MALE AND FEMALE TERMINALIA AS A BASIS FOR TRIBAL CLASSIFICATION OF THE SUBFAMILY OF AMERICA NORTH OF MEXICO (CERAMBYCIDAE , COLEOPTERA)

By

SERGIO AUGUSTO FRAGOSO

A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

1978 ACKNOWLEDGEMENTS

This study was financially supported by Embrapa (Empre- sa Brasileira de Pesquisa Agropecuaria , Brazil) and was

carried through at the facilities courteously provided by

the Attractant, Behavior, and Basic Biology Laboratory,

USDA, and the Division of Plant Industry, Florida Department of Agriculture, Gainesville.

I want to express my thanks to Mr. T. J. Spilman (U.S.

Nat. Museum of Natural History) and Dr. U. R. Martins (Univ.

of Sao Paulo, Brazil) for their permission to dissect rare

specimens belonging to their respective institutions, and

to Drs. E. G. Linsley and J. A. Chemsak (Univ. of California) for their warm reception and helpful suggestions during a

month stay at Berkeley, as well as the donation of most of

the specimens utilized in this study.

I am grateful to my Supervisory Committee of the Uni-

versity of Florida, composed of Drs. R. E. Woodruff (chair- man), R. I. Sailer, S. H. Kerr, G. E. Allen (Dept. of

Entomology and Nematology) and J. Reiskind (Dept. of Zool- ogy) for their guidance on numberless occasions.

To Dr. Robert E. Woodruff and Mrs. Thelma Carlysle (USDA) whose , friendship and kind attention have been a constant support during the last 4 years, I do not know of any words capable of expressing my deep gratitude. TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS ii

ABSTRACT iv

INTRODUCTION 1

MATERIALS AND METHODS 5

GENERAL MORPHOLOGY OF THE TERMINALIA 9

TAXONOMIC ACCOUNT 22

ATLAS 43

Flow-chart of methods 43 Male terminalia of cerdo , diagrammatic. 44 Female terminalia of , diagrammatic 45

Male terminaliae 46

Female terminaliae 56

APPENDIX 1 (HABITUS) 72

APPENDIX 2 (STERNA) 81

APPENDIX 3 (HIERARCHICAL ARRANGEMENT) 84

LITERATURE CITED 86

BIOGRAPHICAL SKETCH 92

iii Abstract of Dissertation Presented to the Graduate Council of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

MALE AND FEMALE TERMINALIA AS A BASIS FOR TRIBAL CLASSIFICATION OF THE SUBFAMILY CERAMBYCINAE OF AMERICA NORTH OF MEXICO (CERAMBYCIDAE COLEOPTERA) ,

By

Sergio Augusto Fragoso

December 1978

Chairman: Robert E. Woodruff Major Department: Entomology and Nematology

A reclassification, at the tribal level, is proposed

for the of Cerambycinae in America North of Mexico,

based on male and female terminalia. The morphology and

terminology of generalized terminaliae are discussed and illustrated in diagrammatic cross -sections . This subfamily was found to have 2 well characterized divisions, based on

the presence or absence of scooplike bristles on sternites preceding the abdominal 9th segment, which implies different ovipositing behavior and habits. Most of the traditional characters (of Lacordaire, 1869) have been disregarded, since they appear to be a parallel evolutionary trend within mono- phyletic groups of tribes.

The following nomenclatural changes are proposed as a result of these investigations:

Division I - Ancilocerina , new status, as a subtribe of

Purpuricenini; , new concept, separated from ;

Nathriini, a new synonym of Obriina (strict sense); Hyboderina, new status, as a subtribe of Obriini.

iv . .,

Division II - Oemini, revalidated, separated from Methiini, Eumichthini , new synonym of ; Tillomorphina and Anaglyptina, new status, both as a subtribe of ; Sphaerionina (revalidated), Elaphidionina Rosaliina, and Dryobiina, new status, all four as subtribes of Hesperophanini

An atlas is provided for the habitus and terminalia (photographically) , and the sterna (diagrammatically)

v .

INTRODUCTION

Cerambycids occur worldwide and have been estimated to

comprise approximately 20,000 species (Linsley, 1959:99).

There is no general agreement among the specialists about

family subdivisions, other than the 3 traditional sub- families Prioninae, Cerambycinae and Lamiinae , which some

authors have even granted full family status. However, 7

other nomenclaturally valid subfamilies are currently re-

cognized (Parandrinae, Aseminae, Spondylinae, Lepturinae, Philinae, Anoploderminae and Oxypeltinae) ; the last 3 do not occur in North America.

Prionines and Lamiines excepted, the remaining groups

taken together are divided into approximately 100 entities,

still basically under the criteria of Lacordaire (1869) The latest worldwide catalogue (Aurivillius , 1912) used

the term "tribes" for all Lacordairean units (which had several denominations) , a taxonomic treatment generally

accepted since. All subsequent published monographs have

been restricted to certain geographical limits.

From 1962 to date, E. G. Linsley (lately coauthored with J. A. Chemsak) has been publishing "The Cerambycidae of North America," which has brought the scattered descrip- tions together, solved many nomenclatural problems, and

1 . .

pointed out many shortcomings on the knowledge of the family

"to inspire further research" (personal communication)

One of these shortcomings is the unsatisfactory, unnatural,

and chaotic arrangement of tribes within the subfamily

Cerambycinae (as well as within the 2 other traditional groups), which I hope my study will help to clarify.

Ideally, a tribal-level study should be cosmopolitan in scope, but the number of specimens, lack of access to specimens, together with limitations of time, and other limiting factors make such study impracticable, at least for the purpose of meeting the academic requirements of a dissertation.

Therefore, I chose to investigate the terminalia

(meaning all structures concerned with coitus and oviposi- tion, as preserved in dry specimens) of the subfamily Cerambycinae, of both sexes , and, for purposes of this study,

I have followed Linsley's definition of the subfamily. Chemsak and Powell (1966) , describing the bionomics of Tragidium armatum LeConte, called attention to the ethological and ecological significance of the morphological adaptations of abdominal somites preceding the so-called

"genital segment" (the ninth) in purpuricenoid cerambycids.

A preliminary survey of the group herein studied convinced me that these adaptations are more varied than initially believed, that the female terminalia may furnish the long sought solid morphological basis for a reclassification of these , and that the group is apparently polyphy- letic . . ;

3

The present dissertation covers 32 tribes, among which

I have included the tribe Cerambicini, which typifies the

family. Some have been synonymized by Linsley in his monograph (e.g., Oemini Sphaerionini , , etc.). The tribes

Trachyderini, Piezocerini, Lissonotini and Rhinotragini

are basically groups of Neotropical occurrence (each

represented in America North of Mexico by only one species)

hence I have left them for a similar future study. A special

effort was made to obtain the type species of each

which typifies each tribe, although a few were unobtainable. These cases are reported under the respective headings in the taxonomic account

Photomacrographs were chosen to illustrate the matter under study because they readily give an idea of the ef-

fectiveness of the methods applied, forbid symmetry "cor- rections," and limit substantially the "creativity" of the

illustrator. Line drawings are unsurpassed for diagrams, and these were used moderately to show generalized sections, etc

Habitus photos of each species studied, as well as line drawings of the sterna, are included in the appendices.

Some of these have never before been illustrated.

I regard the present work as complementary (as well as a tribute) to Linsley' s monograph; hence I have avoided duplication of information (i.e., synonymy, diagnosis, 4

etc.). but whenever some of this information was thought necessary it was included under the proper heading in the taxonomic account. . .

MATERIALS AND METHODS

The great majority of specimens included and figured

in this study were obtained at the E. 0. Essig Museum, Uni-

versity of California (Berkeley) ; the others came from the Florida State Collection of (Gainesville) , the

U. S. National Museum of Natural History (Washington), the

Museu de Zoologia da Universidade de S. Paulo, and my

private collection.

The specimens were cleaned, dusted, and legs and an-

tennae were arranged to compensate for the small depth of

field in order to produce the habitus pictures (appendix I)

A few rare specimens, often very old, had no data on the

label other than a city or a country. The habitus photo-

graphs were taken with the same procedure and apparatus

described in detail in Fragoso (1978)

Male and female terminalia have been processed as

graphically summarized on plate 1, which includes the

average time for each step. The technique is essentially

that of the late Prof. Lauro Travassos, with the exception

of the stain and the melanin elimination steps. The ab-

domen is simply (gently) broken from the pinned dry speci- men, boiled in 10% solution of potassium hydroxide (the

time depending on the degree of sclerotizat ion of the exo-

skeleton), and washed in 3 water baths using large Petri

5 "

6

dishes. At the second wash the abdomen was cut on one side,

opened, and gently pressed with a fine camel-hair brush.

The next step was the elimination of melanin by hair bleach.

This technique (producing excellent results) is from an un-

published, mimeographed sheet of instructions by Yoshiaki

Hokama and Charles L. Judson (Univ. of California, Davis).

They recommend the brand "Clairol" (8 parts "Clairoxide ,

2 parts "Lightening Booster," and 1 part "Lady Clairol

creme"), but others may work equally well. The active ingre-

dient is basically hydrogen peroxide. After- the bleaching,

I found it better to deviate from these instructions and to wash in an aqueous solution of household detergent to elimi- nate the oily component of the mixture. The abdomen was then

transferred to 75% ethanol, where it became very transparent.

To discern the membranous tube (which comprises most of the terminaliae) from other structures (e.g., tracheae, etc.), I found it necessary to stain before dissection.

Aqueous fuchsin (1%) stained sclerotized parts magenta, and, after rinsing in ethanol, a brief dip into a 10% filtered solution of "Parker Ink, permanent black" stained the membranes blue. This unusual stain yields superb re- sults, and I discovered it almost in despair, after un- successful trials with several histological stains.

When transferred to absolute ethanol, part of both stains dissolve, and dissection was made at this step. A final 7

clearing immersion in saturated alcoholic phenol cleared

the structures substantially further, and the photomacro- graphs were taken with the preparations under a cover slip. The thus prepared pieces were then kept in a solution of

10 parts ethanol, 10 parts glycerin, 10 parts water and one part phenol. A few very long ovipositors were mounted on microscope slides using the standard technique, imbedded in Canada Balsam.

The photomacrographs (by definition, without the use

of an eyepiece) were made with Zeiss Luminar macrolenses

(40 and 16 mm focal distances) and an Olympus planachromatic microscope objective of 1.3x (N.A. 0.03), all 3 attached

to the turret of a BH Olympus microscope stand, equipped with an aplanatic condenser (top element removed) . Between the field diaphragm and the condenser, I placed a finely grained frosted "daylight" filter of Leitz manufacture. Over the phototube of the trinocular head I attached a regular Nikon F2 camera (provided with an "M" focusing screen and a 6x magnifying viewfinder) by means of a con- necting tube, which kept the film plane approximately 950 mm from the top of the trinocular head phototube. Plus X (Kodak emulsion 5052, 125 ASA) developed in Microdol (1:2) yielded good exposures within the 2 to 6 seconds range, depending on the lens, the transparency of each individual preparation, and the rheostat setting. Most prints re- quired slight retouching to compensate for certain specimen 8

densities incompatible with the average tonal scale of the

image. This was particularly true of the fine setae at the external borders of the structures illustrated. Some ex- ceptionally long ovipositors had to be photographed in more than one frame, resulting in a final montage (indicated in

the respective legends) . The female terminalia variation made it impossible to arrange them alphabetically, the system followed in male, habitus, and sterna pictures. .

GENERAL MORPHOLOGY OF THE TERMINALIA

The larval cerambycid abdomen has, according to Duffy

(1953), 10 segments, the last comprises the lobes surround-

ing the anus. Iuga & Rosea (1962) consider the larval ab-

domen to be composed of 9 segments plus a terminal "mamelon."

The development of the definitive adult terminalia takes

place during the pupal stage from primordial cells , possibly

kept in those lobes or "mamelon." The pupal abdomen, also

according to Duffy (1953:85, fig. 23), has 10 segments, the

10th located ventrally between the 8th and the 9th (terminal) segment

The adult cerambycid abdomen is said to be cryptogas-

trous, in which segment 2 is fused with segment 3 and the

sternum is membranous (Jeannel & Paulian, 1944). Hence

the first externally visible sternite, proximal to the

thorax, is segment 3. Segments 1 and 2 are hidden and

form the posterior wall of the metacoxal cavities, both

distinctly less sclerotized than segment 3, but not neces- sarily membranous.

The generalized abdomen may be compared with a rubber

glove finger bearing several infoldings, which became larger

toward the abdominal apex. The infoldings are achieved by a distal double exocuticular sclerotized wall of both

dorsal and ventral plates, the internal wall of one segment

9 10

being connected to the external wall of the next by means

of an intersegmental membrane. This design permits several

movements, principally lengthwise, i.e., total or partial

protrusion (or evagination) of the somites, apparently ac-

complished by hemolymph pressure, caused by contraction of alary and peripheral muscles.

Near the distal borders of segment 8, the invagination

is usually greater, due to an extra elongation of the inter-

segmental membrane, which allows for complete concealment

of subsequent segments within the abdominal cavity. The

concealed segments show an increasing specialization from

one segment to the next, to perform the reproductive func- tions (coitus and oviposition) . Segment 8 is always dis- tinct; the proximal fold of its sternite bears a median apodeme in both sexes.

From somite 8 onward, the number and the interpreta-

tion of the subsequent segments of the coleopteran abdomen have been a matter of dispute and disagreement. Snodgrass (1935) in a generalized , discussion of the male genitalia of Coleoptera, mentions segment 10 as "a small projection from the dorsal wall of the entrance chamber, bearing the anus" (In Phyllophaga chiriquiana fig. , 303, p. 596). Later

(1957:31), the same author writes: "It might be suspected, therefore, that the (genital) organs really pertain to the venter of the reduced tenth segment." Thus, the two state- ments are apparently conflicting, but the last agrees with 11

Duffy's pupa figure. Britton (1970:504) categorically affirms

that the "abdomen (of Coleoptera) is composed of 10 segments

in both sexes, but segment 9 is modified as the genital seg-

ment and hidden within the body and segment 10 is greatly

reduced." These views are shared by Jeannel (1949), Handlirsh

(1933-36) and others. Iuga and Rosea (1962:104) state:

"... nous venons de constater que 1' abdomen des cerambycidae adulte comporte 11 segments dans les deux sexes (le derniere vestigial chez les femelles des lamiides)."

Iuga and Rosea' s (1962) paper is the only morphological

work dealing more extensively with both sexes of cerambycids (and comparing them with Hymenoptera) . Snodgrass (1935,

1957) discusses the male genitalia in all insect orders, but barely covers the female coleopteran genitalia. Never-

theless, his work is widely respected and his terminology amply employed. Michener (1956) tries to establish inter- ordinal homologies of male and female segments 8 and 9, concentrates chiefly on the male, redefines the terminol- ogy, discusses briefly the male coleopteran genitalia based on the exceptional Hydrous (Hydrophilidae) , and states: "... that the copulatory organs are new structures (probably of phallic origin) seems far less probable than that they are derived from pre-existing structures." Wood in (1952) a very short paper presents a comparative chart of the terminology of Sharp and Muir, Snodgrass, Michener, and his own, which is basically that of Michener with addition 12

of Snodgrass terms not defined by Michener. There is

no reference to segments, although it may be implied by

the adoption of Michener terminology, but the paper does

not illustrate any cerambycid and adds nothing to the

existing knowledge. Ehara (1954) confined his studies

to the parameres and aedeagus , figures 101 Japanese ceram-

bycids diagrammatical ly (no membrane is shown), and, using

character states as independent characters, confirmed

that the results generally coincided with the traditional

classification of the group. Lindroth (1957) in an un-

pretentious, but paradoxically very useful paper, presents

an extensive chart comparing the terminology used by English,

German, and French authors, then proposes latinization

and defines each term for both sexes. Matsuda (1976) sum- marized the general insect abdomen, order by order, con-

centrating on the embryology of formerly studied species

of Coleoptera Tenebrio , Trox ( , Lytta etc.), , but since

the species covered are not closely related to cerambycids, his work is of little help concerning my purposes. Wan-

dolleck (1906) and Tanner (1927) discussed the female genitalia only, including some cerambycids.

It is beyond the scope of this dissertation to discuss in detail the interpretation of cerambycid terminalia.

The brief description that follows helps to clarify the terminology employed and is based on Cerambyx cerdo 13

Linnaeus, type-species of the genus that typifies the family. Luckily enough, this species is covered by Iuga and Rosea, a female of the same genus is figured by Wandolleck, and above all its terminalia is quite generalized. Male Terminalia

Plate 2 represents a sagittal section of the male

terminalia of Cerambyx cerdo . Figure A maintains the ac- tual relationships among its component parts, and in fig. B these parts have been artificially separated for explana-

tory purposes and labelled sectors , numbered 1 to 3 , inward.

Sector 1 bears dorsally an inverted U-shaped sclerotiza- tion that protrudes from each side of the membranous tube, forming 2 apodemes, a structure that may be wanting in other cerambycid groups. The anal aperture is dorsally located, slightly posterior to this U-shaped sclerite, which articulates to a ventral, ever present, Y-shaped sclerotization at about the same level that the lateral apodemes arise. Essentially the same Y-shaped structure has been named by Snodgrass (1935:596, fig. 303B) as ster- nite 9, and is generally referred to as "spiculum gastrale" by many taxonomists. Iuga & Rosea do not consider these sclerotizations a segment, and use the self-explanatory terms "arc dorsal" and "arc ventral."

Sector 2, generally referred to as "parameres" in most taxonomic papers, is also composed of a dorsal, arcuate sclerite usually provided with 2 lobes (sometimes reduced to one as in and Obrium ) that project inside

the membranous tube and bear sensory hairs at the distal

ends. It also forms laterally an apodeme (not as long

and distinct as in the preceding sector 1) , and fuses

laterally to a V-shaped ventral piece. The arms of the "V" apparently fuse at the vertex in Cerambyx , but show

considerable variation in other groups.

The way in which the sclerotizations of sectors 1 and

2 are linked to the membrane, as well as their construc-

tion, suggests that they are homologous to segment 8. Snod-

grass (1957:30) termed the dorsal lobes "parameres , " the

lateral arms "phallobase" and pictured an internal annulus

(or ring) of an undetermined cerambycid (fig. 8G) which

I have not been able to find in any longhorned . Iuga and Rosea call the dorsal lobes "gonopode 9 , V the lat-

eral arms "hemisternite 9" which is equivalent to interpreting

the entire sector as segment 9.

Sector 3 comprises the "aedeagus" and the "internal sac" of most authors. Snodgrass employs the terms

"aedeagus" (defined as "the distal part of the phallus, usually the principal part of the intromittent organ, typically a sclerotic tube") and "endophallus" for the internal sac. In cerambycids the "sclerotic tube" is not intromittent, and its function appears to be that of opening the plicate female vulva, thus allowing the endo- phallus to evaginate and perform truly intromittent action. 15

Iuga and Rosea interpret the sclerotized aedeagus as segment

10, the tergal plate "tergite 10," the ventral "sternite

10, " both plates fusing together to form the paired

"apodemes 10." The endophallus is usually provided with sclerotizations (spines, hooks, etc.), generally referred as "armature" in taxonomic literature. These serve to fix the membranous endophallus in the vagina during coitus

The distal portion of the internal sac is connected to the ejaculatory duct, and at this point some sclerotic pieces usually occur, which Iuga and Rosea termed "segment

11. " All the components of the terminalia including the ejaculatory duct are of epidermal origin, and resist well the 107o KOH treatment. However, in a few specimens, the distal portion of the internal sac is missing, probably due to uncontrolled circumstances of the drying process.

The remaining parts of the reproductory system are of mesodermal origin and are not preserved in dry specimens.

In the Taxonomic Account that follows I have used the terms dorsal arc and ventral arc for sector 1, para- meres^ for the entire sector 2 and aedeagus and endophallus for sector 3, for this is a convenience, and does not imply the acceptance of any of the interpretations dis- cussed above.

Female Terminalia

Among the Cerambycinae (sensu Linsley) , included in this study, female terminalia present 2 homologous but characteristically distinguishable forms which imply dif- ferent ovipositing habits. These are 16

a) cerambyciform : Capable of considerable protrusion,

and devoid of scooplike bristles;

b) raduform (from radere , Latin meaning to scratch, to touch in passing) : Capable of substantially less

protrusion, and provided with characteristic scoop-

like bristles, curved ventrally at the apex, de-

signed to collect substrate dust which is subse-

quently attached to the viscous egg, laid on the

surface of the host plant (see Chemsak and Powell,

1966) The . raduform terminalia may be further

divided into 3 subforms, according to the segmental distribution of the bristles.

The cerambyciform terminalia is represented in sagittal section on plate based 3, on Cerambyx cerdo . Figure A shows

it invaginated, and fig. B is a diagram of the same terminalia evaginated, where some proportions were disregarded for ex- planatory purposes. Like the male, it is divided into 3 sectors, numbered 1 to 3.

Sector 1 is the tubular intersegmental membrane 8-9, where on each side of the dorsal anal aperture there is a somewhat curved sclerotized rod laterally embedded in the membrane. These rods protrude dorsally and partially reinforce a membranous ventral apodeme at each side, the 2 of which become internal to the tube with the evagination.

The sclerotic rods have been termed "tergit 10" by

Wandolleck, "paraproct" by Tanner, and "gonopode 8" by " ,

17

Iuga and Rosea. The term "paraproct" is defined by Snodgrass

(1935:255) as "two ventrolateral lobes of the 11th segment of lower ," although Tillyard, cited by Torre Bueno

(1939), defines the same term as a "pair of lobes bordering the anus laterally."

Sector 2 has its tubular wall slightly yellowish, and

stains differently from sector 1. Ventrally it bears

paired sclerotizations in the form of a "wishbone" which the shorter arm curves laterally and the longer arm is

substraight, vaguely articulating with a rod of the next sector 3. Wandolleck calls this ventral structure "tergit

9," Tanner "valvifer," and Iuga and Rosea "hemisternite 9." This sector together with sector 3 has been considered as segment 9 by most authors. "Valvifer" is defined by Snodgrass (1935:622) as "the basal plates of the oviposi- tor, probably derived from the coxopodites of the gonopods carrying the valvulae, including first valvifers of the eighth abdominal segment and second valvifers of the ninth

segment .

Sector 3 is separated from sector 2 by a short but distinct fold which stains like sector 1. Dissection shows paired, thin, membranous apodemes deriving from the fold. Distally, at each side, it bears a bulbous struc- ture provided with a distinct articulated appendage, usually referred to as "stylus," which has sensory setae apically. The whole sector, which has been considered as 18

a gonopod of segment 9 by most authors, is reinforced both dorsally and ventrally by paired, longitudinal rods. The ventral rods have been termed "versteifugsstabe" (reinforc< ments) by Wandolleck, "coxite" by Tanner, and "gonopod 9" by most authors.

Between the bulbous apical lobes that bear the styli opens the vulva, and from this point on the generalized tube invaginates permanently inward, forming the vagina. The tubular vagina connects with a transversely plicate uterus, which, at a distal point beyond the middle widens, connects with the common oviduct and bears 2 ventral sclerotized apodemes. The uterus continues beyond this chamber, receives the petiole of the spermatheca (provided with an accessory gland), and forms a pouch (bursa copula- trix), whose walls are not plicate. All these permanently internal (not evaginable) organs are distinct in both dissected females of Cerambyx cerdo , and according to Matsuda (1976) they might be of ectodermal origin, although there is some dispute and variation concerning the deriva- tion of the more internal reproductive organs in different families of Coleoptera. Wandolleck and Tanner did not consider the structures beyond the vulva, while Iuga and Rosea call the vagina "segment 11" and the uterus "segment 10."

In the Taxonomic Account that follows I have resorted to the "sector system" referring to dorsal, lateral, and ventral sclerotizations within each sector, since there 19

is a wide variation in the proposed interpretations by the authors that dealt with female terminalia. From the

vulva inwards , there is a general agreement on the tradi- tional terminology.

The raduform female terminalia may be further divided into 3 different arrangements , according to the segmental occurrence of the scoop-like bristles, as follows:

a) methiform : scoop-like bristles short petiolate,

uniform in size, arranged in a single row at the

V-shaped distal border of sternite 7 (plate 18)

interspersed with elongated, pointed bristles.

b) obriform : scoop-like bristles located in an

arcuate concavity of sternite 4, in several rows

of posteriorly increasing petiole lengths. At

the sides of this concavity there are stout, curved,

blunt-pointed, arcuate bristles that overlap mesally,

and at the antero-central portion of the concavity

there is a tuft of spatulated shorter bristles

(plate 16). A variant of this arrangement, that could be termed hyboderiform , has the lateral

bristles of sternite 4 pointed, and lacks the

anteromesal tuft of spatulated bristles. The

scoop-like bristles have their origin on a fold of

sternite 4, their petioles vary in length, and

are arranged close together, the longer ones form-

ing a brush that ends in a transverse straight .

20

line. Sternites 5 and 6 form a concave debris

pocket bordered by long bristles directed inward

(plate 17)

c) purpuriceniform : scoop-like bristles located at

the distal border of sternite 8, in several rows

of distally increasing petiole lengths (plate 14).

Assuming that the genetic mechanisms necessary to

produce such scoop- like bristles (which imply unique ovi-

positing habits as well as a substantially shorter male

endophallus) characterize some degree of phylogenetic

affinity, it remains to postulate how the 3 raduform ter-

minaliae relate to each other, as well as their relation-

ship with the generalized cerambyciform ovipositor. Since

essentially the same unstudied scoop-like bristles occur

in many tribes of non-North American distribution (e.g., Torneutini, Pteroplatini , some Platyarthrini Plectogas- ,

terini, , Gahanini, Megacoelini, etc.), it is dif-

ficult to ascertain which is ancestral and which is derived.

However, the uniformity and single row arrangement of the methiform bristles appears ancestral compared with the other 2 more complex arrangements, an opinion which is compatible with other characters, usually considered

"primitive," (e.g., the intercoxal process of the sternum, the posterior border of the mesocoxal cavities, etc.). No intermediate form was found that would suggest a link or derivation between the raduform and cerambyciform 21 terminaliae, although the Oemini share some other primiti characters with the Methiini. ,

TAXONOMIC ACCOUNT OF THE SUBFAMILY CERAMBYCINAE LATREILLE 1804

The following account is basically confined to termi- nalia characters, which bear a direct relationship to repro-

duction, and may serve as a general framework for the clas-

sification of cerambycids. However, a sound, detailed

classification at the tribal level cannot be expressed by a single structure, notwithstanding its complexity. It will be a long time before the holomorphy of Hennig (1966) or the omnispective classification of Blackwelder (1967) can be applied to this group. I believe this study will improve the systematics of what is termed below Division I, but the notes and remarks on taxa under Division II should

be viewed as preliminary, since the terminalia structures

are very similar among its tribes.

Traditionally, the characters used to classify the

Cerambycinae are said to present a mosaic distribution (i.e., the same structures appearing in groups of genera not closely related, a coincidence interpreted usually as a sign of past hybridization). Partially, this mosaic is a consequence of superficial observation, or the usual practice of labeling as a "tribe" a homogeneous group of genera (while some older groups are left heterogeneous, ambiguous, and inconsistent), or both.

22 . .

23

The examples of superficial observations (or improper

description) are abundant: The eburneous fasciae or maculae

may be developed as a part of a costa or not, but are usually

referred to only as "ivory marks" or similar expressions;

the mesepimeron always reaches the mesocoxa, whether under

or at the same level of adjoining sclerites, although re-

ferred to as "open or closed to epimeron "; the prothoracic

intercoxal process may function as a fulcrum to coxal move-

ments or not, regardless of its appearance, usually simply

stated as "open or closed behind," etc. The sternal charac-

ters, which have been used heavily (and inconsistently) to

define the tribes dealt with herein, are illustrated diagram-

matically on Appendix II.

To cope with closely related groups which have been

considered as tribes, I resorted to using subtribes, adopting

the commonly used termination ina.

Division I

Terminalia characters: Female with scooplike bristles

on any sternite preceding 9th; ovipositor (sectors 1+2+3)

shortened, usually about the same length as the sternal

plate of segment 8, adapted to lay eggs on host substrate. Male with endophallus proportionately shorter, usually about the same size as aedeagus

Tribe 1: Purpuricenini Fairmaire 1864

Type genus and species: Purpuricenus desfontainii

(Fabricius, 1792), known from Greece, Syria, and North Africa (plate 35A) .

24

Terminalia characters: Female with scooplike bristles arising at the distal border of sternite 8 in several rows of variable petiole lengths; border of tergite 8 divided by

2 notches into 3 apical, flattened lobes (plate 14A) . Male with dorsal lobes of parameres short, but distinct (plate

11C) .

Remarks: Linsley (1962:103), "by present designation"

cites P. kaehleri Linnaeus as the type of Purpuricenus , but McKeown (1947:105) and Gressitt (1970:177) accepted Thomson's

(1864:198) designation of P. desfontainii . Chemsak and

Linsley (1974) transferred to this tribe the following enera: Elytroleptus 8 , Parevander , Parathetesis , and Pteroplatidius from the Pteroplatini ; Chlorida and Chrotoma from the Hesperophanini and , Zenochloris from the Heteropsini. By the same criterion, i.e., the brush of scooplike bristles (Chemsak, personal communication) , Ancylocera should be transferred also. Since several other genera typifying tribes will be transferred to this tribe in the near future,

I have kept Ancylocerina at the subtribal level.

Subtribe 1: Purpuricenina Fairmaire 1864 (strict sense)

Type genus and species: Purpuricenus desfontainii (Fabricius, 1792) Terminalia characters: Female with distal part of sector 3 subparallel (plate 14A) . Male with distal corners of sternite 8 not projecting laterally (plate 11C) . . . . .

25

Remarks: This group is equivalent to the tribe Pur-

puricenini of Linsley, plus the transferred taxa (Chemsak

and Linsley, 1974)

Subtribe 2: Ancylocerina Lacordaire 1869 (new status) Type genus and species : Ancylocera cardinalis Dalman

1822, known from Southeastern Brazil (plate 29D)

Terminalia characters: Female with distal portion of

sector 3 divergent (plate 15). Male with distal corners

of sternite 8 projecting laterally (plate 5A)

Remarks: This tribe is represented in America North

of Mexico by a single species, A. bicolor (Olivier, 1795),

which can be recognized by the black, subcylindrical prono-

tum, truncated red elytra, flattened antennal basal segments,

occurs in Southeastern United States.

Tribe 2: Methiini Thomson 1860

Type genus and species: pusilla (Newman, 1840),

known from Southeastern United States (plate 33C)

Terminalia characters: Female with scooplike bristles uniform in length, short petiolate, arising in a single row on the apical border of sternite 7; styli buttonlike, hemi-

spherical (plate 18A,B,C). Male with tergite 8 lateroapi- cally folded to join the narrow sternite; dorsal lobes of parameres semifused, appearing emarginate (plate 9C)

Remarks: Linsley (1962:13) has combined this group with the "Oemides** of Lacordaire (1869:216), totaling 10 genera, of which only is studied herein, in Division II......

26

The group is under worldwide revision by U. R. Martins (per-

sonal communication)

Tribe 3: Obriini Mulsant 1839

Type genus and species : Obrium cantar inum (Linnaeus

1967)

Terminalia characters: Female with scooplike bristles arising at the distal border of sternite 4 in several rows of unequal petiole lengths; styli cylindrical (plate 16A,B).

Male with a single, mosodorsal lobe; aedeagus conical, pointed distally (plate IOC)

Subtribe 1: Obriina Mulsant 1839 (strict sense)

Synonym: Nathriini Linsley 1963 (new synonym)

Type genus and species: Obrium cantharinum (Linnaeus,

1767) , known from Southern Europe (plate 33D)

Terminalia characters: Female with a tuft of shorter, spatulated bristles preceding the brush composed of scoop- like bristles, which are limited, at each side, by distinctly longer, mesally curved, stout bristles bluntly pointed

(plate 16A.B). Male parameres rounded ventrally (plate IOC). Remarks : The monotypic genus type of ,

Narthriini, is so similar to Obrium in all aspects of the terminalia (plates 10 and 16) , that they clearly belong to the same tribe, although the coxal cavities differ between the 2 genera.

Subtribe 2: Hyboderina Linsley 1940 (new status)

Type genus and species: Hybodera tuberculata LeConte

1873, known from the Pacific coast of North America (plate 33A) . . .

27

Terminalia characters: Female without the tuft of

spatulated setae preceding the brush composed of scooplike

bristles, which are limited at each side, by longer, slender, acute pointed bristles (plate 17) . Male parameres with 2

ventral apodemes (plate 9A)

Remarks: I have used H. debilis LeConte to illustrate

male terminalia since a male of the type species was not available to me.

Division II

Terminalia characters: Female without scooplike bristle

on any sternite; ovipositors usually elongated, adapted to lay eggs into the host (fissures or crevices) . Male endo-

phallus proportionately longer, usually more than twice the length of aedeagus

Remarks: The relationships among the tribes included

in this division are not clearly understood. Their defini-

tions are not as sharp as the tribes bearing scooplike setae, but future studies of worldwide scope will undoubtedly clarify their classification.

Tribe 1. Fairmaire 1864

Type genus and species: (Linnaeus,

1767) , known from Europe (plate 36C) Terminalia characters: Female with sector 1 bearing light sclerotizations of indistinct borders similar to hemisternites basal , apodemes of sector 2 distinctly scler- otized, usual rods wanting; styli elongate, subcylindrical . . .

28

(plate 26A) . Males with sternite 8 deeply emarginate,

dorsal arc indistinct, endophallus bearing sclerotizations

distally to usual hooks, parameres with a single dorsal

lobe, and 2 ventral apodemes (plate 13C)

Remarks: Iuga and Rosea (1962) considered the ovipos-

itor of Stenopterus the most generalized, but they did not

consider any taxa herein under Division I (with the scoop- like bristles preceeding the ovipositor) . The absence of

rods is somewhat exceptional in this Division II, although

the tribe Stenopterini definitely lacks the scooplike bristles

Tribe 2: Oemini Lacordaire 1869 (revalidated)

Type genus and species: Oeme rigida (Say, 1826), known

from the Atlantic Coast of North America (plate 34C)

Terminalia characters: Female with a single rod con-

fined to sector 3, uterine pair of apodemes exceptionally thick, styli subclavate (plate 25A) . Male with a long

sclerotic rod inside endophallus; dorsal lobes of parameres wide and flattened, dorsal arc absent.

Remarks: The normal, short 2nd antennal segment dis-

tinguishes the taxa under this tribe from the .

Methia, considered by Linsley (1963:13-14) as a "specialized

section" among the components of this tribe, belongs to

Division I.

Tribe 3: Opsimini LeConte 1873

Type genus and species : quadrilineatus

Mannerheim 1843, known from the Pacific Coast of North

America (plate 34D) . . . . . ,

29

Terminalia characters: Female with 2 ventral rods at

each side of sectors 2+3 (the internal with a discontinuity

about the fold between Sectors 2 and 3) , styli clavate (plate 22C) Male . with aedeagus bifurcated ventrodistally

endophallus with a long sclerotic rod narrowly branched

distally, dorsal lobes of parameres wide and flattened (plate 11B)

Remarks : Qpsimini can be recognized by the 2nd

antennal segment, which is almost as long as 3rd, and the pronotum, which is "broadly arcuate-emarginate , emargina-

tion filled with thin, corneous plate" (Linsley 1962:2).

Tribe 4: Mulsant 1839

Type genus and species: minuta (Fabricius, 1781) known as an , introduced species from Europe in

Eastern North America (plate 32B)

Terminalia characters: Female with ventral rod of sectors 2+3 continuous, styli clavate (plate 25B) . Male without dorsal arc, dorsal lobes of parameres almost fused, appearing as single emarginate lobe (plate 8 where, on the segment 8 the tergite is accidentally missing)

Remarks: The position and relationships of Gracilia remain obscure, as well as the two other North American genera assigned to the tribe ( Hypexilis and Perigracilia )

Tribe 5: Molorchini Mulsant 1862 Type genus and species: Molorchus minor (Linnaeus 1758) known , from Europe (plate 33D) . . . .

30

Terminalia characters : Female with ventral rods of sectors 2+3 continuous, styli clavate (plate 26B) . Male

without dorsal arc, endophallus without sclerotizations

distally to usual hooks, parameres with both dorsal lobe

and ventral apodemes simple (plate 10A)

Remarks: According to Linsley (1963:157), the type

species of Molorchus is "uncertain," but McKeown (1947:83)

and Gressitt (1970:109) adopted Thomson's (1864:150) designa

tion of M. minor , a criterion adopted here.

Tribe 6: Holopleurini Chemsak and Linsley 1974

Type genus and species: Holopleura marginata LeConte

1873, known from Northern Mexico and Southern United

States (plate 32D)

Terminalia characters: Female with rods of sectors

2+3 continuous, vulva somewhat recessed below the bulbous

apical lobes that bear clavate styli (plate 24A)

Remarks: Holopleura males were not available to

study. Linsley (1962:181) included Holopleura and Elytroleptus in the tribe Pteroplatini . Chemsak and

Linsley (1974) transferred Elytroleptus to the tribe

Purpuricenini and proposing Holopleurini as a new tribe which contains currently a single monotypic genus.

Tribe 7: Lacordaire 1869

Type genus and species: (Say 1826) known , from the Atlantic Coast of North America (plate 36A) . . . .

31

Terminalia characters: Female with apodeme 8 as long as the abdomen, ventral rods of sector 2 and 3 discontinuous and bypassing each other about intersectorial fold (plate 21A). Male without dorsal arc, endophallus exceptionally long, simple (plate 13A)

Remarks Among all : species herein studied, females of

S. cucujiforme have the longest ovipositor. The relation- ships of this small tribe are still obscure, although the

flattened body, terminalia, palpi, etc., are, apparently,

specializations not to be found in forms considered primi-

tive .

Tribe 8: Callidiini Mulsant 1839

Synonym: Eumichthini Linsley 1940 (new synonym) Type genus and species: violaceum (Linnaeus,

1758), known from Europe, Transcaucasia, Siberia, China, Korea, Japan, and Northeastern North America (plate 30B, specimen from Europe)

Terminalia characters: Female with rods of sectors 2+3 discontinuous, but not bypassing each other (plate 20B) Male without a dorsal arc, endophallus with setose hairs distally of usual hooks; dorsal lobes of parameres elongate and thin (plate 5C)

Remarks: Eumichthini (type genus and species: Eumichthus oedipus Leconte 1872, plate 32A, from North America Pacific Coast) has been characterized by the 2nd antennal segment, which is half as long as the 3rd. Since . .

32

some genera currently assigned to Callidiini have the same

segment almost half as long as the 3rd (and most other

characters fit well the tribal diagnosis) , I see no reason

to maintain Eumichthini as a separate group. Superficially, Eumichtus has some affinities with Phymatodes .

Tribe 9: Clytini Mulsant 1839

Type genus and species: arietis (Linnaeus 1758). Terminalia characters : Female with rods of sectors 2

and 3 continuous or discontinuous, styli clavate. Male

without dorsal arc, endophallus with sclerotizat ions dis-

tally to usual hooks, or setose hairs, or both. Remarks Taxa : under the tribe Clytini are apparently

derived from an ancestral form that shared some characters

with modern Callidiini. The Clytini has worldwide distribu-

tion, and in a preliminary way, I have herewith considered

the Tillomorphini and Anaglyptini as subtribes, since they have several characters in common.

Subtribe 1: Clytina Mulsant 1839 (strict sense) Type genus and species: Clytus arietis (Linnaeus, 1758), known from Europe (plate 31A) Terminalia characters: Female with rods of sectors 2 and 3 discontinuous, but not bypassing each other (plate

21B). Male without dorsal arc, aedeagus with sclerotized spines distally to usual hooks (plate 6C)

Remarks: This subtribe can be recognized by the mete- pimeron produced over the angles of 1st abdominal sternite, and enclosing posterior coxae externally. . . . . ,

33

Subtribe 2: Tillomorphina Lacordaire 1869

Type genus and species: Tillomorpha lineoligera

Blanchard 1851, known from Chile (plate 36D)

Terminalia characters: Female with sternite 8 U-shaped, ventral rods of sectors 2+3 continuous (plate 26C) . Male

with the foramen of parameres rounded (no ventral apodeme)

dorsal lobes widely separated, emarginated to their bases;

endophallus with strongly sclerotized spines distally to

the usual hooks (plate 8C)

Remarks: Within Division II, Tillomorpha has the

shortest sectors 2+3 (segment 9 of most authors) of all

species herein studied. All North American genera cur-

rently assigned to this tribe (, , and

Pentanodes ) have a transverse ivory band on each elytron.

The specimen figured on plate 26C has the complex uterus/

vagina unnaturally evaginated, possibly as a consequence of the killing process.

Subtribe 3 : Anaglyptina Lacordaire 1869

Type genus and species: Anaglyptus gibbosus Fabricius

1787, known from Europe (plate 29C)

Terminalia characters: Female with ventral rods of sectors 2 and 3 discontinuous, but not bypassing each other in the usual way (plate 23A) . Male without a true dorsal arc, although the ventral arc arms are folded to form two lateral apodemes; endophallus bearing several scletorized spines distally to the usual hooks, and setose hairs at distal end (plate 4C) . . . .

34

Remarks: This subtribe can be recognized by the basally

gibbose elytra without transverse ivory fascia.

Tribe 10: Cleomenini Lacrodaire 1869

Type genus and species: Cleomenes dihammaphoroides Thomson 1864, known from the Philippines (plate 30D) Terminalia characters: Female with segment 8 cylin- drical, ventral rods of sectors 2+3 continuous (plate 22B)

Remarks: Males of Cleomenes were not available to

study. The single species of this tribe occurring North of Mexico is dispar Chevrolat, which can be

distinguished from by the antennae shorter

than the body in both sexes.

Tribe 11: Blanchard 1845

Type genus and species: Rhopalophora collaris (Germar, 1824) known from , Atlantic Coast of Brazil (plate 35B) Terminalia characters : Female with segment 8 trape- zoidal, ventral rods of sectors 2+3 continuous, sector 3 comparatively short (plate 24D) . Males without dorsal arc, aedeagus bearing angulate sclerotic structures distally to the proximal, usual hooks (plate 12A)

Remarks: If future studies establish the presence of distal sclerotic structures in the aedeagus of Cleomenini, it should be considered, at most, a subtribe of Rhopalophorini

R. longipes meeskei Casey was used to illustrate the female terminalia, since this sex of the type species was unobtain-

able . . . . .

35

Tribe 12: Callichromatini Thomson 1860

Type genus and species: Callichroma auricomum (Linnaeus, 1767) known from , the Atlantic Coast of Brazil (plate 30A) Terminalia characters : Female ventral rods of sectors

2 and 3 discontinuous, bypassing or not each other (plate 23C) Male without . a dorsal arc, with complex sclerotized

structures at the distal end of endophallus, lateral apodemes

distinctly developing from the dorsal lobes of parameres

(plate 5B)

Remarks: This worldwide tribe is represented in

America North of Mexico by a single genus, Plinthocoelium ,

which may be recognized by the metallic integument, large

size (29 to 38 mm), complete elytra, and outer lobe of

maxillae with a brush of hairs at apex.

Tribe 13: Lacordaire 1869

Type genus and species: surinamum (Linnaeus,

1767), known the Atlantic Coast, from Florida to Argentina (plate 29A)

Terminalia characters : Female with ventral rods of

sectors 1 and 2 discontinuous, but not bypassing each other (plate 23B) . Males with distinct dorsal arc, complex

sclerotic structures at distal end of endophallus, apodemes of parameres distinctly developing from the dorsal lobe of parameres (plate 4A)

Remarks: Achrysonini is represented in America North of Mexico by the type species. . . . ,

36

Tribe 14: Curiini LeConte 1873

Type genus and species: Curius dentatus Newman 1840,

known from Southeastern North America (plate 31B)

Terminalia characters: Female with ventral rods of

sectors 2 and 3 discontinuous, bypassing each other (plate

24C) . Male with dorsal arc, distal end of endophallus

bearing setose hairs, lateral apodemes of parameres in-

distinct (plate 6C)

Remarks: This tribe is traditionally characterized

by the clavate femora armed beneath with a .broad tooth.

Tribe 15: Ibidionini Thomson 1860

Type genus and species: Ibidion comantum Serville

1834, known from Brazil.

Terminalia characters: Females with rods of sectors

2+3 continuous (plate 24B) . Males with dorsal arc, distal

end of endophallus bearing sparse setose hairs, lateral

apodemes of parameres indistinct (plate 9B) Remarks : Ibidion is currently a monotypic genus

and, according to U. R. Martins (personal communication),

only three specimens are known to exist in collections.

Thus, I have used Heterachtes ebenus Newman 1840 (plate

33B) to illustrate the tribe.

Tribe 16: Agallissini LeConte 1873

Type genus and species: Agallissus melanioides

Dalman, 1823, known from Mexico. .

37

Terminalia characters: Female with the ventral rods

of sectors 2 and 3 discontinuous, bypassing each other

(plate 20A). Male with dorsal arc, distal end of endophallus

bearing some minute spines and irregularly plicate between

the lateroposterior arms, apodemes of parameres indistinct

(plate 4B)

Remarks: This tribe, traditionally characterized by elytral epipleura deeply emarginate near prominent humeri, is illustrated by A. lepturoides (Chevrolat, 1849), which occurs from Southern Texas to Southern Mexico.

Tribe 17: Hesperophanini Mulsant 1839

Type genus and species: Hesperophanes sericeum

(Fabricius, 1787).

Terminalia characters : Female with ventral rods of sectors 1 and 2 discontinuous. Males with distinct dorsal arc, proximal hooks of endophallus transverse at rest, followed by a series of regular spaced, transverse plicae between aedeagal lateroposterior arms.

Remarks: The following 5 groups are closely related, but traditionally very poorly separated. Knowledge of terminalia of taxa contained under each group will furnish a better understanding of their relationships, but at the present state, it appears to me, the better criterion to adopt is to treat them as subtribes. The characters for each subtribe are tentative only. . . .

38

Subtribe 1: Hesperophanina Mulsant 1839 (strict sense)

Type genus and species: H esperophanes sericeum

(Fabricius, 1787), known from Europe and Northern Africa (plate 32A)

Terminalia characters: Male with endophallus destitute

of sclerotizations and setose hairs; paramere lateral

apodemes developing from dorsal lobes (plate 8B)

Remarks: Linsley (1962:52) distinguished this taxon

from Elaphidionina by the intercoxal process of the pronotum unexpanded behind coxae, (an unreliable character, since it varies within the tribe) , and included in his concept the "Eburiides" of Lacordaire. Females were not available for study.

Subtribe 2: Elaphidionina Thomson 1864 Type genus and species: Elaphidion spinicornis (Drury, 1773) known , from Jamaica (plate 31D)

Terminalia characters: Female with ventral rods of

sectors 2 and 3 discontinuous, curved inward, thus giving the impression of crossing (plate 27). Male with all sclerotized parts narrow and elongate, endophallus with a thin reinforcing rod, more distinct dis tally, paramere lateral apodemes indistinct.

Remarks: Linsley (1963:1) considered under Elaphidionini 25 North American genera, including part of Phoracanthini and all Sphaenionini , groups treated separately by Lacordaire (1869). Phoracantha is a genus of Australian distribution, although one species has been . . . . .

39

introduced into South America. Sphaerion is treated herein

as a separate subtribe (see below)

Subtribe 3: Sphaerionina Lacordaire 1869 (revalidated under new status)

Type genus and species: Sphaerion cyanipenne Serville

1834, known from the Atlantic Coast of Brazil (plate 36B) Terminalia characters : Male with minute spines

subdistally in endophallus, paramere apodemes developing

from its dorsal lobes (plate 13B)

Remarks: This group has been characterized by the

carinate tibia (Lacordaire, 1869), but it remains to be

verified if there is a correlation between this character

and the minute spines of the endophallus. I could not find

any distinctive character on the female terminalia (plate

21C) .

Subtribe 4: Rosaliina Fairmaire 1864

Type genus and species: alpina (Linnaeus. 1758), known from Europe (plate 35C)

Terminalia characters: Female with ventral rods of sectors 2 and 3 slightly bypassing each other (plate 28A)

Male with aedeagus medially widened to permit evagination

of distally bulbous endophallus, bearing a few pointlike, minute sclerotizations subdistally.

Remarks: Rosalia was formerly assigned to (Lacordaire, 1869) , basically a Neotropical group, but Linsley (1963:4) unearthed Fairmaire ' s tribal name rank, which, to my knowledge, includes only 2 genera ( Rosalia and Acrocyrtidus , . . . . ,

40

from Asia). The single representative in North America is

R. funebris Motschulsky 1845 closely related to the type species

Subtribe 5: Dryobiina Linsley 1964

Type genus and species : Dryobius sexnotatus Linsley

1957, known from Eastern North America (plate 31C)

Terminalia characters: Male with endophallus bearing pointlike sclerotizations subdistally, but anterior to a

distal tube, which has its walls covered by small, short

spines (plate 7A)

Remarks: Dryobius has formerly been classed with the Oemini, an obvious misplaced classification. Currently, the group includes only 2 monotypic genera (the type genus and Ornithia) both , occurring in America North of Mexico. I could not find any distinctive character on the female ter- minalia (plate 28B)

Final Remarks

The following 2 tribes, without representatives in America North of Mexico, have been pictured, but not dis- cussed. These are

a) Cerambycini Latreille 1804 (type genus and species: Cerambyx cerdo Linnaeus 1758, known from Europe, (plates 6A, 19, and 30C) which has , some affinities with Hesperophanini although the male terminalia is distinctive. It typifies the family and was the basis of the morphological discussion b) Saphanini Lacordaire 1869 (type genus and species: Saphanus. piceus Laicharting 1784, known from Europe, plates 41

12C, 22A, 35D) of , uncertain current status, and which for- merly included genera later transferred to Opsimini. It was found to have distinctive male terminalia. ATLAS H,0

T=2min - T=2min. T=VAR.

MELANIN ELIMINATION STEP ETHANOL , _ DETERGENT r . "CLAIROL"

T=5min. T=2min.

STAINING STEP CAMERA ETHANOL

DISSECTION

.ate 1. Flow-chart of methods.

43 9

44

I I SNODGRASS I | AEDEAGUS PARA MERES

I ENDOPHALLUS I PHALLOBASE STERNITE 9 1 1 I I I

IUGA a ROSCA I INTERSEG. 10-11 | TERGITE 10 | GONOPOD ARC VENTRAL I | SEG. II J STERNITE I0 HEMISTERNITE 9 ARC DORSAL

Plate 2. Male terminal ia of Cerambyx cerdo : (A) sagittal section, (B) same, diagrammatic, artificially extended. —

45

/ SECTOR ONE 7 TERGITE 8-, 7— SECTOR TWO /SECTOR7 THREE— J APODEME / \ */ VULVA / : J__~ g ===:::^

TANNER

PARAPROCT VALVIFER COXITE

I

I

IU6A 8 ROSCA I I

I / SEG. 10 GONOPODE 8 GONOPODE 9 SEG. II HEMISTERNITE 9

Plate 3. Female terminalia of Cerambyx cerdo: (A) sagittal section (invaginated)7~riri¥nieT~aTagrammatic Cevagmated in ovipositing mode) . 46

4. Segment 8, aedeagus and parameres: (A) Achryson sur inamum , (B) Agal lissus lepturoides ' , XBj Anaglyptus gibbosus :

47

Plate 5. Segment 8, aedeagus and parameres (A) Ancylocera cardinalis , (B) Calichroma auricomum , (C) Callidium violaceum. .

Plate 6. Segment 8, aedeagus and parameres: (A) Cerambyx cerdo , (B) Clytus arietis , (C) Curius dentatus :

49

Plate 7. Segment 8, aedeagus and parameres (A) Dryobius sexnotatus , (B) Elaphidi on spinicorne , (C) Eumichthus oedipus" , ,

50

Plate 8. Segment 8, aedeagus and parameres: (A) Gr acilia minuta (tergite 8 wanting) (B; Hes perophanes serirpng (C) Tiliomorpha lin eoligera . 51 52

10. Segment 8, aedeagus and pararaeres: (A) Molorchus minor, (B) , (C) Obrium cantharinum 53 :

Plate 12. Segment 8, aedeagus and parameres (A) Rhopalophora co llaris , (B) Rosal ia alpjjia, (C) Saphanus piciu^s. 55

Plate 13. Segment 8, aedeagus and parameres (A) Smodicum cueuji forme , (B) Sghaerion cyanlpenne (C) Stenopterus , rufus . 56

Plate 14. Female terminalia: (A) Purpuricenus desfontainii, scooplike~HFistles detached

desfontainii . brush C . ^ of scooplike r S CC) K^Tiana cincta (Purpuriceni) g F M , S.E.M. %iPhoto by T. C. Carlysle. 57 58 :

59

Plate 17. Female terminalia of Hybodera tuberculata (A) abdomen, (B) sternfteTT-anTTr7CTTternite 4. 60

. Female terminalia of Methia pusilla- sternite (A) 7, (B) abdomlHIT^plhrrTCy' ovipositor. 61

Female terminalia of Cerambyx cerdo (montage). 62

B

Plate 20. Female terminalia (montages): (A) Agallissus

lepturoides , (B) Callidium violaceum. 63

Plate 21. Female terminalia (montages): (A) Smodicum cucujiforme (B) , Clytus arietls , (C) Sphaerion cyanipenne (segment 9 detacKedT. 64

Plate 22. Female terminalia (montages): (A) Saphanus giceus (B) Cleomenes dihammaphoroides (C) Opsimus quadrilineatus"; ' a

65

Plate 23. Female terminalia (montages): (A) Anaglyptus gibbosus , (B) Achryson surinamum , (CjC lii chroma . auricomum (A and C with apodeme 8 detacned) 66

•4; /7.;'J

V j

Plate 24. Female termmalia: (A) Holopleura marginata (B) Heterachtes ebenus (TMHionTnT) , (8) Cur ius dentatus (D) , Rhopalophora longipes 67

Plate 25. Female terminalia: (A) Oeme rigi da (montage) ' (B) Gracilia minuta, (C)~E^mic KHiIs oedipus ~ c— (apodeme 8 wanting) . ,

68

Plate 26. Female terminalia: (A) Stenopterus rufus (B) Molorchus minor (C) , Tillomorpha lineolicrera (vagina Q and uterus evaginated) . 69

Plate 27. Female terminalia of Elaphidion spinicorne . 70

Female terminalia (montages) : (A) Rosalia alpina (B) ' , Dryobius sexnotatus . APPENDIX 1 (HABITUS) Plate 29. (A) Achryson surinamum, (B) Agallissus lepturoides (C) Anaglyptus ~ gibbosus .' , (D) Anc7lQC^a carJ£naTTs"

72 73

Plate 30. (A) Callichroma auricomum , (B) Callidium vi olaceum , (C) Ce rambyx cerdo (D) ~Cleomenes dihammaphoroides . 74

(A) Clytus arietis (B) , Curius dentatus (C) Dryobius sexnotatus , (D) Elaphidion spinicorne . (A) Eumichthus oedipus (B) , Gracilia minuta,- (C) Hesperophanes sericeus (D) , Holopli^?!t marginata . 76

(A) Hzbodera . tuberculata , (B) Heterachte ebehurTTBpfioHTni; (C)" , Methia pusllIF: (D) Molorchus minor. ' 77

(A) Nathrius breyipennis , (B) Obrium cantharinum, (C) Oeme ngida , (D) Ops'imus quagrlTineituF: 78

Plate 35. (A) Purpuricenus desf ontainii , (B) Rhopalophora collkris, (C) Rosalia afpiHaT (D) SafHaH^s^eus 79

Plate 36. (A) Smodicum cucuj i forme , (B) Sphaerion " cyanipenne, (C) Stenopterus rufus (D) Tillomorphac lineolisera . — APPENDIX 2 (STERNA) Plate 37

81 Plate 38 )

APPENDIX 3 (HIERARCHICAL ARRANGEMENT HIERARCHICAL ARRANGEMENT

Division I

Tribe 1: Purpuricenini Fairmaire 1864 Subtribe 1: Purpuricenina Fairmaire 1864 Subtribe 2: Ancylocerina Lacordaire 1869

Tribe 2: Methiini Thomson 1860

Tribe 3: Obriini Mulsant 1839 Subtribe 1: Obriina Mulsant 1839 Subtribe 2: Hyboderina Linsley 1940

Division II

Tribe 1: Stenopterini Fairmaire 1864

Tribe 2 : Oemini Lacordaire 1869

Tribe 3 : Opsimini LeConte 1873

Tribe 4: Gracilliini Mulsant 1839

Tribe 5: Molorchini Mulsant 1862

Tribe 6: Holopleurini Chemsak and Linsley 1974

Tribe 7: Smodicini Lacordaire 1869

Tribe 8: Callidiini Mulsant 1839

Tribe 9: Clytini Mulsant 1839 Subtribe 1: Clytina Mulsant 1839 Subtribe 2: Tillomorphina Lacordaire 1869 Subtribe 3: Anaglyptina Lacordaire 1869

Tribe 10: Cleomenini Lacordaire 1869

Tribe 11: Rhopalophorini Blanchard 1845

Tribe 12: Callichromatini Thomson 1860

Tribe 13 : Achrysonini Lacordaire 1869

Tribe 14: Curiini LeConte 1873

Tribe 15: Ibidionini Thomson 1860

84 85

Tribe 16: Agallissini LeConte 1873

Tribe 17 : Hesperophanini Mulsant 1839 Subtribe 1: Hesperophanina Mulsant 1839 Subtribe 2 Elaphidionina Thomson 1864

Subtribe 3 Sphaerionina Lacordaire 1869 Subtribe 4 R.osaliina Fairmaire 1864

Subtribe 5 Dryobiina Linsley 1964 . .

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BIOGRAPHICAL SKETCH

Sergio Augusto Fragoso , born December 20, 1933, in Vila

Isabel, Rio de Janeiro, Brazil, holds a B.S. in Fine and

Graphic Arts from the Escola Nacional de Belas Artes, Uni- versidade do Brasil (1965) and a M.S. in entomology from

University of Florida (1977)

From 1957 to 1975, he has published nine papers in taxonomic entomology as a result of spare time activity, while working as a graphic designer.

From 1975 onward he was granted a scholarship by

Embrapa (Empresa Brasileira de Pesquisa Agropecuaria, Brazil) for study toward the M.S. and Ph.D. at the University of

Florida.

92 I certify that I have read this < study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality as a dissertation for the degree of Doctor of Philosophy

arman Adjunct Professor o tomology

I certify that I have read _ this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality as a dissertation for the degree of Doctor of Philosophy!

George E. Allen Professor of Entomology

I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality as a dissertation for the degree of Doctor of Philosophy'

C ^.^V^ ^

I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality as a dissertation for the degree of Doctor of Philosophy'

\Jcdathan Reiskind ~ Associate Professor of Zoology ,

I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy.

Reece I. Sailer Graduate Research Professor of Entomology

This dissertation was submitted to the Graduate Faculty of the College of Agriculture and to the Graduate Council, and was accepted as partial fulfillment of the requirements for the degree of Doctor of Philosophy.

December 1978

Dean

Dean, Graduate School