although the structure is present on all Leptolycini, its si2 e and form varies from the usual calopterine position. In the Calopterini this antennal prominence is a shelf-like ridge at the top of the frons. In the more derived members of the Leptolycini the antennal prominence is more gibbous and porrect, although it never projects directly to the front. Those that are here termed porrect are directed dorsoanteriorly at approximately a 45 ' angle. The antennal prominence may be distinct from the vertex and frons by transverse impressions or not. Although generally located in the same area, the insertion of the antennal prominence on the vertex may vary from anterior to a line between the forward margins of the eyes when examined dorsally to near a line between the posterior margins of the eyes. The antennal prominence is often sulcate dorsomedially and this sulcus varies from barely visible to a deep impression separating the prominence into two distinct structures. Shape of the prominence varies from bulbous to flat dorsally and flat to concave ventrally. As all these characters have non-discrete shape states and are not amenable to polarization, they were excluded from the analysis.

Antennal socket. The shape of the antennal socket and placement of the antennifer are characters that have not yet been used within the

Lycidae. They are, however, quite useful and studies in progress suggest that they may provide evidence for relationships at the higher levels of universality. For example, the antennal socket in 320

Lygistopterini is low on the frons, round, and more distant than other

lycids. The condition of the antennal socket found in the Calopterini and Leptolycini is high and closely approximate. Its shape varies from

trianguliform, oval, to lenticular.

Antennifer. Again the position of the antennifer, an unused character, varies in the Lycidae from a basomedial position to an elevated mediolateral position. Interestingly, the antenniferae of

some Cantharidae are synapomorphic within that family and appear to be unique within the Coleoptera by placement on the external lateral margin. Preliminary study (Miller, unpub1. data) suggests that there

is a correlation between this state and the ability to independently move the antennae into the mandibles for grooming as documented by

Valentine (1973:65). The antenniferae are internal on the medial margins in the leptolycines and the hypothesized outgroup, but the orientation of the head and antennal prominence gives the more derived

Leptolycini the appearance of a basomedial insertion. Again, the antennifer may be distinct or only slightly differentiated, but variation appears to be a continuum.

Interoccular Distance. See discussion of head shape.

Vertex. The shape of the vertex presents a continuum between the extremes of strongly transversely arcuate to almost flat. The expression of the pretentorial depression anteriorly appears to be independent of the discal orientation or states of the tentorial maculae. Both may be variously expressed as distinct or indistinct and the latter may be separate or coalesced into a single pit. Because of the sparsity of material, I did not examine internal head structure for 321 correlation of the tentorium with the maculae. Crowson (1972:48) mentions that the tentoria of lycids are typically membranous and my studies to date corroborate this hypothesis.

Frons. Traditionally, lycid taxa have been separated by the presence or absence of a rostrum and whether or not the frons is deflexed. However, the "horizontal frons" (eg. Green, 1949:55;

Nakane, 1969:4) is the dorsal surface of the antennal prominence which has shifted to a lower position on the frons. Therefore, Lycus has a shelf-like antennal prominence lower on the front than in many lycids, but the frons is actually similarly directed in life as in other

Lycidae. Generally, there is a concomitant change in head shape with change in direction of antennal prominence in the Leptolycini. The frons (Character 05) becomes more deflexed posteriorly in the more derived leptolycines, but this variation appears to be a continuum.

Again, the frons may be transversely concave or convex in the

Leptolycini and out group and also flat in other Lycidae. Frons shape change (Character 16) is used in the analysis because it appears binary and discrete. The relative dimensions of the frons vary within the tribe and is indicated by a measure of ratio of the frons length from the base of the antennal prominence to the width of the epistomal margin along the midline.

Epistomal Margin. The epistomal margin varies from wide to narrow and may be truncate, slightly sinuous, or emarginate. A measure of the relative size is the ratio of transverse width of the epistoma to the width between the eyes frontally (Character 07).

Mouthparts. The mouthparts of adult Lycidae are usually 322 directed ventrally, but some are directed in an anterioventral position much as in the orientation of larval lycid mouthparts. In the later state the orientation the palps often appears more porrect. The mouthparts of Leptolycini appear to be reduced, sometimes extremely so.

However, their reduction or absence was not described, because they could not be seen clearly under a dissecting microscope. Several species are known only from unique types and/or old specimens and these were not dissected. Because mouthpart character states often require dissection to find meaningful differences, they were not included in the phylogenetic analysis. Future work should investigate the status of mouthparts in these taxa, when additional specimens become available.

Labrum. The labrum in the Lycidae is usually distinct, but may be fused to the epistoma (eg. Lucaina Duges). Shape varies from transverse to square to semicircular. This character system is here deemed useful only as autapomorphies because of variation within various lineages and the inability to resolve shape changes, except as ad hoc support for a hypothesis by position.

Mandibles. The mandibles in the known Lycidae are all unidentate and may be straight, arcuate, falciform, sickle-shaped, or strongly hooked. The apices are acute. The mandibles of Leptolycines are arcuate to strongly hooked. They appear to be reduced or lost in many taxa, but this may be an artifact because of si2e. When unseen, the condition is indicated by a (?).

Maxilla. The lycid maxilla typically consists of distinct to non-differentiated galea and lacinia. The maxillary palpomeres are usually thickened to the apex, and palpomere 1 and 3 are short, 2 longer 323 and thicker, and 4 more or less truncate or expanded. The ultimate palpomere varies from securiform, obovate, cultriform, acuminate, etc.

The investigated taxa within the Leptolycini suggest that development of the maxilla do not vary significantly, except in the palpomeres.

However, it was noted that the terminal palpomere of Leptolycini,

Dexorini, and Lyropaeini all share an acuminate state. It is here hypothesized that the state is the result of paedomorphic development.

Some paedomorphic lampyrids also have similar states.

Labium. The labium in Lycidae is usually short and broad with a small menturn and submenturn, but it may be reduced. Because a number of important taxa were not available for dissection, this character is unstudied in the Leptolycini. The few dissections made did indicate a reduction in size of the mentum and prementum.

The Lycidae usually have 3 labial palpomeres, but reduction in number and size has been noted. The labial palps are usually smaller than the maxillary palps andathe apical palpomere is usually similar in shape to the apical maxillary palpomere. aWithin the Leptolycini there appears to be a trend to reduction to one or no palpomeres and they are acuminate when present. A similar trend is found in Dexoris and

Lvropaeus.

Gula. The gular region in the Lycidae is usually short, quadrate, and transverse as it is in the calopterine outgroup. In the more derived members of the Leptolycini the gula is elongate and trapezoidal in conjunctio with the elongate genal margins. In the more derived Leptolycini the gula is more anteriad of the hind margin of the postgena. 324

Antennae. Although they are easily boken and lost, the antennae

provide some of the best character states for recognition of species and

have been used extensively for classification of the family. These

states often, however, form a series of continuous variation and

analysis is often confounded by sexual dimorphism. The extreme

variation in antennal structure within the Leptolycini is anomalous.

Following taxonomic precedent would suggest more genera than here recognized. Because of this variability and my inability to access

transition series, the resulting homoplasies were the most difficult character states to resolve.

Antennal size within the Lycidae varies from short (reaching hind margin of pronotum) to elongate (3x body length). Antennae of

Leptolycini vary from moderately short to moderately long, but no trends in variation between genera were found.

Antennae vary within the family and its outgroups from compressed to non~compressed and from filiform to serrate and flabel late.

Leptolycini demonstrate essentially the entire range of variation found within the family, but appear discrete and were used in the analysis.

Within the Leptolycini the rami, when present, were always inserted apically, although several genera in the calopterine outgroup have the ramal insertions basal.

Antennal Compression. Lycid antennae are commonly compressed or may be almost round in cross section (Character 02). States found in

Abrolvcus spp. suggest that antennal compression is not correlated with the compression of the legs. In beetle taxa examined with compressed antennae, the pedicel is not compressed. Antennal Flabellation. Although the antennomeres are undoubtedly

serially homologous, there is considerable variation of expression.

Failure to recognize this variation of each antennomere may lead to errors of homology.

There are a number of problems associated with the recognition of flabellation states which may be a result of imprecise terminology.

First, the rami of a number of flabellate antennae of some Calopterini and Leptolycini are loosely lamellate. Thus, the lamellate antennomeres of Scarabaeoidea are not unique, because their "unique" status is just a matter of the approximation of the rami). A number of other lycid and lampyrid genera also have similar states. Insertion, plane of compression relative to antennal axis, and length of rami vary throughout the Lycidae and must be noted in homology decisions. Also, the rami may begin on antennomere 3 or 4 and outside the Leptolycini and

Calopterini, on 5. Antennomeres basal to the first clearly defined ramus may be serrate or transversely filiform. Problems in resolving the relationship between Aporhipis. Acroleptus. and a possible new, monotypic genus rest on the dilemma of the apparent homology of the antennae including the basal antennomeres, but differ in a series that shows most of the variation in the Leptolycini in three species.

Antennal Setation. The setation of the antennae of lycids is usually dense and semi—decumbent, but may be sparse or semi—erect. In addition to this condition, several genera within the Leptolycini have setae with flattened appearance like scales (Character 08). Again,

LePtolvcus has apomorphic elongate bristling setae (Character 07) found nowhere else in the Lycidae or Lampyridae to my knowledge. 326

Scape. The shape and size of the scape were useful at the

species level of universality. The shape in the outgroup and

Leptolycini, like the Lycidae in general, ranged from barrel-shaped to

pyriform to obconic and the insertion of the pedicel from truncate to

obliquely truncate.

Pedicel. The size of the pedicel in relation to the third antennomere (Character 09) has been used for tribal classification (eg.

Green, 1949:56), but here varied within what is undoubtedly the same genus. This variation may have implications for the study of the

Elateroidea, but requires further examination. Again, the pedicel is adnate to the antennomere 3 in Anti 11olcvus (Character 24). This state

is the result of the second antennal segment being "fused" or non­ differentiated from the base of the third antennal segment. This adnation results in the antennae not being flexible at this point of articulation.

Antennomere 3, The relative size of antennomere 3 to 4

(Character 10) varies throughout the family and was used in the analysis. Antennomere 3 was either much shorter than 4 or subequal in the Leptolycini. Antennomere 3 can be either serrate-filiform or flabellate (Character 11). Additionally, antennomere 3 is adnate to antennomere 4 in Ceratoprion much as antennomere 2 is adnate to 3 in

Antillolvcus.

Antennomere 4. Like antennomere 3, antennomere 4 can be either serrate-filiform or flabellate (Character 12). Initially, I thought they were correlated character states, but several genera support the hypothesis that they are not. 327

Antennomere 11. In all species of Lycidae examined, the axis of

antennomere 11 is not distinguishable from its ramus when it is

flabellate. This can be checked by examination of the plane of

compression of the ramus and axis.

THORAX. The soft-bodied malacoderm facies is easily deformed in preservation and during emergence of the adult from the pupal stage.

As a result, thoracic shape can be used for specific diagnosis, but must be examined cum grano salis. Shape is too unreliable except in general si2e relationships for taxonomic recognition and not suited to phylogenetic analysis (see methods).

Prothorax. Like all the cantharoid lineages examined, the prothorax is not tightly coadapted to the pterothorax as in many of the hard—bodied Coleoptera.

Pronotum. The pronotal shape varies from elongate to transverse and it may be quadrate, trapeziform, or rounded. The anterior angles may be acute to rounded and the posterior margins may be acutely produced or not. The disk varies from flat to convex and from elevated frontally and posteriorly to not. The best means of accessing these latter states is in lateral view.

Median Longitudinal Carina. The pronotal carinae have been used extensively to define genera (and higher taxonomic levels) within the

Lycidae. Basically, there are 3 different sets of carinae. There may be a transverse carina which may meet variously along the lateral margins of the pronotal disk at the anterior 0.3x pronotal length to the posterior corners. This transverse carina is absent in all Calopterini and Leptolycini, but present in the Cladophorini and Dexorini among 328 others. There may be a median longitudinal carina which may be divided or not into a pair of parallel carinae that meet at the posterior margin

or meet anterior to the base. In the Leptolycini and some members of

the Calopterini the median longitudinal carina is divided (here hypothesized as a synapomorphy for these taxa). It also occurs in other tribes, but in conjunction with other carinae unless the latter are reduced. Additionally there may be a pair of anterior longitudinal carinae in front of the transverse carina with the median longitudinal carina present or absent when these carinae are present. These carinae are not found in the in- nor the outgroups of interest. Although pronotal carinae appear relatively stable within a lineage, they may be reduced or lost. Such is the case in several species of the more derived Leptolycini and all known Pseudacroleptus Pic, but other character states support the placement of these taxa in the tribe.

These carinae create distinctive depressions on the disk of the pronotum. Nakane (1969:4) called them areolae (singular, areolet), but

Torre Bueno (1973:24) defines this term as "one of the small spaces between veins of net-winged insects; a small wing cell." Therefore, in this discussion they will be referred to simply as cells. Calochromus

Guerin-Menevi1le has a median longitudinal depression for most of the pronotal length with no carinae evident. This may be the result of reduced, divided medial longitudinal carinae with the internal cell still visible as an impression bounded by "folds" analogous to the transverse folds, but this requires more investigation.

Pronotal Stemmata. A character state that may be inappropriately named here is the pronotal stemmata. These small, paired swellings 329

(Character 03) occur dorsaliy toward the lateral edges on or near the anterior and posterior margins. There may be as many as 2 anterior and

2 posterior pairs. Their position varies and is often useful in separating species, but is not used in the keys because they are often difficult to see. I have found pronotal stemmata only in several genera of Leptolycini, but they may appear sporadically in other genera where I have seen only a few species. However, they are here hypothesized to be a synapomorphy of the Leptolycini. Their function, if any, is unknown and light sensitivity is not implied by the term.

Pronotal Margins. The perimeter of the pronotal disk may be swollen along the edge to create a border which has been called

"beaded." Because several systematists have protested the use of this term, I have used the term swollen. While this word is not as descriptive as I would wish, it is better than beaded. The term beaded may be erroneously interpreted to suggest the appearance, like the bead of the book binding industry, of a bead on the spine of a book, which appears more like a rounded carina than a swollen edge. Again, it may be erroneously interpreted as a serial line of bumps in the sense of the welders’ bead. Neither interpretation is descriptive of the state found in lycids. The margins of the prothorax are uniformly swollen along their length, this swelling appearing often on both the dorsal and ventral surfaces and is similar to some bumpers. In the Leptolycini most margins are swollen, but in other lycid genera (eg. Lvcus

Fabricius) all pronotal margins are not swollen. The margin may be, also, simply dorsally directed (ie.erect), explanate, or reflexed laterally near the margin. The lateral margins may appear elevated or 330

subdorsal (Character 13) relative to the disk of the prothorax. The

posterior margin may be elevated or not (Character 14) with respect to

the rest of the pronotum and the anterior portion may be inflated

allowing reception of the base of the head.

Hypomeron. Generally, the hypomeron is concave in the Lycidae

and appears characteristic of the family. The margins of the hypomeron

may be slightly swollen or not in the Leptolycini. Shape is often

useful, but only the maximum vertical height to width along the dorsal

margin is discussed.

Hypomeral Stemmata. These structures are similar to the pronotal

stemmata and are found only on a subset of those taxa with pronotal

stemmata. Again, no light sensitivity is implied by the term.

Prosternum. The prosternal bridge, which is short in front of

the procoxae, is medially short and usually T- or Y-shaped in the

majority of the Lycidae. Differentiation of the two states is often a

matter of interpretation because a T-shaped prosternum may be slightly

incised and, thus, be called Y-shaped by other workers. In the

Leptolycini, both states are found as well as a triangular state in the

more derived members. As the triangular shape is discretely different

and can be distinguished from the T- and Y-shapes, which could not be

separated, it was included in the analysis (Character 01). Although

intercoxal processes extend between the procoxae, they are short and are

not known to reach the mesosternum in the Lycidae. Their length was

not used as a character state, because it varies continuously. On the posterior apex of the intercoxal process the internal furcae are

sometimes visible, but it is unknown if this is an artifact of 331

preservation or an external state, because too little material was

available for dissection. The lack of propleural process behind the

legs precludes closed coxal cavities in all lycids examined.

Mesothorax. The structures of the mesothorax have been used

little in the classification of the Lycidae. Most discussion has been

limited to discussion of the scutellum and the family level character of

the separation of the mesocoxae by the mesosternum.

Scutellum. The scutellum varies throughout the Lycidae in si2e

and shape, particularly the apex. This variation appears to be random

among the various lineages, as it does in the Leptolycini, In the

Lycuidae, including the Leptolycini, and many other beetles the lateral

margins of the scutellum evidently function as a latch to assist holding

the elytra in place.

Mesospiracles. The mesospiracles in the Lycidae may be prominent

with exposed peritremes or they may be hidden and lack peritremes. The

orifice may be constricted in some taxa and they may be hooded as in

some Lvcus Fabricius. The presence of exposed peritremes has been used

in higher classification, but the leptolycine data and concurrent

studies of other taxa within the family suggest that it may not be

useful as a synapomorphy at that level. The mesospiracles are often

prominent in most Leptolycini and its outgroup. They are variously compressed or not, but not hooded.

Mesepisternum. The mesepisternum is elongate in all Lycidae examined to date. Its posterior margin, however, is often concavely arcuate instead of straight. This character is still under study and

is only included for future reference. 332

Mesepimeron. The reflexed mesepimeron is synapomorphic for adult

Lycidae and Lampryridae. Its shape and size appear to be useful at the species level, but varies among congenerics. Again, the length of the mesepimeron to the length of the mesepisternum is of taxonomic value, but varies similarly. The mesepimeron is reduced and not reflexed in some of the more derived members of the Leptolycini.

Mesosternum. The lycid mesosternum may be transverse or elongate. In the leptolycines and outgroup it is usually intermediate.

A median sulcus of the mesosternum, as in Caenia Newman in the

Calopterini, is not found in Leptolycini nor its outgroup.

Mesocoxal Separation. The separation of mesocoxae have been used as a diagnostic character of the Lycidae, but it can vary significantly.

In fact Kleine (1941) argued that Pul iticola Mjoberg was not a lycid largely because the mesocoxae are contiguous (its placement in the family, however, is supported by other adult and numerous larval character states). The separation of leptolycine mesocoxae varies, but none are contiguous. The mesocoxal cavities are always open in the

Lycidae.

Elytra. The obvious dorsal aspect of the elytron is one of the major character suites in traditional taxonomy of the Lycidae, but variation within a lineage is often ignored. Most of the elytral characters were not used in this phylogenetic analysis, because the elytra of paedomorphic taxa are significantly affected by the phenomenon.

Elytral Reticulation. Surface structure of the elytra may be smooth, scabrous, punctate, irregularly reticulate, or uniformly 333 reticulate in the Lycidae. These states may be of value at the generic level of universality, but vary extensively interspecifically.

However, there is also considerable intraspecific variation and, more importantly, reticulation varies within a single elytron. Hence, to be useful, discussion of reticulation must be accompanied by an indication of the part of the elytron examined. Nevertheless, the apices should be avoided because they are the least stable area. The variation of leptolycines demonstrates the entire continuum of states.

Elytral Costae. The number of elytral costae is probably used more than any other elytral character and the relative elevation and degree of expression of the costae has been used traditionally to separate genera. This has resulted in the erection of genera for minor differences that may yield a paraphyletic classification with further study. This character system appears to be a developmental phenomenon and, as in the pronotal configurations found in the Lycidae, there is extreme homoplasy regardless of the polarization. For example, the

Dictvoptera Latreille - Benibotarus Kono - Pvropterus Mulsant lineage has 8, 7, and 4 costae respectively. Similar transitions can be found for nearly every lineage within the family. Additionally, discussion of varying costal expression along its length, besides being of diagnostic significance, might reveal relationships obscured by artificial taxonomic separation. For example, the genus Benibotarus

Kono was erected for those species of Dictvoptera Latreille with only 7 costae versus 8. Examination of both genera demonstrates that the

"missing" costa is variously expressed over an overlapping series of intermediates in both genera. Thus, differentiation into two genera is 334

entirely arbitrary.

Added to the problem discussed above, there are differences in the

costal expression such as costa obsolete, angularly broken in a zigzag pattern (usually not defined or recognized as costae), and in thickness

(alternating elevation of costae). In fact the genus Idi opteron

Bourgeois, as currently characterized, has species with 3 and 4 costae, although costa 3 is weakly present only as an angularly interrupted costa or none. Although this is clearly a transition series, a traditional approach would require splitting the taxon, but this may be premature as other characters do not support such a decision. Costae at the apices of the elytra throughout the Lycidae often terminate in an angularly interrupted pattern.

Part of the problem is a non-standard terminology that varies with taxonomic treatment and ignores homology. If the genus has four costae visible, they are called primary costae 1-4. If a genus has eight, they may be called primary 1—4 and secondary costae 1-4 (even if the secondary cannot be differentiated from the primary costae except by position) or they may be called 1-8. Besides the resulting confusion, the use of primary and secondary implies some assumptions about them that are not warranted, much as the "primary" and "secondary: setae of the lampyrid elytra. No attempt has been made to homologize these states and, in fact, the angularly broken costal state that often produces irregular reticulation (eg. two pair of areolae between

"primary" costae) are usually not considered to be costae. To rectify this problem, I here propose the homologization of the costae as indicated by position. Therefore, using Dictvoptera gorhami Kono as a 335

template (Nakane, 1969:218, fig. A) the costal positions of a

generalized lycid can be plotted from the suture outward. Thus,

Nakane's primary 1=2, his 2=4, his 3=6, his 4=8 and the "secondary" costae are 1, 3, 5, 7, and 9. By comparatively examining the costal

insertions at the elytral base, the homology of any costal state can be ascertained. As can be seen, there are 9 costal positions with one row of reticulate cells between each pair of costal positions. Costa 8 is also called the humeral costa, because it is expressed over the humeral umbone. With this nomenclature, there can only be one row of cells between costae and if two rows occur, a costa is missing. The positions of the cells are hypothesized to be those of the punctation of other Elateroidea.

Although the lateral and sutural veins form "costae" that function similar to the other costae during adult emergence and the sclerotization of the elytra (Miller, unpubl. data), they are not here included in the proposed nomenclature. Their homology outside the

Lycidae is not yet apparent.

The costae of the Leptolycini are hypothesized to form a transition series of 4, 3, and 2. The outgroup states are 8 and 4 costate. Because these states were not used in the phylogenetic analysis, the genera here defined are much more variable with respect to costal number than in other taxonomic work on the family.

Elytral Apices. A few lycids have the elytral apices truncate, but most are independently round and not angulate at the sutural line.

A few that have truncate apices (eg. some species of Lvcus) have their inner apical angles acute. In most of the Leptolycini and its 336 hypothesized outgroup elytral apices are rounded, but some Leptolycini have acute apices.

Elytral Dehiscence. The elytra of most lycids meet along the sutural margin, but in the Leptolycini and some other Lycidae the elytra are divergent apically from the sutural margin. The initiation of dehiscence along the suture varies specifically from near the scutellum to near the elytral apex. Some taxa have elytra reconvergent near the elytral apex, but this was not used in the analysis even though it appears to occur in monophyletic lineages. The expression of dehiscence is hypothesized to be an expression of paedomorphic development, so all states associated with it were discarded.

Elytral Coadaptation. Another character not discussed for lycids is the coadaptation (sensu Crowson, 1967:60) of the elytra with the abdomen. In some genera (eg. Dictvoptera Latreille) the abdomen fits basally within the concavity formed by the paired elytra for about half its length. The elytral apices of all known lycids are not coadapted, but transversely arcuate to relatively flat and rest on top of the abdomen. In some genera the elytra are coadaptive only basally to 0.3x their length. Those that exhibit the latter state may have expanded elytra or a more ligulate shape. The Leptolycini have the ligulate state.

Metathorax. Because of the lateral compression of the thorax, the metathorax of the Leptolycini appears more swollen ventrally and more deep-bodied. Initially, I thought this an artifact of preservation, but examination of the various sutures suggests that it is not. This can most easily be seen by examination of the metacoxae 337

which are discussed below. Most other Lycidae, including the outgroup,

are more transverse and compressed dorsoventrally.

Metathoracic Wings. Lycids have typical cantharoid wing venation

(Forbes, 1923). Lycids are thought to have no closed anal cell

(Nakane, 1969:5), but it has been found in Broxvlus Waterhouse (Miller,

unpubl. data). The venation of the Leptolycini is reduced and this may

be developmental, but it is more likely due to the extremely reduced

size of these lycids. No micropterous leptolycine males were seen,

which supports the latter hypothesis. However, this may be an artifact

because micropterous males (or females) would be less likely captured in

malaise and flight intercept traps.

Metepimeron. The configuration of the metepisternal-metepimeral

suture has been used in the past for family separation of Lampyridae and

Cantharidae (eg. Blatchley, 1910:809), but little attention has been

given to it within the Lycidae. Within the Leptolycini it may vary from straight over most of its length, to arcuate in the more derived members of the lineage. These states are an indication of the change in

orientation of the metathorax. Usually, this suture is straight, but

in the more derived leptolycines with more laterally compressed bodies

it is arcuate.

Metepisternum. Another measure of the lateral compression in the

Leptolycini is the ventral margin of the metepisternum. Usually, in the

Lycidae this suture between the metepisternum and metasternum is more arcuate anteriorly and becomes less arcuate posteriorly. In the

leptolycines the reverse is often the case.

Metasternum. In the lycid metathorax the metasterna1 336 longitudinal sulcus may be complete or incomplete. Leptolycini exhibit the full range of variation of this character.

Legs. The relative size of leg segments changes differentially within various lycid lineages. The more derived members of the

Leptolycini have trochanters more elongate, much as in Thvlodrias. than do basal members. This same transition has been noted in Lvropaeus

Waterhouse. Two measures were taken to illustrate this transition: the trochanter to coxa ratio and the trochanter to femur ratio.

Trochantins. The leptolycine trochantins are free and setiferous in the fore and middle legs as in all Lycidae examined to date.

Coxae. Within the Lycidae the common condition of the trochanteral insertion in the coxae is more or less apical in the procoxae and the mesocoxae. Within the more derived Leptolycini the membranous region surrounding the insertion is more elongate and lateral.

The metacoxae of most lycids are transverse and nearly horizontal, and the metatrochanters insert internally. The coxal cavities are nearly contiguous, but unlike other cantharoids, they are not set within projections of the coxae which extend posteriorly under the abdomen.

Leptolycines share these lycid states, except in the more derived members of the lineage.

Because the laterally compressed thorax is derived in Leptolycini, the metacoxae are predominately obliquely oriented dorso-ventrally instead of horizontal and transverse (Character 17).

Trochanters. The shape of trochanters, particularly the metatrochanters, varies within the Lycidae. Their shape may be 339 triangular or cylindrical and the insertion of the femur may be offset, oblique or truncate (Character 20). All states occur in the

Leptolycini and the outgroups.

Femora. In the Leptolycini the bases of the femora are annulated and this is hypothesized to be a synapomorphy for the lineage (Character

19). This state is often not easily seen in uncleared specimens and, therefore, is difficult to assess in rare species.

The femora and tibiae appear equally compressed and appear to be correlated. They often co-occur with compressed antennomeres, but the

Abrolvcus state implies that their co—occurence is not necessary.

Although limb compression is widespread within the family and is usually thought to be characteristic of various genera, the anomalous Abrolvcus omalysiformis with non-compressed legs within derived Leptolycini requires further study.

Tibiae. The tibiae may be elongate and narrow or more broad.

All Leptolycini have elongate and narrow tibae. In a like manner, the internal margins of the tibae may be straight to sinuate to bisinuate.

These states are often sexually dimorphic and restricted to or better deveveloped on the male prolegs. This may be related to male behavior while grasping the female during copulation.

Tibial Spines. Although the tibial spines may be large and dissimilar in some Lycidae, they are minute and similar, or absent, in the Leptolycini and outgroup. In almost all Lycidae, these states are difficult to see and, thus, of little diagnostic value; however spines are large enough to be used in the subgenera of Lvcus Fabricius sensu lato. where tibial spines are larger, has been used as such by Green 340

< 1949 ).

Tarsi. Tarsomeres may be inserted apically or subapically and dorsally within the Lycidae. Lycidae usually have plantar pads on tarsomeres 1-4, but there may be fewer tarsomeres with these structures.

Plantar pads (Character 16) are usually gradually, uniformly expanded laterally from a narrow first tarsomere to a wider state in tarsomere 4.

Although Calopterini have the usual states, the plantar pads are reduced in size and lateral expansion in the Leptolycini. My first hypothesis was that the lack of expanded plantar pads was a result of allometry, but several species that are comparable in size in other lineages (eg. Haolobothri s spp. ) have plantar pads expanded laterally as in other Calopterini. Whether the state is correlated with paedomorphosis is unknown, but Lvropaeus spp. also has the expanded condi tion.

Ungues. The ungues may be cleft in some Lycidae (eg. Macrolvcus

Waterhouse), but are simple in all Leptolycini and Calopterini examined to date.

ABDOMEN. The abdomen of Lycidae is sexually dimorphic - the males having 8 visible sternites and alate females 7. Leptolycine males do not differ, except the abdomen is often narrower. The only known female leptolycine has the normal larval compliment of 10 segments, including the pygopodium. The abdomen often narrows uniformly, but may narrow more abruptly distally to the base.

Tergites. The tergites of Lycidae are rectangular and tranverse, and in most cases lack explanate margins. When present, these explanate margins are found on the posterior angles as in Lampyridae. 341

There are no known species in the outgroup or Leptolycini with explanate

margins.

Sternites. The sternites of Lycidae are rectangular and tranverse

without explanate margins in most cases. Again, when these explanate

margins are found, they occur on the posterior angles as in Lampyridae.

There are no known species in the outgroup or Leptolycini with explanate

margins.

Abdominal Spiracles. The spiracles are in the membrane, but

their position varies. This character is insufficiently studied

throughout the family for incorporation in this analysis, but the states

are reported so they will be available for future work.

External Male Genitalia. The leptolycine aedeagus varies like

that of other lycid taxa. Some taxa have highly distinctive genitalia at the species level and others do not. Thus, the genitalia of species of Leptolvcus vary little and external character states must be used to

identify them. The use of genitalic character states has been used to support generic concepts, but should be so used with caution. For

instance, Green (1951:18) erected the genus Pseudoplateros for a species of Plateros Bourgeois with parameres because other known congenerics have no parameres. Such reasoning would indicate that Abrolvcus darlingtoni would be placed in a monotypic genus, which is clearly unacceptable based on other states within Abrolvcus. Studies of

Plateros in progress suggest that Pseudoplateros Green should be suppressed.

Internal Male Genitalia. The character states of the internal genitalia were not examined, but preliminary studies indicate that they 342 may provide useful data. They were not included in the analysis, because a database of sufficient si2e for comparative purposes is not yet available. Several species are here figured.

External Female Geniitalia. None seen in the only species available for study.

Internal Female Geniitalia. Not studied, because of the poor preservation of Berlese extracted material in the only species available for study.

Relationships within the Leptolycini. Figure 3 presents a phylogenetic hypothesis of leptolycine genera which accomodates a hypothesis of paedomorphosis throughout the lineage. One immediately perceived limitation of the proposed methodology is the first impression that this hypothesis is not robust. Much of the character state data previously discussed must be discarded in order to provide a hypothesis with the least homoplasy. Alternative analyses including character states that are known to be effected by paedomorphosis generate hypotheses rife with homoplasy or unexamined implicit assumptions. In these cases the high rates of homoplasy observed are not dependent on polarization of character states. Homoplasy does not matter whether the lineage in assumed basal within the Lycidae or highly derived with numerous independent origins. Both assumptions result in high rates.

If paedomorphosis is assumed basal within the Lycidae, then the homoplasy is found as numerous apomorphies of novelty. If it is assumed to be derived, then the apomorphies are of reduction and non­ differentiation. Not apparent in this cladogram, is that several of 343

the traditional characters supporting the higher classification of the

Lycidae are those most effected by paedomorphosis.

A hypothesis of intrageneric relationships is not advanced at this

time, because much of the relevant data requires further study by

methodology not yet available. For example, most of the variation

observed is shape change and polarization of it can be arbitrary.

Again, the extent of intraspecific variation also presents problems.

Bookstein et al. (1985) suggest that ratio data are of little value for

studies of shape change and, hence, different measures constrained by

homology are required for further study. Such an approach has not yet

been attempted with the Leptolycini.

Phylogenetic Conclusions. In summary, the Leptolycini are a relatively

small, but important lineage of Lycidae. This study has resulted in 28 new species and the transfer of 3 described species from the

Calopterini. In new material examined the rate of finding new species

to known species has remained high and, thus, the extent of the lineage

is probably still largely unknown. Additionally, the biology has been studied only inferentially from preserved material. The phylogenetic hypothesis presented here is subject, like all hypotheses, to refutation. Effort at such refutation should be directed at field work, whether merely collecting additional specimens or studies of their biology.

The morphology of the Leptolycini presents an extremely homoplasious distribution of character states, and this has been 344 explained by a hypothesis of paedomorphosis which is corroborated by the anomalous development of females of Leptolvcus heterocornis. Studies of Thvlodrias and other paedomorphic species suggest that both male and female morphology converge to non-differentiated states. Thus, modifications of current phylogenetic methodology to accommodate such paedomorphic taxa are indicated. However, the solutions found to date are somewhat unsatisfactory because too much data cannot be used in the analysis without explicit and or implicit a priori assumptions. I am still looking into a resolution of this problem. BIOGEOGRAPHIC IMPLICATIONS

Because of their expected limited vagility, paedomorphic lineages with flightless females should provide an excellent arena for speculation about vicariance hypotheses. If paedomorphosis can be demonstrated throughout a monophyietic lineage, it is most parsimonious to hypothesize its origin as a unique innovation (ie. apomorphy).

Members of such lineages are, by their dispersal limitations, evidence of a contiguous previcariant origin. Similar arguments have been used by, among others, Ivie (1987, personal communication) in his studies of

West Indian litter-dwelling, flightless beetles such as Archeoglenes

Broun and some monommids, and by Ward (1985:141) in his studies of ants with flightless queens.

All such arguments, however, depend on the tacit assumption that flightlessness is a synapomorphy within the lineage examined. Clearly, the most critical and requisite test of a vicariance hypothesis of this nature is an examination of life cycles. Flightlessness must be demonstrated empirically in all the females of the lineage. It must also be demonstrated that flightlessness is not the result of a simple genetic polymorphism as shown in some carabids (Lindroth, 1946) and curculionids (Jackson, 1928; Stein, 1973). Paedomorphic lineages within the Coleoptera offer an advantage in that they are not known to be polymorphic with respect to flightlessness in females with the

345 346

exception of Micromalthus LeConte, which was studied by Scott <1938).

A number of lineages within the Cantharoidea appear amenable to

such speculation. These include the Phengodidae, the Drilidae, possibly some genera of Lampyridae, and some Lycidae. Among the latter

is the Leptolycini. Although paedomorphosis has been found only in

Leotolvcus heterocornis L. & M. , I have found no winged females in the

museum material examined, including the supposed allotype of the aforementioned species. It, thus, appears that some discussion of

biogeography would be useful.

In the West Indies, leptolycines are known only on Cuba,

Hispaniola, and the Puerto Rican Bank. None are known from Jamaica,

the Bahamian Bank, or the Lesser Antilles. According to Buskirk

(1985), the former is not unexpected as Jamaica is not thought to have ever been adjacent to another island or continental margin since its emergence. Thus, presence of leptolycines there would have to be the result of over-water dispersal which is unlikely for flightless, paedomorphic species. Leptolycini on the Bahamian Bank is unknown, but not unlikely, because of eustasis during the Pleisocene glaciations and much of the Bahamian Bank is thought to have been emergent at that time and contiguous with Cuba. Additionally, leptolycines have not been recovered in the Lesser Antilles, unlike Thonalmus from Montserrat which probably arrived by dispersal (Miller, 1984). As the Lesser Antilles are not thought to have ever been contiguous with the Greater Antilles

(Donnelly, 1988:29), this is not unexpected. Collectively, this evidence of limited dispersal capabilities supports a previcariant contiguous origin of the lineage. 347

Extrapolation of the trajectories of past tectonic configurations

of the Caribbean during the Eocene to the Late Miocene (Buskirk, 1985:

figs. 2a~d; Donnelly, 1986: figs. 2.2-2.4) suggests that Cuba, northern

Hispaniola, and Puerto Rico may have formed an island chain between

North America and South America in the Paleocene or late Cretaceous.

However, Hedges (1982), Savage (1982), Buskirk (1985), and Donnelly

(1986) urge caution in forming hypotheses dependent on a contiguous

island arc. Although some biological evidence supports such a conclusion, there is no geologic evidence to support or refute it.

If, however, such an island chain existed between the continents

and provided stepping stones or a continuous land mass, a mixing of

floral and faunal elements would be expected. Thus, elements of both

North and South American faunas would be expected in the Greater

Antilles, assuming the islands had remained emergent since then. The

West Indian distribution of the paedomorphic Leptolycini strongly

supports such a South American-Anti 11ian vicariance hypothesis despite

this lack of geological evidence. In the Leptolycini only two extant

species of the tribe, Ceratoprion serricorne Gorham and C. chelae new

species, are found outside northern South America and the West Indies.

These anomalous species are members of a genus hypothesized to be derived within the Leptolycines and, hence, the most parsimonious explanation for the observed distribution is that this genus moved into

Central America after the early Tertiary closure of North and South

America. Colonization and subsequent vicariance of the Antillian plate

isolated the Greater Antillian Leptolycini. Further support for this

scenario is found in the distribution of C. serricorne — it is the most 348 widespread leptolycine known and ranges from Ecuador into Panama.

The specific distribution of the Leptolycini within the West

Indies can be explained by two equally parsimonious hypotheses. Either they speciated before fragmentation of the Antillian arc and the present distribution and number of taxa is the result of differential extinction, or they speciated after vicariance events. Appeal to parsimony to resolve this issue is vacuous, even if a phylogenetic hypothesis at the species level were at hand, as all endemic genera are represented on the larger islands, while only one is known to occur on the much smaller Virgin Islands. LITERATURE CITED

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ILLUSTRATIONS

361 362

©JED E S T i

e e s

Figure 1. Theoretically available phylogenetic reconstructions for paedomorphosis Cprogenesisl (after Westheide, 1987:850). A. Case 1, monophyletic lineage. B. Case 2, a monophyletic lineage where Q o non-paedomorphic ancestors produce paedomorphic daughter species. Case 3, non-monophyletic multiple origins of paedomorphic species. More conventional representation of case 2. P = paedomorphic, NP = non-paedomorphic, A = ancestor. 363

A. I

1 1 I B.d* I 3 9 I 1 3 I C. d* I I 3 4 6 7 8 I 1 9 l 3 4 16 7 7 I D.c/ I tp 3 4 ! 6 7 8

9 I \\\ 345 678

Figure 2. Models of possible ontogenetic pathways of a character. Hatched areas are developmental stages of expression. A. Full developmental sequence of character states in the outgroup. B. New states produced by interstitial perturbation between stages II and III. C. Truncation of terminal development. D. Ontogenetic rate accelerated in the female. 364

FOG PCER PSAC FLAB CERA ABRO ANTI LEPT

Al7* bL * 1 1 3 J b2# 3 1 BS* □ 5 Oil e * d 310 0 9 i3 d323 23* 2 4 * 26*

j13

Figure 3. Hypothesized phylogeny of the genera of the Leptolycini. Chararacter states are as discussed in the text and summarized in Table 3. Open rectangles are apomorphies demonstrating homoplasious distribution and solid are those showing non — homoplasious distribtuion. Those character states with an asterisk are the autapomorphies that support this hypothesis without homoplasy. 365

a* b. c.

d.

e.

Figure 4. Abrolvcus bicolor New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Right antenna. e. Right hindwing. 366

a. b.

Figure 5. Abrolvcus cubensis New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. 367

Figure 6. Abrolvcus darlingtoni New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 360

Figure 7, Abrolycus iviei New Species. a — c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. Figure 8. Abrolvcus omalvsiformis New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 370

Figure 9, Abrolvcus sandersoni New Species, a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect of head and prothorax. e. Internal male genitalia. 371

Figure 10. Anti 1loivcus auratosuratus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Last abdomianl sternite. e - f. Ultimate and penultimate abdominal tergites - e. internal view, f. external view. 372

Figure 11. Anti 1lolvcus elongatus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral view of head and thorax. 373

Figure 12. Anti 1lolvcus flavomarginatus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. a. Lateral aspect head and prothorax. 374

Figure 13. Anti 1lolycus semiflavus (Chevrolat). a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. sssr^a

Figure 14. Ceratoprion bordoni New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Right antenna, e. Right hindwing. 376

Figure 15. Ceratoprion chaelae New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. 377

a.

d.

Figure 16. Ceratoprion mandi bularuro New Species. a — c. Aedeagus — a, dorsal aspect, b, lateral aspect, c, ventral aspect. Figure 17. Ceratoprion niarum New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Last abdominal sternite. e - f. Ultimate and penultimate abdominal tergites - e. internal view, f. external view. g. Internal male geni talia. 379

Figure 18. Ceratoprion periosus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. 380

f. Figure 19. Ceratoprion serricorne Gorham. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. e. Left hindwing. f. Left antenna. Figure 20. Flabellocaenia bourgeoisi Pic. a - c. Aedeagus - a. dorsal aspect, b. lateral aspect, c. ventral aspect. d. Lateral aspect head and prothorax. e. Last abdominal sternite. f - g. Ultimate and penultimate abdominal tergites - f. internal view, g. external view. 382

Figure 21. Flabellocaenia elegantulus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 383

Figure 22. Flabel 1ocaenia leticia New Species. a — c. Aedeagus — a, dorsal aspect, b, lateral aspect, c, ventral aspect. 384

Figure 23. Flabellocaenia iolvi New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. e. Frontal view of head. 385

Figure 24. Flabellocaenia pecki New Species. a - c. Aedeagus — a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. Figure 25. Flabellocaenia rimae New Species. a - c . Aedeagus dorsal aspect, b, lateral aspect, c, ventral aspect. 387

a. b. c.

d.

e. f.

Figure 26. Leptolvcus adiaphorus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. e. Last: abdabdominal cergite. f - g. Ultimate and Penultimate abdominal tergites - f. internal view, g. external view. 388

Figure 27. Leptolvcus brunneus New Species. a — c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. e. Internal genitalia. 389

Figure 28. Leptolvcus dominicensis New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. 390

Figure 29. Leptolvcus effeminatus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 391

Figure 30. Leptolvcus flavicol1 is Leng and Mutchler. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 392

wm

Figure 31. Leptolvcus heterocornis Leng and Mutchler. A. Habitus of adult male. 393

Figure 32. Leptolvcus heterocornis Leng and Mutchler. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. e. Right hindwing. 394

Oc

I mm i------

Figure 33. Leptolycus heterocornis Leng and Mutchler. Habitus of adult female. 395

ttjv'f**'!

Figure 34. Leptolycus heterocornis Leng and Mutchler. A. Habitus of larva. igure 35. Pseudacroleptus caeruleus New Species. a c. Aedeagus , dorsal aspect, b, lateral aspect, c, ventral aspect. 397

d.

e. Figure 36. Pseudacro1eptus neblinus New Species. a - c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. t

Figure 37. Pseudacro1eotus neblinus New Species. a. Thorax ~ dorsal aspect on right and ventral aspect on left. 399

Figure 38. Pseudacroleptus neblinus New Species. a. Head — ventral aspect. b. Right maxillary palp. Figure 39. Pseudacroleptus obscuricolor Pic, habitus of adult male (Type). 401

i d.

Figure 40. Pseudoceratoprion bechvne New Species. a - c. Aedeagus “ a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Lateral aspect head and prothorax. 402

a.

I t zr— t

Figure 41. Pseudoceratoprion brasi1iensis New Species. a ~ c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. d. Right antenna. Figure 42. Pseudoceratoprion colombiensis New Species. a — c. Aedeagus - a, dorsal aspect, b, lateral aspect, c, ventral aspect. Lateral aspect head and prothorax.