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Contributions

of the

American Entomological Institute

Volume 22, 1984-1986

No. 1. Michener, C.D. & R.W. Brooks. Comparative study

of the glossae of bees. July 15, 1984. 173 pages.

No. 2. Wahl, David. The nigriscapus group of Venturia

(Hymenoptera: Ichneumonidae), Aug.25, 1984. 35 pages.

No. 3. Fairchild, G.B. The Tabanidae of Panama. March 1,

1986. 138 pages.

No. 4. Wirth, W.W., A.L. Dyce & B.V. Peterson. An atlas

of wing photographs with summary of the numerical

characters of the Nearctic species of Culicoides, |

(Diptera: Ceratopogonidae). Nov.25,1985. 46 pages.

No. 5. Fairchild, G.B. & R.C. Wilkerson. A review of the

Neotropical genus Stypommisa (Diptera: Tabanidae).

July 27, 1986. 61 pages.

No. 6. Sanborne, Michael. On the correct application of

Rhimphoctona Foerster (Hymenoptera: Ichneumonidae).

July 27, 1986. 6 pages.

No. 7. Sanborne, Michael. The subgenera of Xylophylax

Kriechbaumer (Hymenoptera: Ichneumonidae) with de-

scriptions of two new Eastern Palearctic species.

July 27, 1986. 4 pages.

No. 8. Sanborne, Michael. Revision of the Nearctic species

of Xylophylax Kriechbaumer (Hymenoptera: Ichneu-

monidae). July 27, 1986. 44 pages.

No. 9. Sanborne, Michael. A revision of the Nearctic species

of Cymodusa Holmgren (Hymenoptera: Ichneumonidae).

July 27, 1986. 26 pages.

No. 10. Sanborne, Michael. A revision of the Nearctic species

of Cymodusopsis Viereck (Hymenoptera: Ichneumonidae).

July 27, 1986. 24 pages.

No.11.Sanborne, Michael. A revision of the world species of

Sinophorus Foerster (Hymenoptera: Ichneumonidae):

Supplement 1. July 27, 1986. 8 pages.

pec

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Contributions

of the

American Entomological Institute /

Volume 22, Number 1, 1984

5.5L LOR

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Ree ee ———

COMPARATIVE STUDY

of the

GLOSSAE OF BEES (APOIDEA)

Charles D. Michener and Robert W. Brooks

A COMPARATIVE STUDY OF THE GLOSSAE OF BEES (APOIDEA)

Charles D. Michener

and

Robert W. Brooks

Snow Entomological Museum and Department of Entomology

University of Kansas

ABSTRACT

Glossae of 290 species representing nearly all suprageneric taxa of

bees, and many genera within some such taxa, were examined using the

scanning electron microscope as well as cleared material under light

microscopes. Emphasis was placed on apical structures. Homologies of

various parts are postulated. Major features are an annulate, mor-

phologically anterior surface bearing annular hairs and a disannulate,

morphologically posterior surface bearing disannular hairs, most of

which are seriate hairs (in two rows) except in Colletidae and

Stenotritidae. In long-tongued bees the disannulate surface is in-

vaginated and forms the glossal canal, in the wall of which is the glos-

sal rod, which sometimes partly encloses the smaller bacular canal. In

Colletidae the disannular surface is elaborated into the glossal lobes

bearing the glossal brush; in long-tongued bees and some others it is

elaborated into the flabellum. New characters usable at the family

level are not numerous. One such is the presence of preapically bifid

or modified annular hairs in Halictidae (except most Dufoureinae); such

hairs do not occur in other families. Group characters are more

numerous at subfamily, tribal, and generic levels. Examples are the

bipartite flabella of Ctenioschelini, the transversely grooved serial

ridges of Perdita and the coarsely cobblestoned but otherwise placoid

flabella of the Eucerini. The similarity of meliponine, apine, and bom-

bine glossae, all very different from those of euglossines, does not

support the current classification of the Apidae. The tendency for

reduced glossal complexity in parasitic bees suggests that part of the

glossal attributes may be associated with nest construction and

provisioning rather than feeding.

CONTENTS

ENLPODUCTION Ser A Fat Renee earn tiee ee eae hea ee eames aie meee Ee oo 1

Acknowledgements ----------r nn nnn ncn nnn nn ne eee ne nnn ee e- 1

Materials and Methods------------------------------------------------ 1

ii Contrib. Amier. ent. Inst.) vol. 22,0. 1, 7984

MeEONOINGY aid Tenninciogy (2s e8s os sere ora reac eee ee ee Me

COmOat et Ve. ACCOUNT Soha tesa eer at eee Cane eee = 10

Lan Reba Tet rr ae Oo ae HR Pe He Siren tom eae 10

i! MSL Le SUT ACE See ter ta sag ee meee aS ee ete he ie

o. AnRUlar Neale seme eras san Coe te hte Pre eee eee eee = 15

Aone SDISONNUNALe SUGAR eS ert ar eae Pe = ht hme mm ee mt ee tei 7

So) DISanMil ale Walppes oss Sea he ot ee Ke ea eo nde eee 20

6. Distal speciatization (mon labellate)-----se+-----H ese ee 22

ET a ales pc ala Reni ae maa laa 25

Systematic Considerations------------------------------------------- 29

Pala y SLORGET mes pone tee er aa Aa eee re ee ema ae 29

Pate: COG Caeat ss estas see ee: STR R eres eee tema wee 31

Parl yer eae te ne te eee ee er ee ae Sheers Tore he Sie Se

SU eG Heil ae Sons een ee ee se ie eee Pree ot 32

FEN Ter Ite oe iene een ie eg a me ie > ein canine cule 32

SUDteniy DeeOUreilige= tants Aart os tees eee see tee e rier 34

be we 208 81 2) oleae ae i ae Sora bee aaa aia Tah an nan fala 34

PaO ae BOO eae ae oiee ie Re eee rin es eH Re a cain ee ie gm ele 39

Sepa RU heed aerate besogtc us Creatas. vias legals tare taal achalasia at ca eee a ln a ent 39

Fay I enOirecrriaae. “asp rer en near an RE ee meth atin met se 4]

FRA SE Oke rae me a ot lt eer a a me or oS et ge 41

heey Piel veleersch ates eter ease eee ae So hdeene eee ine neo 43

RN OTST ET aa oe ove Sam ee a a ao ae a a al 43

POM OD ORC degh ara Re rans oer Te ee ees ee ees Se ee 44

SUSTaniny AN LRe Der nage essa on rae ors seer mg etc ne snes #4

eee pater ts Res tae ro Se eet <= ters te a

he Sere e hits Clete hr eet eee treet oe mae een 47

Age en re res oh ae te ah eo eo 47

Pee ee ce ee yma Sie ir Sit mininnhy ete siate 4]

Rae 8 CCN Ot a ile sal lise a oiiee aod! oan ne theta oe ala 90

TIDE ACP ies re Tet et hee mee oi ee ee a

BR reek Tot | edi RG ae ae aa oo Nena eu eae cea alata 31

PRET Ee rs em re ene ae a en ero else ie moe 53

TVIOS Cant tae ree sar hor deen ahem adi thee 93

Subfamily: Nomad Qae ++ +ea-r Hs renin te ree ene ee creme ne heen es ete 34

BUOTAMITY AVIOCO GINS eG tS n oar nee er nino ae ee a ee 36

Tbe: CERI se ee haere nninitnin eine cio ein pai 36

Tribe Xylocopini-------- errr rrr ccc rrr e rrr rtrc rrr cnn 58

Michener and Brooks: Glossae of Bees iii

Family Aploge ts == nat sa ta ae oe hai aie reer e ene Oc ee a aes 28

Subfamily Bompinaet =e tes oan eae ee nee nee eee ena a eae ee wn em 38

Tribe Euglossini------------ 9 rrr rer n rrr rrr 58

Tribe Bombini---------- oon nnn nr rn enn eee 61

Subfamilies Meliponinae and Apinae------------------------------ 61

DISCUSSION orn rn ee en nn nn en eee eee 63

|) nee 70

Literature Cited----------------------------------- 2-2-2 oe =e 73

Michener and Brooks: Glossae of Bees 1

INTRODUCTION

Mouthparts of bees have been extensively used in developing the.

current classification of these insects. Indeed some of the best

familial characters are in the mouthparts, inconveniently hidden from

easy examination in most dried specimens. The mouthparts, however, con-

tain many interesting characters as yet unstudied. The investigation

here reported concerns the glossa, the characteristics of which, both

as seen by the light microscope and by the scanning electron microscope

(SEM), are valuable for indicating relationships. There are also in-

teresting convergences, especially among parasitic forms, that suggest

the important roles that the glossa must perform.

ACKNOWLEDGEMENTS

We wish to thank Dr. Ronald J. McGinley for making the SEM

photographs of glossae of colletid bees and, for comparative purposes,

sphecoid wasps and other Hymenoptera. Brian H. Smith made some of the

SEM photographs of other bees for us. (Most were made by Brooks.) All

the SEM work was done at the Electron Microscope Facility of the

University of Kansas with the help of Lorraine Hammer. For expert typing

and editorial assistance we thank Joetta Weaver.

The study was possible thanks to grant no. DEB 77-23035 from the

National Science Foundation.

MATERIALS AND METHODS

The proboscides have been examined for 290 species representing all

families and subfamilies and nearly all tribes of the Apoidea. Those of

various other Hymenoptera, especially sphecoid wasps, have also been

studied. After separation from the head, for study with light

microscopy, each proboscis was placed in a solution (about 10%) of

sodium hydroxide (usually over night at room temperature) until all

muscle tissue was dissolved or removable. It was then transferred to

water acidulated with acetic acid for at least 10 minutes, and thence

to glycerine in which the parts were permanently preserved. While still

in water the proboscis was maximally extended by holding the prementum

and gently pulling the glossa with a needle or fine forceps. Examina-

tion was with a binocular dissecting meeree rene and, for finer details,

a compound microscope.

Proboscides of 1/6 species were examined and photographed with a

Philips 501 SEM. The dry glossae were placed in warm water containing

a little detergent (Triton-xX) and cleaned in a three ounce ul-

trasonicator for about three minutes. They were then transferred to

xylene, sonicated again, air dried, and attached with casein (Elmer's)

glue to the stud for microscopy. Coating was with 200 A of gold-

palladium and scanning with 15 KV and 200 spot size. SEM photographs

of colletids and sphecoid wasps made with the same instrument by Dr. R.

J. McGinley (see McGinley, 1980) were also available and a few are

reproduced here.

2 Contrib. “Amer. Ent: Imst., vol. 22;-no.. i, 1984

Cross-sectional views were obtained by cutting the cleared glossa

(in glycerine), often two or three times, near the middle with a pair

of fine scissors and examining the resultant short sections end on, with

transmitted light, using dissecting and compound microscopes. This

method is adequate for seeing the general shape, extent of the annulate

surface, and shape of the glossal rod if present, but in long-tongued

bees, especially small species, it is often difficult to determine the

extent of the invaginated disannulate surface which is dotted in the

diagrams.

The species studied are listed after the account of each taxon,

with the notations C for cleared preparation and SEM for scanning elec-~

tron micrograph. For practical reasons not all preparations and not all

species were examined for every character listed in the descriptions.

The SEM photographs were primarily of the apices of the glossae. Cer-

tain structures are better studied with SEM photographs than with

cleared preparations while for other structures the reverse is the case.

Good understanding of the structures can often be achieved only by using

both methods.

MORPHOLOGY AND TERMINOLOGY

The terms used are based on those of Snodgrass (1956), McGinley

(1980), and Winston (1979). New terms introduced here are marked with

asterisks.

The proboscis is regarded as projecting downward from the head.

Thus the anterior surface of the glossa is the surface that would be

called dorsal if the proboscis were considered to project forward, and

the posterior surface would be called ventral in that case.

Unlike most insects, there is only one "tongue" in the Hymenoptera.

It is believed to be a fusion product of the two glossae of most insects

(Snodgrass, 1935, 1956; Michener, 1944), and it is called the glossa.

In most Hymenoptera other than bees it is a rather short, blunt, trun-

cate, or commonly bilobed structure. I!t is at least superficially

similarly truncate or bilobed in most members of the bee family

Colletidae, but is pointed and often elongate and slender in most other

bees. The parts of the glossa are labelled in Figures 1-10.

The anterior surface of the glossa bears transverse rows of hairs

whose bases spread and commonly join laterally to form dark ridges.

Because in long-tongued bees these ridges continue nearly all the way

around the glossa, and are thus ringlike, they are termed annuli. The

hairs arising from the annuli are annular hairs.* These hairs are not

setae; they do not arise from sockets but are broadly attached to the

annuli (Figs. 31A; 32E, H; 95E; 99A). They may be greatly enlarged

spicules, similar to but larger than those found on various sclerites

and especially membranes.

While the morphological anterior surface is largely or wholly an-

nulate, in many bees there are modifications. For example, this surface

may be expanded and the posterior surface invaginated so that the word

anterior is not descriptive. For this reason the area covered with an-

nuli is called the annulate surface.* It is homologous to the anterior

surface of wasps, except that at the base of the anterior surface of the

Michener and Brooks: Glossae of Bees

,BASIGLOSSAL SCLERITE

~. PREAPICAL FRINGE

*. GLOSSAL BRUSH” 2

Oe Mine Ee apres ANNULATE SURFACE

DISANNULATE SURFACE ------~., Cy

MARGINAL HAIRS

------ BASIGLOSSAL SCLERITE

[em --->*BASIGLOSSAL SENSILLA SERIATE LINE -----

MARGINAL LINE -----

ANNULATE SURFACE------- :

any ANNULATE SURFACE SURFACE

DISANNULATE SURFACE ----- :

ANNULAT RFAC

a raid E SERIATE HAIRS ------

7 ‘

DISANNULATE SURFACE’ 6 ‘SERIATE HAIR

Figures 1-6. Diagrams of colletid and halictid glossae, with

structures labelled. 1, 2 Anterior and posterior surfaces of glossa

of a colletid. 3. Cross-section of same, anterior surface above. 4,

5. Anterior and posterior surfaces of glossa of a halictid, with struc-

tures labelled. 6. Cross-section of same, anterior surface above.

4 Contrib. Amer. Ent. lhst., vole 22> ne. 1, 1964

_-BASIGLOSSAL SCLERITE

=====BASIGLOSSAL SENSILLA

~--- PREANNULAR AREA ,ROD

DISANNULATE SURFACE GLOSSAL GROOVE --------

ANNULATE SURFACE

BACULAR PLATE

uo? SETAE FLABELLUM --------

~~~. FLABELLUM ae | a

10 SERIATE HAIR

MARGINAL LINE

MARGINAL HAIRS

Figures 7-10. Diagrams of the glossa of a long-tongued bee, with

structures labelled. 7, 8. ‘Anterior’ and posterior surfaces. 9.

Cross-section of same, anterior surface above. 10. Inner surface of

a portion of the glossal canal and adjacent edge of annulate surface,

flattened out.

glossa is an area without annuli called the preannular area, basal to

which is the Bbasigiossal sclerite. “In short-tongued bees, ‘an irregular

line of sensilla, the basiglossal sensilla, lies approximately along the

line between the basiglossal sclerite and the preannular area. If the

preannular area is somewhat sclerotized and joined to the basiglossal

sclerite, as in Pseudaugochloropsis, the basiglossal sensilla seem to

be in the middle of the basiglossal sclerite. In Ctenoplectra, various

colletids, and others this line is broadly broken medially and in long-

tongued bees as well as Diphaglossinae, the basiglossal sensilla are in

two clusters, one on each side, on the basiglossal sclerite.

Michener and Brooks: Glossae of Bees 5

The morphological apex of the glossa is at the distal margin of the

annulate surface, i.e., where the anterior and posterior surfaces meet

distally. This statement is based on wasps and other Hymenoptera in

which the annulate surface reaches the glossal apex. In Colletidae,

however, the morphological apex is marked by a distinct transverse

preapical fringe (Figs. 1, 11D), and is surpassed by the commonly

bilobed extension of the posterior surface, called the glossal lobes,

from which the glossal brush arises. The flabellum, when present in

other families of bees, is also largely or wholly beyond the annulate

surface and may be similar in derivation to the glossal lobes of col-

letids.

‘The morphological posterior surface of the glossa is bare,

uniformly hairy, or with hairs in various patterns, but it lacks annuli.

It is therefore called the disannulate surface.* The hairs on this sur-

face, like the annular hairs, are not setae and do not arise from

sockets (Figs. Sil; 32C, D, G; 381; 52F; 59C; etc.): they are called the

disannular hairs.* These hairs are exposed in most short-tongued bees

but in long-tongued bees are mostly minute and largely hidden inside of

the glossal canal which is formed by invagination of the disannulate

surface. :

In long-tongued bees the lateral margins of the annulate surface,

instead of being on the lateral margins of the glossa, are drawn toward

the median posterior line of the glossa, forming between them the

glossal groove* (Fig. 8). The disannulate surface, in this case, is in-

vaginated to form the glossal canal (Fig. 9), which extends for most of

the length of the glossa but is open for its full length along the glos-

sal groove. The glossal canal has also been called the salivary canal

(Snodgrass, 1956).

In many bees the hairs along the lines (called marginal lines*)

where the annulate and disannulate surfaces meet appear to be par-

ticularly dense. The marginal lines may be vague because in some bees

the annuli fade away instead of ending sharply. The hairs arising near

the marginal lines are called marginal hairs.* In short-tongued bees;

they are usually prominent only near the base of the glossa. The name

“marginal' seems particularly appropriate in long-tongued bees because

they margin and tend to fill the glossal groove. In some long-tongued

bees the annuli do not reach the glossal groove. This is probably

because of reduction of the annuli in a zone on either side of the

groove. The hairs along the lips of the groove are here termed marginal

hairs, even though in this case they are not at the margin of the ac-

tually annulate surface.

The marginal hairs are not all alike. Some of them are annular

hairs, often smaller but sometimes larger than other annular hairs.

Marginal hairs may be especially dense because in some long-tongued bees

the ends of the annuli bend basad so that the annuli are closer together

in the marginal area than elsewhere. In long-tongued bees the hairs may

appear dense when the glossa is viewed from the posterior surface

because the margins of the annulate surface are often inflexed into the

glossal groove (Figs. 63, 65, 76, etc.), so that the line of sight in

this area is tangential to the surface or nearly so. Finally, disan-

nular hairs are often involved. When they are similar in form and size

to nearby annular hairs, and when the hairs are dense, it may be almost

6 Contrib. Ameri: Ent. Inst.; vole<22, nox: +; 1964

Figure 11. Hairs of anterior surfaces of sphecoid and colletid

glossae (photographs by R. McGinley). Scale line = .01 mm, applies to

all four photographs. A, Annular hairs of Philanthus ventrilabris. B,

Annular hairs (near Daas. of glossa) of Policana a ana albopilosa. C, Annular

hairs of Hylaeus episcopalis, preapical fringe (F) at lower margin. D,

Hylaeus episcopalis; top to bottom: annular hairs, preapical fringe

(F), and glossal lobe.

impossible to decide where the often weakened ends of the annuli are,

especially if the margins of the annulate surface are inflexed. Thus

the term marginal hairs may be only one of convenience, perhaps per-

taining to dissimilar things, but its use is nonetheless practical.

In many bees the disannulate surface bears two longitudinal rows of

hairs, running more or less from the base of the glossa, where the rows

are well separated, to the distal region, where the rows are close

Michener and Brooks: Glossae of Bees "4

together or fused, sometimes diverging again just before the apex of the

glossa. These are the seriate hairs.* In many bees these hairs are

short and simple, but in some short-tongued bees they are enormous (the

largest hairs on the glossa) and branched (Figs. 16; 26; 31E, |; 32B,

etc.). In some bees which lack or nearly lack marginal hairs, the

seriate hairs are very close to the margins of the annulate surface

(Figs. 16, 43). In long-tongued bees the seriate hairs are minute

(sometimes absent) and close to the glossal rod or on ridges (seriate

lines) of the glossal rod (Figs. 9, 10, 62, etc.). Some bees in which

the seriate hairs are limited, for example to the apex of the glossa,

or which lack seriate hairs, nonetheless have a pair of lines or ridges

to which the seriate hairs would be attached if they were present. Such

lines or ridges, whether haired or not, are termed the seriate lines.*

In short-tongued bees, if there are only two longitudinal lines on the

posterior surface of the glossa, it is sometimes not easy to tell

whether they are the seriate lines or the marginal lines. If one can

follow such lines, commonly toward the apex of the glossa, to where each

bears a row of often branched, seriate hairs, they are the seriate

lines. If one can follow them, commonly toward the glossal base, to

where each supports an area (rather than a row) of marginal hairs, then

they are marginal lines. Of course the annuli should end along the mar-

ginal lines. Forms exist, like Pseudopanurgus aethiops (Fig. 45) and

especially some Andrena (Fig. 43), in which the space between the mar-

ginal and seriate lines is only a narrow strip. And in some Andrena,

except near the base of the glossa, the ends of the annuli appear to

reach or nearly reach the seriate lines so that the seriate and marginal

lines are fused.

The integument of the disannulate surface of short-tongued bees is

frequently delicate and membranous (indicated by dotted lines in cross-

sectional diagrams) but often the seriate hairs are on weak longitudinal

ridges. This is especially true when the integument of the zone between

the two rows of seriate hairs is slightly sclerotized. There are all

degrees of such sclerotization, from that just sufficient to maintain

the weak concavity of the surface between the rows of seriate hairs to

that sufficiently strong that if one tears the glossal integument, this

zone can be stripped from the rest of the glossa intact and often car-

rying the seriate hairs along its margins. When noticeable, this

sclerotized zone, margined by the seriate hairs, is called the glossal

thickening.*

In long-tongued bees the rows of seriate hairs are close together

and the glossal thickening is usually intensified, forming the glossal

rod (Figs. 9, 10, 62-91). This rod is often narrow, so that the seriate

hairs are separated from the rod by unsclerotized integument (Figs. 70,

74, 81). Sometimes (e.g., in Megachilidae, Fig. 65) sclerotization

along the ridges from which the seriate hairs arise forms a secondary

rod on each side of the median rod proper, the seriate hairs arising

from the secondary rods. Sometimes (e.g., in Apis, Fig. 89) the rod en-

compasses the whole space between the rows of seriate hairs, which

therefore arise from lateral seriate ridges on the glossal rod itself.

These ridges may be high and enclose a channel, the bacular canal,*

which like the rod is on the anterior surface of the glossal canal, and

like the latter is incompletely closed along its posterior surface

8 Contrib. Amer. Ent. Inst:, vol: 22); no..1, 1984

Soe

—

—_——

————

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—_——=

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————

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Wilh

Michener and Brooks: Glossae of Bees 9

Figure 21. Glossa and paraglossae of Meroglossa torrida, male, an-

terior view (photograph by R. McGinley).

(Figs. 76, 88, 99). The commonly converging seriate hairs partly close

the bacular canal, as the marginal hairs partly close the glossal canal.

In most Apidae the sclerotization (or at least darkening) of the rod is

strongest laterally so that the rod appears double (Figs. 86-91).

Figures 12-20. Diagrams of colletid glossae. 12, 13. Anterior

and posterior surfaces, Amphylaeus morosus, male. 14, 15. Same,

female. 16. Posterior surface, Meroglossa torrida, male. 17. Cross-

section of same. 18, 19. Anterior and posterior surfaces, Palaeorhiza

parallela, female. 20. Posterior surface of same, male. The pair of

transverse lines across a glossa (Figs. 16 and 20) represent two annuli,

the lines being broken where they are on anterior surface. Annular

hairs and other annuli are omitted. For Figure 17 the conventions are

the same as for sectional diagrams, Figures 22-30.

Gi nen ec NE NNER

10 Contrib. Amer. Ent. inst., vol.-22,. now), T9604

At least in many dead long-tongued bees the median part of the

glossal rod becomes evaginated, together with the rest of the disan-

nulate surface, through the glossal groove (Figs. 65, 69, 72, 75, etc.).

Whether this can ever happen in life is uncertain, but in living Bombus

pennsylvanicus we were unable to bend, stretch, or compress the glossa

in such a way as to cause evagination of the rod. It is clear that if

the rod were evaginated the glossal canal would no longer exist and

could not carry saliva.

At its distal end the glossal rod commonly expands to form the

posterior surface of the flabellum. The expansion tapers basally toward

its union with the rod, usually extending considerably farther basally

than the base of flabellum, and is termed the bacular plate* (Figs. 8,

92E, 93B, 96C, 99D). If one tears the glossal rod from the glossa, the

posterior surface of the flabellum often comes off with the rod.

An interesting parallel to the glossal rod is found in some

colletids. Thus in, Caupoticana,. Colletes, etc., there. is a rod-like or

strip-like thickening in the anterior surface on each side, arising from

the basiglossal sclerite, and extending lateroapically, supporting the

sides of the broad, bilobed glossa.

In many bees there is a distal specialization* at the apex of the

glossa. In the Colletidae this takes the form of the glossal lobes and

brush. In many other short-tongued and a few long-tongued bees it is

merely a small apical area, sometimes with the surface slightly dif-

ferentiated, and bearing the distal seriate hairs; usually it lacks an-

nuli. In these respects it resembles the glossal lobes of Colletidae,

with which it is probably homologous. In a few short-tongued and most

long-tongued bees, there is a constriction at the base of the distal

specialization; the latter may be expanded and elaborately shaped. When

such a constriction is present the area beyond it is termed the

flabellum (labellum of some authors, e.g., Snodgrass, 1956, but not

homologous to the labellum of Diptera). In the Ctenioschelini it is

divided into a basal preflabellum* and a distal postflabellum*

(Fig. 96E-G).

In addition to the annular and disannular hairs there are in many

bees a few short and often blunt glossal setae (Figs. 52B, 60G, 93E,

etc.), mostly near the apex of the glossa. They arise from distinct,

commonly protuberant bases or sockets. It seems likely that they are

chemoreceptors but in the absence of data on their functions, they are

merely called setae. Fortunately there are no other types of setae on

the glossa.

COMPARATIVE ACCOUNT

The paragraphs in this section are numbered, to facilitate cross

referencing in other sections of this paper.

1. General form: 1.1 Major differences in the form of apoid

glossae have been known for a long time (Reaumur, 1742) and have long

been used in classifying bees (Kirby, 1802; Fabricius, 1804). Much of

the diversity was illustrated by Saunders (1890) and by various other

authors. Since the general form is well known for each group of bees,

only a brief summary is included here.

Michener and Brooks: Glossae of Bees 1a

1.2 In most bees the glossa ends in an acute apex (often modified,

e.g., with a flabellum, and not acutely pointed when examined at higher

magnifications). An acute apex is characteristic of all long-tongued

bees (families Fideliidae, Megachilidae, Anthophoridae, and Apidae).

(For an account of the mouthparts of these families see, Winston, 1979.)

Most short-tongued bees (Andrenidae, Oxaeidae, Halictidae, Melittidae,

Ctenoplectridae) also have an acute glossal apex and the glossa is

strongly flattened, several to many times as broad as thick. The same

shape is characteristic of males of the Australasian colletid genera

Meroglossa and Palaeorhiza (Figs. 16, 19, 10, 21). Males of Amphylaeus,

another Australian colletid genus, have a small median apical glossal

projection (Fig. 12, 13) and are thus intermediate between the

Meroglossa-Palaeorhiza condition and that of other colletids in which

there is no sexual dimorphism in glossal form. Most Colletidae, i.e.,

all females and males of all genera except those mentioned above, have

a broad glossa, the distal margin of which is gently convex, truncate,

or usually bilobed (Figs. 1, 2, 14, 15, 18, 20). Because of the

similarity of this shape to glossae of other Hymenoptera, the colletids

have been widely regarded as the most primitive bees, but the truncate

or bilobed glossa of colletids may be a reversion; see the Discussion

section. The distal margin of the annulate surface is marked by the

preapical fringe in most colletids; beyond this fringe are the glossal

lobes which produce the broad and often emarginate glossal apex.

Sometimes (Diphaglossa) the two glossal lobes are greatly extended so

that the glossa is as long as in some long-tongued bees. The apex of

the glossa in the Stenotritidae is rounded (Fig. 39).

1.3 The long-tongued bee families have a slender, somewhat flat-

tened glossa, often less than twice as wide as thick (although oc-

casionally much flattened, four or more times as wide as thick, as in

Euglossini), usually parallel-sided in the basal region and tapering

apically. The glossa is more slender and longer than in most short-

tongued bees, but in spite of the terminology there are "short-tongued"

bees with a glossa as long as in some long-tongued forms. The crucial

difference between long- and short-tongued bees is not in the glossal

length itself but in the labial palpi. In the long-tongued families the

first two segments of these palpi are long, broad, sheath-like, and

along with the galeae ensheath the glossa, forming a sucking apparatus

with the protrusable and retractable glossa in the center. The third

and fourth segments of the labial palpi are much shorter, usually more

or less cylindrical and directed laterally. In the short-tongued

families all four segments of the labial palpi are usually similar,

cylindrical, and not sheath-like. Occasionally in some relatively long-

tongued Panurginae (Andrenidae), the first segment of the labial palpus

is much longer than the others and flattened; in the genus Melitturga

the first two segments are long and flattened. Other differences

between the long-tongued and short-tongued bee families are enumerated

by Michener and Greenberg (1980). There is a general difference in

glossal shape between short- and long-tongued bees that is partly in-

dependent of glossal length. In the former (except most colletids) the

glossa is broad near the base, then narrows rapidly, but the apical part

is usually drawn out, attenuate. In nearly all long-tongued bees the

glossa narrows gradually toward the apex, but there is no particularly

Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

28 30

S — SE Ww

Michener and Brooks: Glossae of Bees 13

attenuate apical region.

1.4 Long acute glossae probably arose from short acute glossae

several times. Among "short-tongued bees" long glossae are found in

such andrenid genera as Callonychium, Plesiopanurgus, Melitturga, and

various species of other genera such as Perdita. Halictids like

Thrinchostoma (especially the subgenus Diagonozus) and Rophites also

have long glossae. Finally, the glossa is moderately long in the

Ctenoplectridae, the sister group of the long-tongued bees proper

(Michener and Greenberg, 1980).

2. Annulate surface: 2.1 The annulate surface is entirely or al-

most entirely anterior in Colletidae (except males of Meroglossa and

Palaeorhiza) and Stenotritidae. At the sides of the glossa in these

forms, annuli only curve part way toward the posterior surface and do

not encroach on that surface (Figs 3, 40). In all other bees the an-

nulate surface curves around the lateral parts of the glossa onto the

posterior surface.

2.2 In the short-tongued bees this curvature results in the disan

nulate surface being narrowed, but at least largely exposed (Figs. 16,

17, 19, 22-30, 33-37, 39-46). In some preparations in the preapical

part of the glossa, the lateral margins of the annulate surface meet or

nearly meet along the posterior median line, virtually hiding the disan-

nular surface in this region. This is especially so in dry material

such as SEM preparations. For example in Fig. 51F, G (Calliopsis and

Nomadopsis) a deep median groove is evident just basal to the flabellum.

This ts formed by the margins of the annulate surface drawing together.

In a cleared preparation (Figs. 47, 48) the margins of the annulate sur-

face are well apart, exposing the disannulate surface. In other species

the seriate lines are seemingly fused for much of the length of the

glossa so that the disannulate surface is limited to a basal and

sometimes a small apical portion (Figs. 22, 31H, 32A). In certain

Panurginae such as Psaenythia bergi, on the other hand, the margins of

the annulate surface, at least in some preparations, curl into the glos-

sa (except near the base and apex) so that the disannulate surface is

depressed. In some preparations of Melitturga the lateral margins of

the annulate surface in the middle part of the glossa approach one

Figures 22-30. Diagrams of halictid glossae, posterior surfaces

and cross-sections. 22, 23. .Halictus quadricinctus. 24, 25. Corynura

chilensis. 26, 27. Pseudaugochloropsis graminea. 28, 29. Nomia

(Acunomia) melanderi. 30. Nomia (Epinomia) nevadensis. The pair of

transverse lines across a glossa represent two annuli, the lines being

broken where on anterior surface. Annular hairs and other annuli are

omitted. For long glossae a median portion is omitted; the structures

continued from basal to apical parts are joined by broken lines. For

posterior views seriate hairs (or other disannular hairs) are shown only

on the left, marginal hairs (when well developed) and setae (when

present) are shown on the right. For sectional diagrams, annular sur-

faces are shown as solid lines, disannular as dotted lines or as a weak

solid line (in other figures) if there is a glossal thickening. Disan-

nular hairs are shown in sectional diagrams but annular hairs are

omitted.

~ .2.—$—$$

14 Contrib. Amer <-Ent. Insts; aol..22, nou) 1964

another so closely that there is a glossal groove, partly closed over

by the lateral annular hairs; in this case the disannulate surface in

this part of the glossa is invaginated as in long-tongued bees (Fig.

50).

2.3 In most long-tongued bees the annulate surface almost sur-

rounds the glossa (Figs. 62-91). In many cases the margins of the an-

nulate surface are deeply invaginated into the glossal groove which is

therefore closed over by ordinary annular hairs, in addition to marginal

hairs. 9 This is true of Lithurge, Megachilinae,’ ‘some °“Centridini

(Epicharis), Anthophorini, some Exomalopsini (Tapinotaspis), Bombus, and

others (Figs. 63, 65). In other cases, even within some of the same

tribes or subfamilies, the lateral margins of the annulate surface are

external, and the glossal groove is margined or closed over only by mar-

ginal hairs. This is the condition found in Fideliidae, Xylocopa,

Nomadinae (Thalestria, Triepeolus), some Centridini (Centris), some

Eucerini (Thygater), Melitomini (Melitoma), some Exomalopsini

(Ancyloscelis), Apis, Euglossini, and Meliponinae. (The degree of in-

vagination of the lateral margins of the annulate surface varies in dif-

ferent parts of the glossa, and in different preparations of the same

species, so that it is not always a useful character.) In various

parasitic long-tongued bees (Melectini, many Nomadinae, Nasutapis and

Eucondylops in the Xylocopinae), the disannulate surface is not or

scarcely invaginated, so that the lateral margins of the annulate sur-

face are widely separated from one another as is usual in short-tongued

bees (Figs. 79-81).

2.4 The annuli are generally moderately fine, numerous, and

reasonably uniform. In most Colletidae, however, they are excessively

fine and close together, either throughout or in a particular zone.

Thus in genera such as Colletes, Scrapter, Euryglossa, Ptiloglossa,

Hyleoides, and. Leloproctus as. well ‘as’ females .of | Meroglossa,

Palaeorhiza, and Amphylaeus, the annuli are fine and close, sometimes,

as in Hyleoides (see McGinley, 1980), divided into distinctly different

basal and apical zones. In contrast, Chilicola plebeia and males of

Meroglossa, Palaeorhiza, and Amphylaeus, have moderately coarse annuli

like those of other short-tongued bees. In Ptiloglossa and Diphaglossa

the annuli are limited to a narrow zone basal to the preapical fringe.

In contrast to most colletids, the annulation is especially coarse

(i.e., annuli far apart) in the very long glossae of Euglossini (in the

distal part of the glossa the distance between annuli may be over half

the glossal width), in the reduced glossa of the parasitic allodapine

Eucondylops, and in the Stenotritidae.

2.5 As noted in the section on Morphology and Terminology, the ex-

tremities of the annuli (= lateral margins of the annulate surface,

along the marginal lines) vary among various groups. They may be abrupt

or fade away. Often they are difficult to see, being hidden by the mar-

ginal hairs. Sometimes they reach the seriate lines so that the seriate

and marginal lines are the same except near the base of the glossa.

Sometimes in the middle part of the glossa the ends of the annulli

closely approach the seriate hairs so that there is only a narrow zone

between the seriate and marginal lines. In most bees, however, the an-

nuli end far short of the seriate lines.

Michener and Brooks: Glossae of Bees 7S

2.6 The annulate surface in Melectini as well as in Caenoprosopis

in the Nomadinae appears to be minutely papillate (Figs. 96B, 98J).

3. Annular hairs: 3.1 In many bees the hairs arising from the

annuli are slender, tapering from the bases to the sharply pointed

apices. Deviations from that type are described below. There is a

widespread tendency for the hairs to be broadened, usually medially or

postmedially, although acutely pointed. This lanceolate shape, espe-

cially when only moderately broad, sometimes escapes notice on superfi-

cial examination because hairs are most easily examined along the sides

of the glossa, where they are seen against a background light. Because

the hairs there are seen on edge, their expansions are invisible. Lan-

ceolate annular hairs occur at least near the apex of the glossa in some

Dufoureinae (Fig. 38A-G) (but not other Halictidae), bees like Panurgus,

Perdita, Protandrena bancrofti, Panurginus polytrichus, Callonychium

flaviventre (but not other Andrenidae), and in most long-tongued bees.

The hairs all appear to be tapering, not lanceolate, however, in Apis,

Bombus, Melipona, Lestrimelitta, and some Trigona (in the Apidae) as

well as in the parasitic allodapines Nasutapis and Eucondylops. In the

Fideliidae and most Melectini and Ctenioschelini the hairs are also

tapering except that some of the distal ones are slightly broadened

(Figs. 60B; 96A-D). Most long-tongued bees have the lanceolate hairs

limited to the distal part of the glossa, sometimes the area near the

flabellum (e.g., in Exomalopsis solani which has only a few lanceolate

hairs), and sometimes the distal fourth, half, or more. Variation often

occurs even in related bees. Thus in Xenoglossa lanceolate hairs extend

nearly to the base of the glossa while in Svastra they are limited to

the distal part. The lanceolate hairs gradually grade into tapering

hairs basally, so that as one looks along the glossa from base to apex

it is impossible to say at what point they become lanceolate.

3.2 Many Nomadinae (Holcopasites, Nomada, Isepeolus, etc.) have

broad, thin, imbricated scales forming a lamella along each side of the

glossal groove (Cif present) or disannulate surface (Figs. 79, 80); fre-

quently these lamellae overlap and hide the disannulate surface (or

glossal groove if present) (Figs. 97G, |, K; 98A, E, 1). These lamellae

are often continuous, but sometimes seem to be made up of separate

scales which may be modified marginal hairs; it is not clear whether or

not they arise from the ends of the annuli. The apices of the scales

can be simple and acute, blunt, or ending in two to several spines.

Ancyloscelis in the Exomalopsini has similar lamellae (Fig. 92E). The

Euglossini are unusual in that annular hairs are absent or greatly

reduced on much of the anterior surface of the glossa, except near the

base and apex. However, around the lateral margins of the glossa there

are tapering hairs, becoming very large and lanceolate on the posterior

parts of the annuli, sometimes with a slender hair before each annulus

fades away toward the glossal groove. The large lanceolate hairs are

directed somewhat mesally and cover over a channel formed by the gently

concave posterior surface of the glossa (Figs. 86, 87).

3.3. Although males of the colletid genera Meroglossa and

Palaeorhiza have tapering annular hairs, other colletids (including

males of Amphylaeus) have glossal hairs which are minute, dense, short,

flattened, truncated or emarginate, and usually capitate, with the

apices commonly bent forward (Fig. 11). ‘Such hairs are also found in

Contrib. Amer. Ent. last., vol. 22; no. 1, 1984

irs of

ines represen

idae).

ict

(Hali

ices of glossae and (C and D) annular ha

Scale

and Nom

Figure

ictin

Hal

Michener and Brooks: Glossae of Bees 17

the Stenotritidae, but not in other bees; similar hairs, however, are

widespread in the Hymenoptera, from Tenthredinidae (Tenthredo) and

Ichneumonidae (Cryptus) to sphecoid wasps such as Sphex, Philanthus, and

Oxybelus. Such hairs may therefore be plesiomorphic.

3.4 The remaining group of bees which commonly has annular hairs

that deviate from the simple, tapering type is the Halictidae. The

simple, tapering type is found in the Dufoureinae, with lanceolate hairs

preapically in some (Fig. 33E, G). In Dufourea, however, at least some

of the hairs in the middle part of the glossa are divergently bifid

preapically. In Nomia (Nomiinae) at least some hairs on the preapical

part of the glossa are divergently bifid preapically (Figs 22H)... 1n

Halictinae most of the annular hairs are preapically bifid, or they are

expanded preapically with two divergent prongs and often one to several

points between (Figs. 31A, C, D, H, |; 32A, E).

4. Disannulate surface: 4.1 This is the converse of the annulate

surface. In bees in which the annulate surface does not wrap around the

glossa, the disannulate surface is fully exposed, while in those whose

annulate surface wraps around, the disannulate surface is exposed only

at the base and apex. In short-tongued bees the disannulate surface is

usually broadly exposed. Commonly it is gently concave (strongly so in

some forms like Rophites, Melitturga, and sometimes in Dufourea and

Nomia) forming a longitudinal depression at least in the basal part of

the glossa. In the distal part of the glossa, where it is attenuate,

the disannulate surface usually occupies the entire posterior surface.

4.2 In most long-tongued bees, except for its extreme base and

apex, the disannulate surface is invaginated and forms the glossal canal

(Figs. 9, 63, etc.). The surface seems to be incompletely invaginated

in Fideliidae (Fig. 62) and Tapinotaspis (Fig. 68) but this may be an

artifact of preparation. In many common long-tongued bees the canal and

glossal rod can be evaginated in the middle part of the glossa, as noted

earlier and illustrated in sectional diagrams (Figs. 65, 69, 72, 75, 83,

85, 91). In the parasitic allodapine Nasutapis, the invagination is in-

complete, the glossal groove being quite wide in the basal part of the

glossa, and in Eucondylops there is no invagination, the disannular sur-

face being a mere flat membrane as in many short-tongued bees. The same

is true of Nomada and some other Nomadinae as well as Melectini

(Figs. 73, 78-81), although the glossa is quite elongate.

4.3. In short-tongued bees the disannulate surface is un-

sclerotized, transparent in cleared specimens. However, in certain

species the zone between the seriate lines, i.e., the glossal

thickening, is somewhat tougher than the cuticle lateral to it and can

be torn out more or less intact from the posterior surface of the glossa

in specimens prepared in KOH. The seriate hairs remain attached to such

a fragment, at least on one margin. Bees having such a recognizable

glossal thickening include Perdita (at least P. acapulcona), Panurgus,

H = seriate hair, A = annular hair. A, Lasioglossum (Hemihalictus)

lustrans, anterior view. B, Lasioglossum (Sphecodogastra) texanum,

posterior view. C, Same species. D, Sphecodes gibbus. E, Halictus

ligatus, posterolateral view. F, Sphecodes gibbus, posterior view. G,

Thrinchostoma productum, anterior view. H, |, Pseudagapostemon

divaricatus, posterior views. J, Nomioides divisa, posterior view.

18 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

Rophites, and Systropha. In Rophites the structure looks much like a

true glossal rod (see below). Other forms, such as Andrena and

Melitturga, have a weak glossal thickening which is narrow and not as-

sociated with seriate hairs. Calliopsis and some allied panurgines have

distinct glossal thickenings but no seriate hairs. (Glossal thickenings

of short-tongued bees, when present, are shown as solid rather than

dotted lines in the sectional diagrams.) Some andrenids, halictids, and

melittids have the seriate lines approximate or fused distally or for

most of the length of the glossa (Fig. 31H), bearing long and commonly

branched seriate hairs with broad, flattened bases on at least the

united part of the seriate lines (Figs. 31E, !; 32B, G). In such cases

the united or approximate seriate lines are also somewhat tough and can

be torn out from the posterior integument of the glossa.

4.4 A glossal thickening such as is found in Panurgus is shallowly

trough-shaped, with the seriate hairs along the lateral margins

(Fig. 51D). This is similar to the glossal rod of many long-tongued

bees except that the latter is sclerotized, much more slender, and ex-

tends for the full length of the glossa. The fideliids (Cat least

Neofidelia) have such a glossal rod, not or scarcely invaginated into

the glossal groove in our preparations, with the entire space between

the seriate lines sclerotized and part of the rod (Fig. 62). The same

(but with deep invagination) is true of Lithurge, Ancyloscelis, the

Eucerini, Anthophorini, Centris, etc., except that in most of these the

amount of sclerotization (darkening) is less, so that there is a median

rod occupying the bottom of the trough, lateral to which are the un-

sclerotized seriate lines (ridges). Tapinotaspis is similar but with

these ridges quite high (Fig. 68); they are still higher and partially

enclose a distinct bacular canal in Epicharis and especially in

Xylocopa, Melipona, and Apis (Figs. 77, 82, 88, 89). In many

Megachilinae the sclerotized rod is limited to the bottom of the trough,

with the hair-bearing summits of the seriate lines (ridges) being also

but less strongly sclerotized, so that there seem to be three rods, the

median one stronger than the others (Fig. 65).

4.5 In the Apidae the rod tends to look double because of

thickening on each side of the mid line (Figs. 86-91). In Bombus and

Euplusia the median strip is pale, with a dark band on either side; in

Euglossa there is a weak median thickening in addition to lateral ones.

In the Meliponinae and Apinae the seriate ridges nearly close over the

bacular canal, which is further enclosed by the inward-directed, minute

seriate hairs. In the same taxa as well as Xylocopa the seriate ridges

are sclerotized or at least prismatic, and become part of the rod, which

therefore nearly encloses the bacular canal.

Figure 32. Apices (and preapical part, C) of glossae of

Augochlorini and Nomiinae (Halictidae). Symbols as for Figure 31. A,

B, Paroxystoglossa transversa, posterior view, three preapical serial

hairs broken off. C, Pseudaugochloropsis graminea, posterior view of

preapical part. D, Same species, posterolateral view. by

Augochloropsis metallica, anterior view. F, Same, posterior view, a

seriate hair broken off. G, Nomia nevadensis, posterolateral view. H,

|, Nomia nortoni, anterior and posterior views.

ene

Glossae of Bees

Michener and Brooks

20 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

4.6 Finally, there are a few long-tongued bees, all parasitic

Anthophoridae, that have lost the rod or retain at most a feeble glossal

thickening. These are Eucondylops, Nasutapis, and various Nomadinae

such as Nomada and Biastes, and Melectini such as Xeromelecta.

5. Disannulate hairs: (This section does not describe the disan-

nulate hairs on flabella or flabellum-like apical areas.) 5.1 In some

bees the disannulate surface is hairy. For example, in Stenotritidae

(Fig. 39) it is covered with long hairs, longer than the annular hairs,

except for a bare basal triangular area. In Colletidae there are many

intermediates from hairy to bare (Figs. 2, 13, 15, 20), except that the

glossal lobes always bear long hairs forming the glossal brush. Thus

in Amphylaeus the surface is densely hairy, with a weak longitudinal

band of denser hair in the male; denser hairs that might be called mar-

ginal hairs are present lateroapically. In Chilicola the surface is

also largely hairy with a slender, triangular, longitudinal median, bare

area, widest at the base of the glossa; midapically the hairs grade into

large, branched hairs like those of the glossal brush. In Euryglossa

the surface is covered with short, rather sparse hairs, sparsest before

the glossal brush. In most other colletids examined the surface is half

half to largely bare, often with small hairs on the basal third of the

glossa and extending apically as a broad median zone, but with a

longitudinal median bare strip (for example in the female of

Meroglossa), widened basally, or with a basal bare triangle, sometimes

depressed (e.g., in Colletes).

5.2 The lateral margins of the bare median triangle or strip found

in various colletids may represent the seriate lines of other families.

No such lines are present in stenotritids. The seriate lines of other

short-tongued bees are widely separated basally but converge toward the

apex of the glossa; sometimes they meet forming a basal triangular area

suggesting that often seen in colletids. Just before the apex, the

seriate lines often diverge, spreading to the lateral margins of any

flabellum-like structure that may be present.

5.3 In Eucondylops (a parasitic allodapine anthophorid whose glos-

sa suggests that of a short-tongued bee) there seem to be no seriate

lines or hairs and the disannular surface is bare. The same is true of

some andrenids such as Calliopsis (Fig. 47).

5.4 In melittids, oxaeids (Fig. 41), Nomia nevadensis, and some

andrenids, the seriate lines appear to be absent except on the distal

part of the glossa, where they bear enlarged seriate hairs. In some

such cases, the seriate lines may be fused with the marginal lines, as

is shown for example in Macropis by the basal separation of these lines,

leaving a bare laterobasal area between the two (Fig. 55)... Even in

Melitturga, in which the disannulate surface of the long glossa is

strongly invaginated and largely hidden in the glossal groove, no

seriate lines are recognizable, perhaps because they are united with the

marginal lines (Fig. 49). |

5.5 In many other panurgines (Panurgus, Heterosarus, Perdita,

Psaenythia, Metapsaenythia, Pterosarus, Protandrena, Pseudopanurgus )

(Fig. 45), and in Dufourea (Fig. 33), Ctenoplectra (Fig. 5/7), and

others, the seriate lines are recognizable and separate from the mar-

ginal lines for much of the length of the glossa, each frequently

bearing a row or narrow band of hairs. In Macropis these lines are

Michener and Brooks: Glossae of Bees 21

easily recognizable for their full lengths, even where they are united

with the marginal lines, by the series of long hairs on them (Fig. 55).

5.6 In Nomia melanderi the seriate lines are distinct for almost

the full length of the glossa (Fig. 28), but hairless except for the

large hairs at the glossal apex. The male of Meroglossa is similar but

for their full lengths the seriate lines give rise to large, branched

hairs (Fig. 16). In the Halictinae as well as in most Dufoureinae, the

seriate lines are likewise distinct, and each bears a continuous fringe

of long, usually simple hairs which grade, in Halictinae, into long,

branched, seriate hairs near the glossal apex. An exception to this

statement is found in Thrinchostoma in which there is a gap between

simple seriate hairs near the glossal base and branched ones apically.

9.7 In the Halictinae, in Melittidae (except Dasypoda and

Macropis), and in Nomia nevadensis, Heterosarus (some preparations),

Psaenythia, and Metapsaenythia the seriate lines come very close

together in the distal, attenuate part of the glossa and commonly fuse

indistinguishably, sometimes to the extent that the bases of the large,

branched, seriate hairs form a single row with alternate hairs directed

toward opposite sides. Such fusion or near fusion is often limited to

the apical half or much less of the glossa. However, in Halictus

(Fig. 22), Lasioglossum (s.l.), Homalictus, and Agapostemon the seriate

lines are approximate except for the basal sixth to tenth of the glossa

where they diverge toward the glossal base. The same is true of

Augochlora, Pseudaugochloropsis (Fig. 26), Megalopta, Paroxystoglossa

(Fig. 32A), etc.; moreover, in these genera the large, branched, seriate

hairs like those of the glossal apex extend nearly to its base.

29.8 When the seriate lines are close together, especially near the

apex of the glossa, the interval between them is commonly represented

by a groove, which usually terminates abruptly at about the base of any

distal glossal specialization. This groove is shown in many of the SEM

illustrations of glossal apices (e.g., Figs. 38D, F; 51F, J, etc.).

5.9 In the genus Perdita the seriate lines form strong, rounded

ridges running the length of the glossa. These ridges are divided into

short segments by numerous transverse grooves (Figs. 52B, C), so that

they sometimes appear as a series of scales (hair bases?), and in most

species examined (not P. texana) each segment is drawn out laterally in-

to a point or short hair. In P. texana the basal parts of the seriate

ridges are not divided by grooves and bear numerous short seriate hairs.

In the same species, robust, pointed setae are scattered along the

lateral parts of the seriate ridges in the distal part of the glossa,

where these setae are larger and more conspicuous than the annular hairs

which are restricted to the anterior surface. In P. acapulcona, also,

there appear to be large setae occupying the positions of marginal hairs

near the apex of the glossa, but no such setae are visible in the SEM

photographs of smaller species; a few, short, probable setae are in this

position in P. albipennis.

5.10 In long-tongued bees the seriate hairs, when present, are

minute, simple, and directed distomesally. (in short-tongued bees they

are directed laterally or distolaterally.) They are absent, as are the

seriate lines, in Eucondylops, and the hairs are absent or nearly so in

various Nomadinae such as Holcopasites, Caenoprosopis, Biastes, Nomada,

and Isepeolus. They are present in Triepeolus and Thalestria, although

22 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

absent from the distal part of the glossa in the former. Sometimes, as

in Centris, the seriate hairs seem to be fused into a longitudinal

sheet, sometimes broken (naturally or in preparation?) into separate

hairs. The same is true to greater or lesser degrees in various other

Anthophoridae. In Anthophora and Amegilla the seriate hairs are large

for long-tongued bees, and each arises from a laterally expanded hair

base suggesting similar hair bases in Perdita. In Perdita each scale-

like hair base gives rise to an extremely short, laterally directed hair

while in the Anthophorini each gives rise to a much larger, mesally

directed hair.

5.11 Although in short-tongued bees other than colletids and

stenotritids it is rare to find hairs on the disannulate surface other

than the seriate hairs and some marginal hairs, in long-tongued bees

minute hairs in the space between the marginal and seriate lines are

present in xXylocopa, Triepeolus, Centris, Anthophorini, Eucerini,

Exomalopsini, Bombinae, some Meliponinae, and Apinae. These hairs

project into the glossal canal but are usually rather sparse and trivial

in length compared to the diameter of the canal. Even when present,

such hairs may not be found all along the glossa. Thus they do not ap-

pear in some sectional diagrams (Figs. 62-81) of species that possess

them.

6. Distal specialization (nonflabellate): 6.1 In a rather small

number of bees there is no striking distal specialization. As in most

sphecoid wasps, the annulate and disannulate surfaces meet at the apex

with no special features. This is approximately the situation in

Stenotritidae (Fig. 39), in which the apex of the short glossa is

rounded, with some rather long hairs, a continuation of those of the

disannulate surface, a few of them deeply forked. The parasitic an-

thophorid bees Nasutapis (Fig. 1001) and Eucondylops have slender,

pointed glossae, without any indications of specialized apical areas or

of special types of hairs in that region. The panurgine genus Panurgus

also shows no appreciable distal specialization (Figs. 51D, E). The

same is true for Perdita texana (but not of other species of Perdita ex-

amined), except that it has at least one blunt seta arising near the

glossal apex, in addition to tapering setae scattered along the distal

part of the glossa on the posterior surface.

6.2 Most bees have some sort of apical glossal structure, or at

least hairs noticeably different from those of more proximal parts of

the glossa. This statement is debatable for the groups discussed below

in paragraph 6.4 and the first part of 6.5; in these forms the distal

specialization is not defined but involves only progressive changes in

the hairs and forward curvature of the slender glossal apex. In

Colletidae (except males of Meroglossa and Palaeorhiza) the distal

specialization consists of the glossal lobes (Fig. 1) which form the

broad and commonly bilobed apex of the glossa. An exception is in male

Amphylaeus, in which there is only one rather narrow, median apical lobe

(Figs. 12, 13). The glossal lobes bear on both anterior and posterior

surfaces disannular hairs forming the glossal brush. These hairs are

commonly branched but are merely bifid (some of them simple) in female

Meroglossa and appear simple in Euryglossinae.

6.3 In Colletes, Hylaeus, and male Amphylaeus (but not in other

colletids) the hairs of the glossal brush are arranged to form weak an-

Michener and Brooks: Glossae of Bees 23

Figures 33-37. Diagrams of dufoureine glossae, posterior surfaces

and cross-sections (Halictidae). 33, 34. Dufourea marginata. 35.

Micralictoides altadenae. 36, 37. Systropha curvicornis. Explanation

as for Figures 22-30.

nuli, each broken into short sections in Hylaeus. Annuli on the glossal

lobes could indicate that we are incorrect in attributing these lobes

to the disannular surface. However, they are distinctly different from

annuli on the annulate surface and the hairs are similar to those on the

seriate lines of many other short-tongued bees. We believe that the an-

nuli on the lobes are secondary features, not homologous to the normal

annuli. This possibility is supported by the observation that hairs are

arranged in transverse rows (annuli) on the paraglossae of some bees

(e.g., Meganomia); thus transverse rows of hairs and therefore annuli

could arise independently on different areas.

6.4 In most of the remaining short-tongued bees the apex of the

glossa is pointed, often curved anteriorly, usually attenuate, slightly

convex posteriorly, with annuli recognizable nearly to the apex on the

anterior surface but absent from the posterior surface, which is largely

occupied by the distal parts of the seriate lines, often unrecognizably

fused. The hairs on the posterior surface are therefore the distal

seriate hairs. As noted previously, there is often a groove

(representing the space between the seriate lines) that ends basal to

or within the most slender, terminal part of the glossa. The seriate

hairs, diverging laterally and usually curving apicad and branched, are

large, usually robust and flattened at their bases. When the seriate

24 Contrib. Amer. Ent. Inst., vol. 22;-no..1, 1964

lines are united, and when the hair bases are large, the hairs almost

form a single row, those extending to one side alternating with those

extending toward the other.

6.5 Glossal apices of the general type described in 6.4 are found

in males of Meroglossa and Palaeorhiza (but distal seriate hairs not as

large as those nearer the glossal base), all Halictinae, Nomiinae,

Melittidae, Oxaeidae, Andreninae, and the following genera of

Panurginae: Xenopanurgus, Heterosarus, Metapsaenythia, Pterosarus,

Protandrena, Cephalurgus, Liphanthus, and Epimethia. In some other

panurgines, however, such as Panurginus, Psaenythia, Protandrena

bancrofti (slightly, Fig. 51B), Pseudopanurgus aethiops (slightly,

Fig. 45), and P. herbsti, the seriate lines and hair rows diverge

markedly preapically, partially enclosing a convex apical area on the

posterior side of the glossa suggestive of a flabellum. The same is

true of the halictid, Nomioides (Fig. 31J). The glossa as a whole is

not constricted at the base of this area; hence we do not call such an

area a flabellum. However, the disannulate surface may be narrow or a

mere groove, which does widen to form the apical area on the posterior

surface of the glossa, enhancing the flabellum-like aspect. In

Panurginus and perhaps some others listed above, specimens cleared in

KOH show the disannulate surface moderately broad preapically, not nar-

rowed basal to the apical area, while SEM shows the seriate lines ap-

proximate preapically, diverging to enclose the apical area. Dasypoda

has a gradual distal glossal broadening, with a broad "cobblestoned"

posterior surface (Fig. 59A, B), which we arbitrarily do not call a

flabellum because of its length and the numerous annuli on its anterior

surface.

6.6 Certain other panurgine bees such as Nomadopsis scutellaris

and Hypomacrotera are similar but the posterior, bare, convex, apical

area is broader, the seriate hairs being extensions of the margins of

the area. These hairs are part of the same series as the other seriate

hairs when the latter are present. Although some are branched, at least

at the extreme apex of the glossa, they are smaller than the large

seriate hairs of halictids and others. Spinoliella and Arhysosage are

similar but the marginal hairs of the apical area are reduced to mere

small, unbranched, thorns.

6.7 Dufourea, Rophites, and Systropha also have a flat or convex

apical area on the posterior surface of the glossa, this region being

not or only slightly wider than the glossa basal to it. It is formed

by termination of the groove between the seriate lines and reduction in

the size of all the apical hairs of the glossa so that hair bases do not

interrupt the flat or convex surface (Figs. 38A-G). The seriate hairs

are small and arise from the margins and apex of the apical area, or in

Rophites they are absent or nearly so; these hairs are branched in

Dufourea, simple in Systropha. Possibly some of the simple hairs on the

anterior surface of the glossal apex are seriate hairs but probably they

are annulate hairs, differing from those of the preapical part of the

glossa by being slender and simple.

6.8 The panurgine genera Camptopoeum and Plesiopanurgus have a

similar glossal apex, with reduced lateral hairs, which are, however,

perhaps setae. The apical area is broadly connected to the space

between the seriate lines, which is thus broad for the whole length of

Michener and Brooks: Glossae of Bees 25

the glossa. The seriate hairs are small and simple. —

6.9 Some long-tongued bees also lack flabella. In the nomadine

genera Epeolus, Triepeolus, and Isepeolus the extreme apex of the glossa

is simply irregular and bare, with a few setae projecting apically from

both upper and lower surfaces (Fig. 97A, F). Doeringiella and espe-

cially Thalestria are similar, but with the area flattened with a

distinct edge and thus more like a flabellum (Fig. 97B, E); Epeolus ap-

proaches Doeringiella in this feature.

6.10 In the anthophorine genera Habropoda and Emphoropsis there is

no flabellum (Fig. 95E, F). The annulate part of the glossa abruptly

narrows; annular hairs which are broadly lanceolate or truncate arise

to its very apex on the anterior surface. The posterior surface of the

abruptly narrowing part of the glossa, which is beyond the apex of the

glossal groove, is flat and bare. Setae have not been recognized in

this region, but the broad annular hairs cover so much of the surface

that SEM photographs might not show them. Setae are, however, scattered

among the annuli on the posterolateral parts of the glossa (considerably

lateral to the groove) from near the apex of the groove to the middle

of the glossa in Emphoropsis at least.

7. Flabellum: 7.1 The flabellum is a type of distal specializa-

tion recognized by its basal constriction, separating it from the rest

of the glossa. All intergradations exist between flabellate and non-

flabellate glossae, and sometimes one must make an arbitrary decision

as to whether to call a given distal specialization a flabellum or not.

The posterior surface of the flabellum is convex, hairless (hairs,

setae, or processes sometimes present along distal and lateral margins),

and its margins converge basally to connect with the rest of the seriate

lines, so that the posterior flabellar surface is continuous with the

cuticle between the seriate lines. In long-tongued bees this means that

the posterior flabellar cuticle connects with the glossal rod; if one

tears out the glossal rod, the posterior but not the anterior surface

of the flabellum often comes out attached to the rod. From this

finding, and from the observation that the hairs of the concave anterior

surface of the flabellum are disannular hairs, similar to those of the

disannulate surface of the rest of the glossa, it appears that the

flabellum (or only part of it in Meliturgula and Ctenoplectra) is an

elaboration of the disannulate surface. It is thus comparable to the

glossal lobes of Colletidae. Annuli are absent on the flabellum, with

the exception of a few forms such as Thalestria, Meliturgula, and

Ctenoplectra. In Thalestria the flabellum-like, posterior, bare, glossal

area is similar to that of its close epeoline relatives, but is

elongated apparently by the basad extension of the bare area. This

results in annuli being on what appears to be the anterior surface of

the "flabellum." In a few other bees without or with only a weakly

defined flabellum, the same thing may have occurred. This may include

Meliturgula and Ctenoplectra, which have annuli on the anterior surface

of an elongate flabellum.

7 hee Some panurgine genera are. similar to Panurginus,

Pseudopanurgus, etc., but with the apical, bare area clearly broader

than the glossa at the base of that area, and thus forming, by defini-

tion, a flabellum margined by long processes. These processes or hairs,

however, are simple and tapering, sometimes united in pairs (or threes)

26 Contrib. Amero Ent: Insts) vol..22, nos 1, 1984

at their bases, so that they could be interpreted as branched. The con-

cave, anterior side of the flabellum has numerous small, simple, disan-

nular hairs. Examples of such glossae (Fig. 47; 51F-J) are found in

Liopoeum trifasciatum, Nomadopsis zebrata, and Calliopsis. In Liopoeum

hirsutulum the marginal processes are very short and rather sparse.

7.3. In Callonychium the flabellum is short, broader than long,

with only minute, short, marginal processes. However, a series of setae

arises from the concavity on the anterior surface of the flabellum,

among a few much smaller, disannular hairs. The row of well spaced

setae continues basally on the posterior surface, lateral to the nar-

rowing base of the bare, posterior, flabellar surface (Fig. 52D, E).

Perdita, except for P. texana which lacks a flabellum, is similar to

Callonychium but the margins of the flabellum are simple (Fig. 52A-C).

7.4 Melitturga has a peculiar flabellum with a narrow base, short

hairs on the anterior surface and thick, blunt hairs (or tapering pegs)

on the posterior surface, some of the largest, lateral pegs being bifid

(Fig. 49). These structures are not setae; perhaps they are modified

disannular hairs.

7.5 Another unusual glossal apex is that of the panurgine

Meliturgula. It is only slightly broadened, thus doubtfully a flabellum,

and is elongate with five or six annuli recognizable on its anterior

surface. There are no disannular hairs on the distal area and its mar-

gins are lobed as in some species of Ctenoplectra.

7.6 The Ctenoplectridae (Ctenoplectra) have a flabellum shaped

like that of Meliturgula, with annuli on the anterior surface, and with

lobed margins, the lobes sometimes acute and possibly derived from

disannular hairs (Fig. 59F, 1). On the concave anterior side of the

glossal apex, near the margins, there are several distinct setae.

7.7 >In many long-tongued bees the flabellum is of a type here

called placoid, i.e., it is smooth and convex on the posterior surface,

with the distal and lateral margins entire (Fig. 92). The lateral mar-

gins continue as converging lines on the posterior surface of the glossa

and appear to be continuous with the seriate lines along the glossal

rod. The anterior surface bears a few to numerous small, disannular

hairs, usually so short as to be invisible in posterior view. Similar

small hairs are often continued around the sides of the base of the

flabellum and are continuous with the disannular hairs of the main part

of the glossa. A series of setae (sometimes only two or four) crosses

the glossa just basal to the flabellum on the anterior surface and

sometimes extends along each side onto the posterior glossal surface and

basally for a short distance lateral to the distal end of the glossal

groove. These setae are often long enough that their apices radiate

conspicuously beyond the flabellum as seen in posterior view (Fig. 981).

Placoid flabella occur in all Fideliidae, Megachilidae, Exomalopsini,

Melitomini, and in the allodapine genera Allodape and Braunsapis

(Ceratinini). The panurgine genera Perdita (except texana) and

Callonychium have similar flabella (the glossal groove is absent).

7.8 A minor modification of the placoid type of flabellum occurs

with reduction of the distal part of the flabellum (or perhaps it is

distad migration of the transverse row of setae). The flabellum looks

typically placoid as seen from the posterior side, including the

presence of a few setae radiating beyond the flagellar margin. From the

Michener and Brooks: Glossae of Bees 27

anterior side, however, it can be seen that the flabellum extends only

a short distance beyond the setal bases, so that the small, simple,

disannular hairs arising from the anterior surface of the flabellum |

beyond the setae are restricted to a narrow preapical zone. (Sometimes,

as in Nomada, some of these hairs, as well as the setae, extend beyond

the flabellar apex.) Such flabella are found in Nomada (Nomadini)

(Fig. 98C) and, Manuelia (Fig. 99F) (Ceratinini) and others. Macrogalea

(Fig. 99J) is similar but the rather elongate flabellum lacks the small

disannular hairs on its anterior surface.

7.9 A further step in the same direction, in which the disannular

hairs on the anterior surface of the flabellum are absent or nearly so

and the setae arise just beneath the apex of the flabellum, occurs in

the following Nomadinae: Caenoprosopis, Holcopasites (Fig. 98F),

Morgania (Fig. 98D), Biastes, Ammobatoides, Neolarra (Fig. 98K, L)

(bases of two of the lateral setae cause indentations in the converging

basolateral margins of the posterior surface of the flabellum),

Epeoloides (Fig. 97L), and Leiopodus (Fig. 97H).

7.10 Pseudodichroa is similar to the "pasitine' bees listed in the

above paragraph except that a few setae appear to have moved mesad onto

the basal part of the posterior surface of the flabellum, so that the

main part of the flabellum is surrounded by a ring of setae (Fig. 98H).

Osiris has the setae rather scattered instead of in a neat row, on the

apical part of a thickened flabellum that lacks a thin apical edge and

has some transverse apical wrinkles (Fig. 971, J, M).

| 7.11 Certain other Nomadinae (Epeolus, Triepeolus, Doeringiella,

and Isepeolus) do not have a flabellum but their distal specialization

is apparently a result of further flabellar reduction and migration of

some of the setae onto its posterior surface (Fig. 97A-F). Thalestria

is similar but a bare, flabellum-like area extends basad on the

posterior surface; on the corresponding anterior surface there are

numerous annuli and annular hairs. |

7.12 A modification of the placoid type of flabellum is found in

the Bombini, Meliponinae, and Apinae (Apis) (Figs. 100F-H; 101). In

these forms the flabellum is rather short and broad, its lateral margins

converge strongly basad, together forming a convex or nearly transverse

base; apically the flabellum extends far beyond the setae which form a

row across its base on the anterior surface; only in some Trigona do the

setae extend beyond the lateral (not distal) margins of the flabellum.

The small, anterior, disannular hairs of the flabellum are numerous and

some of them arise at the flabellar margin and extend beyond it; these

hairs are mostly simple but the marginal ones are branched in Apis.

Basally the small hairs continue in a usually dense mass across the

preflabellar area of the posterior side of the glossa, especially in

Bombini (Fig. 100F) and Apis. The flabellar margin is irregularly shal-

lowly lobed rather than entire in Bombus, Lestrimelitta, and Apis.

7.13. Canephorula (Canephorulini) has a rather elongate flabellum,

the posterior surface smooth, the apex irregularly lobed (Fig. 94G-1).

Its remarkable feature is the anterior surface, on which the numerous

small disannular hairs are slender at the flabellar base but have large

bases with short, attenuate apices medially and distally, along the

lateral margins becoming coarse and peg-like; some such hairs project

as flattened pegs beyond the apical flabellar margin. A few setae are

28 Contrib Amer. Ent. inst., volec2; now VF; 1964

present sublaterally, mixed with small hairs, to a point beyond the

middle of the flabellum rather than across its base. |

7.14 Coarsely pebbled or cobblestone-like posterior surfaces of

flabella characterize the Eucerini (Fig. 94A-F). Otherwise they are

placoid (unusually elongate in Thygater). The small disannular hairs on

the anterior side are numerous, at least the distal ones mostly

branched, some arising from the very apex of the flabellum as in Apis.

As in Canephorula, setae arise among the small hairs of the anterior

surface, sublaterally, usually to a point beyond the middle of the

flabellum; there are no setae across the base of the flabellum as

described above for placoid flabella. Similarly cobblestone-like

posterior surfaces characterize flabella of the Centridini (Fig. 96H-J).

However, in this tribe the flabellum is elongate, the small hairs on the

anterior surface are simple and reduced toward the apex, and setae are

present near the base, but not on the flabellum. The same remarks apply

to Deltoptila (Anthophorini) (Fig. 95D, G), in which the distal and

lateral small hairs of the anterior surface are blunt and capitate.

7.15 Cobblestone-like posterior flabellar surfaces also occur in

Melectini (Fig. 96A-D), especially in Melecta, but in Thyreus the

"stones" are in transverse rows and in Xeromelecta the surface is only

irregularly transversely wrinkled. As in Nomada, Manuelia, etc., the

distal part of the flabellum appears to be reduced so that the setae

arise on its anterior surface near the apex; they do not, however,

radiate beyond the flabellar margin except laterally, but they are ar-

ranged across the anterior surface preapically and around the sides to

the posterior surface lateral to the apex of the glossal groove as in

most forms with placoid flabella. Small disannular hairs on the an-

terior surface, often flattened and toothed at the apices, are limited

to a narrow zone beyond the setae.

7.16 Beautifully cobblestoned posterior surfaces (curved over onto

lateral surfaces) characterize the elongate fflabella of the

Ctenioschelini (Fig. 96E-G). The most peculiar feature of this tribe

is the preapical, anterodistal projection, the postflabellum, extending

beyond the apex of the main part of the flabellum (or preflabellum), and

bearing several short setae on its posteroapical margin; a few small

setae also arise laterally among the small, simple, disannular hairs of

the anterior surface of the preflabellum.

7.17. An elongate (about as long as the flabellum), preflabellar,

nonannular zone, hairless except for setae, characterizes Xylocopa

(Fig: 99A, B), Bombini, and the Euglossini (Fig. 100). The setae in

Xylocopa form a row across the apex of this zone on the anterior side,

while in Bombini and Euglossini there are several setae scattered on

both surfaces of this zone. The flabellum is elongate in Euglossini,

a little longer than broad in Xylocopa, and its posterior surface with

a cobblestone pattern at least distally. Its anterior surface bears

numerous small, simple, disannular hairs but no setae. The apical mar-

gin is irregularly, weakly lobed in Eufriesea, simple or essentially so

in other Euglossini and in Xylocopa.

7.18 In Anthophora (including Micranthophora), Amegilla, and

Clisodon the flabellum is distinct, elevated basally above the level of

adjacent glossa, often curled anteriorly at the sides giving a falsely

elongate appearance, at least in dry specimens. Except in Clisodon and

Michener and Brooks: Glossae of Bees 29

some species of Anthophora the apex has one to several deep sinuses,

forming elongate lobes (Fig. 95A-C). The small, simple, disannular

hairs of the anterior surface are usually rather sparse. Setae are ab-

sent on or close to the flabellum, but are scattered among the annular

hairs on the posterolateral surfaces of the glossa from the level of the

apex of the glossal groove basad.

7.19 Finally, in Tetrapedia (Fig. 93A, D-F) the elongate, rather

thick flabellum is smooth on its posterior surface, with a series of

setae around its apex and with small simple hairs on the anterior sur-

face. Thus these small hairs, instead of being distal to the row of

setae as in the many other forms having such a row, are basal to the

setae. If the setal row is homologous to the usual one, then the usual

flabellum has been lost and is replaced by a presetal enlargement, but

if the small hairs are homologous to the usual ones and in the usual

position, then the setal row is a new development or has migrated

distad.

SYSTEMATIC CONSIDERATIONS

No changes in taxonomic positions are made in this paper. Such

changes should be based on consideration of all known characters, not

merely on glossal attributes. Some taxa discussed below can readily be

characterized by glossal characteristics. Others contain genera having

such diverse features that characterization of higher taxa is im-

possible, a situation that in some cases may indicate lack of close

relationship among included genera. The numbers refer to the paragraphs

of the Comparative Account, where more details are provided.

FANALY STENOTRITIDAE

(Fias. 39, 40)

Glossa short, thick, scarcely exceeding paraglossae, rounded,

without distal specialization except that some bifid hairs are inter-

mixed among simple ones. Disannulate surface as broad as annulate sur-

face, hairy (5.1) except for triangular basal, median, bare area; hairs

with tendency to be in longitudinal bands. Setae sparsely scattered

among hairs of posterior surface and laterally among annuli of anterior

surface. Annular hairs short, capitate, ending bluntly or truncate as

in Colletidae (3.3); but annuli coarse, not fine and close as in most

colletids (Figs. 30, 40).

The glossa is not as different from that of colletids as McGinley

(1980) believed. Among the longest hairs on the glossa are those at its

apex; these are disannular hairs like those of the glossal brush of col-

letids, and some are bifid, suggesting the branched hairs of the ‘col

letid glossal brush. McGinley correctly noted that the basiglossal

sclerite of stenotritids is divided medially, forming two sclerites.

However, contrary to his observations, the same is true of many col-

letids (Colletinae, Diphaglossinae and Hyleoides in the Hylaeinae), so

that this characteris mot unique to stenotritids. lt is,0in fact, a

feature of those colletids that have a broad glossa and fold it

longitudinally when it is retracted.

J) of glossae of

Scale lines represent .01 mm

part,

= thes,

. | )) ae A

(and preapical

; Amer... Ent. Inst., voli 22, no. 1, 1984

Contri

Apices

a6.

Figure

Dufoureinae (Halictidae) and Andreninae.

Michener and Brooks: Glossae of Bees 31

Species studied: Ctenocolletes smaragdinus (C); Stenotritus

SO (),

FAMILY COLLETIDAE

(Figs) 'T*3,12+21)

Glossa short, truncate, bilobed, or bifid (but see next paragraph).

Disannulate surface as broad as annulate surface because marginal lines

are lateral; disannulate surface forming pair of large, apical, glossal

lobes which bear conspicuous, branched or simple hairs forming the glos-

sal brush (5.1); largely because of these glossal lobes, apex of glossa

truncate or usually bilobed (1.2, 6.3), lobes long, attenuate, and

directed laterad in Diphaglossinae; rest of disannular surface hairy or

with substantial hairy areas (5.1). Setae forming transverse row across

anterior surface basal to preapical fringe, but hidden by annular hairs,

usually visible in cleared material from posterior view; setal row

usually broken medially, represented only by one or two setae at extreme

sides in some forms (e.g., Amphylaeus), and setae apparently absent in

Chilicola and Palaeorhiza. Diphaglossinae also with some setae on

posterior side and on basal parts of glossal lobes. Annuli, or one

series of them, usually fine and close; annular hairs usually minute,

their ends blunt or capitate (Fig. 11B-D); annular surface ending at

preapical fringe (Figs. 1, 11C, D).

Almost none of the above listed characters apply also to males of

Meroglossa and Palaeorhiza (6.5). They have a pointed glossa without

glossal lobes (Figs. 16, 19); the disannular surface is narrower than

the annular surface and the seriate hairs are large (5.6) but the disan-

nular surface is otherwise bare. In these features they resemble the

Halictidae. In males of Amphylaeus, also, the glossa is not bilobed, but

has a small, hairy, median, apical projection of presumably disannulate

origin, although it shows annuli (6.2, 6.3; Figs. 12, 13).

Because the colletid glossa is treated by McGinley (1980 and in

prep.), minimum attention is paid to it in this paper.

Species studied: Colletinae. Colletes inaequalis (C), thoracicus

(C, SEM); Leioproctus caerulescens (SEM), erythrogaster (SEM), herbsti

(C); Lonchopria similis (SEM); Parapolyglossa sp. (C); Scrapter sp. (C,

SEM). Diphaglossinae. Cadeguala occidentalis (C); Caupolicana hirsuta

(C); Diphaglossa gayi (C); Policana albopilosa (SEM); Ptiloglossa guin-

nae (C). Euryglossinae. Brachyhesma incompleta (C); Euryglossa sub-

sericea (SEM); Euryglossula chalcosoma (C); Hyphesma sp. (C).

Hylaeinae. Amphylaeus morosus (C, SEM). Hylaeus basalis (SEM),

episcopalis (SEM); Meroglossa torrida (C, SEM); Palaeorhiza parallela

(C). Xeromelissinae. Chilicola ashmeadi (C, SEM).

H = seriate hair, A = annular hair, S = seta. A, B, C, Rophites

trispinosus, anterior, posterior, and lateral views. D, E, Systropha

curvicornis, posterior and lateral views. F, Dufourea marginata,

posterior view. G, Same, apical view, apex curved anteriorly so that

view is posterior for apex proper. H, |, Andrena erythrogaster,

posterior and anterior views. J, Andrena helianthi, posterior view of

preapical part.

32 Contrib. Amer. Ent. Inst. , vol..22, no. 1, 1984

FAMILY HALICTIDAE

(Figs. 4-6, 22-32)

Glossa short to rather long, tapering to attenuate apex, without

flabellum, with weak apical enlargement in some Dufoureinae (6.7).

Disannulate surface somewhat narrower than annulate surface because mar-

ginal lines are on posterior lateral surfaces of glossa (2.2), or nar-

rowed to a median posterior line along much of glossa (Figs. 22, 31H).

Disannular surface largely bare but seriate lines usually with long hair

(5.7). Setae absent, a few among annular hairs in Rophites.

Subfamily Halictinae (tribes Augochlorini, Halictini, and

Nomioidini). Annular hairs tapering to near apices, there abruptly

broadly expanded or abruptly, divergently bifid, trifid, etc. (3.4,

Fig. 31). Seriate hairs long (shortest in Corynura), sometimes simple

except near apex of glossa (as in Sphecodes, Halictus, etc., Figs. 22,

31D), often broad-based and branched especially near apex of glossa,

such hairs sometimes continued almost to glossal base as _ in

Pseudaugochloropsis (Fig. 26) but more often replaced by simple hairs

basally; seriate lines fused distally (5.7, 5.8) and sometimes

(Pseudaugochloropsis) almost to base of glossa ei 22, 31H).

Species studied: Augochlorini. Augochlora pura (C, SEM); Augoch-

lorella striata (C), persimilis (SEM); RiGee nis psis metallica (C,

SEM); Corynura chilensis (C), chloris (C); Megalopta genalis nalis (C, SEM);

Neocorynura pubescens escens (C, SEM); Par Paroxystoglossa transversa (C, SEM);

Pseudaugochloropsis graminea (C, SEM); Temnosoma metallica (C, SEM).

Halictini. Agapostemon radiatus (SEM), texanus (C); Caenohalictus

azarae (C); Habralictus trinax (C); Halictus hesp hesperus (C), ligatus (C,

SEM), maculatus (C), quadricinctus (C);. Lasioglossum calceatum (C),

imitatum (C), leucozonium (C), lustrans (C, SEM), nigripes (C), rohweri

(SEM), sisymbrii, texana (C, SEM); Pseudagapostemon divaricatus ; (SEM),

Pseudagapostemon sp. (C); Ruizantheda mutabilis (C); Sphecodes gibbus

(C, SEM); Thrinchostoma producta (SEM), Thrinchostoma sp. (C); Zonalic-

tus sp. (SEM). Nomioidini. Nomioides divisa (SEM), minutissima (C).

Subfamily Nomiinae. There are striking differences between the two

subgenera of Nomia examined, Acunomia and Epinomia, supporting the view

expressed elsewhere (Michener, 1965) that these subgenera should

probably be placed in separate genera.

Annular hairs mostly simple, some bifid near apex of glossa

(Acunomia) or mostly rather abruptly bifid (Epinomia) (3.4, Fig. 32H,

|). Seriate ridges weak, hairless except for small, branched seriate

hairs at apex of glossa (Acunomia, Fig. 28), these ridges separate

nearly to apex of glossa; alternatively, seriate hairs large, mostly

trifid or bifid, present only near apex of glossa (Epinomia, Fig. 30),

seriate ridges fused in distal third or fourth of glossa and otherwise

absent.

Figures 39-50. Diagrams of posterior surfaces and cross-sections

of glossae of Stenotritidae, Oxaeidae, and Andrenidae. a0, a0.

Stenotritus elegans. 41, 42. Protoxaea gloriosa. 43, 44. Andrena

helianthi. 45, 46. Pseudopanurgus qurgus aethiops. 47, 48. Calliopsis

andreniformis. 49, 50. Melitturga clavicornis. Explanation as for

Figures 22-30. | co AES MME ety 2 ES ARR

Michener and Brooks: Glossae of Bees

34 Contrib. Amer. Ent. Insti; vol.:22, no..1, 1984

Species studied: Nomia melanderi (C), nevadensis (C, SEM), nortoni

(SEM).

Subfamily Dufoureinae. Annular hairs usually slender, tapering,

sometimes slender lanceolate (3.4), in Dufourea hairs of middle annuli

abruptly bifid, more distal ones expanded apically with several teeth.

Seriate hairs short and sparse, present only on basal half of glossa

(Systropha), moderately long and simple (Rophites and Micralictoides)

or long, some with a short branch (Dufourea). Seriate ridges separated

nearly to apex (Figs. 33-37). In Rophites and Systropha there is a

glossal thickening which suggests a glossal rod (4.3) and a suggestion

of such a thickening is sometimes visible in the distal part of the

glossa of Dufourea. Distal specialization as described in 6.7

(Fig. 38A-G) (absent in Micralictoides).

Although as indicated in the illustrations, there is considerable

variation in the Dufoureinae, this subfamily shows some distinctive

common features, several of them shared with the Nomiinae and with

Halictinae. Especially noteworthy are the abruptly bifid, trifid, or

expanded apices of at least some of the annular hairs in Dufourea.

Other genera of Dufoureinae are different in various ways and would not

necessarily have been placed in the Halictidae on the basis of glossal

characters.

Species studied: Dufourea marginata (C, SEM); Micralictoides al-

tadenae (C); Rophites quinquespinosus (C), trispinosus (SEM); Systropha

curvicornis (C, SEM).

FAMILY ANDRENIDAE

(Figs: 38H-J, 43+52E )

Glossa short and tapering (Andrena, Figs. 381, 43; Epimethia) to

rather long (1.5) and sometimes parallel-sided except distally, apex at-

tenuate, with or without flabellum. Disannulate surface largely bare,

narrower than annulate surface because marginal lines are on posterior

lateral surface of glossa; in a few genera marginal lines close together

on posterior surface of glossa (2.2) and in Melitturga (Fig. 49) so

close, and with the surface between them so invaginated, as to delineate

a glossal groove. Marginal line and seriate line of each side sometimes

approximate or apparently fused so that annuli reach or nearly reach

seriate lines (2.5); marginal and seriate hairs in such areas become in-

distinguishable. Flabellar presence (7.1-7.5) not necessarily a generic

character, thus distinct in most species of Perdita but scarcely recog-

nizable in P. maculigera and totally absent in P. texana; distinct in

Nomadopsis zebrata but absent (merely an attenuate apex, without basal.

constriction) in N. scutellaris. Glossal apex without setae except in

Arhysosage, Perdita, and Callonychium (7.7) and perhaps Camptopoeum and

Figure 51. Apices of glossae of Panurginae (Andrenidae). Scale

lines represent .01 mm; H = seriate hair. A, B, Protandrena bancrofti,

anterior and posterior views. C, Heterosarus illinoiensis, posterior

view. D, E, Panurgus calcaratus, posterior views. F, G, Calliopsis

andreniformis, posterior and = anterior views. HH, “Caltepsrs

coloradensis, anterior view. 1, Liopoeum trifasciatum, posterolateral

view. J, Nomadopsis australior, posterior view.

ar ae ae rages tans ee EEEEEESEEEEEEEEEEE cl

Glossae of Bees

Michener and Brooks

36 Contrib, Amen. Ent. Inst., vol; 22, no. 1, 1984

Figure 52. Apices of glossae of Panurginae (Andrenidae) and

Oxaeidae. Scale lines represent .01 mm; H = seriate hair, A = annular

hair, S = seta. A, B, Perdita zebrata zebrata, posterior and anterior

views. C, Perdita tridentata, posterior view. D, E, Callonychium

flaviventre, posterior and anterior views. F, Mesoxaea nigerrima,

posterior view.

Plesiopanurgus (6.8). Annular hairs usually slender and tapering,

rarely narrowly lanceolate on distal part of glossa (e.g., in Panurginus

polytrichus but not occidentalis, Protandrena bancrofti but not

mexicana), largely lanceolate in some species of Perdita (3.1).

Interesting features of certain groups are as follows: A group of

presumably primitive forms (6.5) have the seriate hairs of most of the

glossal length simple and slender (5.5) but those of the distal part of

the glossa coarse, bifid or sometimes with more branches, resembling

those of Halictinae. Such forms are Andrena helianthi (Fig. 43) (not

A. erythrogaster in which the seriate hairs are either simple or split

to their bases, Fig. 38H), Pseudopanurgus (Fig. 45), Xenopanurgus,

Panurginus, Protandrena (Fig. 51B), Pterosarus, Heterosarus, Psaenythia,

and Metapsaenythia. The two series of seriate hairs are well separated,

joined only near the extreme apex of the glossa, leaving an elongate,

Michener and Brooks: Glossae of Bees ay

bare, apical area, sometimes narrow, on the posterior surface of the

glossa. Similar glossal apices with large seriate hairs, but these

hairs mostly simple, are found in Cephalurgus, Liphanthus, and

Epimethia. In Panurgus the basal third of the glossa is similar to that

of Pseudopanurgus (Fig. 45), but the rest has the seriate hairs closely

associated with the marginal hairs, and the space between the seriate

lines cobblestoned, slightly strengthened (4.4), broad and concave all

the way to the apex so that the apical region is not differentiated,

with the disannular hairs at the apex minute and simple (Fig. 51D, E).

Panurgus also has two or three setae scattered among the annular hairs

laterally, on the apical part of the glossa.

In a considerable group of genera such as Calliopsis (Fig. 47),

Nomadopsis, Hypomacrotera, Spinoliella, Arhysosage, Liopoeum, and

Acamptopoeum there appear to be no disannular hairs except for small

ones preapically (Fig. 51J) and around the edges of the flabellum or its

equivalent (6.6). (It is often difficult to distinguish a finely

wrinkled disannular surface from a series of hairs. We believe,

however, that hairs are absent in all of these genera except near the

glossal apex.) The distal specialization, as in the preceding groups,

does not usually involve recognizable setae. In Hypomacrotera and

Nomadopsis scutellaris there are long, marginal hairs around the at-

tenuate apex which is bare on the posterior surface. In Spinoliella and

Arhysosage these hairs are reduced to small spicules and in Arhysosage

there are four setae on the anterior surface near the apex. In Acamp-

topoeum and Liopoeum hirsutulum a preapical constriction differentiates

an elongate flabellum which bears small spicules similar to those of

Spinoliella; setae are absent. Finally, in Calliopsis, Liopoeum

trifasciatum, and Nomadopsis zebrata, there is a well formed flabellum

with long marginal processes or hairs (Fig. 51F-J).

Melitturga also lacks seriate hairs. The glossa is elongate and

tubular, with an elongate flabellum similar to that of Acamptopoeum but

with the spicules coarse and blunt (Fig. 49), the anterior side with a

few small hairs. A few setae are among annular hairs at the sides,

basal to the flabellum.

In Callonychium and Perdita (Fig. 52A-E) there are distinct

Ppreapical setae. Except for some species of Perdita, there is a

distinct, transverse flabellum, the margin of which has small, sharp

projections or spicules in Callonychium but is smooth in Perdita. The

flabellum is placoid, similar to that of many long-tongued bees, even

to the presence across its base on the anterior surface of a series of

several setae. In Perdita (5.9) the seriate lines are broad and

rounded, each divided by grooves into a series of transverse, rounded

ridges. These ridges probably represent expanded bases of seriate hairs

for (except in P. texana) each tapers laterally into a short hair or

point. In P. texana the basal parts of the seriate ridges are not

transversely ridged, but bear scattered (ij.e., not in a row), short

seriate hairs. |

A weak glossal "rod" appears in some preparations of Calliopsis,

Panurgus, etc. This is a weak integumental thickening or darkening,

sometimes flat and sometimes a trough resulting from the meeting of

sloping planes of the integument (4.3, 4.4).

38 Contrib.: Amer. Ent. 'nst.,:vol.°227 nov 1, 1964

—=_

—

———

Sn ewe oe

—

>=;

—_—

Figures 53-58. Diagrams of posterior surfaces and cross-sections

of glossae of Melittidae and Ctenoplectridae. 53, 54. Dasypoda

Ppanzeri. 55, 56. Macropis europaea. 57, 58. Ctenoplectra fuscipes.

Explanation as for Figures 22-30. |

Species studied: Andreninae. Ancylandrena larreae (SEM); Andrena

erythrogaster (C, SEM), halli (SEM), helianthi (C, SEM), illinoiensis

(C, SEM), miserabilis (SEM), nigrae (SEM), perplexa (SEM), prunorum

(SEM); Megandrena enceliae (SEM). Panurginae. Acamptopoeum pruni (C);

Arhysosage ochracea (C); Calliopsis coloradensis (SEM), andreniformis

(C, SEM); Callonychium flaviventre (SEM); Camptopoeum frontale (SEM);

Heterosarus_ illinoiensis (SEM), neomexicanus (C); Hypomacrotera

subalpinus (C, SEM); Liopoeum trifasciatum (SEM); Melitturga clavicornis

(C, SEM); Meliturgula sp. (C); Metapsaenythia abdominalis (C);

Nomadopsis australior (SEM), scutellaris (SEM); Panurginus occidentalis

(SEM), polytrichus (SEM); Panurgus calcaratus (C, SEM); Perdita

acapulcona (C), albipennis (C), bishoppi (C), maculigera maculipennis

(C), texana (C), tridentata (SEM), zebrata zebrata (SEM); Protandrena

bancrofti (SEM), mexicana (SEM); Psaenythia bergi (C); Pseudopanurgus

aethiops (C), herbsti (SEM); Pterosarus ornatipes (SEM).

Michener and Brooks: Glossae of Bees 39

FAMILY OXAEIDAE

(Figs. 41, 42, 52F)

Glossa rather short, tapering to an attenuate apex, without flabel-

lum. Disannulate surface largely bare (Fig. 41), narrower than annulate

surface because marginal lines are on posterior lateral part of glossa.

Marginal hairs present (probably all annular hairs); seriate hairs large

and branched, present only on distal part of glossa (5.4). Median and

basal part of glossa with peg-like setae scattered among posterior ex-

tremities of annuli. Distal part of glossal surface and bases of

seriate hairs with surfaces (Fig. 52F) finely granular or "mealy." An-

nular hairs simple, tapering.

The only character that distinguishes the glossa consistently from

that of melittids, andrenids, and the few halictids with simple annular

hairs, is the presence of setae in its median and basal part. Among

other short-tongued bees, setae are present in Stenotritidae,

Colletidae, and in a few dufoureine and panurgine genera.

Species studied: Mesoxaea nigerrima (SEM); Oxaea flavescens (SEM);

Protoxaea gloriosa (C, SEM).

FAMILY MELITTIDAE

(Figs. 53-56, 59A-E, G, H)

Glossa short (Macropis, Fig. 55) to moderately elongate, tapering,

apex attenuate, without flabellum although with gradually broadened apex

in Dasypoda (6.5, Fig. 59A). Disannulate surface largely bare, narrower

than annulate surface because marginal lines are on posterior lateral

surfaces of glossa. Marginal hairs often abundant, especially basally.

Seriate hairs distinct on distal part of glossa, moderately large, at

least distal ones branched. Basad, seriate hairs long and simple, and

seriate lines diverging from marginal hairs in Macropis (Fig. 55); in

others seriate hairs absent basally (possibly indistinguishable from in-

nermost marginal hairs), or small as in Meganomia or minute as in

Dasypoda. On distal attenuate part of glossa, seriate lines and

therefore bases of seriate hairs adjacent (Fig. 59C, H) or separated so

that elongate smooth or "cobblestoned" strip is present, basal to which

seriate lines (if recognizable) are usually close together for some

distance before diverging basally. Setae absent or in Macropis scat-

tered among annular hairs on lateral and anterior surfaces. Annular

hairs simple, tapering.

There are no consistent familial characters by which the melittid

glossa differs from that of Andrenidae, etc. Dasypoda and Meganomia are

unusual in that there are two longitudinal bands of small to minute

hairs on the basal part of the disannulate surface. These hairs do not

intergrade with the large seriate hairs on the distal part of the glos-

sa. If, as seems likely, the small or minute hairs are seriate hairs,

then seriate hairs and lines are simply absent on the basal part of the

glossa except in Macropis, Dasypoda, and Meganomia.

Species studied: Meganomiinae. Meganomia gigas (C, SEM).

Melittinae. Macropis europaea (C, SEM); Melitta leporina (C), tricincta

(SEM). Dasypodinae. Capicola braunsiana (SEM); Dasypoda panzeri (C,

SEM); Haplomelitta ogilviei (SEM); Hesperapis carinata (SEM), pellucida(C).

Contrib. Amér. Ent. nist: “volioe2, mo.:+, 1964

]

ify

dae.

and

oda

as,

erolateral

Das

iew.

ior V

Meganomia

i, post

iew.

poster

ior view.

ior V

ta,

anzer

poster

ttidae and Ctenoplectr

carina

inc

Dasypoda

ta,

esperapis

view.

, Mel

iew.

Apices of glossae of Mel

A, B

C tta tri

lateral

|, Ctenoplectra albolimbata, poster

igure 52

ior V

Figure 59

Symbols as

ior...

anzeri, anter

Ctenoplectra

ior views.

albolimbata,

lews

poster

Michener and Brooks: Glossae of Bees 41

FAMILY CTENOPLECT RIDAE

(Figs, 5/, 96, SYF, FE)

Glossa moderately elongate, tapering, apical part not conspicuously

attenuate, its narrowing being continuous with that of rest of glossa;

flabellum present, longer than broad (sometimes much longer), weakly

constricted at base, annuli present on its anterior surface (7.1).

Disannulate surface considerably invaginated (Fig. 58), largely bare,

narrower than annulate surface because marginal lines are on posterior

lateral surfaces of glossa. Annuli often not reaching marginal lines;

seriate lines well separated from marginal lines, with converging short,

fine, simple seriate hairs, not enlarged toward glossal apex, not evi-

dent in flabellar area. Flabellum with a few tapering setae on lateral

margins of anterior surface, extending laterally beyond sometimes

toothed or lobed lateral flabellar margins; preapically two or four

setae forming row across flabellum on anterior surface; a seta on each

side basal to flabellum on posterior lateral surface of glossa. Annular

hairs slender, tapering, some distal hairs (marginal or annular) in some

species broadened, bifid or trifid.

Other ctenoplectrid characters in agreement with those of long-

tongued bees are listed by Michener and Greenberg (1980). The same

authors illustrate some of the variation in flabellar structure within

the genus Ctenoplectra.

Species studied: Ctenoplectra albolimbata (C, SEM), fuscipes (C),

bequaerti (C).

LONG-TONGUED BEES

(Figs. 7-10, 60-101)

Members of this group, which includes the families listed below,

share numerous characteristics of the glossa, if various parasitic forms

are ignored. The characters listed here are those of nonparasitic

forms, although comments on parasitic groups are incorporated into ac-

counts of the relevant taxa.

Glossa elongate, commonly parallel-sided basally and tapering

apically, not conspicuously attenuate, its narrowing being continuous

with that of rest of glossa (1.3); flabellum present with a few excep-

tions, weakly to strongly constricted at base, usually without annuli

on anterior surface. Disannulate surface of glossa, except at extreme

base and apex, commonly invaginated to form glossal canal (4.2) which

is sufficiently closed to form glossal groove, commonly filled by mar-

ginal hairs; seen externally, annuli usually seem to go all the way

around glossa except for glossal groove (2.3); marginal hairs abundant,

much smaller than most annular hairs, usually larger than seriate hairs;

sclerotized but flexible glossal rod (4.4) in roof of glossal canal.

(In prepared material and perhaps in life the canal is sometimes

evaginated in middie part of glossa so that rod is exposed.) Seriate

lines on or lateral to rod; seriate hairs (5.10) simple, usually minute,

mostly directed mesodistally instead of laterodistally as in short-

tongued bees, exposed and sometimes branched only at base of glossa and

on anterior side of flabellum (if such hairs are correctly identified

as seriate). Setae present, usually at base or anterior surface of

flabellum, but sometimes only lateroposteriorly on glossa among annular

42 Contrib.Amer. Ent: tast:, volhce2y, no), 1964

Figure 60. Apices of glossae of Fideliidae and Megachilidae.

Scale lines represent .0| mm; S = seta. A, Parafidelia pallidula, an-

terolateral view. B, Neofidelia profuga, posterior view. C, Fidelia

villosa, anterior view. D, Trichothurgus dubius, anterior view with

flabellum curved so that posterior surface of its apex is visible. E,

Trichothurgus wagenknechti, anterior view. F, Anthidium emarginatum,

anterior view. G, Dianthidium curvatum sayi, anterior view.

hairs basal to flabellum; a few setae among annular hairs nearly always

present. Annular hairs sometimes tapering, often lanceolate, sometimes

in part expanded and scale-like.

Michener and Brooks: Glossae of Bees 43

FAMILY FIDELIIDAE

(Figs, S0A-C, 62)

In available preparations glossal groove open, fully exposing glos-

sal rod and disannulate surface lateral to it (4.4). (lf this is

characteristic of the family, it must be a primitive feature.) Rod

V-shaped in cross-section (Fig. 62), minute seriate hairs on ridges

(seriate lines) formed by ends of "V." No other hairs on disannulate

surface, but membrane minutely papillate in some areas. Flabellum

placoid, rounded, broader than long, with numerous short unbranched

hairs on anterior surface. Setae present basal to flabellum on both an-

terior and posterior surfaces (Fig. 60B), at least laterally, not

forming complete row across base of flabellum. Annular hairs tapering

(3.1) or very narrowly lanceolate.

The glossa sheds no new light on whether or not this family should

be included within the Megachilidae; there are no glossal characters

that clearly distinguish Fideliidae from Megachilidae.

Species studied: Fidelia villosa (SEM); Neofidelia profuga (C,

SEM); Parafidelia pallidula (SEM).

FAMILY MEGACHILIDAE

(Figs. 60D-G, 61, 63-65)

Glossal rod U-shaped in cross-section (Lithurge, Hoplitis) or len-

ticular, seriate lines along summits of the U or as separate sclerotized

ridges a short distance lateral from rod, so that there appear to be

three rods (Fig. 65); seriate hairs minute, no other hairs on disan-

nulate surface. Flabellum placoid, rounded, broader than long,

posterior surface smooth, anterior surface concave, with short, un-

branched hairs (or a few branched in some Anthidiini). A few setae

scattered among annulate hairs on posterolateral surfaces of distal part

of glossa. Several setae at level of base of flabellum at sides forming

row (sometimes broken medially) across base of flabellum on anterior

side (Figs. 60F, G; 61A, C), often (e.g., Hoplitis, Megachile, and

Parevaspis) extending around onto posterior surface at base of flabellum

(Fig. 61D, E). Annular hairs tapering or lanceolate.

Species studied: Lithurginae. Lithurge apicalis (SEM), gibbosus

(C); Trichothurgus dubius (C, SEM), wagenknechti (C, SEM).

Megachilinae. Tribe Anthidiini. Allanthidium (SEM); Anthidiellum (C,

SEM); Anthidium (C, SEM); Aztecanthidium (C, SEM); Callanthidium (C,

SEM); Capanthidium (SEM); Carinanthidium (SEM); Dianthidium (C, SEM);

Euaspis (C, SEM); Heteranthidium (C); Hypanthidium (C); Immanthidium

(C); Nananthidium (C); Notanthidium (C); Odontostelis (C);

Pachyanthidium (C); Paranthidium (C); Parevaspis (C, SEM); Serapista

(SEM); Spinanthidium (C, SEM); Stelis (C, SEM). Tribe Dioxyini. Dioxys

(C, SEM). Tribe Megachilini. Anthocopa (C); Ashmeadiella (C, SEM);

Chalicodoma (C); Chelostoma (C); Chelostomopsis (C); Coelioxys (C);

Creightonella (C); Heriades (C, SEM); Hoplitis (C); Megachile (C, SEM);

Noteriades (C); Osmia (C, SEM); Proteriades (C). (To save space, only

the genera are listed for Megachilinae. )

44 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

Figure 61. Apices of glossae of Megachilidae. Scale lines

represent .01 mm; S = seta. A, Anthidiellum perplexum, anterolateral

view. B, C, Spinanthidium volkmanni, posterior and anterior views. D,

Parevaspis carbonaria, posterior view. E—, Megachile melanophaea,

posterior view. F, Ashmeadiella bigeloviae, anterior view.

FAMILY ANTHOPHORIDAE

(Figs. 66-85, 92-99, 1001)

This large family, like some of the included subfamilies, is

diverse in glossal characteristics. Aside from the characteristics of

long-tongued bees listed above, there are no unifying attributes of the

glossae, and there exist parasitic anthophorids which do not agree with

the features listed for long-tongued bees.

Subfamily Anthophorinae. Tribe Exomalopsini. Glossal rod rather

flat or lenticular in cross-section (Fig. 67, 68); seriate lines near

rod, forming distinct and sometimes high ridges; seriate hairs numerous

and minute. Ends of annuli not reaching margins of glossal groove, some

Figures 62-91. Cross-sections through median parts of glossae of

long-tongued bees. 62. Neofidelia profuga. 63. Lithurge gibbosus.

64. Hoplitis albifrons. 65. Creightonella frontalis (rod everted).

66. Melitoma taurea. 67. Ancyloscelis panamensis. 68. Tapinotaspis

caerulea. 69. Xenoglossa strenua (rod everted). 70. Same, rod not

everted. 71. Canephorula apiformis. 72. Anthophora occidentalis (rod

everted). 73. Xeromelecta californica. 74. Mesocheira bicolor. 75.

— $$

Michener and Brooks: Glossae of Bees 45

Centris poeciia. 76. Epicharis elegans. 77. Thalestria spinosa. 78.

Triepeolus verbesinae. 79. Nomada_ annulata. 80. Leiopodus

lacertinus. 81. Isepeolus viperinus. 82. Xylocopa virginica. 83.

Same (rod everted). 84. Ceratina laeta. 85. Macrogalea candida. 86.

Eulaema cingulata (bases of large, lanceolate annular hairs at sides).

87. Eufriesea violacea (notation as for 86). 88. Melipona fasciata.

89. Apis mellifera. 90. Bombus pennsylvanicus. 91. Same (rod

everted). Conventions are the same as for sectional diagrams, Figures

22-730.

46 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

Figure 92. Apices of glossae of Exomalopsini (Anthophoridae).

Scale lines represent .01 mm; S = seta. A, Exomalopsis solani, anterior

view. B, C, Exomalopsis zexmeniae, anterior and posterior views. D,

E, Ancyloscelis apiformis, anterior and posterior views. aw

Caenonomada bruneri, anterior and posterolateral views. H, Tapinotaspis

caerulea, anterior view. 1, J, Paratetrapedia lugubris, anterolateral

and anterior views.

small hairs usually scattered between ends of annuli and groove, as well

as on surface in glossal canal (5.11) between seriate lines and margins

of groove. Glossa only feebly narrowed at base of flabellum. Flabellum

placoid, posterior surface smooth, margin simple, rounded or in

Michener and Brooks: Glossae of Bees 47

Paratetrapedia subtruncate but rounded laterally; anterior surface with

short, simple hairs beyond setal row. Setae present basal to flabellum

at sides, and forming row across base of flabellum on anterior surface,

sometimes almost encircling it. Annular hairs tapering or distal ones

scarcely lanceolate in Exomalopsis (3.1, Fig. 92A-C), broadly lanceolate

in other genera studied (Fig. 92D-J).

In Ancyloscelis (Fig. 92D, E) the flabellum is unusually strongly

curved anteriorly at the apex, so that in most views the setal row seems

to be near the apex of the flabellum. The short, simple hairs on the

anterior side of flabellum are few, minute. Also in this genus, as in

many Nomadinae, there is a lamella perhaps made of fused hairs on either

side of and closing over the glossal groove (3.2).

Species studied: Ancyloscelis apiformis (SEM), panamensis (C);

Caenonomada bruneri (C, SEM); Exomalopsis solani (SEM), zexmeniae (C,

SEM); Paratetrapedia lugubris (SEM); Tapinotaspis caerulea (C, SEM);

Trigonopedia sp. (C).

Tribe Tetrapediini. Similar to Exomalopsini but seriate hairs

relatively long; no noticeable hairs on disannular surface except

seriate hairs. Flabellum (7.19) elongate, thick, not or scarcely curved

anteriorly, no setae at base or a lateral seta near base but distal end

with several setae, variable in length (Fig. 93D), forming a row across

apex of flabellum (Fig. 93E, F); anterior surface with short hairs. An-

nular hairs mostly tapering but distal ones on both surfaces or on

posterior surface only broad and commonly three or four dentate at

apices (Fig. 93D-F).

Species studied: Tetrapedia sp. (C, SEM), sp. (SEM).

Tribe Melitomini. Glossal rod flat in cross-section or evenly con-

cave on anterior surface (Fig. 66), rather broad; seriate lines on

ridges arising lateral to rod; seriate hairs small to minute; disan-

nulate surface otherwise probably without hairs. Flabellum placoid

(Fig. 93B, C), glossa scarcely constricted at base of flabellum but nar-

rower than apical part of flabellum, apex strongly curved anteriorly,

subtruncate, posterior surface smooth or in Melitoma coarsely trans-

versely wrinkled, anterior surface strongly concave with short simple

hairs in space beyond setae; setae forming transverse row across an-

terior surface of base of flabellum, others scattered at sides of apical

part of glossa. Annular hairs frequently broad lanceolate or foliose,

some or all along anterior surface in Diadasia and Melitoma foliose with

three or four apical points. Annuli, especially in Melitoma, well

separated, perhaps because of glossal elongation characteristic of this

genus.

Species studied: Diadasia afflicta (C), australis (SEM); Melitoma

euglossoides (C), taurea (SEM); Ptilothrix bombiformis (SEM).

Tribe Eucerini. Glossal rod lenticular to semicircular in cross-

section (Figs. 69, 70); seriate lines weak ridges, usually close to rod

but widely separated from it in Xenoglossa; seriate hairs small to

minute; disannulate surface lateral to seriate lines with few (Svastra)

to many (Xenoglossa) small hairs (5.11), such hairs probably absent in

Melissodes, etc. Flabellum (7.14, Fig. Q94A-F) broader than long to

elongate oval, glossa not constricted at base of flabellum but narrower

than flabellum, apex of flabellum curved anteriorly, rounded or subtrun-

cate apically, posterior surface coarsely cobblestoned; anterior surface

48 Contrib. Amer. Ent. Inst., vol. 22, no.'1, 1984

Figure 93. Apices of glossae of Tetrapediini and Melitomini

(Anthophoridae). Scale lines represent .01 mm; S = seta. A, Tetrapedia

sp., posterior view. B, Diadasia australis, posterior view. C,

Melitoma taurea, anterior view. D, E, F, Tetrapedia sp., posterior,

apical, and anterior views.

concave with short to rather long simple hairs all or mostly distal to

setae; some of these hairs at apex of flabellum usually branched,

forming transverse series of small branched hairs (Fig. 94C, D), but

these hairs simple in Gaesischia, Alloscirtetica, Xenoglossa, etc.

Setae scattered along sides of apical part of glossa; one to four

similar setae on each side of anterior surface of flabellum, more distal

ones sometimes beyond middle of flabellum; in Xenoglossa setae numerous

and forming U-shaped row around sides and apex of flabellum just basal

to apical series of hairs; in other genera setae not arranged in row

across flabellum. Annular hairs often mostly tapering or very narrowly

lanceolate, broadly lanceolate on distal part of glossa, on Xenoglossa

broadly lanceolate hairs extending nearly to base.

Species studied: Alloscirtetica tristrigata (SEM); Eucera

chrysopyga (C); Florilegus condignus (SEM); Gaesischia exul (SEM);

Martinapis luteicornis (SEM); Melissodes agilis (C), lupina (SEM);

Michener and Brooks: Glossae of Bees 49

Figure 94. Apices of glossae of Eucerini and Canephorulini

(Anthophoridae). Scale lines represent .01 mm; S = seta. A, B,

Peponapis pruinosa, posterior and anterior views. C, Svastra obliqua,

anterior view of extreme apex; does not show concave anterior surface

of flabellum. D, Melissoptila (Ptilomelissa) sp., posterior view. E,

Thygater analis, posterior view. F, Svastra obliqua, posterior view.

G, H, |, Canephorula apiformis, posterior, anterior with apex of flabel-

lum toward observer, and anterior.

Melissoptila (Ptilomelissa) sp. (SEM); Peponapis crassidentata (C),

CC CRASH a RINE RRR

pruinosa (SEM); Svastra atripes (C), obligua (SEM); Synhalonia hamata

50 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

Figure 95. Apices of glossae of Anthophorini. Scale lines

represent .01 mm; S = seta. A, B, Anthophora cockerelli, posterior and

anterior views. C, Anthophora walshii, anterior view. D, Deltoptila

n. sp., posterolateral view. E, Emphoropsis miserabilis, anterior view.

F, Emphoropsis pallida, posteroapical view. G, Deltoptila n. sp, an-

terior view.

(SEM); Thygater analis (C, SEM); Xenoglossa strenua (SEM), fulva (C).

Tribe Canephorulini. Glossal rod rhomboid in cross-section

(Fig. 71); seriate hairs abundant, covering low membranous ridges im-

mediately lateral to rod; disannulate surface lateral to seriate lines

apparently hairless. Flabellum (7.13, Fig. 94G-1I) elongate oval, glossa

constricted at base of flabellum, apex of flabellum curved anteriorly,

generally rounded but margin with several small lobes from beneath which

project several small, flattened, blunt hairs, base of flabellum arched

posteriorly; posterior surface of flabellum smooth, anterior surface

concave with numerous short hairs which have large bases but taper

abruptly to short attenuate apices, these hairs laterally and distally

Michener and Brooks: Glossae of Bees 51

are blunt and merge with blunt hairs projecting from beneath apical

flabellar lobes. Setae absent at or near base of flabellum, not forming

transverse row; about three setae on each side of anterior surface of

flabellum among small hairs, distal setae beyond middle of flabellum;

a few setae scattered among annular hairs of distal part of glossa. An-

nular hairs partly tapering but those of distal part of glossa mostly

lanceolate, the lateral ones broad.

The glossal characters reinforce the _ distinctness of the

Canephorulini from the Eucerini.

Species studied: Canephorula apiformis CC USE),

Tribe Anthophorini. Glossal rod rather weak, flat to shallowly

U-shaped in cross-section (Fig. 72); seriate lines weak ridges near rod;

seriate hairs rather long, at least in Anthophora and Amegilla arising

from broad, scale-like bases. Ends of annuli nearly reaching margins

of glossal groove which bear dense marginal hairs, some of which at apex

of glossa are branched in Anthophora occidentalis. Small scattered

hairs on disannular surface between marginal and seriate lines (5.11),

in Emphoropsis such hairs near seriate lines arising from incomplete

flat scales, one hair per scale. Setae scattered along distal half or

third of glossa, on posterior lateral surfaces, not forming row across

base of flabellum (7.18). Annular hairs lanceolate, the more distal

ones broader and leaf-like except in Deltoptila. Distal specialization

highly variable, as follows:

In Anthophora (including Micranthophora), Clisodon, Amegilla, and

their relatives, flabellum (7.18) distinct, thin, strongly constricted

and arched posteriorly at base, posterior surface smooth, apical margin

rounded (Clisodon and some Anthophora) or with one to several deep

sinuses separating apical projections (Fig 95A-C); anterior surface con-

cave, with scattered small, simple hairs of variable lengths.

In Emphoropsis and Habropoda, flabellum absent (6.10, Fig. 95E, F);

glossa with apical part (last four or five annuli) narrowed to bluntly

pointed apex, posterior surface of this region flat and bare. Annular

hairs before apical, narrowed part of glossa unusually large, broadly

lanceolate or foliose, often hiding glossal apex; annular hairs of nar-

rowed distal part of glossa smaller, especially distally, those of more

apical annuli expanded apically and truncate, several of most distal

ones arising from extreme apex of glossa.

In Deltoptila, flabellum distinct (7.14, Fig. 95D, G), elongate

oval, strongly constricted and arched posteriorly at base, posterior

surface with cobblestone pattern; apical margin rounded; anterior sur-

face gently concave with numerous small, unbranched hairs, the distal

and marginal ones expanded apically and through shortening, merging into

cobblestone pattern of posterior surface.

Species studied: Amegilla comberi (C, SEM); Anthophora cockerelli

(C, SEM), occidentalis (C), urbana (SEM), walshii (SEM); Clisodon

terminalis (SEM); Deltoptila n. sp. (SEM); Emphoropsis miserabilis

(SEM), pallida (SEM); Habropoda deiopea (SEM).

Tribe Melectini. Disannulate surface not invaginated (Fig. 73).

Glossal rod weak, flat, scarcely sclerotized. Seriate lines and hairs

absent. Ends of annuli not reaching margins of glossal groove which are

not to sparsely haired except near apex of glossa where marginal hairs

are present and in Xeromelecta lanceolate; disannular surface hairless.

. Amer. Ent. Inst.) vol. 22; no. 1, 1984

Contri

ini, and

1 mm for

iews. C,

ioschel

mm (0

ion

Cten

me)

ini,

ior and anter

D, Xeromelecta californica, posterior

ines represen

poster

Scale

ior View.

Melecta armata,

ices of glossae of Melect

idae)

i CAnthophor

seta. A, B

igure 96.

Bos S.=

Thyreus ramosus, poster

Centr

;

Michener and Brooks: Glossae of Bees 53

Flabellum (7.15, Fig. 96A-D) thin, curved anteriorly, narrowed basally,

posterior surface strongly cobblestoned (Melecta), less strongly so

(Thyreus), or merely coarsely transversely wrinkled (Xeromelecta), apex.

rounded or subtruncate; anterior surface with small simple hairs. Setae

scattered among annular hairs near sides of distal part of glossa;

similar setae forming transverse row or arc across anterior side of

flabellum rather near its apex, this row extending basally on each side

so that bases of one or more setae are visible in posterior view near

flabellar base. Glossal surface distally, minutely granular or papil-

late. Annular hairs largely tapering but slender-lanceolate on apical

part of glossa.

Species studied: Melecta armata (SEM); Thyreus ramosus (C, SEM);

Xeromelecta californica (C, SEM).

Tribe Ctenioschelini. Glossal rod lenticular in cross-section

(Fig. 74). Seriate lines distinct, well separated from margins of rod,

each bearing a row of seriate hairs. Ends of annuli sometimes not or

scarcely reaching margins of glossal groove which is margined with

simple hairs, or ends of annuli deeply invaginated into glossal groove

which is therefore closed by annular hairs. Disannular surface hairless

except for seriate hairs. Flabellum (7.16, Fig. 96E-G) consisting of

two parts, an elongate preflabellum with thin, rounded apex curved an-

teriorly, and a_esmall, shorter, rounded postflabellum arising

preapically from anterior surface of the preflabellum; bases of both

preflabellum and postflabellum narrowed; preflabellum with posterior and

lateral surfaces cobblestoned, anterior surface with numerous curved,

simple, small hairs arranged in irregular, transverse rows suggestive

of annuli but hairs much smaller than annular hairs; postflabellum cob-

blestoned on all surfaces. Setae scattered lateral to hairy zone on an-

terior side of preflabellum, the series continued apically onto

posterior surface of postflabellum where they form an arched row near

apex; a few setae among annular hairs of distal part of glossa. Annular

hairs tapering, or near apex of glossa narrowly lanceolate, more broadly

so in Mesoplia.

Species studied: Ericrocis arizonensis (SEM), lata (C); Hopliphora

funerea (SEM); Mesocheira bicolor (C, SEM); Mesoplia azurea (SEM).

Tribe Centridini. Glossal rod flat in cross-section (Fig. 75), in

Epicharis thickened laterally so that it appears to be a partly double

rod (Fig. 76); seriate lines very strong ridges rising from margins of

rod and in Epicharis curved toward one another at summits, largely en-

closing round bacular canal; seriate hairs of moderate length but mostly

fused to form a lamella (5.10) here and there broken into hairs, rising

from summit of seriate ridge. (An alternative explanation is that there

is a lamella, sometimes torn into hair-like units, in place of seriate

hairs.) Disannulate surface lateral to seriate lines with scattered very

minute hairs (5.11). Flabellum (7.14, Fig. 96H-J) elongate, flattened,

strongly constricted and arched posteriorly at base, posterior surface

with cobblestone pattern which is exaggerated at margins which are

view. E, Hopliphora funerea, posterior view. F, Mesocheira bicolor,

lateral view. G, Mesoplia azurea, anterior view; some hairs of the

basal half of the surface are broken off. H, |, Centris atripes,

posterior and anterior views. J, Epicharis rustica, anterior view.

54 Contrib.-Amer. Ent. inst), vol. 22) noe. , 1984

therefore finely lobed, these lobes grading into small hairs of anterior

surface; apex of flabellum weakly curved anteriorly, anterior surface

with numerous short simple hairs. Seta present on posterior lateral

surfaces of glossa basal to flabellum, one or two on each side among an-

nular hairs, also forming row across anterior surface basal to flabel-

lum. Annular hairs mostly tapering but distal ones often lanceolate.

Species studied: Centris atripes (SEM), poecila (C); Epicharis

elegans (C), rustica (SEM).

Subfamily Nom: Nomadinae. Disannulate surface not invaginated as in

Nomada, to deeply so like that of most long-tongued bees in Thalestria

(Figs. 77-81). Glossal rod usually weak, absent or only a feeble dark

zone in Nomada, Biastes, etc., but well sclerotized in Caenoprosopis,

lsepeolus, and Leiopodus, distinct and V-shaped in cross-section in

Thalestria. Seriate lines absent or nearly so, except in Triepeolus and

Thalestria which have small ridges adjacent to and on each side of rod.

Seriate hairs usually absent or very sparse and minute, but abundant and

minute in Triepeolus and Thalestria, absent on distal part of glossa in

Triepeolus. Disannulate surface otherwise hairless or with a few minute

hairs, more numerous in Triepeolus (5.11). Annular hairs slender and

tapering on basal part of glossa, for example in Triepeolus, and Nomada,

but apically broadened, lanceolate, often broadly so, in Morgania, —

Isepeolus, Protepeolus, and Leiopodus many of them with two or three

apical points, in Osiris many of those of anterior surface greatly ex-

panded with about five to seven apical points. Hairs(?) on either side

of disannular surface often expanded and sometimes fused into a thin

broad lamella on each side, this lamella may project away from glossa

but usually lies over and completely hides disannular surface (3.2);

such structures present in Leiopodus (Figs. 80, 97G), Osiris (Fig. 97G),

Nomada (Figs. 79, 98E), Epeoloides (Fig. 97K), Isepeolus, Caenoprosopis,

Holcopasites, and Morgania (Fig. 98A), but apparently absent in Biastes,

Thalestria, Triepeolus, Epeolus, and Doeringiella.

Distal specialization (Figs. 97, 98) variable, sometimes (e.g., in

Nomada, Epeoloides, Ammobatoides, Morgania, Holcopasites, and

Caenoprosopis) essentially a placoid flabellum, but with glossa not or

scarcely narrowed at its base; flabellum distinctly wider than glossa

at flabellar base in Ammobatoides and Holcopasites; posterior surface

of flabellum smooth or with weak coarse wrinkles in Epeoloides and

Ammobatoides; apical margin of flabellum rounded, curved anteriorly,

projecting little beyond transverse row of setae (7.8, 7.9); flabellum

without small hairs on the very limited anterior side (except annular

hairs, the distal ones of which may arise near base of distal

specialization) or in Nomada with a narrow zone of small, simple or lan-

Figure 97. Apices of glossae of Nomadinae (Anthophoridae). Scale

lines represent .01 mm; S = seta. A, Isepeolus viperinus, posterior

view. B, C, Thalestria spinosa, anterolateral and anteroapical views,

posterior surfaces at lower right in both. D, E, Doeringiella bizonata,

posterolateral and anterolateral views. F, Triepeolus verbesinae, an-

terolateral view. G, H, Leiopodus lacertinus, posterior and anterior

views. I, J, Osiris sp., posterior and posterolateral apical views. K,

L, Epeoloides pilosula, posterior and anterior views. M, Osiris sp.,

anterolateral view. ——$—$—$—

Glossae of Bees

Michener and Brooks

56 Contrib. Amer. Ent: tnste, vol. 22; not, 1984

ceolate hairs arising between row of setae and apical flabellar margin

(Fig. 98C). Setae present, one or two at each side of apical part of

glossa and transverse row or arch extending across anterior flabellar

surface preapically, these setae often quite large and extending beyond

flabellar margin (Fig. 98F). Osiris and Leiopodus (Fig. 97G-J, M) are

similar but the flabellum is thick, not or little curved anteriorly,

transversely wrinkled apically in some Osiris, and the setae are small

and widely separated, in Osiris scattered and not in a neat row (7.10).

A few genera have distal specializations similar to those of

Holcopasites, etc. (i.e., placoid), but the transverse arch of setae ex-

tends basally onto posterior surface, these setae arising along the mar-

gins of the flabellum (Biastes, Neolarra, Fig. 98L) or on the basal part

of posterior flabellar surface (Pseudodichroa) (7.10, Fig. 98H).

Distal specializations of Isepeolus, Protepeolus, Doeringiella,

Epeolus, and Triepeolus are thickened, not placoid, scarcely curved an-

teriorly, blunt, with a few short setae on both anterior and posterior

surfaces (6.9). While the setae are apparently derived from setae of

the transverse arch, no such row of setae is recognizable (7.11). In

Thalestria the structure is similar but the bare flabellar area on the

posterior surface is elongated basad, so that it is elongate oval; many

annular hairs are on the corresponding anterior surface (7.1, 7.11).

Species studied: Epeolini. Doeringiella bizonata (SEM); Epeolus

pusillus (SEM); Thalestria spinosa (C, SEM); Triepeolus verbesinae (C,

SEM). Nomadini. Nomada annulata (C, SEM). Osirini. Osiris obtusus

(SEM). Protepeolini. Isepeolus viperinus (C, SEM); Leiopodus

lacertinus (C, SEM). Epeoloidini. Epeoloides pilosula (SEM).

Ammobatini. Morgania histrio transvaalensis (SEM); Pseudodichroa

capensis (SEM). Caenoprosopidini. Caenoprosopis crabronina (C, SEM).

Holcopasitini. Holcopasites arizonicus (SEM), _ heliopsis (C).

Biastini. Biastes brevicornis (C, SEM). Ammobatoidini. Ammobatoides

abdominalis (SEM). Neolarrini. Neolarra verbesinae (SEM).

Subfamily Xylocopinae. Tribe Ceratinini. The following is based

on the nonparasitic Ceratinini; parasitic forms are described later.

Glossal rod distinct, crescentic in cross-section (Figs. 84, 85).

Seriate lines weak ridges lateral to margins of rod. Seriate hairs

small, abundant, in a row. Disannulate surface lateral to seriate lines

hairless. Flabellum (Fig. 99D-1) tapering basally to rod, apex rounded

or subtruncate, curved anteriorly, posterior surface smooth, anterior

surface concave, in allodapine genera extending well beyond arch of

setae and with small simple hairs (absent in Macrogalea, few in

Exoneurella), in Ceratina, Pithitis, and Manuelia extending not so far

beyond arch of setae and with only narrow band of hairs arising in space

between setal bases and apex of flabellum (7.8). Setae often recog-

nizable on anterior surface of glossa among annular hairs, basal to

flabellum; a transverse arch across anterior surface of flabellum con-

sists of large setae, often only four but six or more in Macrogalea.

Annular hairs lanceolate, especially laterally and on distal part of

glossa.

Among parasitic genera, Inquilina has a glossa similar to that of

other allodapine bees, but Nasutapis and Eucondylops (5.3) have greatly

reduced glossae, described as _ follows: Glossa rather short, in

Eucondylops tapering with the apex attenuate as in short-tongued bees.

Glossae of Bees

Michener and Brooks

9) ae

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poster

58 Contrib. Amer. Ent. Inst., vol. 22, no. 1, 1984

Disannular surface exposed, depressed to form a channel in Nasutapis,

not depressed in Eucondylops. Glossal rod and seriate lines and hairs

absent; disannulate surface with scattered tiny hairs in Nasutapis,

which also has moderately dense marginal hairs. Flabellum absent (6.1);

glossal apex with long annular hairs like those in more proximal posi-

tions (Fig. 1001). Annuli few and widely separated especially in

Eucondylops. Setae absent. Annular hairs slender, tapering.

Species studied: Allodape ceratinoides (SEM), stellarum (C);

Allodapula dichroa (SEM); Braunsapis calidula (SEM), facialis (C);

Ceratina calcarata (C), laeta (C), neomexicana (SEM), rupestris eons

(Ceratinidia) sp. (C); Eucondylops reducta (C, SEM); Exoneura hamulata

(SEM); Exoneurella lawsoni (SEM); Halt Halterapis nigrinervis ‘inervis (SEM);

Inquilina excavata (SEM); Macrogalea candida (C, SEM); Manuelia gayi (C,

SEM); Nasutapis s s straussorum (C, SEM); Pithitis binghami (C, SEM).

Tribe Xylocopini. Glossal rod shallowly lt U-shaped in cross-section

(Figs. 82, 83); seriate lines on very strong ridges rising from lateral

margins of rod and approaching one another at summits, thus largely en-

closing bacular canal; seriate hairs numerous but minute, on summits of

ridges. Disannulate surface lateral to seriate lines with scattered

minute hairs (5.11). Distal annulus far short of base of flabellum,

leaving bare shank (7.17) which is abruptly constricted at base of

flabellum. Flabellum broadly oval, flat, strongly constricted at base,

posterior surface with weak cobblestone pattern, apex rounded and weakly

curved anteriorly, anterior surface with numerous, small but moderately

long, curved hairs, simple in Xylocopa augusti, with one or two branches

each in X. virginica. Transverse row of about eight setae arising from

shoulder at distal end of shank; no setae on flabellum, no setae recog-

nizable among annular hairs. Annular hairs tapering.

Several of the main features were also illustrated for X. capitata

by Louw and Nicholson (1983).

Species studied: Lestis aeratus (C); Xylocopa augusti (SEM),

varipuncta (C), virginica (C).

FAMILY APIDAE

(Figs. 86-91, 100A-H, 101)

Members of this family share the characters listed above for long-

tongued bees. The glossa of the Apidae is quite variable; for that

reason and because of the enormous variability in anthophorid glossae,

there are no diagnostic glossal features of the Apidae. However, the

glossal rod is usually thicker than in anthophorids and has two or even

three longitudinal darkened lines, so that it appears double or triple.

Such rods are uncommon in Anthophoridae.

Subfamily Bombinae. Tribe Euglossini. Glossa flattened, three to

six times as wide as thick (Figs. 86, 87). Glossal rod (4.5) consisting

of two thickenings in floor or sides of glossal canal, often with a weak

thickening (reduced distally) between the two, so that there appear to

be two or three closely parallel rods; glossal canal small. Seriate

lines just lateral to lateral rods, and bearing minute seriate hairs

Michener and Brooks: Glossae of Bees 59

Figure 99. Apices of glossae of Xylocopinae CAnthophoridae).

Scale line represents .01 mm (0.1 mm for A and B); S = seta. A, B, C,

Xylocopa augusti, posterolateral, anterior, and anterior views. D, E,

Ceratina neomexicana, posterior and anterior views. F, G, Manuelia

gayi, anterior and posterior views. H, |, Allodape ceratinoides,

posterior and anterior views. J, Macrogalea candida, anterior view.

«Amer. Ent: bnst.; vol..22, nord, 1984

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Michener and Brooks: Glossae of Bees 61

flabellum, leaving bare shank (7.17) which is constricted before base

of flabellum. Flabellum several times as long as wide, flat, strongly

constricted at base, posterior surface with weak cobblestone pattern,

apex narrowly rounded and moderately to strongly curved anteriorly, an-

terior surface with numerous small hairs (Fig. 100A-E). Several setae

scattered on anterior surface of shank and near apex of shank on

posterior surface; no setae on flabellum, none seen among annular hairs.

Annular hairs (3.3) reduced or absent on much of anterior surface; enor-

mous and broadly lanceolate along lateral margins, these large hairs

partly closing over channel along concave posterior surface of glossa;

almost no hairs between these large hairs and glossal groove, although

this zone is crossed by weak extensions of annuli.

A possible alternative explanation of the glossal structure of

Euglossini is that the disannulate surface is broadly exposed, the weak,

hairless annuli across it being secondary thickenings, and the largely

enclosed tube being the bacular canal rather than the glossal canal.

This explanation seems unlikely in view of the condition in related bees

and the presence of what appear to be marginal hairs along the margins

of the glossal groove. However, the glossal canal is singularly small,

intermediate in size between what one would expect for a glossal canal

and for a bacular canal, and its floor and walls are extensively

sclerotized by parts of the rod; in this respect it suggests a bacular

canal.

Species studied: Eufriesea surinamensis (SEM), violacea (C),

sp. (C); Euglossa chalybeata (SEM), cordata (C); Eulaema cingulata (C),

meriana (SEM); Exaerete smaragdina (C).

The distinctive glossa of the Euglossini does not support placement

of this tribe in the same subfamily as the Bombini. The possibility of

such a relationship exists, however; the familiar euglossine features

are synapomorphies of members of that tribe and do not establish any

other relationship. :

Tribe Bombini. Glossal rod nearly flat in cross-section, darkened

on both sides of midline so that it appears double, a broad, thick pris-

matic strip (in which the dark bands lie) on floor of glossal canal, the

whole probably functioning as a large rod (Figs. 90, 91). Seriate lines

in form of strong ridge on each side of rod; seriate hairs: numerous,

small. Disannulate surface lateral to seriate lines with numerous

minute hairs (5.11). Distal annulus far short of base of flabellum,

leaving bare shank (7.17) which is constricted just before base of

flabellum; flabellum (7.12) broader than long, flat, placoid

(Fig. 99F-H), apex rounded (slightly lobed), curved anteriorly,

posterior surface smooth, anterior surface with numerous small hairs,

mostly simple but largest ones with one or two branches, these hairs

densest toward flabellar base which they surround and cover (Fig. 99F).

Setae forming row (partly hidden among short hairs) across apex of shank

on anterior side; a few small setae on anterior surface of shank; scat-

tered setae among annular hairs near sides of posterior surface of

distal part of glossa. Annular hairs slender, tapering.

Species studied: Bombus pennsylvanicus (C), morrisoni (SEM);

Psithyrus variabilis (C).

Subfamilies Meliponinae and Apinae. Glossal rod darkened on both

sides of midline (clear in Apis) so that in Meliponinae it appears

62 Contrib: Amer. ft. Met vol 22;on0o 1, 1984

Figure 101. Apices of glossae of Meliponinae and Apinae (Apidae).

Scale lines represent .01 mm; S = seta. A, B, C, Trigona (Partamona)

musarum, anterior, anterior, and posterior views. D, E, Dactylurina

staudingeri, posterior and anterior views. F, Meliponula bocandei, an-

terior view. G, Apis mellifera, posterior view. H, Lestrimelitta

limao, anterior view with flabellum curved forward to show posterior

surface. |, J, Apis mellifera, posterior and anterior views.

double, rod itself much larger than dark areas, prismatic, and nearly

surrounding: round bacular canal (Figs. 88, 89). Seriate lines on sum-

mits of lateral extensions of rod that enclose bacular canal; seriate

hairs numerous, small, directed mesad and closing over bacular canal.

Michener and Brooks: Glossae of Bees 63

Disannulate surface lateral to seriate lines with numerous minute hairs

(5.11) in some areas, sometimes minutely papillate. Flabellum (7.12,

Fig. 101) as broad as or broader than long (longer than broad in male

Apis), flat, placoid, apex rounded (slightly lobed in Lestrimelitta and

Apis), curved anteriorly, posterior surface smooth, anterior surface

with numerous (few in Lestrimelitta and male Apis), small, simple hairs

(branched near margin in worker Apis), densest toward flabellar base and

often spreading as a narrow collar around base of flabellum, distal to

annular hairs. Setae two or four (more in Apis), in transverse row just

basal to flabellum on anterior side, none on flabellum; scattered setae

among annular hairs on posterior surface of distal part of glossa. An-

nular hairs tapering to slender lanceolate.

The similarities of glossal structures of Meliponinae, Bombini, and

Apinae are impressive. They do not support the isolated position given

the Meliponinae by Winston and Michener (1977).

Species studied: Apinae. Apis dorsata (C), florea (C), mellifera

(C, SEM). Meliponinae. Dactylurina staudingeri (C, SEM); Lestrimelitta

limao (C, SEM); Melipona fasciata (C), marginata (C), rufiventris

brachychaeta (C, SEM); Meliponula | bocandei (C, SEM); Trigona buyss« buyssoni

(C), capitata zexmeniae (C), chanchamayoensis CC). trontare (Cy,

musarum (SEM), schrottkyi (C), (Hypotrigona) sp. (C), (Plebeia) sp. (C).

DISCUSSION

The glossal structure provides a wealth of morphological details

which are of significance in understanding the systematics and evolution

of the bees. The major division of the bees into short-tongued and

long-tongued families (Michener and Greenberg, 1980) is in general sup-

ported. Rather curiously, tongue length is not reliable in support of

this distinction. The following glossal characters are of interest in

this connection:

(1) The short-tongued families have the disannular (morphologically

posterior) surface of the glossa exposed (rarely invaginated, as in

Melitturga, Fig. 50) and not or only feebly strengthened by a glossal

rod. Because of the exposed disannulate surface the annuli do not go

all the way around the glossa. In long-tongued families, however, the

disannulate surface is invaginated along with the glossal rod (which is

a median thickening of this surface); the annuli therefore seem to go

most of the way around the glossa. Sometimes the surface and rod are

everted and exposed in dead specimens. In some parasitic genera the in-

vagination and rod have been lost; incomplete invagination in specimens

of Fideliidae and Tapinotaspis (Figs. 62 and 70) could represent reten-

tion of a primitive feature but we suspect it is merely a result of par-

tial eversion.

(2) When present, the seriate hairs of short-tongued bees are

divergent and the hairs themselves commonly large and some of them

branched. [Recognizable seriate hairs are absent in Colletidae and

Stenotritidae, hairs being widely scattered over the disannulate surface

of the glossa; in some members of other families the seriate hairs have

been nearly or entirely lost (Figs. 28, 47).] In contrast, the seriate

hairs of long-tongued bees, when present as they usually are, are

convergent, small to minute, simple, and arise from a ridge along each

64 Contrib. Amer. Ent. Imst., vol. 22; no. 1, 1984

edge of or near the glossal rod. The Ctenoplectridae, a family of

short-tongued bees with many features of long-tongued bees, has conver-

gent seriate hairs like long-tongued bees (Fig. 58).

(3) Short-tongued bees usually have an apically attenuate glossa

(i.e., with the lateral apical margins concave due to the apex being

drawn out), with the annulate and disannulate surfaces both reaching the

apex, the former bearing annular hairs, the latter seriate hairs. In

the Colletidae (except certain males) the disannulate surface is ex-

panded beyond the morphological apex of the glossa to form the glossal

lobes (Fig. 1). In most long-tongued bees the disannulate surface is

expanded beyond the morphological apex of the glossa to form the flabel-

lum. Both flabella and the colletid glossal lobes have disannulate

hairs on their anterior surfaces, suggesting similar origins for these

structures. Elsewhere the disannulate surface is entirely posterior (or

invaginated ).

Further consideration of flabella is necessary, however, for they

are neither restricted to long-tongued bees nor characteristic of all

such bees. No bee with a really short glossa has a flabellum or any

similar distal specialization. A few unrelated groups of short-tongued

bees with rather long glossae have distal specializations suggestive of

flabella, or have true flabella, sometimes much like those of long-

tongued bees. Thus while short-tongued bees with rather long glossae

and no flabella (Fig. 26) do exist, as well as long-tongued bees without

flabella, there is a tendency for an association between glossal length

and presence of a flabellum.

Various steps in flabellar development among short-tongued bees are

described in the Comparative Account, paragraphs 6.5-6.8 and 7.2-7.6.

This is not a phylogenetic series. Some species of Panurginae

(Andrenidae), Dufoureinae (Halictidae), and all Ctenoplectridae are in-

volved, with an interesting broadened glossal apex also in Dasypoda

(Melittidae). In the Panurginae every gradation exists from no sugges-

tion of a flabellum (Fig. 51A-C) to a well formed flabellum (Fig. 51F-J)

to a flabellum incredibly like that of long-tongued bees (Fig. 52A-E,

Perdita and Callonychium). The similarity between the placoid flabellum

of Perdita or Callonychium and that of long-tongued bees involves not

only position, but also shape, the smooth, convex, posterior surface,

the forward curvature, the curved row of setae across the base of the

anterior surface, and the small disannular hairs on the anterior surface

distal to the setal row. It is hard to suppose that the development of

such similar and complex placoid flabella could be independent in

Panurginae and long-tongued bees. Yet Panurginae have so many common

derived features (e.g., lack of the gonobase, two subantennal sutures)

not shared with long-tongued bees that it hardly seems believable that

long-tongued bees and the small group of Panurginae having flabella are

sister groups. The remarkable flabellar similarities must have arisen

from convergence, along with some elongation of the basal segments of

the labial palpi, a feature of many Panurginae that is more fully

developed in long-tongued bees.

Flabella are lost in certain groups that normally have them. The

flabellum was quite probably lost in ancestors of Perdita texana al-

though that species could represent an ancestral preflabellar condition.

The flabellum was certainly lost in the Habropoda-Emphoropsis group of

Michener and Brooks: Glossae of Bees 65

Anthophorini; other members of this tribe and all related tribes possess

flabella. Numerous stages of flabellar reduction can be found in the

parasitic anthophorid subfamily Nomadinae (see Comparative Account,.

paragraphs 7.9-7.11). These do not form any single series; reduction

must have occurred independently in different nomadine phyletic lines.

Loss of flabella also occurred in the parasitic allodapine xylocopines,

Eucondylops (derived from the nonparasitic genus Allodapula which has

a flabellum) and Nasutapis (derived from the nonparasitic genus

Braunsapis which has a flabellum) (Michener, 1970).

Various Nomadinae as well as Eucondylops and Nasutapis not only

show reduction or loss of flabella, but also reduction or loss of the

glossal rod and the glossal canal, so that the disannulate surface is

no longer invaginated (Figs. 78-81). Another parasitic group, the

Melectinae, while having a flabellum, shows similar reduction of the

glossal rod and canal (Fig. 73). Although some parasitic groups retain

glossal structure normal for their families or subfamilies, the reduc-

tion in glossal complexity in several parasitic groups suggests that

much of the glossal complexity may involve adaptations for nest con-

struction or provisioning rather than merely for feeding. Parasitic

bees presumably feed like any other bees. (Nasutapis and Eucondylops

are not known to go to flowers, but the other groups mentioned visit

flowers freely to take nectar.) They do not, however, have to construct

and provision nests, as do females of nonparasitic forms. Glossae are

Known to be involved in waxing or varnishing cells and the like, and it

is likely that some of their features are correlates of such behavior.

One would expect the glossa to be well equipped with organs of

taste and or tactile sensilla. This may be so, but the only structures

that we have identified as presumably sensory are the setae, which are

small and sparse. In many sphecoid wasps setae form an apical row

across the distal glossal margin. In Colletidae they commonly form a

preapical row on the anterior surface (Fig. 1). They are usually ab-

sent, however, in short-tongued bees, but occur in the following: many

Colletidae (anterior surface, also posterior surface in Diphaglossinae),

Stenotritidae (anterior and posterior surfaces), Halictidae (only seen

among annular hairs of Rophites), Andrenidae (only seen near glossal

-apices of a few genera of Panurginae), Oxaeidae (among extremities of

annuli in middle and basal parts of glossa), Melittidae (among annular

hairs of Macropis only), and Ctenoplectridae (near flabellum and on its

anterior surface). Among long-tongued bees, setae are usually present

among annular hairs and especially at the base of or on the flabellum.

The descriptions and illustrations of taxa give more detailed accounts

of the locations of setae. Since setae are sometimes hard to find among

the much longer and usually broader annulate hairs, we may have missed

them in some forms that possess them.

In order to provide a visually helpful summary of part of our data,

we have made a cladistic analysis using the Wagner 78 computer program

of S. J. Farris as implemented on the University of Kansas academic com-

puter. Table 1 lists the synapomorphies which we have recognized with

the corresponding plesiomorphies in parentheses, and in most cases a

brief indication of the reason for our decision as to the polarity of

each variable. The taxa are at the tribal level and above. The data

come only from the species listed earlier in this paper; the use of

66 Contrib. Amer.cEnt. last.) -vol. 22, m0. 1, 1984

names of family~group taxa does not indicate a broader survey of genera

and species. Taxa for which data are incomplete were omitted. There

are too few glossal variables with known polarity to make this summary

a reliable cladistic pattern for bees. It should be seen as a device

for making a body of glossal data more easily accessible and for

promoting discussion of some interesting problems. Unfortunately the

many fascinating characters of the flabellum exhibit much convergence

and it is difficult to determine the polarities of the variables.

Therefore the cladistic pattern for long-tongued bees as presented here

is not very useful.

The variables used include some that are almost duplicates. These

result from the major sexual differences in several glossal features in

the genera Meroglossa and Paleorhiza of the Hylaeinae (Colletidae), so

that the females and males would be placed in different families on the

basis of glossae if they were not known to be conspecific. One "taxon"

included in the study is Hylaeinae-MP, the letters referring to the two

genera mentioned which are distinguished from all other colletids by the

Andrena-like male glossa. There are six pairs of variables recorded

separately for females and males (Table 1) because of this situation,

as. follows: 1; 2) 3, 4; 5,6, 12,137.16, 7} and 21, 22... in each pair

the first is for females, the second for males. The cladograms in

Figures 102 and 103 are based on all characters but Figure 104 is based

on female characters only, to correct the over weighting of some

characters that results from treating them separately for each sex.

On the cladograms, autapomorphies are omitted. A character is

shown on the stem leading to a single taxon if the character appears

elsewhere in the cladogram. In Figures 103 and 104 the right hand por-

tions of the cladogram, representing the Ctenoplectridae and long-

tongued bees, are omitted since they are similar to the comparable part

of Figure 102.

A feature of all the cladograms is the numerous characters

isolating the Colletidae and Stenotritidae from the rest of the bees.

Elsewhere the number of characters on any given stem is small. An in-

teresting feature is the branch leading to several parasitic groups, the

Melectini and various nomadine tribes. The position and content of this

branch result from loss of characters like 19 and 26, and from secondary

loss of characters 7 and 8 such that apomorphies (losses) within long-

tongued bees look like (and for the cladogram are interpreted as)

plesiomorphic characters of bees as a whole.

The Colletidae requires attention because the family has been

recognized principally on the basis of its glossal shape. This family

has traditionally (e.g., Michener, 1944) been regarded as the most

primitive family of bees, largely because of the broad and commonly

bilobed glossa, sometimes merely truncate or rounded at the apex. This

shape resembles that of many other Hymenoptera including the sphecoid

wasps from which bees quite clearly arose. The southern distributions

of most colletid groups (Michener, 1965, 1979), suggestive of the dis-

junct southern ranges of many primitive groups of organisms, together

with the possession of other (i.e., nonglossal) plesiomorphies, also

support the idea that colletids may be the most primitive bees. They

could be the sister group of all other bees, or a paraphyletic group

from which other bees arose.

Michener and Brooks: Glossae of Bees 67

EUCERINI -CENTRIDINI

CTENIOSCHELINI

ISEPEOLINI \.%

TETRAPEDINI vo X,

CAENOPROSOPIDINI eee CANEPHORULINI

PANURGINAE - DUFOUREINAE

8 1 >

NEOPASITINI NOMADINI oy Y ANTHOPHORINI -

ANDRENINAE - MELITTIDAE MEGACHILIDAE

Gy, CERATININI

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HYLAEINAE - MP eel ie gah ash

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102

ANDRENINAE - MELITTIDAE ANDRENINAE - MELITTI DAE

HALICTINAE -N HALICTINAE- NOMI INAE

adi PANURGINAE - PANURGINAE -

DUFOUREINAE COLLETIDAE WITH DUFOURE!NAE

HYLAEINAE - MP

STENOTRITIDAE STENOTRITIDAE

COLLETIDAE

103 104

Figures 102-104. Cladograms based upon glossal characters of both

sexes. Character numbers and polarities correspond with those in Table

1; those indicated by an X are reversions. Autapomorphies are omitted.

102. Cladogram for all higher taxa. The out-group used was all

plesiomorphies, a condition also found in the Andreninae-Melittidae.

"Colletidae" includes all subfamilies, even Hylaeinae other than

Hylaeinae-MP (Meroglossa, Palaeorhiza). "Andreninae-Melittidae" in-

cludes also Oxaeidae. "Anthophorini-Megachilidae" includes’ also

Fideliidae, Exomalopsini and Melitomini. These taxa do not differ in

the characters here analyzed. 103. Basal part of above cladogram.

Polarities are reversed to correspond with "traditional" views for

characters 3, 4, 5, 6, 12, 13, 16, and 17 (see text). The out-group was

all plesiomorphies, a condition found in some Sphecidae. 104. Basal

part of above cladogram based on glossal characters of females only.

Explanation as for Figure 103.

68 Contribs Amer. Ent.-inst., vol. 22, no. 1, 1884

Perkins (1912) and McGinley (1980), however, cast doubt on the

traditional idea that an obtuse glossa is primitive among bees, and sug-

gested that the acute glossa of certain male colletids (Meroglossa and

Paleorhiza) may be more primitive, with the obtuse or bilobed glossa

being a "special development" in the words of Perkins. According to

this view the most primitive bees must have had a short, acute glossa

like most short-tongued bees. Female colletids then evolved a broad,

obtuse glossa, presumably in connection with their use of the glossa to

paint their distinctive lining material onto the cell and sometimes the

burrow walls. Males initially would have retained an acute glossa and

two genera still do, but perhaps because the acute glossa had no special

advantages for males and required maintenance of separate genetic

machinery, it disappeared in most male colletids. According to this

idea the obtuse or bilobed glossa is an apomorphy among bees, a superfi-

cial reversion toward the ancestral sphecoid glossa. Like Michener

(1981), we have accepted this view and elaborated on it in determining

the polarity of the characters listed in Table 1 and in preparing Figure

102.

It seems clear that the Colletidae have certain glossal apomorphies

at the family level. These are the glossal lobes (1 and 2, Table 1),

the glossal brush which is on the lobes and is not separately listed in

Table 1, and the preapical fringe (21 and 22, Table 1). These

characters are not found in sphecoid wasps nor in any other family of

bees. Although lacking in the males of the Hylaeinae-MP group, these

characters indicate that the Colletidae is a holophyletic unit,

presumably the sister group to most or all other bees, and not a

paraphyletic unit from which other bees arose. (Of course the colletid

apomorphies could have been lost in the evolution of all other bees,

making Colletidae paraphyletic, but this seems improbable.) The col-

letid apomorphies do not establish by themselves whether the truncate

or emarginate glossa of typical colletids is derived from an acute apoid

glossa or from a truncate or emarginate sphecoid glossa.

The typical colletid glossa, even without the glossal lobes, is

blunt, truncate, or emarginate (3 and 4, Table 1). This shape, ac-

cording to the Perkins-McGinley view, has been derived from the acute

glossa like that found in Andreninae or Halictinae. The shape could

have arisen as the base for supporting colletid apomorphies, glossal

lobes and the preapical fringe (1, 2, 21, 22, Table 1). This all makes

good sense when one postulates, as suggested above, that the colletid

glossa arose as a correlate of nest construction by females, leaving

males at first (as in Hylaeinae-MP) with acute glossae.

To accept the traditional view that the typical colletid glossal

shape is ancestral and that acute glossae arose from it requires that

in ancestors of Hylaeinae-MP, an acute glossa evolved first in males,

and later became characteristic of both sexes as in Andreninae,

Halictinae, etc. This idea is hard to accept because the male glossa

has no known special function differing from that of the female. It is

the female's glossa that has a special function in nest construction,

unlike that of the male. It is for these reasons that for variables 3

and 4, Table 1, we list the pointed glossa as plesiomorphic (for bees;

of course it is an apomorphy among Hymenoptera or looked at from the

condition of sphecoid wasps).

Michener and Brooks: Glossae of Bees 69

This view of polarity in variables 3 and 4 is not without problems,

however. There are three other morphologically and probably func-

tionally independent variables that are distributed among taxa in the

same way as 3 and 4, i.e., the sphecoid features are found in the col-

letids except for males of Hylaeinae-MP, while the latter and other bees

(except Stenotritidae) have the alternative features. These variables

are (a) the extent of the annulate surface on the posterior glossal sur-

face (5, 6, Table 1), (b) the shape of the annular hairs (12, 13, Table

1), and (c) the scattered vs. seriate positioning of the disannular

hairs (16, 17, Table 1). It is not legitimate to assume reversion in

glossal shape to the colletid form from an acute antecedent unless one

also accepts the reversion in the other three variables. We have ac-

cordingly coded the three other variables also as though they reverted

in colletids to the sphecoid condition, i.e., the apomorphies among bees

look the same as the plesiomorphic conditions in sphecoids. We em-

phasize here the gross difference in nearly every attribute between male

and female glossae of Meroglossa and Palaeorhiza (Figs. 19 and 20). The

male structures can easily be compared and the parts homologized with

those of most short-tongued bees while the female glossa is like that

of other colletids.

We are faced with two improbabilities--(a) evolution of an acute

glossa (and the three accompanying features) first in males, later to

be transferred to females, or (b) parallel reversion of four different

characters in colletids from bee-like to sphecoid-like. The correct

choice may become evident later. For the present, however, we leave the

question open. We reversed the coding of characters 3, 4, 5, 6, 12, 13,

16, and 17 (really only four characters if one ignores the sexual dif-

ferences in Hylaeinae-MP) and made an alternative cladogram (Fig. 103)

based on the traditional view that colletid characters are mostly

plesiomorphic. This diagram seems to be as informative as Figure 102,

although different. Figure 104 is based on female characters only,

coded traditionally, and sidesteps the Hylaeinae-MP problem. The

cladogram based on female characters only, coded as in Table 1, had a

branching pattern like that of Figure 102. Of course a cladogram based

on male characters only would put Hylaeinae-MP entirely away from the

Colletidae and with the Andreninae.

The endemic Australian family Stenotritidae is important in this

connection. It has the colletid glossal characters (3, 4, 5, 6, 12, 13,

16, 17) except for the obvious apomorphies, 1, 2, 21, 22 (Table 1).

Thus its glossa could be similar to the type from which the typical col-

letid glossa evolved. If this were true, one would have to accept the

traditional view and believe that the pointed glossa and associated fea-

tures arose first in males.

Given the fact that males of a few colletids have glossae that are

in all features like those of acute-tongued, noncolletid, short-tongued

bees, one may expect to discover bees that are in most features col-

letids, but that have acute glossae in both sexes, or at any rate glos-

sae that are not of the usual colletid style. Possibly such bees should

be included in the Colletidae, since obviously the colletid-style glossa

is not an essential feature of the family, at least in males. Can-

didates for possible inclusion in the Colletidae are the Oxaeidae and

Stenotritidae. This idea is attractive because of the similarity of

70 Contribs Amer. Entutuinst.; vol. Ze, no. 1, 1984

these families in various features to the colletid subfamily

Diphaglossinae.

TABLE 1

Synapomorphies (with corresponding plesiomorphies in parentheses )

for some glossal characters of bees. Numbers in parentheses refer to

paragraphs where the characters are more fully explained. For variable

26, two supposedly sequential, derived characters are indicated by the

numbers 1 and 2. Variables 15 and 28 are purposely omitted since they

were autapomorphic.

1.Glossa of female with glossal lobes so that it is truncate or

bilobed (1,2;°6:2).

(Glossa of female without glossal lobes. Glossal lobes are

absent in Sphecidae. )

2.Glossa of male with glossal lobes so that it is truncate or

bitobed. (1:2).6.2).

(Glossa of male without glossal lobes. Glossal lobes are ab-

sent in Sphecidae. )

3.Glossa of female (excluding glossal lobes if present) rounded,

truncate, or emarginate at apex (1.2).

(Glossa of female pointed.See Discussion. )

4.Glossa of male (excluding glossal lobes if present) rounded,

truncate, or emarginate at apex (1.2).

(Glossa of male pointed.See Discussion. )

5.Annulate surface of female entirely or almost entirely on an-

terior side of glossa (2.1).

(Annulate surface of female curling onto posterior surface of

glossa. See Discussion. )

6.Annulate surface of male entirely or almost entirely on anterior

side of glossa (2.1).

(Annulate surface of male curling onto posterior surface of

glossa.See Discussion. )

7.Disannulate surface of glossa largely hidden (invaginated) (2.2,

Paes ao).

(Disannulate surface of glossa exposed, as in Sphecidae. )

8.Glossal rod present (4.4, 4.5).

(Glossal rod absent, as in Sphecidae. )

9.Glossal rod appearing double (4.5).

(Glossal rod absent or if present appearing simple. The rod

is absent in Sphecidae and many bees, and is not double in

most bees that have a rod.)

10.Bacular canal nearly closed (4.5).

(Bacular canal absent or wide open. There is no such canal in

forms that lack a rod, as well as in most forms that have a

rod. )

11.Margins of disannulate surface with thin, broad lamellae Couey.

(Margins of disannulate surface without lamellae. Such lamel-

lae are present only in a few otherwise strongly apomorphic

bees. )

12.Annular hairs of glossa of female minute, short, blunt or

capitate (3.1, 3.3).

13%

14.

16;

i ae

18.

19.

rae

lle

24.

oo.

26.

a7,

Michener and Brooks: Glossae of Bees 71

(Annular hairs of glossa of female elongated, ending in one or

more points.See Discussion. )

Annular hairs of glossa of male minute, short, blunt or capitate

Wate eee

(Annular hairs of glossa of male elongate, ending in one or

more points.See Discussion. )

Annular hairs of glossa, or some of them, divergently branched

preapically (3.4).

(Annular hairs not branched, as in Sphecidae and most bees. )

Disannular hairs of female scattered (5.1-5.10).

(Disannular hairs of female in two rows (the seriate hairs),

or sometimes absent.See Discussion. )

Disannular hairs of male scattered (5.1-5.10).

(Disannular hairs of male in two rows (the seriate hairs), or

sometimes absent.See Discussion. )

Disannular hairs convergent, seriate, rarely absent (5.1-5.10).

(Disannular hairs divergent and seriate, or scattered, rarely

absent.See Discussion. )

Disannular hairs absent (5.10).

(Disannular hairs present.Such hairs are present in

Sphecidae. )

Glossa with apical specialization (not including glossal lobes

of Colletidae) (6.5).

(Glossa without apical specialization, as in Sphecidae and

many bees, or with apical lobes in Colletidae. )

.Glossa of female with a preapical fringe (1.2).

(Glossa of female without a preapical fringe, as in

Sphecidae. )

Glossa of male with a preapical fringe (1.2).

(Glossa of male without a preapical fringe, as in Sphecidae. )

Flabellum present (7).

(Flabellum absent, as in Sphecidae and many bees.)

Posterior surface of flabellum cobblestoned (7.14, 7.15).

— (Posterior surface of flabellum not cobblestoned or flabellum

absent. )

Apical specialization or flabellum with setae on posterior as

well as anterior surface (7.10, 7.11).

(Apical specialization or flabellum without setae or flabellum

absent. This is as in Sphecidae and nonparasitic bees. )

(1) Flabellum reduced so that setae basal to flabellum arise

near its apex but a zone of disannular hairs is present on an-

terior surface between setae and apex (7.8).

(2) Flabellum more reduced, so that no zone of disannular

hairs intervenes between setae basal to flabellum and its apex

C7 is,

(Flabellum without setae seeming to arise near its apex, or

flabellum absent. Sphecidae and many bees lack a flabellum.

Among bees that have one, a widespread and presumably

plesiomorphic pattern is a flabellum extending far beyond its

base and with a large anterior surface occupied by hairs. )

Setae arising on anterior surface of flabellum proper (7.13).

V2 Contrib: Amer. oEnt. lmst. > vol.. 22 mo. 1, 1984

(Setae, if present, basal to flabellum, if present. This is

as in nearly all bees. )

29.Elongate, preflabellar, nonannular, hairless zone present

CPeTE).

(No such hairless zone, the flabellum, if present, arising

from apex of hairy part of glossa, as in nearly all flabellate

bees. )

30.Basiglossal sclerite rather long with a pair of posterior basal

processes (hence the term bonnet-like sclerite).

(Basiglossal sclerite a transverse ribbon or band, as in

Sphecidae. )

Michener and Brooks: Glossae of Bees 73

LITERATURE CHIED

Fabricius, J. C. 1804. Systema Piezatorum. xiv+439+30 pp.,

Brunsvig, C. Reichard.

Kirby, W. 1802. Monographia apum Angliae. Vols. 1 and 2, London,

J. White.

Luow, G. N. and §S. W. Nicholson. 1983. Thermal, energetic and

nutritional considerations in the foraging and reproduction of

the carpenter bee Xylocopa capitate.” Jour. Entom. Soc.

Southern Alr., 46: 227-240.

McGinley, R. J. 1980. Glossal morphology of the Colletidae and

recognition of the Stenotritidae at the family level. Jour.

Kansas Entom. Soc., 53:539-552.

Michener, C. D. 1944. Comparative external morphology, phylogeny,

and a classification of the bees. Bull. Amer. Mus. Nat.

Hist., Gev1ot=426.

Michener, C. D. 1965. A classification of the bees of the

Australian and South Pacific regions. Bull. Amer. Mus. Nat.

Hist. , 13071-3062, Dis. 1715:

Michener, C. D. 1970. Social parasites among African allodapine

bees. Zool. Jour. Linnean Soc. [London], .49:199-215.

Michener, C. D. 1979. Biogeography of the bees. Ann. Missouri

Bot. Gard., 66:27 7-347.

Michener, C. D. 1981. Comparative morphology of the middle coxae

of Apoidea. Jour. Kansas Entom. Soc., 54:319-326.

Michener, C. D. and L. Greenberg. 1980. Ctenoplectridae and the

origin of long-tongued bees. Zool. Jour. Linnean Soc.

[London], 69:183-203.

Perkins, R. C. L. 1912. Notes, with descriptions of new species,

on aculeate Hymenoptera of the Australian region. Ann. Mag.

Nat. Hist., (8)9:96-121.

Reaumur, R. A. F. 1742. Mémoires pour servir a l'histoire des in-

sectes. Vol. 1, Paris, Imprimerie Royale.

Saunders, E. 1890. On the tongues of British Hymenoptera An-

thophita. Jour. Linnean Soc. London, Zo0l,, 23:410-4a2.

Snodgrass, R.E. 1935. Principles of insect morphology. ix+667

pp., New York and London, McGraw-Hill.

Snodgrass, R. E. 1956. Anatomy of the honey bee. xiv+334 pp.,

Ithaca, N. Y., Cornell Univ. Press.

Winston, M. L. 1979. The proboscis of the long-tongued bees: a

comparative study. Univ. Kansas Sci. Bull., 51:631-667.

Winston, M.,L. and €. DD. Michener., 1977. Dual origin of manly

social behavior among bees. Proc. Nat. Acad. Sci. (U.S.A.),

74: 1135-7 Tae;

THE NIGRISCAPUS GROUP OF VENTURIA

CHYMENOPTERA: ICHNEUMONIDAE)

David B. Wahl

Department of Entomology

University of Kansas

ABSTRACT

Descriptions, figures, and a key are given for fifteen species of

the Nigriscapus species group of Venturia, all of which are confined to

the New World, the majority being Neotropical. Fourteen species cv.

bicarinata, catarinensis, catoptron, citriscapus, compressa, depressa,

genalis, longicauda, musae, nitida, plaumanni, tetragona, tezcatlipocai,

townesorum) are described as new, and the remaining species, nigriscapus

(Viereck) is redescribed. The characters of the group are analyzed and

a tenative phylogeny presented.

ACKNOWLEDGEMENTS

| express my gratitude to C. D. Michener of the University of Kan-

sas for his advice, encouragement, and suggestions throughout this

study. My thanks also go to George W. Byers of the University of Kansas

for carefully reading the manuscript. The computer skills of Robert W.

Brooks of the University of Kansas were of great help in the preparation

of camera-ready copy. | am especially appreciative of the help given

by Henry and Marjorie Townes of the American Entomological Institute,

Ann Arbor, Michigan, for initial discussions concerning the investiga-

tion and for lending most of the specimens used. | am indebted to Syd-

ney Cameron for her advice and support during the investigation.

INTRODUCTION

The Nigriscapus Group of the genus Venturia (Ichneumonidae:

Porizontinae) consists of fifteen species, all restricted to the New

World, and all but one tropical. Venturia nigriscapus (Viereck) is the

only species previously described. Available information indicates that

they are parasitoids of moderate-sized Macrolepidoptera. Although lack

of detailed collection records precludes an analysis of habitats, they

are absent in deserts and at least one species occurs in seasonally cold

areas.

Preliminary results indicate that the Nigriscapus Group (charac-

terized below) is monophyletic (sensu Hennig). Until Venturia is ex-

amined on a world-wide basis and its phylogeny better known, the taxon

is best regarded as an informal but monophyletic assemblage of species.

2 Contrib. Amer. Ent. inst, , vol. 22, no. 2, 1984

MATERIALS, METHODS, AND TERMINOLOGY

Most specimens examined for this study were borrowed from the fol-

lowing collections; | am indebted to the curators listed (acronyms used

are either from Arnett and Samuelson (1969) or consistent with their

style):

California Academy of Sciences (CASC): San Francisco, California

(Dro PoA. Arnaud, oF:

Canadian National Collections (CNCI): Ottawa, Ontario (Dr. John Bar-

ron).

Fundacion e Instituto Miguel Lillo (FIML): Universidad Nacional de

Tucuman, San Miguel de Tucumdn, Argentina (Dr. A. Willink).

Museum of Comparative Zoology (MCZC): Harvard University,

Cambridge, Massachusetts (Dr. R. J. McGinley).

Henry and Marjorie Townes (HMTC): 5950 Warren Road, Ann Arbor,

Michigan.

National Museum of Natural History (USNM): Washington, D.C. (Dr.

Arnold Menke).

The terminology used for structures is mostly that of Townes

(1969). Gena is used in place of "temple"; malar space is similarly

substituted for "cheek". The mesopleural depression immediately an-

terior and ventral to the speculum is referred to as the scrobal groove

(figs. 4-5), assuming that the "mesopleural fovea" is homologous to the

episternal scrobe of other Hymenoptera (C. D. Michener, 1944, and pers.

comm.). Mesosoma and metasoma are used to refer to the apparent thorax

and abdomen respectively; the term first metasomal segment applies to

the second true abdominal segment. Reference to the orientation of the

various parts of the body follows Townes (1969), in that the legs are

considered to be stretched out horizontally at right angles to the body.

For the purposes of this paper, the vertex does not include the ocellar

triangle and is considered that region between the lateral ocellus and

the summit of the compound eye.

The surface sculpture between punctures on the head and mesosoma is

of the type referred to by Harris (1979) as granulate. This varies in

strength, from strongly developed to weak (with only shallow grooves

present) to absent (appearing smooth and polished even at high (80x)

magnification). Coarse punctures have a diameter of about 0.30 times

the greatest diameter of the lateral ocellus; moderate-sized punctures

have a diameter of about 0.15 times the greatest diameter of the lateral

ocellus.

Coloration of the body parts, especially that of the metasoma, can

be affected by the amount of oils on the specimen (from internal or

other sources); oily specimens are generally darker and often exhibit

irregular fuscous areas on the normally brownish-red portions of the

metasoma. The color of the metasomal ventral intersegmental membranes

is yellowish. The color of the veins of the fore and hind wings, both

dorsally and ventrally, is dark brown, although occasional specimens are

ventrally brownish-white.

In males, the carinae and general surface sculpturing of the

propodeum are more pronounced; the areola and petiolar area are narrower

than in the female. While there are differences in the outlines of the

propodea, the problem of viewing each specimen in the same orientation

Wahl: Nigriscapus Group 3

and the difficulty of attaining a defined edge (due to the sloping sides

of the structure) should be kept in mind.

Most measurements are self-explanatory. The malar space is

measured from the upper articulation of the mandible to the lower margin

of the compound eye. Width of the gena relative to the eye is measured

in lateral view; the resulting value is only approximate. The length

of the ovipositor, if its base is hidden by enveloping terga, is

estimated relative to the position of the apices of the valvifers and

is believed to be fairly accurate. Mesopleural punctation is measured

in an area immediately below the speculum (see circled area in fig. 4).

When the lengths of the body, wing, and ovipositor are given, the values

in parentheses are those of the holotype or allotype.

RELATIONSHIPS

Venturia (synonyms: Idechthis, Devorgilla, Balcarcia, Notamor-

phota, Exidechthis) has been generically described in detail by Townes

(1969) and Gupta and Maheshwary (1977) and it is unnecessary to repeat

their descriptions and generic synonomy. The features that readily

distinguish Venturia from other porizontines are:

1. Eye not emarginate opposite antennal socket.

2. Gena wide, not receding immediately behind eyes (as in Casinaria

or Charops).

3. Propodeum without median longitudinal depression, its apex

usually reaching at least middle of hind coxa; second Jateral

area usually defined by carinae.

4. First metasomal segment: postpetiole without carinae; petiole

cylindrical, suture separating first tergum from sternum in

middle of lateral aspect; first tergum without lateral groove

(glymma), or with only very faint traces.

5. Ovipositor usually long, 2-3 times apical depth of metasoma.

The existence of a distinct group of largely Neotropical Venturia,

with nigriscapus (Vier.) as the only described representative, has been

recognized by ichneumonid specialists for some time (H. K. Townes, pers.

comm.). The only literature reference is that of Carlson (1979, >;

634), who noted the existence of such a group.

Polarities of characters for the Nigriscapus Group were determined

by out-group comparison (examination of related taxa-- see Wiley (1981)

and Watrous and Wheeler (1981) for extended discussion). Although the

phylogeny of Venturia and related genera has not been analyzed in

detail, Venturia isan apparently monophyletic group (sensu Hennig) that,

like Sinophorus, probably originated in the very large ye and paraphyletic

genus Campoplex. Finlayson (1975) cites several larval characters that

suggest a possible relationship with Sinophorus. Until these can be

analyzed more fully in relation to other porizontine characters, | will,

on the basis of my observations of adult morphology, consider the

sister-group of Venturia to be within Campoplex, as currently recog-

nized. Accordingly, | have used Campoplex, Sinophorus, and other

species of Venturia as outgroups. The following is a list of characters

considered to be apomorphic for the Nigriscapus Group, with comments on

the bases for decisions on character polarities.

4 Contrib. Amer. Ent. Inst., vol. 22, no. 2, 1984

1. Face and clypeus coarsely granulosopunctate. (Facial and

clypeal punctation fine, not coarse, in outgroups. )

2. Malar space 0.5-0.6 times basal width of mandible. (Usually

0.2-0.3 in other Venturia species, often 0.5 or larger in Cam-

poplex and Sinophorus. )

3. Gena wide and flat, 1.4-1.6 times as wide across middle as

across eye, receding in dorsal view as in fig. 6. (1.7-3.0

times as long as eye in other Venturia species, near latter

value in majority of species; gena receding in dorsal view as in

figs. 7-8; gena often wide in Campoplex and Sinophorus, but not

flattened. )

4. Fore wing with nervulus distal to basal vein by about 0.3 times

its length (fig. 1). (Opposite or slightly distal in out-

groups.)

5. Postnervulus intercepted at upper 0.3 by subdiscoideus (fig. 1).

(Intercepted at or near middle in outgroups. )

6. Lower division of metapleurum coarsely rugosopunctate with large

shallow punctures, at least on the lower 0.3-0.5. (Almost al-

ways closely punctate on shining or granulate surface in out-

groups; if rugae are present next to submetapleural carina, they

are weak, not high and strong.)

7. First metasomal segment long and straight (or upcurved in catop-

tron) in lateral profile (fig. 1) with width of postpetiole not

much greater than that of petiole (fig. 9). (Campoplex,

Sinophorus, and most Venturia usually have postpetiole wider

than petiole (fig. 10); most Venturia species have the petiole

relatively shorter and postpetiole more convex (fig. 2).)

8. Second metasomal segment long and narrow with width of apex

0.3-0.4 times length of segment. (Most Venturia species, and

all Campoplex and Sinophorus, have the segment much shorter and

wider; remaining Venturia not so narrow or relatively elongate. )

9. Metasoma narrowly compressed, as in Ophion or the Anomaloninae.

(This condition approached by only a few Venturia. )

10. Ovipositor 1.7-2.1 times length of hind femur. (COvipositor

usually 2.3-3.0 times length of hind femur in most Campoplex,

Sinophorus, and other Venturia species, with most nearer latter

value. )

11. Body length 9.2-14.2 mm, with most species 10.5-12.0 mm. (Most

Campoplex and Sinophorus 5-7 mm in length; other Venturia range

from 3.9-9 mm in length, most being 6-7 mm.)

Only characters 1, 5, 6, and the upper range of body size are

unique to the Nigriscapus Group. Characters 2 and 3 may be found in

perhaps one-quarter of the species of Venturia. The remaining

characters were each found only in several species. No Venturia species

outside this group possesses all of the above characters. Species may

be found that have one or two of them but they will be readily excluded

by the lack of the others.

The relationship of the Nigriscapus Group to the rest of the genus

is not clear. Possibly it is related to several as yet undescribed

Nearctic species, but a more comprehensive analysis of the genus must

be undertaken.

10,

Ths

Wahl: Nigriscapus Group 5

Characters that are apomorphic within the Nigriscapus Group are:

Malar space 0.6-0.7 as long as mandibular width. (0.5-0.6 in

Campoplex and Sinophorus; usually about 0.3 in other Venturia

species; on the basis of the distribution of a malar space of

0.5, | assume this to be the primitive state for the group, al-

though an initial state of 0.6 and subsequent reversal to 0.5 is

equally possible. )

Occipital carina medially straight. (Arched in outgroups. )

Transverse rugae on polished lateral area of pronotum adjacent

to collar weak or absent. (Rugae usually present and strong in

outgroups; if absent, pronotum granulate, not polished. )

Upper posterior corner of lateral area of pronotum with areas

between punctures smooth or weakly granulate. (Areas between

punctures strongly granulate in outgroups. )

Spaces between punctures on mesopleurum polished and smooth.

(Spaces usually distinctly granulate in Venturia, Campoplex, and

Sinophorus. )

Upper anterior region of mesopleurum with area immediately ad-

jacent to scrobal groove having transverse rugae absent or

reduced and few or no punctures; surface granulation absent so

that this area appears to be an extension of speculum (see

outlined area in fig. 5 for indication of total expanse of

polished area). (In other members of the Nigriscapus Group and

in other Venturia, Campoplex, and all but one of the examined

Sinophorus, the area in question has extensions of the scrobal

groove's transverse rugae plus a normal complement of

mesopleural punctation and surface sculpture. )

Scutellum coarsely punctate, surface polished. (This is a par-

tial reversal of one of the apomorphies of the Nigriscapus

Group; other species of Venturia and Campoplex have small punc-

tures on a granulate surface; a few Sinophorus have small punc-

tures on a smooth surface. )

Mesopleurum with large shallow punctures separated by about 0.2

times their diameters or confluent. (Outgroups have small deep

punctures usually separated by more than 0.2 times their

diameters. ) 7

Scape, pedicel, tegula, coxae, and hind femur black or fuscous;

see discussion of nigriscapus (Vier. ) for reasons why it is con-

sidered to possess this character. (In other groups of Venturia

and Campoplex, the majority have the fore and middle coxae yel-

low or, if all coxae are black, the scape, pedicel, and tegula

usually are not dark; the joint possession of dark coxae, scape,

pedicel, and tegula is very rare in Venturia; Sinophorus tends

to have the above parts fuscous-black, but rarely has the hind

femur darkened as well. )

Hind femur fuscous. (This character seems to be independent of

character 11; fuscous coloration of the hind femur is rare when

the fore and middle leg coloration is yellowish. )

Metapleurum with lower division with at least upper half

coarsely punctate, rugae weak or absent.

6 Contrib. Amery Ent.<Imst., vol. 22,>no. 2, 1984

(The plesiomorphic condition, which is one of the apomorphies of

the Nigriscapus Group, is to have the rugae more noticeable than

the punctures, forming a series of parallel low vertical

ridges. )

12. Propodeal carinae pattern as in figs. 23-25 and 28: median and

longitudinal carinae tending to be weak and irregular in height

and outline; basal transverse carina curved laterally, not

angular; basal section of areola, between median longitudinal

carinae, relatively wide (compared to, for example, figs. 19 or

27); median longitudinal carinae parallel, not diverging

laterally at their midpoints. (This particular configuration is

not found in outgroups. )

13. Second lateral area of propodeum short and rectangular (figs.

12,15,20 and 23). (This configuration is limited to V. genalis,

tetragona, and males of nigriscapus and townesorum,; it has ap-

parently evolved twice. )

A cladogram for the group (fig. 29) was generated using the "Wagner

78" parsimony program of James S. Farris. This was identical to one

previously done by hand.

A weak point of the analysis results from the lack of male

specimens for musae and longicauda. It is not known if musae males ex-

hibit a squaring of the second lateral area, or if males of longicauda

possess large and shallow punctures on the mesopleurum. | have, in the

absence of information, treated each in the negative; further informa-

tion could change the cladogram.

Autapomorphies that distinguish each species are not shown on the

cladograms except when the character appears elsewhere within the

species group.

A considerable number of the species recognized herein are

described from only one specimen or a limited series. Decisions as to

specific status were made on the basis of character variability in those

species represented by relatively abundant material.

Some interesting patterns become evident when distribution data are

added to the cladogram, which | view as a statement of immediate common

ancestry. V. nigriscapus, found in eastern North America (and possibly

Mexico) is the sister species to a Central American species, townesorum.

These species are most closely related to two Central American species,

musae and longicauda. This pattern is found in a wide number of plant

and animal taxa (Rosen, 1978). Unfortunately, as Rosen points out,

present knowledge does not allow any temporal boundaries to be set to

these patterns; they may not even be due to a common event or events.

V. nitida, a member of a subgroup that is otherwise South American, ap-

pears to have dispersed through North and Central America, whose fauna

otherwise consists only of the relatively primitive nigriscapus, musae,

townesorum, and longicauda. V. tezcatlipocai, known at present from

only one location in Mexico, is perhaps derived from nitida, or a

similar common ancestor.

Wahl: Nigriscapus Group 7

Key to the Species of the Nigriscapus Group

Fore and middle coxae with basal 0.7 black or dark brown.

se peo gn en ego oe tga ny a NT PN ga fl, ety ek Say ie ot Peay on oped Mas on ny LMU en ecey Une aegis vane aT Bice ipgiescf ee ie 9 Dy Sg SERNA Ung oes i Peg 2

Fore and middle coxae predominantly yellow, although with varying

amounts of basal dark coloration.

saa at a si a a oe a a eS ae ie ae a Gt a rasa aa ces ree act ae een ay eet ea ak eC Can aaa aA eH 6

Males (males of bicarinata and musae unknown).

Mi i nin ttn uk a tS a a aa a a a ce i a sent a Ve A lle cs ca aa ee a rt ae eal oe eat cae a gc an aa i eae ag ee oe ce 3

Females.

Py ny rae gaa en FOE ey neg hay poy Ey pay re oh omen (OE Cran ang enggy cot nen maap hone en Ep styenes ity Joey [een pesey mr wey mey pa yy ig ea ety ee ST OSE ea rg ea Ty a 4

Postpetiole brownish-red, same color as metasomal segments 3-7;

middie femur yellowish, apical 0.2 of dorsal surface brownish.

me ee ani ne eee pr i Da pt nee townesorum (p. 14)

Postpetiole fuscous or deep brown; middle femur dark brown except

for yellow median dorsal stripe.

icioeidehatiea eh Ge ila a bee eee nigriscapus (Viereck) (p. 10)

Scutellum with strong lateral carina extending its length; lateral

areas of pronotum with transverse rugae present only as traces at

lateral margins, central area smooth and shining; propodeal pattern

as in fig. 24.

Sm FS a tm oa tc tm no bicarinata (p. 23)

Scutellum with lateral carinae distinct only to basal 0.3; lateral

area of pronotum with strong transverse rugae; ventral surface of

scape dark brown or fuscous; propodeal carinae pattern with junc-

ture of areola and petiolar area indicated by a constriction, not

as in fig. 24.

Propodeal carinae weak apically, petiolar area with reticulate

rugae; mesopleural surface with weak granulation; middle femur yel-

lowish, apical 0.2 of dorsal surface brownish; postpetiole

brownish-red, same color as metasomal segments 3-7.

Doce r prone tmere carrer cartes reer ret et ete townesorum (p. 14)

Propodeal carinae uniformly strong, petiolar area with strong

transverse rugae; mesopleural surface with strong granulation;

middle femur dark brown except for yellow median dorsal stripe;

postpetiole fuscous or deep brown.

BUT Trt hr een eT ThA TET oP er eneote nigriscapus (Viereck) (p. 10)

Scape ventrally yellow or reddish-yellow.

Contrib. Amer. Ent. Inst., vol. 22, no. 2, 1984

Scape ventrally dark brown or fuscous, sometimes with lighter

apical area.

Central region of lateral area of pronotum extremely smooth and

polished, with at most faint traces of transverse rugae; upper

posterior lateral corner punctate, granulation absent between punc-

tures.

BaeY ecm Cin nee gill te Meth eet Meaty Yel Sure An aa tgs thy Rig cate Pe On MN Fey WOR SED a eee NERA OC ANON Re gee ag 8

Central region of lateral area of pronotum with at least moderately

distinct rugae; upper posterior lateral corner coarsely rugosopunc-

tate, often with granulation between punctures.

te a sce as ne eee a ee as mee ao coke cen eas lw a ee ee a ere ee ee 9

Under edge of first metasomal segment strongly upcurved in profile

(fig. 3); pronotum broadly triangular (fig. 4).

Acre hoe Ca We Sie eee bee eee eee el catoptron (p. 27)

Under edge not strongly upcurved in profile; pronotum in_ lateral

view long and narrow; propodeal carinae pattern as in fig. 27.

Rg 2s cdlisc i gc < Glace eal a ade een compressa (p. 29)

First metasomal segment brownish-red, petiole with darker areas

bordering suture and basal 0.5 of dorsum.

First metasomal segment with petiole and at least part of post-

petiole fuscous, postpetiole occasionally deep brownish-red.

Mesopleural punctures coarse and shallow, separated by about 0.2

times their diameter or confluent; apex of areola short and squared

(fig. 20); malar space about 0.7 as long as basal mandibular width;

petiolar area with strong transverse rugae, not reticulate or

striate-reticulate; fore wing about 8.0 mm long. :

a wn nnn nnn nen ne e+ genalis (p. 19)

Mesopleural punctures smaller and deeper, separated by 0.3-0.8

times their diameter; apex of areola larger and narrowed (fig. 23);

malar space about 0.6; petiolar area with weak rugae, either trans-

verse or reticulate or combination of both; fore wing 4.7-6.0 mm

long. ;

Fa hit es a llc? un snes isk ad tdi taleri acta gaslacgekans ges citriscapus (p. 22)

12%

14.

15.

16,

Wahl: Nigriscapus Group g

Upper half of lower division of metapleurum coarsely rugosopunc-

tate; fore wing 5.0-6.1 mm long.

plaumanni (p. 16)

Upper half of lower division of metapleurum coarsely punctate; fore

wing 4.6-5.0 mm long.

oe ATG OR OED GEO A Se ee SE RN ee OR OE a eS ee OS es Te eS ee Ge eS Gee ee ee nen Ame ow GS GE mm Sl Sey Goh GES om MED SD GE Gs We oe a oe

Propodeal carinae as in fig. 25: juncture of areola and _ petiolar

area not constricted (or only weakly so in some males), median-

longitudinal carinae roughly parallel and often weak and irregular

in height and outline; mesopleural surface occasionally weakly

granulate.

some specimens of nitida (p. 24)

Propodeal carinae as in fig. 26: juncture of areola and_ petiolar

area constricted, median longitudinal carinae strong and regular in

height and outline; mesopleural surface not granulate.

PUTT MTs Soest a Tee et eS t ekhe ote sense tezcatlipocai (p. 26)

First metasomal segment with at least petiole and sometimes part of

postpetiole black or piceous, postpetiole brownish-red either com-

pletely or just at apex.

First metasomal segment brownish-red, although basal 0.6 can have

traces of darkening.

Basal transverse carina strong, median and lateral longitudinal

carinae weak and hard to distinguish from sculpture of petiolar and

lateral areas; areola-petiolar area reticulate.

arene cde we oe ay dae a eds a) ediettrtat = vim es cag saa oe depressa (p. 21)

All propodeal carinae strong and easily distinguishable from sculp-

ture of adjoining areas; areola-petiolar area with strong trans-

verse rugae.

Propodeal carinae as in fig. 22, second lateral area short and rec-

tangular; upper 0.5 of lateral region of pronotum with rugae

reduced, producing smooth and occasionally punctate surface;

metapleurum coarsely rugosopunctate.

tetragona (p. 20)

10 Contrib... Amer. "Ent. tnst:, vols 22, no. 2, 1984

-~- Propodeal carinae as in figs. 18 and 25, second lateral area more

elongate; upper 0.5 of lateral region of pronotum usually with

QIStCL ae ee ee eee eee eae eee 17

17. Basal outer corner of second lateral area prolonged (fig. 18); in-

tersection of areola and petiolar area constricted, especially in

males; areas between mesopleural punctures moderately granulate.

is oka eo Ke eg ee a catarinensis (p. 17)

-- Basal outer corner of second lateral area not prolonged; intersec-

tion of areola and petiolar area not constricted; areas between

mesopleural punctures not granulate or only weakly.

Sea eS er ee ee ae ee ae eae me eS nitida (p. 24)

18. Propodeal carinae strong basally, becoming weaker apically (fig.

16); petiolar area of propodeum reticulate; mesopleural punctures

coarse and shallow, separated by about 0.2-0.5 their diameter;

ovipositor 1.8 as long as hind femur; fore wing 6.6 mm long.

oe kW tn wee oA leant ws apr us gees dalam leone eaaciulanchas musae (p. 13)

-- Propodeal carinae strong for length of propodeum; petiolar area of

propodeum with transverse rugae; mesopleural punctures moderate,

separated by about 0.3-0.7 times their diameter; ovipositor about

2.1 as long as hind femur; fore wing 4.9 mm long.

Ce eer ea ee eek. ee longicauda (p. 15)

19. Hind femur usually fuscous or dark brown; mesopleurum with spaces

between punctures smooth or weakly granulate; median longitudinal

carinae usually weaker than basal transverse carina; midlines of

metasomal segments 3-7 fuscous.

Pere aerate ap ede ee ite AT ne entry oe me Rg ns me eR en re a EI nitida (p. 24)

-- Hind femur brownish-red; mesopleurum with spaces between punctures

moderately granulate; median longitudinal carinae usually about as

strong as basal transverse carina; metasomal segments 3-7 brownish-

red, midlines sometimes with fuscous hints.

ey nen or ene Ce ae renin Rea nigriscapus (Viereck) (p. 10)

Venturia nigriscapus (Viereck)

(figs. 6,9,11-12)

Idechthis “nigriscapus Viereck,® 1921! Psyche 28:77. Q (MCZC)

Idechthis mimicus Viereck, 1926. Roy. Soc. Canada, Proc. and Trans.

(3) 20°(5)2 1850 CF CONC)

DIAGNOSIS. The medially straight occipital carina and lack of an

apparent specular extension differentiates nigriscapus from the majority

of species; separable from the related musae and longicauda by the

fuscous ventral surface of the scape, black first metasomal segment with

the apex of the postpetiole brownish-red, and strong propodeal carinae.

Most North American specimens have the basal 0.5 of the fore and middle

Wahl: Nigriscapus Group 4

coxae yellow, and the middle and hind femora brownish-red; specimens

with fuscous coxae from the northeastern United States, southeastern

Canada, and Mexico also have the middle and hind femora dark brown. The

related townesorum also has fuscous coxae, but has the middle femur yel-

low and the postpetiole brownish-red.

FEMALE. Structure: 1. Malar space about 0.5 as long as basal

mandibular width. 2. Antenna with 40-44 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with linear depression perpendicular to median ocellus, its length about

0.7 times greatest diameter of lateral ocellus. 5. Pronotum with upper

margin and upper posterior corner of lateral region coarsely rugosopunc-

tate, punctures separated by about 0.3 times their diameter; remainder

of lateral region impunctate, shining, with strong transverse rugae. 6.

Scutellum with anterior half rugosopunctate. 7. Mesopleurum punctate,

punctures in central region immediately below and in front of speculum

moderate-sized, separated by 0.3-0.5 their diameter, on shining,

strongly granulate surface. 8. Metapleurum with lower division

coarsely rugosopunctate. 9. Propodeal carinae (fig. 11) high and

strong and, except for basal transverse carina, slightly irregular in

height and outline; lateral longitudinal carina usually weaker than

median longitudinal carina. Areola impunctate and shining, with few

rugulae intruding laterally; petiolar area with strong transverse rugae.

11. Ovipositor about 1.8 times as long as hind femur. Color (for

typical form in eastern United States): Black, the following pale yel-

low: mandible except for brown of apex and base; palpi; tegula; apical

0.5 of fore and middle coxae and about apical 0.3 of hind coxa;

trochanters except for (usually) dark brown of hind first trochanter;

fore and middle tibiae and femora except for lateral faint brown stripes

on middie femur and tibia. Fore tarsus light brownish-yellow. Middle

tarsus light brown. Scape and pedicel light brown to fuscous, except

for lighter apices. Hind femur brownish-red, occasionally yellowish on

ventral surface and at extreme apex. Hind tibia and hind tarsus dark

brown. Hind tibial spurs yellow. First metasomal segment ranging from

black to piceous, extreme apex of postpetiole sometimes deep brownish-

red. Basal 0.3 and midline to basal 0.8 of second tergum, basal 0.3 and

midline to basal 0.5 of third tergum, and occasionally midlines of

remaining terga, fuscous. Terga otherwise light brownish-red. Length:

9.4-11.3 mm (10.9 mm); fore wing 4.9-6.0 mm (5.5 mm); ovipositor 3.1-3.8

mm (3.5 mm).

MALE. Structure: Similar to female except: 7. Mesopleural punc-

tures moderate to coarse in size, separated by about 0.3 times their

diameter, on strongly granulate surface. 9. All propodeal carinae

strong and high, irregular in height and outline; transverse rugae of

areolar area almost same height and width as carinae; propodeal carinae

pattern as in fig. 12, combined areola-petiolar area narrower, and with

tendency for second lateral area to become squarish in outline. Color:

Similar to female, including same range of variation. Claspers same

brownish-red as abdominal terga. Length: 9.0-11.2 mm; forewing 4.4-5.5

mm.

SPECIMENS EXAMINED. 77 females, 31 males.

12 Contrib. Amer. Ent. Inst., vol. 22, no. 2, 1984

TYPE MATERIAL. Holotype female of nigriscapus, VIRGINIA

(Arlington Co., Glencarlyn, 30 June, on Ceanothus, Nathan Banks) (MCZC

type no. 11187). Four female paratypes, VIRGINIA (same data as

holotype, except collected 8 June and 2 July; Fairfax Co., Great Falls,

25 June, Nathan Banks; Fairfax Co., Falls Church, 11 June, Nathan Banks)

(MCZC). Note: paratypes labelled as such but not formally designated

in original description. Holotype male of mimicus, CANADA (Ontario:

Georgetown, 27 Aug. 1893) (CNCI type no. 1768). | examined the types

of nigriscapus and mimicus in 1982.

DISTRIBUTION. Eastern North America (fig. 30) as far west as Ar-

kansas (the latter fide Carlson, 1979; p. 634), and possibly Mexico

(Oaxaca).

COMMENT. The nature and amount of dark coloration on the fore and

middle coxae varies considerably, ranging from the dark area being

brownish-black and covering the basal 0.5 of the hind coxa and basal 0.7

of the middle coxa, to having the dark area light brown and covering

only the basal 0.3 of both coxae. Intermediates may also be found with

black and brown areas intermixed in the same coxa. The normally black

or fuscous areas of the hind coxa and metasomal terga may be dark

brownish-red. Also, the brownish-red of metasomal terga 2-3 may oc-

casionally be either quite deep or very pale in color.

The difference in the mesopleural punctation between males and

females can be quite striking.

The cephalic sclerites of the final-instar larva are figured in

Finlayson (1975) and Short (1978).

Townes (1945) placed Idechthis mimicus Viereck as a synonym of

nigriscapus. The type of |. mimicus is representative of individuals

of nigriscapus in Michigan, New York, Quebec, and Ontario. This nor-

thern form differs from the more typical nigriscapus in the following:

1) the light area of mandible is brownish-yellow; 2) the basal 0.7-0.9

of the fore and middle coxae is black-fuscous; 3) the middle femur is

brown to fuscous except for the extreme apex and a narrow yellow dorsal

stripe; 4) the middle tibia ventrally and tarsus are dark brown; 5) the

hind femur, except for the extreme base and apex, and tibia are fuscous.

Due to the existence of intermediate specimens from the following

locales, | believe that this northern form is only a regional color

variant: 1) 2 males, St. Charles, Michigan (fig. 30, a); 2) 1 female,

Barry Co., Michigan (fig. 30, b); 3) 1 female, Front Royal, Virginia

(fig. 30, c). V. sokanakiakorum (Viereck) shows a similar geographic

pattern of color variation. Specimens from Ohio, Michigan, and Canada

have the mandible, and fore and middle coxae fuscous-dark brown, in con-

trast to the usual light yellow. In one Michigan locale, specimens are

found exhibiting the extremes of color, as well as an intermediate con-

dition of having the basal 0.7 of the coxa fuscous. Specimens of this

species from a location in Ohio have the males possessing fuscous coxae,

while females have light yellow ones. This is apparently paralled in

reverse by nigriscapus specimens from Ann Arbor, Michigan.

My preference is not to recognize these forms as subspecies; my

views on the formal designation of subspecies are those of Wilson and

Brown (1953).

A specimen from Mexico (Oaxaca: 85.5 km SW of Tuxtepec, 20 Oct.

1962, H. and M. Townes (HMTC)), is tenatively placed in this species.

Wahl: Nigriscapus Group bx

The size, mesopleural punctation, and propodeal carinal pattern are

similar to nigriscapus. The mandible, scape, tegula, coxa, middle first

trochanter, middle tibia, and hind femur are fuscous; metasomal color

pattern is: the: same>as in migriscapus... Except for the coler of the

mandible and trochanter, the pattern is the same as that of the northern

dark form of nigriscapus.

Dates of collection are from late spring to mid-fall. The fol-

lowing are the earliest and latest dates: 25-28 April 1973 at Rapides

Parish, Louisiana; 25-31 April 1976 at Athens, Georgia; 15-18 Oct. 1967

at New Concord, Ohio; 31 Oct. 1950 at Southern Pines, North Carolina.

Label data from specimens in the Canadian National Collection and

the National Museum of Natural History state that this species has been

reared from pupae of the following Noctuidae: Acronicta interrupta

“form elizebetha", Bomalocha sp., Catacola sp., Flathypena scabra (F. )

and Zale sp.

Venturia musae, sp. nov.

(fig. 16)

DIAGNOSIS. Lack of an apparent specular extension separates this

species from the majority of the Nigriscapus Group. It may be

distinguished from nigriscapus, longicauda, and townesorum by the yellow

of the fore and middle coxae, yellow ventral surface of the scape,

coarse mesopleural punctation, weak median and lateral longitudinal

propodeal carinae, and brownish-red first metasomal segment.

FEMALE. Structure: 1. Malar space about 0.5 as long as basal

mandibular width. 2. Antenna with 44-42 flagellomeres. 3. Occipital

carina with short median part straight. 4. Ocellocular distance about

equal to greatest diameter of lateral ocellus; ocellar region coarsely

granulosopunctate with linear depression composed of confluent punctures

perpendicular to median ocellus, its length about 0.8 times greatest

diameter of lateral ocellus. 5. Pronotum with upper margin and upper

posterior corner of lateral region coarsely rugosopunctate, punctures

separated by about 0.3 times their diameter; remainder of lateral region

impunctate, shining, with strong transverse rugae. 6. Scutellum with

anterior half of scutellum coarsely rugosopunctate. 7. Mesopleurum

with coarse shallow punctures, punctures in central region immediately

below and in front of speculum separated by 0.2-0.5 times their

diameter, on shining, strongly granulate surface. 8. Metapleurum with

lower division coarsely rugosopunctate. 9. Propodeal carinae (fig. 16)

as follows: basal transverse carina, and median longitudinal carina

basad of basal transverse carina, strong; other carinae weaker or hard

to distinguish from strongly rugose propodeal surface. Anterior half

of areola smooth with lateral rugulae; remainder of areola and petiolar

area reticulate. 11. Ovipositor about 1.8 times as long as hind femur.

Color: Black, the following pale yellow: mandible except base and apex;

palpi; both scape and pedicel except for brown dorsal surface; tegula;

fore and middle coxae except for dark brown of basal 0.3; hind coxa at

apical 0.3 and median dorsal stripe; trochanters; fore and middle femora

and tibiae except for faint lateral brown stripes; front tarsus except

for brown apical tarsomere. Middle tarsus yellowish-brown. Hind femur

14 Contrib. Amer. Ent: inst.,; vol. 22, no. 2, 1984

and tibia brownish-red with pale yellow ventral area of femur extending

0.5 of way up posterior lateral face. Hind tibial spurs yellowish-

brown. First metasomal segment with petiole and postpetiole light

brownish-red red except for yellowish apex of postpetiole. Basal 0.8

of second tergum and basal 0.2 of third tergum fuscous. Terga otherwise

yellowish-red. Length: 12.9 mm (13.7 mm); fore wing 6.9 mm (6.3 mm);

ovipositor 4.3 mm.

MALE. Unknown.

TYPE MATERIAL. Female holotype, MEXICO (Veracruz: "Penuelas",

with notation "at Laredo (Texas), 10 Dec. 1947, 45956, 48-1029, ex Lep

pupa on banana") USNM). Female paratype, GUATEMALA (22 July 1935,

with notation "at light in banana hold; Phila., PA, no. 27453") CUSNM).

COMMENT. The paratype differs from the holotype in having the hind

coxa completely black except for a thin band of yellow at the extreme

apex, and in the darker color of the rest of the hind leg. Both

specimens were apparently collected by plant quarantine inspectors.

ETYMOLOGY. The specific name is derived from the generic name of

the banana, Musa, on which the unknown lepidopterous host was found.

Venturia townesorum, sp. nov.

(figs. 13-15)

DIAGNOSIS. Lack of an apparent specular extension differentiates

townesorum from the majority of species. The fuscous coxae and ventral

surface of the scape, moderate mesopleural punctation (in the female),

and dark petiole distinguish this species from musae and longicauda.

See the nigriscapus diagnosis to differentiate these two species.

FEMALE. Structure: 1. Malar space about 0.5 as long as basal

mandibular width. 2. Antenna with 41 flagellomeres. 3. Occipital

carina medially arched. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region moderately

granulosopunctate. 5. Pronotum with upper margin and upper posterior

corner of lateral region coarsely rugosopunctate, punctures separated

by about 0.3 times their diameter; remainder of lateral region impunc-

tate, shining, with strong transverse rugae; rugae narrowly connected

in area adjacent to upper posterior corner. 6. Scutellum with anterior

half coarsely rugosopuctate. 7. Mesopleurum punctate, punctures in

central region immediately below and in front of speculum separated by

about 0.5 times their diameter, superimposed on shining, weakly

granulate surface. 8. Metapleurum with lower division coarsely

rugosopunctate. 9. Propodeum (fig. 13) with basal transverse carina

and median longitudinal carina, from base of propodeum to apex of

areola, strong; rest of carinae weak or absent. Areola smooth and im-

punctate, with transverse rugulae; petiolar area reticulate, becoming

transversly striate toward apex. 11. Ovipositor about 1.7 times as

long as hind femur. Color: Black, the following pale yellow: apical

0.2 of fore and middle coxae; trochanters except for brown of second

trochanter of hind leg; fore and middle femora except for dark brown of

apical 0.4 of dorsal surface of middle femur; fore tibia except for

faint brownish stripe. The following yellowish-brown: mandible except

for dark brown of base and apex; palpi; and fore tarsus. Scape,

Wahl: Nigriscapus Group 15

pedicel, tegula, middle tibia, and middle tarsus, brown, except for dor-

sal yellow stripe extending length of tibia. Hind leg deep brownish-red

except for pale yellow band at extreme base of femur. First metasomal

segment with basal 0.7 of petiole piceous, rest of petiole and post-

petiole brownish-red. Basal 0.5 of second tergum and extreme base of

third tergum, dark brownish; terga otherwise brownish-red. Length:

10.6 mm; fore wing 5.2 mm; ovipositor 3.5 mm.

MALE. Structure: Differs from female as follows: 2. Antenna

with 42-45 flagellomeres. 5. Pronotum with transverse lateral rugae

weak. 7. Mesopleurum with punctures in central region immediately

below and in front of speculum large and shallow, separated by about 0.3

times their diameter, on shining, strongly granulate surface. 9.

Propodeal carinae (figs. 14 and 15) strong, irregular in height and

width. Areola and petiolar area with strong transverse rugae. Second

lateral area squarish in outline. Color: As in female, except mandible

completely dark brown; basal 0.8 of first metasomal segment black; basal

0.7-0.8 of second tergum, basal 0.4 and extreme apex of third tergum,

fuscous; midlines of terga 4-7 with fuscous hints. Length: 10.7-12.7

mm; fore wing 5.5-6.3 mm. so

TYPE MATERIAL. Female holotype, MEXICO (San Luis Potosi: nr. El

Naranjo, approximately 64 km W. Nuevo Morelos, 7 Oct. 1962, in oak

forest, H. and M. Townes) (HMTC). Allotype, PANAMA (Chirigui:

Bambito, nr. Volcan, 21 March 1965, S. and W. D. Duckworth) (USNM).

Male paratype, MEXICO (Veracruz: Catemaco, 8 July 1969, R. E. Crutwell,

with notation “emerged 2-VII1-1969 ex A. insulata") (USNM).

COMMENT. Based upon the sexually-based morphological differences

in nigriscapus, | have associated the two male specimens with the

holotype female of townesorum. The males' coarse and close mesopleural

punctation, and the high and irregular propodeal carinae, suggest the

male nigriscapus morphology relative to that of the female.

The shape of the second lateral area differs somewhat between the

two male specimens (figs. 14 and 15). A similar range of variation is

found among nigriscapus males.

The brownish-red coloration of the allotype's metasoma is deeper

than that of the other two specimens. This is probably due to post-

mortem changes.

The paratype is recorded as parasitizing an arctiid, Pareuchaetes

insulata (Walker) (= Ammalo insulata). The host was collected on

Chromolaena odorata (L.) R. M. King and H. Robison, a composite native

to the Neotropics.

ETYMOLOGY. This species is named for Drs. Henry and Marjorie

Townes in recognition of their studies of the Ichneumonidae.

Venturia longicauda, sp. nov.

(fig 261)

DIAGNOSIS. Lack of an apparent specular extension differentiates

longicauda from the majority of species. This species is separable from

musae, nigriscapus, and townesorum by the yellow fore and middle coxae,

yellow ventral surface of the scape, moderate mesopleural punctation,

strong propodeal carinae, and brownish-red first metasomal segment.

16 Contrib. Amer. Ent. Inst., vol. 22, no. 2, 1984

FEMALE. Structure: 1. Malar space about 0.5 as long as basal

mandibular width. 2. Antenna with 43 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with linear depression composed of several confluent punctures perpen-

dicular to median ocellus, its length about 0.3 times greatest diameter

of lateral ocellus. 5. Pronotum with upper margin and upper posterior

corner of lateral region coarsely rugosopunctate, punctures separated

by about 0.3 times their diameter; remainder of lateral region impunc-

tate, shining, with strong transverse rugae; rugae narrowly connected

in a few areas. 6. Scutellum with anterior half coarsely rugosopunc-

tate. 7. Mesopleurum punctate, punctures in central region immediately

below and in front of speculum separated by 0.3-0.7 times their

diameter, on shining, strongly granulate surface. 8. Metapleurum with

lower division coarsely rugosopunctate. 9. Propodeal carinae (fig. 17)

strong and distinct. Areola and petiolar area with strong, narrowly

separated transverse rugae. 11. Ovipositor about 2.1 times as long as

hind femur. Color: Black, the following pale yellow: mandible except

dark brown of base and apex; palpi; both scape and pedicel except for

brown dorsal surfaces; tegula; fore and middle coxae except dark brown

extreme bases; remainder of fore and middle legs except for brownish-

yellow fore tarsus and brown middle tarsus; hind coxa at apical 0.4 and

median dorsal stripe; hind trochanters. Hind tarsus brown. Hind femur

brownish-red with ventral area pale yellow, extending 0.4 of way up

lateral faces. Hind tibia, except for ill-defined pale annulus oc-

cupying median 0.6, and tarsus, brown. Hind tibial spurs light brown.

First metasomal segment with petiole and postpetiole light brownish-red,

apex of postpetiole yellowish. Second terga with 0.5 of segment fol-

lowing basal 0.3, and basal 0.1 of third terga, brownish. Terga other-

wise brownish-red. Length: 9.6 mm; fore wing 4.7 mm; ovipositor 3.5

mm.

MALE. Unknown. .

TYPE MATERIAL. Female holotype, PANAMA (Colon: Margarita,

June 1960, S. Breeland) (HMTC).

ETYMOLOGY... from the Latin longus, long; plus cauda, tail, refering

to the relatively long ovipositor. It is a noun in apposition to the

generic name.

Venturia plaumanni, sp. nov.

(fia. 19)

DIAGNOSIS. Similar in size and coloration to catarinensis, but may

be distinguished by the following: ventral surface of the scape brow-

nish to reddish-yellow; hind first trochanter brown; hind coxa fuscous

except for extreme apex; basal outer corner of the second lateral area

not prolonged; areola-petiolar area not as constricted medially.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 41-43 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region coarsely

granulosopunctate. 5. Pronotum with upper margin and upper posterior

Wahl: Nigriscapus Group V7

corner of lateral region coarsely rugosopunctate, punctures separated

by about 0.3 times their diameter; remainder of lateral region impunc-

tate, shining, with moderate transverse rugae, rugae absent immediately

adjacent to epomia. 6. Scutellum with anterior half coarsely

rugosopunctate. 7. Mesopleurum punctate, punctures in central region

immediately below speculum separated by 0.3-0.8 times their diameter,

on shining, weakly granulate surface; upper anterior region of

mesopleurum immediately adjacent to scrobal groove having transverse

rugae absent or weak and with few punctures, surface granulation absent

so that area appears to be extension of speculum. 8. Metapleurum with

lower division coarsely rugsopunctate. 9. Propodeal carinae (fig. 19)

strong, with longitudinal carinae irregular in height and outline.

Areola fairly smooth with several rugulae; petiolar area reticulate to

striate. 11. Ovipositor about 1.8 times as long as hind femur. Color:

Black, the following pale yellow: mandible except for dark brown of base

and apex; palpi; fore and middie coxae except for dark brown of basal

0.5 of ventral surfaces; extreme apex of hind coxa; remainder of fore

and middle legs except for brownish-yellow of middle tarsi. Ventral

surfaces of scape and pedicel range from dark brown to reddish-yellow.

Tegula brown to brownish-yellow. Hind first trochanter entirely or dor-

sally brownish, second trochanter yellow. Hind femur, tibia, and tar-

sus, dark brown. Hind tibial spurs yellow. First metasomal segment

piceous except for deep brownish-red of petiolar apex and postpetiole.

Second tergum except for apical 0.2, basal 0.5 and midline of third ter-

gum, and midlines of remaining terga fuscous; terga otherwise deep

brownish-red. Length: 9.6-12.9 mm (12.3 mm); fore wing 5.0-6.1 mm (6:

mm); ovipositor 3.1-3.9 mm (3.9 mm).

MALE. Structure: Similar to female, except that propodeum with

carinae much stronger and higher. Color: As in female, except second

through third metasomal terga, plus midlines and upper lateral areas of

fourth through eighth terga, and gonoforcep, fuscous. Length: 9.3 mm;

fore wing 4.7 mm.

TYPE MATERIAL. Female holotype, BRAZIL (Santa Catarina: Nova

Teutonia, Oct. 1970, F. Plaumann) CHMTC). Allotype, BRAZIL (Santa

Catarina: Nova Teutonia, Dec. 1967, F. Plaumann) (HMTC). Four female

paratypes, same data as holotype, except collected: 15 Nov. 1952, 3

Dec. 1952, 22 Dec. 1958, 20 Jan. ‘1960:-(CNCI).

COMMENT. Two female specimens, apparently belonging to this

species, are from Argentina (Tucuman: Burruyacu, V. Padre Monte, 17

Jan.-7 Feb. 1948, R. Golbach (FIML)).

ETYMOLOGY. This species is named after the late Fritz Plaumann, a

professional collector in Santa Catarina, Brazil, who obtained many of

the specimens used in this study.

——_

Venturia catarinensis, sp. nov.

(Tia 1716)

DIAGNOSIS. This species may be distinguished from the similar

plaumanni by the following: tegula, ventral surface of the scape,

apical 0.2 and dorsal median stripe of the hind coxa, hind first

trochanter, yellow; basal outer corner of the second lateral area

18 Contrib. Amer. Ent. bnst.j.vol<.22,.no.. 2/1904

prolonged; areola-petiolar area constricted medially.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 44 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about 0.8 times

greatest diameter of lateral ocellus; ocellar region coarsely

granulosopunctate. 5. Pronotum with upper margin and upper posterior

corner of lateral region coarsely rugosopunctate, punctures separated

by about 0.3 times their diameter; remainder of lateral region impunc-

tate, shining, smooth to moderately striate toward upper and lower

posterior corners. 6. Scutellum with anterior half coarsely

rugosopunctate. 7. Mesopleurum punctate, punctures in central region

immediately below speculum separated by 0.2-0.5 times their diameter,

on shining, weakly granulate surface; upper corner of mesopleurum im-

mediately adjacent to scrobal groove having transverse rugae weak or ab-

sent and with few punctures, surface granulation absent so that area ap-

pears to be extension of speculum. 8. Metapleurum with lower division

coarsely rugosopunctate. 9. Propodeal carinae (fig. 18) strong.

Second lateral area with basal outer corner prolonged anteriorly.

Areola smooth and shining with 2-5 transverse rugulae; entire petiolar

region with strong transverse rugae; areola-petiolar area often con-

stricted medially. 11. Ovipositor about 1.8 times as long as hind

femur. Color: Black, the following pale yellow: mandible except dark

brown of base and apex; palpi; scape except for brown dorsal surface;

pedicel on ventral surface; tegula; fore and middle legs, except for

dark brown of middle tarsus and extreme bases of coxae. Hind

trochanters yellow except for faint brown of anterior face of first

trochanter. Hind femur brownish-red with pale area on middle 0.3 of

ventral face, sometimes extending nearly to base and apex. Hind tibia

and tarsus dark brown to fuscous. Hind tibial spurs yellowish-brown.

Petiole and approximately basal 0.7 of postpetiole piceous. Approx-

imately basal 0.7 of second tergum, basal 0.3 and narrow dorsal

longitudinal strip of third tergum, midlines of remaining terga, and

ovipositor sheath, fuscous. Apical 0.3 of postpetiole and rest of ter-

ga, brownish-red. Length: 10.6-12.6 mm (12.8 mm); fore wing 5.5-6.5

mm (6.5 mm); ovipositor 3.6-4.1 mm (4.1 mm).

MALE. Structure: Differs from the female in the following: 5.

Lateral region of pronotum with rugae much more pronounced, especially

toward upper and lower posterior corners. 9. Propodeum with carinal

patterns similar to female, but carinae stronger and higher. Color: As

in female except brown and fuscous regions deeper in color; hind first

trochanter entirely brown; dorsal fuscous regions of third through eigth

terga extending at least one-third of way down lateral faces of terga.

Gonoforcep fuscous except for brownish-red of basal median edge.

Length: 11.3 mm (11.2 mm); fore wing 5.5 mm (5.4 mm).

TYPE MATERIAL. Female holotype, BRAZIL (Santa Catarina: Nova

Teutonia, 300-500 m, 25 March 1960, F. Plaumann) (HMTC). Allotype,

BRAZIL (Guanabara: Représa Rio Grande, August 1966, M. Alvarenga)

(HMTC). Twenty-one female paratypes, one male paratype, BRAZIL (same

data as for holotype except collected: 18 Dec. 1953, 27 Nov. 1957, 22

Oct. 1958, 22 Nov. 1958, 30 Nov. 1958, 16 Dec. 1958, 23. Dec. 1958, 26

Dec. 1958, 27 Dec. 1959, 4 Jan. 1960, 28 Jan. 1960, 14 Feb. 1960, 5 Mar.

1960, 25 Mar. 1960, 24 Apr. 1960, 5 May 1960, Jan. 1968, one specimen

Wahl: Nigriscapus Group 19

without date) (CNCI, HMTC). Additional specimen, SURINAM (Suriname:

Republiek, 45 km S. Paramaribo, 13-19 Oct. 1963, D. C. Geijskes) (HMTC).

COMMENT. In females, the yellow stripe on the venter of the hind

femur is sometimes quite large, occupying the lower half of the anterior

face. The petiole is sometimes black, without traces of brownish-red.

The postpetiole is occasionally entirely brownish-red. Occasionally the

hind tarsus has yellow bands at the extreme bases of the tarsomeres.

ETYMOLOGY. The specific name derives from the type locality, the

state of Santa Catarina, Brazil.

Venturia genalis, sp. nov.

(hig. 20)

DIAGNOSIS. Differs from depressa, plaumanni, catarinensis, and

tetragona by the reddish-yellow ventral surface of the scape, malar

space 0.7 as long as basal mandibular width, brownish-red petiole,

coarse mesopleural punctation, strong median and lateral longitudinal

carinae, strong transverse rugae of areola-petiolar area, and short and

rectangular second lateral area.

FEMALE. Structure: 1. Malar space about 0.7 as long as basal

mandibular width. 2. Antenna with 49 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with linear depression perpendicular to median ocellus, its length about

0.8 times greatest diameter of lateral ocellus. 5. Pronotum with upper

margin and upper posterior corner of lateral region coarsely rugosopunc-

tate, punctures separated by about 0.3 times their diameter; remainder

of lateral region impunctate, shining, with moderately distinct trans-

verse rugae which are weaker in region adjacent to epomia. 6. Scutel-

lum with anterior half coarsely rugosopunctate. 7. Mesopleurum punc-

tate, punctures in central region immediately below speculum coarse and

shallow, separated by 0.3-0.5 times their diameter, on shining, weakly

granulate surface; upper anterior corner of mesopleurum immediately ad-

jacent to scrobal groove having transverse rugae weak or absent and with

few punctures, surface granulation absent so that area appears to be ex-

tension of speculum. 8. Metapleurum with lower division coarsely

rugosopunctate. 9. Propodeal carinae (fig. 20) high and strong,

slightly irregular in height and outline. Areola impunctate and shining

with rugae intruding laterally; petiolar area with strong transverse

rugae. 11. Ovipositor about 1.9 times as long as hind femur. Color:

Black, the following pale yellow: mandible except for dark brown of apex

and base; palpi; tegula; fore and middle legs except for dark brown of

extreme apices of coxae and light brown of apical tarsomeres; apex and

dorsal stripe of hind coxa; hind trochanters. Scape and pedicel

reddish-yellow except for brown of dorsal surfaces. Hind femur

reddish-brown except for narrow yellow ventral stripe. Hind tibia and

tarsus dark brown, tibia with lighter brown median annulus about 0.6 of

its length. Hind tibial spurs yellow. First metasomal segment

brownish-red except for fuscous tinges along basal 0.5 of petiole.

Basal 0.7 of second tergum, basal 0.3 of third tergum, and midlines of

remaining terga, fuscous; terga otherwise brownish-red. Length: 14.2

20 Contrib. Amer. Ent. inst., vol. 29% no. 2, 1984

mm; fore wing 8.2 mm; ovipositor 4.7 mm.

MALE. Unknown.

TYPE MATERIAL. Female holotype, BRAZIL (Santa Catarina: Nova

Teutonia, 300-500 m, 12 Dec. 1952, F. Plaumann) (HMTC).

COMMENT. This species is unusual in having coarse mesopleural

punctation and a squarish second lateral area present in the female,

features that are usually found only in the males of nigriscapus and

townesorum.

ETYMOLOGY. The specific name is from Latin and refers to the

unusually long malar space.

Venturia tetragona, sp. nov.

(fia. 22)

DIAGNOSIS. Differs from depressa, plaumanni, catarinensis, and

genalis by the yellow ventral surface of the scape, black petiole,

moderate mesopleural punctation, strong median and lateral longitudinal

carinae, transverse rugae of areola-petiolar area, and short and rec-

tangular second lateral area.

FEMALE. Unknown.

MALE. Structure: 1. Malar space about 0.6 as long as basal man-

dibular width. 2. Antenna with 43-45 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about 0.9 times

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with linear depression composed of confluent punctures perpendicular to

median ocellus, its length about equal to greatest diameter of lateral

ocellus. 5. Pronotum with upper margin and upper posterior corner of

lateral region coarsely rugosopunctate, punctures separated by 0.2-0.5

times their diameter; remainder of lateral region shining and polished

with weak transverse rugae present along lateral margins, but absent in

central area, forming smooth and punctate surface. 8. Scutellum with

anterior half coarsely rugosopunctate. 7. Mesopleurum punctate, punc-

tures in central region immediately below speculum moderate, separated

by 0.3-0.5 times their diameter, on shining, weakly granulate surface;

upper anterior region of mesopleurum immediately adjacent to scrobal

groove having transverse rugae weak or absent and with few punctures,

surface granulation absent so that area appears to be extension of

speculum. 8. Metapleurum with lower division coarsely rugospunctate,

with several large rugae extending into center from lower anterior edge.

9. Propodeal carinae (fig. 22) high and strong, slightly irregular in

height and outline. Areola impunctate and shining with rugulae in-

truding laterally; petiolar area with strong transverse rugae. Color:

Black, the following pale yellow: ventral surface of scape and pedicel,

mandible except for dark brown of base and apex; palpi; tegula; fore and

middie legs except for dark brown of coxal base and yellowish-brown of

tarsi (darker on apical segments); apex and dorsal stripe of hind coxa;

hind first trochanter; second hind trochanter varying from yellow to

dark brown. segment, which varies from yellow to dark brown. Hind

femur brown with ventral surface yellow, extending 0.3 of way up lateral

faces. Hind tibia and tarsus dark brown. Hind tibial spurs yellow.

First metasomal segment with basal 0.8 of petiolar portion of tergum

Wahl: Nigriscapus Group 21

piceous; corresponding portion of sternum with more red; rest of segment

brownish-red. Basal 0.6 of second tergum, basal 0.3 and apical 0.1 of

third tergum, and basal 0.3 of fifth through eigth terga, brown. Terga

otherwise light brownish-red. Gonoforcep brownish-red with brown

streaking. Length: 10.9-11.2 mm (10.2 mm); fore wing 6.0 mm (6.1 mm).

TYPE MATERIAL. Male holotype, ARGENTINA (Tucumdn: 11 km.

W. Las Cejas, 8-30 Dec. 1967, L. Stange) (HMTC). Two male paratypes,

ARGENTINA (Buenos Aires: La Plata, 30 Jan. 1966, H. and M. Townes; Mis-

siones: -San-Javier,:t2:Nov. 1977, Cx Porter) CHMTGC, FIML):

COMMENT. The specimen from La Plata has more fuscous markings upon

its metasomal terga than either of the other specimens. This is

probably due to a vagary of preservation.

Extrapolation from nigriscapus and townesorum might lead one to ex-

pect that the second lateral area would be more elongate in the as yet

unknown female of tetragona. The presence of a similarly modified area

in the female specimen of genalis (apparently closely related to this

species) and the lack of extreme sexual dimorphism in species from this

branch of the cladogram makes me believe, however, that the propodeal

carinae pattern is probably the same in both sexes of tetragona.

ETYMOLOGY. The specific name is from Greek and refers to the

squarish second lateral area cf the propodeum.

Venturia depressa, sp. nov.

(fig. 21)

DIAGNOSIS. Related to plaumanni, catarinensis, tetragona, and

genalis (see cladogram), but separable by the yellow ventral surface of

the scape, moderate mesopleural punctation, black petiole, weak median

and lateral longitudinal carinae, and reticulate rugae of the areolar-

petiolar area.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 44 flagellomeres. 3. Occipital

carina medially straight. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region coarsely

granulosopunctate. 5. Pronotum with upper margin and upper posterior

corner of lateral region coarsely rugosopunctate, punctures separated

by about 0.3 times their diameter; remainder of lateral region impunc-

tate, shining, with low transverse rugae which are absent in area adja-

cent to epomia. 6. Scutellum with anterior half coarsely rugosopunc-

tate. 7. Mesopleurum punctate, punctures in central region immediately

below and in front of speculum moderate in size and separated by 0.3-0.7

times their diameter, on shining, smooth surface; upper anterior region

of mesopleurum immediately adjacent to scrobal groove having transverse

rugae weak or absent and with few punctures, surface granulation absent

so that area appears to be extension of speculum. 8. Metapleurum with

lower division coarsely rugosopunctate. 9. Propodeal carinae (fig. 21)

as follows: basal transverse carina strong, somewhat irregular in

height and outline; median and lateral longitudinal carinae barely

distinguishable from rugae of lateral areas and areola-petiolar area.

Areola having basal half with a few transverse rugae; rest of areola and

petiolar area reticulate. 11. Ovipositor about 1.8 times as long as

ce Contrib. Amer. Ent.°inmst.; vol. 22, no. 2, 1984

hind femur. Color: Black, the following pale yellow: mandible except

for brown of base and apex; palpi; ventral and outer lateral surfaces

of scape; tegula; fore and middle legs except for pale brown of extreme

base of coxae, brownish-yellow of fore tarsi and dark brown of middle

tarsi; hind coxa on apical 0.3 and median dorsal stripe; trochanters ex-

cept for brown spot on anterior face of hind first trochanter. Hind

femur medially brownish-red, becoming darker at base and apex; ventral

surface yellowish. Hind tibia and tarsus fuscous. Hind tibial spurs

brownish. First metasomal segment black, becoming dark brownish-red on

postpetiole; apex of postpetiole brownish-red. Second tergum except for

apical 0.2, basal 0.4 and midline of third tergum, and midlines of

fourth through seventh terga, fuscous. Terga otherwise light brownish-

red. Length: 8.5 mm; fore wing 6.5 mm; ovipositor 3.8 mm.

MALE. Unknown.

TYPE MATERIAL. Female holotype, VENEZUELA (Lara: Yacambu,

7 May 1981, 1200 m, H. K. Townes) CHMTC).

ETYMOLOGY. The specific name is from Latin and refers to the weak

nature of most of the propodeal carinae.

Venturia citriscapus, sp. nov.

(figs: 23)

DIAGNOSIS. Related to bicarinata, nitida, and other species (see

cladogram); may be distinguished by the reddish-yellow ventral surface

of the scape, moderately granulate mesopleurum and scutellum, brownish-

red hind femur and petiole, and rugosopunctate lower division of the

metapleurum.

FEMALE. Structure: 1. Malar space about 0.6 times as long as

basal mandibular width. 2. Antenna with 41-45 flagellomeres. 3. Oc-

cipital carina medially arched. 4. Ocellocular distance about equal

to greatest diameter of lateral ocellus; ocellar region coarsely

granulosopunctate with linear depression composed of confluent punctures

perpendicular to median ocellus, its length about 0.7 times greatest

diameter of lateral ocellus. 5. Pronotum with upper margin and upper

posterior corner of lateral region coarsely rugosopunctate, punctures

separated by about 0.3 times their diameter; remainder of lateral area

impunctate, shining, usually with strong transverse rugae in upper

posterior region, those in median area adjacent to epomia either weaker

or without traces of rugae. 6. Scutellum with anterior half coarsely

rugosopunctate. 7. Mesopleurum punctate, punctures in central region

immediately below speculum separated by 0.2-0.8 times their diameter,

on shining, weakly to moderately granulate surface; upper anterior re-

gion of mesopleurum immediately adjacent to scrobal groove having trans-

verse rugae weak or absent and with few punctures, surface granulation

absent so that area appears to be extension of speculum. 8.

Metapleurum with lower division coarsely rugosopunctate. 9. Propodeal

carinae (fig. 23) as follows: basal transverse carina, and portions of

median longitudinal carina apicad to it, strong and regular; remaining

carinae varying in appearance from nearly as distinct as basal trans-

verse carina to resembling rugae of petiolar area, in all cases ir-

regular in height and outline. Areola basally smooth with few to many

Wahl: Nigriscapus Group 23

transverse rugulae, occasionally with longitudinal rugae, area im-

mediately adjacent to carinae depressed relative to center; petiolar

area ranging from either containing all transverse rugae, or being

basally reticulate and apically striate; see remarks for synapomorphy

no. 12. 11. Ovipositor about 1.8 times as long as hind femur. Color:

Black, the following pale yellow: mandible except dark brown of base

and apex; palpi; tegula; fore and middle legs except dark brown of ex-

treme bases of coxae, and yellowish-brown tarsi; hind trochanters.

Scape and pedicel ventrally reddish-yellow. Hind coxa with extreme apex

yellow, and brownish-yellow dorsal median stripe. Hind femur and tibia

brownish-red, femur lighter in color ventrally and tibia occasionally

with lighter areas medially. Hind tibial spurs light brown. Hind tarsi

brown. First metasomal segment brownish-red, petiole with darker areas

bordering suture and basal 0.5 of dorsum. Apical 0.7 of second tergum,

basal 0.4 of third tergum, and midlines of remaining terga, fuscous;

terga otherwise light brownish-red. Length: 9.6-13.0 mm*(11.°9°mm);

fore wing 4.7-6.1 mm (5.9 mm); ovipositor 3.1-4.3 mm (4.0 mm).

MALE. Unknown.

TYPE MATERIAL. Female holotype, BRAZIL (Parana: Campina

Grande, 17 Feb. 1966, H. and M. Townes) (HMTC). Twenty-two female

paratypes, BRAZIL ( Guanabara: Représa Rio Grande, May, June, Oct.,

Dec. 1967, Nov. 1968, Oct.-Nov. 1970, M. Alvarenga; Parana: same data

as holotype, except collected 15 Feb. 1966 and 19 Feb. 1966, H. and M.

Townes; Santa Catarina: Nova Teutonia, 300-500 m, 15 May 1939, 7 Dec.

1952; °25 “Novis: 14953, - Weadanert9S6.014-: Fett: 1956.19) den 999, .F:

Plaumann; Sao Paulo: Teodoro Sampaio, Aug. 1973, F. M. Oliveira) (CNCI,

HMTC).

COMMENT. In some specimens, the median longitudinal carinae weakly

diverge at their midpoints.

The following variations in color have been observed: the fore and

middle coxae may have the ventral 0.5 light brown; the ventral yellow

regions of the hind coxa may be very reduced; the hind first trochanter

may be brownish-yellow; the basal 0.5-0.8 of the petiole may be deep

reddish-black; the fuscous midlines of terga 4-7 may be considerably ex-

tended laterally.

Differences in the state of preservation seem to influence the

color of specimens: those appearing oily or greasy will show darker

brownish-red areas on the metasoma. The color of the petiole is espe-

cially sensitive to this influence.

ETYMOLOGY. The specific name is formed from the Latin words,

citrus, orange, plus scapus, referring to the antennal scape. It is a

noun in apposition.

Venturia bicarinata, sp. nov.

(fig. 24)

DIAGNOSIS. Related to citriscapus, nitida, tezcatlipocai, and

other species (see cladogram); the reddish-yellow ventral surface of the

scape, weakly granulate mesopleural surface, scutellum with irregular

lateral carinae extending its length, punctate lower division of the

metapleurum, largely black fore and middle coxae, and pattern of the

24 Contrib. Amers Ent. inst.,; vol..22, no. 2, T1964

propodeal carinae, distinguish this species from its relatives.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 44 flagellomeres. 3. Occipital

carina medially arched. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; cellar region coarsely

granulosopunctate with linear depression composed of confluent punctures

perpendicular to median ocellus, its length about equal to greatest

diameter of lateral ocellus. 5. Pronotum with upper margin and upper

posterior corner of lateral region coarsely rugosopunctate, punctures

separated by 0.3-0.5 times their diameter; remainder of lateral region

shining and impunctate, without transverse rugae except for traces at

edges. 6. Scutellum with anterior half coarsely rugosopunctate, and

irregular lateral carinae extending its length. 7. Mesopleura punc-

tate, punctures in central region immediately speculum moderate in size

and separated by 0.5-0.7 times their diameter, on shining, weakly

granulate surface; upper anterior of mesopleura immediately adjacent to

scrobal groove having transverse rugae weak of absent and with few punc-

tures, surface granulation absent so that area appears to be extension

of speculum. 8. Metapleura with lower division coarsely punctate. 9.

Propodeal carinae (fig. 24) with basal transverse carina somewhat

stronger than others; areola and petiolar area with strong transverse

rugae; see remarks for synapomorphy no. 12. 11. Ovipositor about 2.1

times as long as hind femur. Color: Black, the following pale yellow:

mandible except for brown of base and apex; palpi; tegula; trochanters,

apical 0.3 of fore and middle coxae; femora and tibiae of fore and

middle legs; apical 0.2 of hind coxae. Fore and middle tarsi brownish-

yellow. Ventral surfaces of scape and pedicel reddish-yellow. Hind

femur and tibia brownish-red except for ventral yellow stripe of femur

and ill-defined pale sub-basal annulus of tibia. (Hind tarsi missing).

Hind tibial spurs brownish-yellow. First metasomal segment black; apex

of postpetiole deep brownish-red. Apical 0.7 of second tergum, basal

0.2 and midline of third tergum, and midlines of fourth through seventh

terga, fuscous. Terga otherwise deep brownish-red. Length: 10.6 mm;

fore wing 5.7 mm; ovipositor 3.8 mm.

MALE. Unknown.

TYPE MATERIAL. Female holotype, PARAGUAY (Pirapo: 29 Dec. 1971,

L. Pena) CHMTC).

COMMENT. Color of terga probably deeper than in life, due to post-

mortem changes.

ETYMOLOGY. The specific name is from Latin and refers to the

scutellar carinae.

Venturia nitida, sp. nov.

(figs 25)

DIAGNOSIS. Related to citriscapus, bicarinata, tezcatlipocai, and

other species (see cladogram); it may be distinguished by the (usually)

striate lateral region of the pronotum, weakly (Cif at all) granulate

mesopleurum, smooth surface of scutellum, coarsely punctate lower divi-

sion of the metapleurum, fuscous hind femur, and shape of the first

metasomal segment.

Wahl: Nigriscapus Group 25

FEMALE. Structure: 1. Malar space about 0.5 as long as basal

mandibular width. 2. Antenna with 39-42 flagellomeres. 3. Occipital

carina medially arched. 4. Ocellocular distance about equal to

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with impunctate shining groove perpendicular to median ocellius, its

length about equal to greatest diameter of lateral ocellus. 5.

Pronotum with upper margin and upper posterior corner of lateral region

coarsely rugosopunctate, punctures separated by about 0.3 times their

diameter; remainder of lateral region with moderate to weak transverse

rugae, some populations with rugae absent adjacent to upper posterior

corner. 6. Scutellum with anterior half coarsely punctate, surface

granulation absent. 7. Mesopleurum punctate, punctures in central re-

gion immediately below speculum separated by 0.3-0.5 times their

diameter or occasionally confluent, on shining, weakly granulate or

smooth surface; upper anterior region of mesopleurum immediately adja-

cent to scrobal groove having transverse rugae weak or absent and with

few punctures, surface granulation absent so that area appears to be ex-

tension of speculum. 8. Metapleurum with upper 0.6 of lower division

coarsely and deeply punctate, punctures separated by about 0.3 times

their diameter; lower 0.4 rugosopunctate. 9. Propodeal carinae (fig.

25) as follows: basal transverse carina strong and regular in height

and outline; other carinae (excluding pleural carina) weaker and ir-

regular in height and outline, but always distinguishable from other

propodeal sculpturing; in some populations, median and _ lateral

longitudinal carinae absent in parts; see remarks for synapomorphy no.

12. Areola impunctate with some rugulae extending from lateral edges;

petiolar area reticulate, sometimes apically with strong transverse

rugae. 11. Ovipositor about 1.7 times as long as hind femur. Color:

Black, the following pale yellow: mandible except for brown of base and

apex; palpi; tegula; fore and middle legs except for dark brown of ex-

treme bases of coxae, and brownish-yellow tarsi. Scape and pedicel

fuscous, in some cases becoming reddish- yellow or yellow at apices or

even lateral and outer ventral surfaces. Hind first trochanter yellow,

sometimes with brown on anterior and dorsal surfaces. Hind femur,

tibia, and tarsus usually dark brown except for yellow on extreme base

of femur; some populations with femur and tibia light brownish-red, or

with femur light brownish-red and tibia dark brown. Hind tibial spur

yellow. First metasomal segment black, postpetiole with apical 0.3

brownish-red. Basal 0.8 of second tergum, basal 0.3 and midline of

third tergum, and midlines of remaining terga fuscous; terga otherwise

light brownish-red. Length: 9.0-9.4 mm (8.2 mm); fore wing 4.4-5.0 mm

(4.4 mm); ovipositor 2.7-3.7 mm (2.8 mm).

MALE. Structure: Similar to female except that relative strengths

of propodeal carinae stronger and more well developed; areola and

petiolar area with well-developed transverse rugae. Color: Similar to

female and subject to same variations in color, except that all of

second tergum (except extreme lateral corners) and third tergum, plus

midlines and portions of dorsal lateral areas of remaining terga,

fuscous. Gonoforcep brownish-red. Length: 9.3-10.7 mm (8.7 mm); fore

wing 4.1-5.2. mm (4.1 mm).

TYPE MATERIAL. Female holotype, UNITED STATES (Arizona: Cochise

Co., Parker Canyon Lake, 22 Aug. 1974, H. and M. Townes) (HMTC). AlI-

26 Contrib. Amer. Ent.ohysts, vol. -22y no. 2, 1994

lotype, same data as holotype (HMTC). Three female and one male

paratypes, same collection data as holotype except collected 20 Aug.

1974 and 22 Aug. 1974, and one male paratype collected from Portal

(Cochise Co.), 13 Aug. 1974 (HMTC). Other specimens from ARGENTINA

(Tucumdn: San Pedro de Calalao, Jan. 1968, L. Stange); BRAZIL (Pernam-

buco: Caruaru, April-June 1972, 900 m, J. Lima; Santa Catarina: Nova

Feutonia, 300-500 -m;) 2% Mar.:-1947,°21 April 1952, 19:Febe 1954, °F.

Plaumann); COLOMBIA (Valle del Cauca: Cali, Oct. 1971, M. J. West-

Eberhard); COSTA RICA (Guanacaste: Santa Rosa Park, 3 July 1977, 12

Septic 1977 PAyeSept.. 19777 6 Oet..41977,.11 Jume 1978; 28 July:1978 ,- "dry

hill", D. H. Janzen); MEXICO (Michoacan: 9 miles NW Hidalgo, 20 August

1962, 7100 ft., R. H. and E. M. Painter); VENEZUELA (Zulia: Tucuco,

23 April 1981, H. K. Townes) (HMTC).

DISTRIBUTION. Southern Arizona to Brazil and northern Argentina.

COMMENT. In males, the lower division of the metapleurum can be

weakly rugosopunctate.

ETYMOLOGY. The specific name is from Latin and refers to the

weakly granulate surface of the mesopleurum.

Venturia tezcatlipocai, sp. nov.

(fig. 26)

DIAGNOSIS. Differentiated from the related bicarinata, nitida,

citriscapus, and other species (see cladogram) by the reddish-yellow

ventral surface of the scape, smooth surfaces of the mesopleurum and

scutellum, striate lateral region of the pronotum, coarsely punctate

lower division of the metapleurum, fuscous hind femur, and propodeal

pattern.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 39 flagellomeres. 3. Occipital

carina medially arched. 4. Ocellocular distance about 1.2 times

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with shining linear depression perpendicular to median ocellus, its

length about 0.8 times greatest diameter of lateral ocellus. 5.

Pronotum with upper margin and upper posterior corner of lateral

coarsely rugosopunctate, punctures separated by 0.3-0.5 times their

diameter; upper posterior corner with spaces between punctures without

granulation. Remainder of lateral area shining and impunctate with low

transverse rugae; rugae in area adjacent to epomia and upper posterior

corner fused, area appearing smooth and shining. 6. Scutellum with an-

terior half coarsely punctate, surface granulation absent. /.

Mesopleurum punctate, punctures in central region immediately below

speculum separated by 0.5-0.7 times their diameter, on smooth and

shining surface; upper anterior region of mesopleurum immediately adja-

cent to scrobal groove having transverse rugae weak or absent and with

few punctures, surface granulation absent so that area appears to be ex-

tension of speculum. 8. Metapleurum with upper 0.6 of lower division

coarsely punctate, punctures separated by about 0.3 times their

diameter; lower 0.4 coarsely rugosopunctate. 9. Propodeal carinae

(fig. 26) strong and regular except for weak lateral longitudinal

carina. Areola impunctate and shining, with few rugae intruding

Wahl: Nigriscapus Group ar

laterally; petiolar area with strong transverse rugae except in median

strip in basal 0.3, which is depressed and shining. 11. Ovipositor

about 1.9 times as long as hind femur. Color: Black, the following

pale yellow: palpi; mandible except for dark brown of base and apex;

tegula; fore and middle legs except for dark brown of apical 0.2 of cox-

ae and yellowish-brown of apical tarsomeres; apical 0.3 of hind coxa;

hind trochanters except for brown of dorsal and anterior faces of first.

Scape and pedicel ventrally reddish-yellow with brown streaks. Hind

femur, tibia, and tarsi, brown. Hind tibial spurs yellow. First

metasomal segment black, postpetiole with apical 0.5 deep brownish-red.

Second tergum except apical 0.1, third tergum except lower lateral re-

gions, and midlines of fourth through eight terga, fuscous; terga other-

wise light brownish-red. Length: 9.3 mm; fore wing 4.7 mm; ovipositor

3.1.mm.

TYPE MATERIAL. Female holotype, MEXICO (San Luis Potosi: Tamazun-

chale, 23 Nov. 1946, E. S. Ross) (CASC).

ETYMOLOGY. This species is named for the Aztec god Tezcatlipoca;

the translation "Smoking Mirror" is an oblique reference to the polished

black mesopleurum.

Venturia catoptron, sp. nov.

(fig. 3,4,23)

DIAGNOSIS. Related to compressa and other species (see cladogram),

it may be recognized by the yellow ventral surface of the scape, ab-

sence of striae on the lateral region of the pronotum, smooth

mesopleural surface, coarsely punctate lower division of the

metapleurum, fuscous hind femur, and convex sternum of the first

metasomal segment.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. Antenna with 41 flagellomeres. 3. Occipital

carina medially arched. 4. Ocellocular distance about 0.7 times

greatest diameter of lateral ocellus; ocellar region granulosopunctate

with linear depression composed of confluent punctures perpendicular to

median ocellus, its length about 0.7 times greatest diameter of lateral

ocellus. 5. Pronotum with upper margin and upper posterior corner of

lateral region punctate, punctures separated by about 0.3 times their

diameter; upper posterior corner having spaces between punctures without

granulation; remainder of lateral area shining and impunctate, without

transverse rugae, except for weak traces in lower posterior corner. 6.

Scutellum with anterior half punctate, surface granulation absent. 7.

Mesopleurum punctate, punctures in central region immediately below

speculum separated by 0.3-0.5 times their diameter, occasionally con-

fluent or separated by their diameter, on extremely smooth and shining

surface with little or no granulation; upper anterior region of

mesopleurum immediately adjacent to scrobal groove having transverse

rugae weak or absent and with few punctures, surface granulation absent

so that area appears to be extension of speculum. 8. Metapleurum with

upper 0.7 of lower division coarsely punctate, punctures separated by

about 0.5 times their diameter; lower 0.3 rugosopunctate. 9. Propodeal

carinae (fig. 28) as follows: basal transverse carina always present,

28 Contrib. Amer. Ent.-inst., vol. 22, no. 2, 1984

somewhat irregular in height and outline; other carinae weaker although

always present and distinguishable from other propodeal sculpture.

Areola smooth and without rugulae and with deep punctures present along

lateral margins; petiolar area with transverse rugae, although sometimes

reticulate in basal half; see remarks for synapomorphy no. 12. 10.

Sternum of first metasomal segment convex in profile (fig. 3). 11.

Ovipositor about 1.9 times as long as hind femur. Color: Black, the

following pale yellow: mandible except for brown of base and apex; pal-

pi; ventral surfaces of scape and pedicel; tegula; fore and middle coxae

except for pale brown of extreme base; hind coxa at apex and median dor-

sal stripe; trochanters except for dark brown of hind first trochanter.

Fore and middle legs with femora, tibiae, tibial spurs, and tarsi (ex-

cept brown of basal 0.3 of ventral surface of middle femur and apical

segment of middle tarsus), yellow with brownish tinge. Hind femur,

tibia, and tarsus, fuscous. Hind tibial spurs yellow. First metasomal

segment black except for brownish-red at apical 0.2 of postpetiole.

Second tergum except for apical 0.15, basal 0.3 and midline of third

tergum, and midlines and dorsal 0.3 of lateral surfaces of third through

seventh terga, fuscous. Terga otherwise deep brownish-red. Length:

9.8-10.6 mm (10.4 mm); fore wing 4.4-4.7 mm (4.6 mm); ovipositor 3.3-3.6

mm (3.3 mm).

MALE. Structure: Similar to female except all propodeal carinae

of equal height and strength; combined areola-petiolar area narrower

than in female. Petiolar area with strong transverse rugae. Color: As

in female, except all of second tergum, and dorsal 0.5 of lateral sur-

faces of third through seventh terga, fuscous. Gonoforcep fuscous.

Length: 9.3-9.8 mm (10.1 mm); fore wing 4.3 mm (4.6 mm).

TYPE MATERIAL. Female holotype, BOLIVIA (Santa Cruz: Robore, Obi-

quitos, Oct. 1959, 300 m, F. H. Walz) (HMTC). Allotype, same collection

data as for holotype, (HMTC). Four female and two male paratypes, same

collection data as for holotype except one collected in November (HMTC).

COMMENT. The amount of fuscous coloration on the metasoma varies,

apparently due to the state of preservation of the specimens. Several

of the specimens in the series are quite oily in appearance and this

seems to increase the amount of dark coloration in otherwise brownish-

red portions of the terga.

A specimen with the same collection data as the holotype is

provisionally placed in this species. It differs in the reddish-yellow

color of the ventral surface of the scape, mostly yellow hind first

trochanter, weakly wrinkled mesopleural surface, narrower areola-

petiolar region, and less convex sternum of the first metasomal segment.

ETYMOLOGY. The specific name is a noun in apposition; it is from

the Greek for mirror, in reference to the polished surface of the

lateral area of the pronotum.

Venturia compressa, sp. nov.

(Fige S27)

DIAGNOSIS. Related to catoptron and other species (see cladogram),

it may be recognized by the yellow ventral surface of the scape, absence

of striae on the lateral region of the pronotum, smooth mesopleural sur-

Wahl: Nigriscapus Group 29

face, long and narrow lateral region of the pronotum (fig. 5), coarsely

punctate lower division of the metapleurum, fuscous hind femur, and

propodeal pattern.

FEMALE. Structure: 1. Malar space about 0.6 as long as basal

mandibular width. 2. (Antennae with flagella incomplete.) 3. Oc-

cipital carina medially convex. 4. Ocellocular distance about 0.8

greatest diameter of lateral ocellus; ocellar region coarsely

granulospunctate with linear depression composed of confluent punctures

perpendicular to median ocellus, its length about 0.6 greatest diameter

of lateral ocellus. 5. Pronotum with upper margin and upper posterior

corner of lateral region punctate, punctures separated by about 0.3

times their diameter; upper posterior corner with area between punctures

without granulation; remainder of lateral region shining and impunctate,

without transverse rugae except for traces at edges; pronotum in lateral

view long and narrow (fig. 5), as opposed to broader triangular condi-

tion (fig. 4). 6. Scutellum with anterior half coarsely punctate, sur-

face without granulation. 7. Mesopleurum punctate, punctures in cen-

tral region immediately below speculum moderate in size and separated

by 0.3-0.5 times their diameter, on shining, smooth surface; upper an-

terior region of mesopleurum immediately adjacent to scrobal groove

having transverse rugae weak or absent and with few punctures, surface

granulation absent so that area appears to be extension of speculum. 8.

Metapleurum with upper 0.3 of lower division coarsely punctate, lower

0.7 rugosopunctate. 9. Propodeal carinae (fig. 27) with basal trans-

verse carina strong and regular, remaining carinae somewhat weaker, with

portion of lateral longitudinal carina forming outer side of second

lateral area almost absent. Areola shining and impunctate, with a few

small lateral rugae; petiolar area with transverse rugae. 11.

Ovipositor about 1.8 times as long as hind femur (broken in paratype).

Color: Black, the following pale yellow: mandible except dark brown

of base and apex; palpi; tegula; fore and middle legs except for brown

of extreme bases of coxae, and brownish-yellow tarsi; extreme apex and

dorsal median stripe of hind coxa; distal segment of hind coxa; hind

second trochanter. Ventral surfaces of scape and pedicel reddish-

yellow. Hind first trochanter dark brown. Hind femur entirely dark

brown , or with anterior face dark brown, ventral surface yellowish,

becoming darker yellowish-brown on posterior face. Hind tibia and tar-

sus dark brown. Hind tibial spurs yellowish-brown. First metasomal

segment black, dorsal apex and apical 0.5 of lateral faces of post-

petiole brownish-red. Apical 0.7-0.9 of second tergum, midlines of

third through seventh terga, fuscous; terga otherwise light brownish-

red. Length: 8.5 mm (9.2 mm); fore wing 4.5 mm (5.0 mm); ovipositor

(3.3 ram),

MALE. Unknown.

TYPE MATERIAL. Female holotype, PERU (Cuzco: Quincemil,

nr. Marcapata, 10-15 Nov. 1962, 750 m, L. Pena) (HMTC). Female

paratype, BRAZIL (Rondonia: Vilhena, Nov. 1973, M. Alvarenga) (HMTC).

ETYMOLOGY. The specific name derives from Latin and refers to the

compact shape of the lateral area of the pronotum.

30 Contrib. Amery-Enticinsr., wol.22, no. 2, 1984

LITERA PURE CITED

Arnett, R. H., Jr., and G. A. Samuelson. 1969. Directory of Coleoptera

Collections of North America (Canada through Panama). Cushing-

Malloy, Ann Arbor, Michigan. 123 pp.

Carlson, R. W. 1979. Family Ichneumonidae. In K. V. Krombein, P. D.

Hurd, D. R. Smith, and B. D. Burks (eds.), Catalog of Hymenoptera

eee ee

Washington, D.C. 1198 pp.

Finlayson, T. 1975. The cephalic structures and spiracles of final-

instar larvae of the subfamily Campopleginae, tribe Campoplegini

(Hymenoptera: Ichneumonidae). Mem. Entomol. Soc. Canada 94: 1-137.

Gupta, V. K. and S. Maheshwary. 1977. Ichneumologia orientalis, 1V:

the tribe Porizontini. Oriental Insects Monograph 4: 1-203.

Harris, R. A. 1979. A glossary of surface sculpturing. Occas. Papers

Entomol. 28: 1-31. State of California, Dept. of Food and

Agriculture, Div. of Plant Industry, Sacramento.

Hennig, W. 1966. Phylogenetic Systematics. University of Illinois

Press, Urbana, Illinois. 263 pp.

Michener, C. D. 1944. Comparative external morphology, phylogeny, and

a classification of the bees. Bull. American Mus. Nat. Hist. 82:

1317326.

Rosen, D. E. 1978. Vicariant patterns and historical explanations in

biogeography... Syst. Zook. 27: 159-188.

Short, J. R. T. 1978. The final larval instars of the Ichneumonidae.

Mem. American Entomol. Inst. 25: 1-508.

Townes, H. 1944-45. A catalogue and reclassification of the Nearctic

Ichneumonidae. Mem. American Entomol. Soc. 11: 1-925.

. 1969. Genera of Ichneumonidae, part 1. Mem. American En-

tomol.. inst..i14: 1-300.

. 1970. Genera of Ichneumonidae, part 3. Mem. American En-

Comal... dae. 61361807;

Watrous, L. E. and Q. D. Wheeler. 1981. The out-group comparison

methed of character analysis. Syst::Zoo). :30: 1-11.

Wiley, E. O. 1981. Phylogenetics. Wiley-Interscience, New York. 439

Pp.

Wilson, E. O. and W. L. Brown, Jr. 1953. The subspecies concept and

its. faxpnoemic application: Syst. Zool....2: 97-111.

Nigriscapus Group

Wahl:

Kas

= i casociaeemmene > sek

SN “Aou ‘ds sisusuiueyeo “A jo snqiqeH “1 eunbi4g

ee...

a2 Contrib. Amer: Ents ist.) vel. 22, no. 2, 1964

®@

ae @

@ @

9 10

Figures 2-10. 2, Lateral view of first metasomal segment of Venturia

sp. 3, same of V. catoptron sp. nov. 4, Lateral view of mesosoma of

V. catoptron sp. nov. 5, Same of V. compressa sp. nov. 6, Dorsal view

of head of V. nigriscapus (Vier.). 7-8, Dorsal views of heads of Ven-

turia spp. 9-10, Dorsal view of first metasomal segments of V.

nigriscapus and Venturia sp.

Wahl: Nigriscapus Group

1112 Ve

Propodea of Nigriscapus Group (dorsal view).

13=157 V . townesorum sp. now,

female holotype, allotype, male paratype. 16, V. musae sp. nov. 17,

Vv. longicauda sp. nov. 18, V. catarinensis sp. nov. 19, V. plaumanni

sp. nov.

Figures 11-19.

nigriscapus (Vier.), female and male.

34 Contrib. Amer: Ent. inst.;. vol. 22, no. 2, 1964

Figures 20-28. Propodea of Nigriscapus Group (dorsal view). 20, V.

genalis sp. nov. 21, V. depressa sp. nov. 22, V. tetragona sp. nov.

23, V. citriscapus sp. nov. 24, V. bicarinata sp. nov. 2byi Ve netida

sprounow: ~ 26, Vi tezoathipocel sp. nov. 2/7, V. compressa sp. nov. 28,

V. catoptron sp. nov.

Wahl: Nigriscapus Group Be

SA @ @ @ © ® & @ © ® ®

CA ® oS & @ ©

NA S e

tow nig mus lon pla cata dep tet gen cit bic nit tez cato com

2\/8 1 -

13 10

9 5

‘ 13 11 29

2 6

/ t j a os

H ar, Vet” b

ey AE

¥, r on

é es

ai. a¢5

Figures 29, 30. 29, Cladogram of Nigriscapus Group (NA = North America,

CA = Central America, SA = South America). 30, Distribution of ce

nigriscapus (Vier.) [triangles indicate localities where northern dark

form is present; see text for explanation of letters].

w 7

BENS

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Pan

PAPAIN Oi

Contributions

of the

American Entomological Institute

Volume 22, Number 3, 1986

THE TABANIDAE OF PANAMA

G. B. Fairchild

Florida State Collection of Arthropods

Gainesville, Florida 32602

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ean

THE TABANIDAE OF PANAMA

G.B. Fairchild

Research Associate

Florida State Collection of Arthropods

P.O. Box 1269

Gainesville, FL 32602

INTRODUCTION

In 1942 I published the last of 10 papers on the Tabanidae of the Republic

of Panama, which summarized what was then known of the fauna (Fairchild

1942). In that paper I listed 25 genera and subgenera and 91 species, subspecies

and varieties as having been collected within the Republic. During the sub-

sequent years until the end of 1969, I continued to collect Tabanidae in Panama

and published short papers describing new species. Upon retirement in 1969, I

eontinued work on Neotropical Tabanidae, but published only one paper specif-

ically on Panamanian species, though including occasional such species in re-

visionary publications.

The present effort is designed to be a synopsis of the family as it occurs in

Panama, including keys to genera and species, geographic distribution, hosts,

biting habits, seasonal distribution and everything available bearing on the habits

and ecology. The discerning reader will quickly observe that little except the

taxonomy is included, largely because little else is available in published form.

Only 3 publications are concerned with the early stages; 1 has no descriptive

eontent, 1 treats of a single species, the last treats all early stages of known

Neotropical species, 47 species, of which 21 are Panamanian (Goodwin and

Murdoch 1974). There is one limited publication on seasonal distribution,

another on arboreal species, and a dozen or so seattered observations on biting

and host preference inserted in largely taxonomic publications.

It is hoped that this review will furnish a taxonomic backstop for anyone

wishing to pursue studies on the biology and ethology of these large and con-

Spicuous flies. Such aspects as mating habits, oviposition and larval habitats, lar-

val food, occurrence of autogeny, host seeking behavior, including choice of host

and preferred sites on hosts for feeding, length of various stages, range of

flight, and preferred habitats of adults all suggest themselves as virgin fields for

study.

ACKNOWLEDGEMENTS

I am gratefully indebted to the officers and staff of the Gorgas Memorial

Laboratory of the Gorgas Memorial Institute of Tropical and Preventive Med-

icine and Public Health, Ine., who individually and collectively furnished me with

financial and other material support, advice, criticism and encouragement for 32

years. My thanks are also due to the many persons who have furnished the very

‘large numbers of specimens which have made this work possible. Of special

importance in this respect have been my former colleagues Harold Trapido and

Florida Agricultural Experiment Station, Journal Series no. 6654.

2 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Pedro Galindo, whose field crews collecting mosquitoes in connection with their

long term studies of Sylvan Yellow Fever have contributed most of what is

known of the arboreal Tabanidae in Panama. Much of the material reported as

collected at light or in horse baited mosquito traps was furnished by Marvin

Keenan, the long time civilian sanitary engineer with the Environmental Health

office of the U. S. Army on the Isthmus, and the successive Army entomologists

working at that facility, the late Col. Stanley J. Carpentor, Maj. Gordon Field,

Col. F. S. Blanton and Maj. Wallace Murdoch. During the early 1960's the Of-

fice of Interoceanic Canal Surveys made extensive collections, using Malaise

traps and other methods, on several of the routes proposed as alternative sea-

level canals. Gorgas Memorial Laboratory was contracted by OICS to determine

the organisms from mammals to viruses, likely to be of public health interest.

Much of the personnel on this survey were Army entomologists under the lead-

ership of Maj. Bruce Eldridge. He and Lt. David G. Young collected long series

of Tabanidae from areas never before sampled. During the early years of my

stay in Panama, the late Prof. Joseph Bequaert provided indispensable taxonomic

guidance, a position later (and still) occupied by Drs. C. B. Philip and L. L.

Pechuman. Finally my colleague Richard Wilkerson has provided indispensable

logistie support in the final stages of preparation of this review.

This work supported in part by NSF grant BSR 8211808. Florida Agricultural

Experiment Station Journal Series No.6654.

EXTENT OF COLLECTIONS AND COLLECTING METHODS

The material on which this study is based was, for the most part, collected

in a somewhat haphazard manner over a period of about 30 years. My predeces-

sor at Gorgas Memorial Laboratory, L. H. Dunn, did some collecting and his col-

lection, determined by J. S. Hine and J. Bequaert, formed the starting point for

my own collecting. From 1938 to 1942 most specimens were netted by myself

in the vicinity of the Canal Zone, or on short excursions to accessible sites such

as El Valle de Anton, Cerro Campana, and Boquete, Chiriqui. Only 1 systematic

collection was made during this period (Fairchild 1942), consisting of 26 fort-

nightly collections of 2 days each. Collections were made from bait animals,

horses and cattle, and totalled 11,997 specimens belonging to 35 species. The

collecting site was in an area of second-growth forest adjacent to swamps bord-

ering the Chagres River, and is the only large collection available from this

lowland habitat. During this period I also examined the Hine Collection at Ohio

State University, and the collections at U. S. National Museum and Museum of

Comparative Zoology at Harvard, from which I abstracted reeords of Panama-

nian species. This early work was reported in a series of papers, summarized in

a resume (Fairchild 1943) which listed 89 species as occurring in Panama.

With the commencement of World War II, intensive work on Tabanidae

closed. Time and transportation for collecting became unavailable, so that what

little collecting was done was incidental to other work. Various colleagues, both

in and out of the Army, continued to secure specimens in areas that had not

been sampled before, mostly in connection with malaria mosquito surveys at

scattered Army outposts.

During the war I had become associated with Marshall Hertig, and we had

embarked on a study of the Phlebotomine sand flies. This work continued into

the 1960's and the Phlebotomines being of more public health significance than

Tabanidae, occupied much of my time, although enabling incidental collections

of Tabanids to be made as opportunity offered. An outbreak of Sylvan Yellow

Fever in late 1949 turned the attention of my colleagues, Harold Trapido and

G. B. Fairchild: Tabanidae of Panama i

Pedro Galindo, towards study of possible arboreal mosquito vectors of this dis-

ease. To study these mosquitoes, platforms were built in the forest canopy at

heights of up to 90 feet, and collectors stationed on them. The collectors were

instructed to take all insects attracted to them, and in this way a remarkable

assortment of arboreal Tabanidae was secured. Collections were made at 9

localities: Progresso, Chiriqui Prov., elevation 300-400 feet, June-December,

1950; Santa Fe, Veraguas Prov. elevation 2000 feet, May-August, 1950; Cerro

Campana, Panama Prov., elevation 400, 1200, and 2100 feet, February 1949-Jan-

uary 1950; Almirante, Bocas del Toro Prov., elevation 600 feet, 1951-1953. At

each locality from 1 to 4 tree platforms were used, and simultaneous ground

level and canopy collections made on a weekly basis. In addition to these

Panamanian localities, similar stations were manned near Villa Somoza,

Nicaragua; Lancetilla, Honduras; and Teapa, Tabasco, Mexico. These yielded a

wealth of comparative material, the Honduras stations being operated for nearly

a year.

Similar collecting stations were also operated, but only for short periods, in

several localities in the Canal Zone. All the stations were necessarily in areas

of heavy forest. The catches from the Panamanian stations were summarized in

1953 (Fairchild 1953). Sinee the purpose of this work was to collect diurnal

mosquitoes, the stations were generally manned only from 8 am to 4 pm so that

possibly crepuscular or nocturnal species were seldom taken. The station at

Almirante yielded the most material, as it was run almost continuously for two

years, and for shorter periods in subsequent years.

Later, in the 1950's and 1960's, in connection with continuing surveys for

Sylvan Yellow Fever, and surveys for ectoparasites, camps were set up in a

number of isolated areas in Darien and Bocas del Toro provinees. These were

operated, with tree stations, for periods of several months. They were supplied

by helicopter, and permitted sampling of areas at higher elevations, practically

unreachable without this support. While most of the collecting was done by

hand with small nets and human bait, light traps of several sorts were used at

times, and the Shannon trap was used extensively. This is a box-like affair of

netting, open below and with a central panel reaching the ground. It had no de-

vice for holding insects, so had to be visited at short intervals to remove the

temporarily trapped insects.

During the years subsequent to about 1946, Army entomologists on tours of

duty with what came to be known as the Environmental Health Branch, under-

took surveys for biting insects in the Canal Zone and Panama. Their work in-

eluded extensive use of horse-baited mosquito stable traps and New Jersey light

traps. Field teams and trained laboratory technicians sorted the large numbers

of insects collected, turning over to us the Tabanidae and Phlebotomine sand-

flies. The great majority of specimens of Tabanidae were taken in the horse-

baited stable traps, and since most of these were being used to measure popula-

tions of Anopheles mosquitoes close to inhabited areas, the bulk of Tabanidae

belonged to a few common lowland species. The light traps caught far fewer

tabanids, but a greater variety, including males of a number of species not oth-

erwise taken.

The problem of storing the large collections secured was not satisfactorily

solved until within a few years of my leaving Panama. Until some years after

WWII, no air conditioning was available at Gorgas Memorial Laboratory so that I

took the bulk of the collections to the U. S. A. every 1 or 2 years, depositing

most of them at the Museum of Comparative Zoology at Harvard University.

When air conditioning was installed, specimens seemed to last well, and most

material was kept either pinned in Schmitt boxes, or stored dry in pill boxes.

4 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Later the bulk of the collection was kept in 48 glass topped drawers,

duplicates in about 50 Schmitt boxes and several thousands more unpinned in pill

boxes of various sizes stored in tin cans.

At first, most specimens were pinned; later only the rarer ones or those

from unusual habitats, the remainder being recorded and discarded or a sample

saved unpinned. Holotypes of nearly all species are deposited in M. C. Z., the

remainder still in my collection for eventual deposit in M. C. Z. and Florida

State Collection of Arthropods. Paratypes have been sent to various museums

and colleagues, though on no consistent basis. Though no exact tabulation has

been made, it is believed that about 200,000 specimens from Panama have been

seen and identified.

CHARACTERISTICS OF THE FAUNA

In this treatment 152 named forms of Tabanidae are included as occurring in

the Republic of Panama, including the former Canal Zone. These are distributed

in 3 subfamilies 5 tribes and 35 genera and subgenera. This is a comparatively

rich fauna; but since intensive collecting has not been done in comparable areas,

comparisons may reflect effort to some extent rather than actual diversity.

Trinidad (Fairchild and Aitken 1960) has 45 species, and Costa Rica 90 (Hogue

and Fairchild 1974), while Wilkerson (1979) reports 158 species from the three

northwestern departments of Colombia. The whole United States boasts 3 sub-

families, 7 tribes, 31 genera and 282 species (Philip in Stone et al 1965), consid-

erably less than twice the number of species found in Panama though with an

area many times as great.

In 1939 when collecting really started in Panama, only 30 species were

known. Of the 122 species subsequently found, 59 proved new and were de-

scribed almost wholly from Panama specimens. In spite of this richness, the

fauna is not a peculiar one. The percentage of precinctive species is quite low,

about 15%, and most of these are species likely to be found elsewhere when

searched for in their restricted habitats.

Comparison with the Neotropical fauna as a whole, shows the Panama fauna

to be deficient in Pangoniinae, particularly Pangoniini. Eleven percent of the

total fauna are Pangoniinae versus 26% for the whole Neotropical fauna.

Chrysopsinae are about the same, 8% for the Neotropics, 9% for Panama. In

the Tabaninae Panama is correspondingly richer, 78% vs. 65% for the Neo-

tropics. Of the 2 tribes, Diachlorini and Tabanini, the former greatly out-

numbers the latter in the Neotropics as a whole, while in Panama there are only

about twice as many Diachlorini as Tabanini. I have the impression that the

relationship between the 2 tribes continues to change in favor of Tabanini as

one moves west and north. Certainly in America north of Mexico the Diachlor-

ini form an insignificant percentage (0.6) of the Tabaninae.

Perhaps a more interesting way of looking at the fauna as a whole is by

habitats. Though the data do not lend themselves to very detailed consideration,

certain broad correlations seem evident (see map p. 137).

The forest environment, especially the wetter areas along the Atlantic coast

and in the mountains over 1000 ft. elevation, appear to have the largest number

of species, 105 or about 71% of all species. Of the 17 Pangoniinae, all but 3

species of Esenbeckia are forest species; all but 3 of the 14 Chrysopsinae are

primarily forest species; only 15 of the 56 Diachlorini are not primarily forest

inhabitants, but only 18 or 50% of the 36 Tabanini are forest species, and many

of these are taken outside the forest with some frequency.

G. B. Fairchild: Tabanidae of Panama 5

Consideration of the non-forest species shows many to be restricted to a .

single well-defined habitat. For example, 1 Chrysops, 5 Stenotabanus and 6

Tabanus species are beach or mangrove haunting species. Another dozen

Species, none Tabanini, are closely associated with freshwater swamps. The

species which occur widely in many habitats, especially open country, either

natural or man-made, include 2 Esenbeckia, 2 Chrysops, 1 Diachlorus and 13

Tabanus species. All but 2 or 3 of these are widespread, common, lowland

Species occurring throughout much of tropical America.

The distribution and composition of the group of 22 precinctive species is

also somewhat curious. Only 6 of these are species of Tabanus, but of the 6, 5

are closely associated with mangrove swamps and the remaining one may be

also. The 3 precinctive Pangoniinae are all forest species, one of the 2 pre-

einective Chrysops is a forest species, the other strictly confined to mangrove

Swamps. The remaining 11 precinctive species are all Diachlorini, 2 certainly

and 2 probably from beach or mangrove environments, the remainder all forest

Species. I suspect that the high proportion of precinctive halophilous species is

due to neglect of this habitat by collectors, but the lack of any Diachlorini in

this habitat other than species of the chiefly Caribbean Stenotabanus

(Aegialomyia) is surprising.

To sum up. The Pangoniinae are mostly forest species and mostly relatively

widespread, only 3 or 4 being so far confined to Panama. The Chrysopsinae are

moderately well represented, mostly by forest species, and none are very abun-

dant as individuals. Only 1 or 2 are at present confined to Panama. The bulk

of the fauna belongs to the Diachlorini, very predominantly forest species, but

only a few species can be considered really abundant. Many are arboreal and a

fair number nocturnal or crepuscular. Diachlorini also tend to be rare in col-

lections, either through true scarcity or because of undiscovered habits or

habitats. Of the 56 species, 13 are known from 5 or fewer specimens, while

among the 35 Tabanini only 3 are known from so few specimens. Only about

half as many species of Tabanini occur, but they will generally vastly outnumber

the Diachlorini in numbers of individuals, especially in collections made in open

or cultivated country. Numerically the Tabanini are dominant, except in dense

forest, the bulk of the specimens belonging to a few common and widespread

species.

ARBOREAL SPECIES

Included in this list are those species which are primarily arboreal, that is,

most of the available specimens were taken by collectors stationed on platforms

built in the forest canopy, from 40 to 90 feet above ground level. Collections

were made in this habitat sporadically over a number of years (as detailed

previously under collecting methods) for the purpose of collecting canopy fre-

quenting mosquitoes, so that Tabanids were collected only incidentally. These

collections, with few exceptions, were made during daylight hours, so that cre-

puscular or nocturnal species were not usually taken. In 1953 (Fairchild 1953) I

published a short note on this subject, with a list of the arboreal species then

known from Panama. Subsequent collecting has confirmed most of the species

there listed as arboreal, but added only a few others.

6 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Fidena eriomeroides Lutz. Of 7 specimens seen, 4 were taken in the forest

canopy and at least 2 of the others may have been.

Fidena trapidoi Fehld. In 1953 I reported this as 42.3% arboreal; subsequent

collections suggest this figure is low, and the species better than 50% ar-

boreal.

Stibasoma panamensis Curran. Of 24 specimens on which habitat information

was recorded, 16 were taken in the forest canopy.

Stibasoma apicimacula Fehld. Twenty-six of the 28 specimens with adequate in-

formation were taken in the canopy.

Stibasoma flaviventre Viacqg. Where information is available, specimens were

usually recorded as taken in the forest canopy. The subspecies pulla Fehld.

& Aitken from Trinidad is almost entirely arboreal (Fairchild and Aitken

1960).

Stibasoma fulvohirtum Wied. Highly arboreal, at least 90% of all available

specimens from forest canopy.

Dichelacera regina Fchld. About half the available specimens were taken in the

forest canopy.

Dichelacera crocata Fehld. The most highly arboreal species of Dichelacera.

Dichelacera submarginata Lutz. Frequently taken in the forest canopy, but also

at ground level along forest margins.

Catachlorops (Psalidia) umbratus Hine. Nearly all specimens seen taken in for-

est canopy.

Stypommisa maruccelii Fehld. Almost entirely arboreal.

Stypommisa jaculator Fehld. Predominantly arboreal.

Philipotabanus grassator Fchld. The few specimens seen were all taken in the

forest canopy.

Philipotabanus magnifiecus Kroeber Predominantly a ground level species, though

often taken in the canopy.

Philipotabanus (Melasmatabanus) fascipennis Macq. The few Panama specimens

were all taken in the treetops.

Philipotabanus (Mimotabanus) inauratus Fchld. Probably largely arboreal.

Philipotabanus (Mimotabanus) phalaropygus Fchld. The few known specimens

from Panama were from the canopy.

Tabanus xenorhynehus Fehld. Predominantly arboreal.

Tabanus defilippii Bell. Highly arboreal.

Tabanus polyphemus Fehld. Probably arboreal, though data inconclusive.

The species listed above suggest that arboreal habits are rather widespread

in Panama Tabanids, as the list represents nearly 14% of the known Panama

species. There seems also a tendency for arboreal species to be brightly col-

ored, and to resemble Hymenoptera or Diptera of other families. The bulk of

the arboreal species are Diachlorini, with only 2 Pangoniinae, no Chrysops, and

but 3 Tabanini. Only the genus Stibasoma seems to be exclusively arboreal. In

this connection, I must mention the work of Goodwin and Murdoch (1974) who

found that larvae of 4 species of Stibasoma were taken only in Bromeliads, in 3

eases only epiphytic species.

NOCTURNAL AND CREPUSCULAR SPECIES

The data on which the following list is based are exceedingly fragmentary,

as little collecting designed to secure this type of information has been done,

nor has the time of day when specimens were collected often been recorded. A

G. B. Fairehild: Tabanidae of Panama 7

small series of night collections were made at the Almirante Yellow Fever Sta-

tion, a great deal of material taken in New Jersey light traps was studied, and

scattered observations were made as opportunity offered. The taking of males

in light traps is not considered evidence that the females are also nocturnal, as

evidence is available that males of many species whose females are known to be

active by day are frequently thus taken, i. e. Tabanus lineola (Richardson and

Wilson, 1969) in the U. S. A. It also appears to be true that some species,

abundant by day, will continue to attack late into the evening, while primarily

crepuscular species may be active on overcast days, or in deeply shaded habi-

tats. Many species taken only in stable traps, Malaise or Shannon traps may be

crepuscular or nocturnal but information is not usually available as to times

when traps were emptied.

It is perhaps notable that of 17 species of Pangoniinae known from Panama,

6 or about 33% are crepuscular, and of 77 Diachlorini 14, or 18% are crepus-

cular. None of 14 species of Chrysopsinae and only 2 of 37 species of Tab-

aninae are crepuscular. Pangoniinae and Diachlorini are poorly represented

outside the tropics, while the preference of temperate species of Tabanus and

Chrysops for bright sunny weather has frequently been noted and the subject re-

viewed by Burnett and Hayes (1974). Although daylight temperatures in temper-

ate regions are frequently as high or higher than in the tropics, night tempera-

tures are far more likely to fall below the threshold of insect activity outside

the tropics.

Esenbeckia osornoi Fchld. Definitely nocturnal and crepuscular.

Esenbeckia translucens Macq. Specimens were taken on one occasion late in the

evening biting a horse.

Scione rufescens Ric. Definitely crepuscular and nocturnal, flying just after

sunset and before dawn.

Fidena schildi Hine. Twice taken before dawn, and possibly primarily ecrepus-

cular.

Fidena rhinophora Bell. Taken biting man and pigs on two occasions before

dawn, with F. sehildi.

Pityocera festae Giglio.-Tos. Crepuscular:

Chlorotabanus inanis Fab. Nocturnal. Both sexes frequently at lights.

Chlorotabanus mexicanus Linn. Nocturnal. Both sexes in light traps.

Cryptotylus unicolor Wied. Nocturnal.

Cryptotylus chloroticus Phil. & Fehld. Nocturnal.

Bolbodimyia erythrocephala Big. One was taken between 6 and 7 P.M., most of

the remainder in stable traps. This and the other 2 Panamanian species may

be crepuscular or nocturnal, but data are not adequate for certainty.

Selasoma tibiale Fab. Nocturnal.

Phaeotabanus phaeopterus Fchld. Nocturnal. Both sexes at lights.

Phaeotabanus longiappendiculatus Macq. Nocturnal.

Philipotabanus pallidetinctus Kroeber. Crepuscular.

Philipotabanus chrysothrix Fehld. Crepuscular and/or nocturnal.

Philipotabanus magnificus Kroeber. Often crepuscular or nocturnal.

Philipotabanus pterographicus Fchld. Probably nocturnal.

Philipotabanus ebrius 0. S. Crepuscular and/or nocturnal.

Philipotabanus (Melasmatabanus) criton Kroeber Probably nocturnal.

Tabanus quinquepunctatus Hine. Taken only in light traps.

Tabanus unipunctatus Big. Crepuscular and taken biting mules at night.

8 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

HOSTS

So far as I am aware, all the Tabanidae known from Panama are

haematophagous in the female sex. If non-blood feeding species occur, they

must be either very rare or with habits or habitats not easily or often sampled

by the numerous entomologists who have visited Panama.

The sourees of blood for any kind of haematophagous insect other than true

ectoparasites must usually be gleaned from scattered field observations, as only

in the eases of known or suspected vectors of serious disease is it worth the

time and effort needed to utilize techniques yielding information from the blood

meals contained in captured females. In Panama this has not been done, so that

our knowledge as to host preferences is fragmentary and largely inferred. It is

known that most arthropods which feed on vertebrate blood are attracted by

CO,, and Tabanids are no exception. But the observation that a species of

Tabanid was taken flying about a collector or a bait animal- sourees of CO,—is

no guarantee that that species will habitually or even occasionally feed on that

host. Conversely, if a species of Tabanid is regularly taken within an animal

baited trap in a replete condition, it can usually be assumed to have fed on the

bait animal. In the following lists the species actually collected while feeding

on a host (1) taken in an animal baited trap (2), or netted while flying about the

bait animal or the collector (3), are indicated by the appropriate numbers

(Species not listed below are without host data or taken in Shannon, Malaise, or

light traps only).

Esenbeckia (Proboscoides)

ecuadorensis chagresensis(3) horse, cattle

Esenbeckia (Esenbeckia)

translucens (1, 2) horse, man

Esenbeckia (E.)

prasiniventris (1, 2) horse, cattle, man

Esenbeckia (E.)

osornoi (2) horse

Fidena eriomeroides (3) man

Fidena rhinophora (1,3) pig, man

Fidena auribarba (1,3) man

Fidena flavipennis (1) snake, cayman

Fidena schildi (1,3) pig, man

Fidena trapidoi (3) man

Fidena howardi (3) man

Scione maculipennis (1,3) man, horse

Scione costaricana (3) man

Scione rufescens (3) man

Scione ablusus (1) man

Pityocera festae (1,2,3) horses, man

Chrysops alleni (3) man

Chrysops auroguttatus (1,2,3) man, horse

Chrysops calogaster (3) man

Chrysops chiriquensis (3) man

Chrysops melaenus (1,3) man, horse

Chrysops sealaratus (1,3) man, horse

Chrysops soror (3) man

Chrysops variegatus (1,2,3) man, horse

Stenotabanus obseurus (3) man

G. B. Fairchild: Tabanidae of Panama

Stenotabanus (Aegialomyia)

changuinolae (1)

Stenotabanus (A.) littoreus (1)

Stenotabanus(A.) paitillensis (1,3)

Stenotabanus (Brachytabanus)

longipennis (1)

Diachlorus curvipes (1,2,3)

Diachlorus jobbinsi (1,2,3)

Bolbodimyia philipi (2)

Selasoma tibiale (2,3)

Chlorotabanus inanis (2)

Chlorotabanus mexicanus (1,2)

Phaeotabanus

longiappendiculatus (1)

Dichelacera (Dichelacera)

erocata (1,3)

Dichelacera (D.) faseiata (1,2,3)

Dichelacera (D.) marginata (2,3)

Dichelacera (D.) regina (3)

Dichelacera (D.) rex (3)

Dichelacera (D.)

seapularis and var. (2,3)

Dichelacera (D.)submarginata (3)

Dichelacera (Desmatochelacera)

transposita (3)

Catachlorops (Psalidia)

umbratus (3)

Catachlorops (P.)

fulminea and f. ocellata (3)

Dasychela badia (1,3)

Stibasoma apicimacula (3)

Stibasoma chionostigma (1,2)

Stibasoma flaviventre (3)

Stibasoma fulvohirtum (3)

Stibasoma panamensis (3)

Cryptotylus unicolor (1,2)

Cryptotylus chloroticus (1,2)

Philipotabanus (P.) ebrius (1,3)

Philipotabanus (P.) elviae (3)

Philipotabanus (P.) grassator (3)

Philipotabanus (P.) magnificus (1,3)

Philipotabanus (P.) medius (3)

Philipotabanus (P.) nigrinubilus (3)

Philipotabanus (P.)

pallidetinetus (2,3)

Philipotabanus (P.)

pterographicus (1,2)

Philipotabanus (Mimotabanus)

inauratus (3)

Philipotabanus (M.)

phalaropygus (3)

Philipotabanus (Melasmatabanus)

eriton (2,3)

man

man, horse

horse

man, horse

horse

pig, cattle, horse

horse, cattle

man

man, horse,sloth

horse, man

man

man, horse

man

horse

man

cattle, horse

horse, cattle

horse, man

man

man, sloth

man

man, horse

horse

man

horse, man

10 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Philipotabanus (M.)fascipennis (3) man

Stypommisa maruccii (3) man

Leucotabanus canithorax (3) horse

Leucotabanus exaestuans (1,2,3) eattle, horse

Lepiselaga crassipes (1,2,3) man, horse

Poeciloderas 4-punctatus (2) horse

Tabanus albocirculus (2) horse

Tabanus bigoti (3) man

Tabanus occidentalis

dorsovittatus (1,2,3)

Tabanus

Tabanus stenocephalus (1,2) horse

Tabanus colombensis (1,2) horse

Tabanus commixtus (1,2) horse

Tabanus defilippii (1,3) horse, man

Tabanus enanus (2) horse

Tabanus erebus (1,3) horse, tapir, man

Tabanus importunus (1,2) horse, cattle

Tabanus lacajaensis (3) man

Tabanus macquarti (3) man

Tabanus nebulosus (1,2) horse, cattle

Tabanus nereus (2) | horse

Tabanus nondescriptus (2,3) horse, man

Tabanus oculus (3) mule

Tabanus platycerus (2) horse

Tabanus polyphemus (3) man

Tabanus praeteritus (1,2) horse

Tabanus pseudoculus (1,2) —eattle, horses

Tabanus punctipleura (1) horse

Tabanus pungens (2) horse

Tabanus rhizophorae (3) man

Tabanus surifer (1,3) man

Tabanus vittiger

guatemalanus (2) horse

Tabanus unistriatus (3) man

Tabanus unipunctatus (1,2) mule, man

Tabanus xenorhynchus(3) man

o. modestus (1,2,3)

horse, cattle

SEASONAL DISTRIBUTION

Although information as to the seasonal distribution as reflected in the col-

lections made in Panama is given under each species, it may be of interest to

present this information in a more accessible form. Below are listed by months

all species for which data on 20 or more specimens are available, 89 species in

all. The remaining 61 species are each represented by less than 20 specimens.

As ean be seen below, the conclusions expressed previously (Fairchild 1942)

in a limited study of Tabanids taken during a year's collecting at a single lo-

cality, are in general confirmed. That is, the species fall roughly into 3 groups:

those with a short flight season of about 3 months, those flying only in the rainy

or dry seasons, and those on the wing throughout the year. There is a limited

eorrelation between the type of flight season and taxonomic position. Thus all

species of Esenbeckia have relatively short flight seasons, as does

G. B. Fairchild: Tabanidae of Panama 11

Pityocera and most species of Fidena, while of 3 species of Secione, 2. fly

throughout the year. All the species of Chrysops seem to be on the wing at all

seasons. Diachlorini have all types of flight season, while most Tabanini fly for

extended periods. The data are not to be trusted in all cases, as long series of

some species may have been collected at a certain locality, but the locality only

visited onee for a short period. In general, more collecting was done in the

months from May through August, least in October through December. |

Since a number of the names used in my earlier publicaton on seasonal dis-

tribution (Fairchild 1942) have been changed for various reasons, I give here the

current equivalents for those that are different for ease of comparison.

Fairchild 1942 Names Present Usage

Esenbeckia illota Esenbeckia osornoi osornoi

Esenbeckia sp. Esenbeckia ecuadorensis chagresensis

Dichelacera analis Dichelacera fasciata

Psalidia fulminea Catachlorops (Psalidia) fulminea var.

ocellata

Cryptotylus limonus Cryptotylus chloroticus

Cryptotylus luteoflavus Phaeotabanus longiappendiculatus

Leucotabanus leucaspis Leucotabanus exaestuans

Stenotabanus plenus Philipotabanus (Mimotabanus) plenus

Tabanus (Phaeotabanus) medius Philipotabanus pallidetinctus

T. (Hybomitra) quadripunctatus Poeciloderas quadripunctatus

T. (Neotabanus) lineola var. carneus T. occidentalis var. dorsovittatus

T. (N.) lineola var. stenocephalus T. stenocephalus

T. (N.) maya T. commixtus

T. (N.) angustivitta T. pungens

T. (N.) amplifrons T. colombensis

T. (N.) fumatipennis T. platycerus

T. (Tabanus) ferrifer T. nebulosus

Seasonal distribution of Panama Tabanidae. Approximate number of

specimens examined for these data follows the scientific name.

MONTH Now kde ERM RAD MY J £4on 23.. O ean

PANGONINAE

Esenbeckia

translucens ol ie ete ee Xx

prasiniventris O11 Se, ay 4 X

osornoi 105 Pe ae

chagresensis 87 bee Soe.

Pityocera

festae 64 Xie k ni:

MONTH

Sceione

maculipennis

eostaricana

rufeseens

Fidena

auribarba

flavipennis

rhinophora

sehildi

trapidoi

CHRYSOPSINAE

Chrysops

alleni

auroguttatus

calogaster

melaenus

mexicanus

nexosus

sealaratus

soror

variegatus

DIACHLORINI

Dasychela

badia

Stibasoma

apicimacula

fulvohirtum

panamense

Cryptotylus

unicolor

ehloroticus

Catachlorops

fulminea

f. ocellata

umbratus

Dichelacera

crocata

fasciata

marginata

regina

rex

scapularis

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

No.

PS PS PS PS PS PS PS PS

ms PX

Boyce: Mo d

MxM MM

rs PX

ax xKM KM

ms PS

rs sS

PS PS PS PS PS PS PS PS OS

PS PS PS

rs PX

MPN MMM Ps PS PS PS

ms ms XS

PS PS OX PS

PS PS PS PS

ms ms PS XM

rs PX

PX MN SS

PS PS PS PS PS PS PS rs PS PS

rs PS PS

PS PS PS PS

PX PS PS

PPS PS PS OS

PS PS

Mr PS PS

rm XS

~~ PS

~*~ XS

ms PS

mn xX

ms XS

MONTH

submarginata

subcallosa

Himantostylus

intermedius

Lepiselaga

crassipes

Diachlorus

curvipes

jobbinsi

Chlorotabanus

inanis

mexiecanus

Phaeotabanus

longiappen-

diculatus

Stenotabanus

incipiens

minusculus

changuinolae

Leucotabanus

exaestuans

flavinotum

Stypom misa

captiroptera

jaculator

lerida

marucclii

pequeniensis

Philipotabanus

ehrysothrix

ebrius

magnificus

medius

nigrinubilus

pallidetinetus

pterographicus

Poeciloderas

quadripunctatus

G. B. Fairchild:

No. J

100

193 X

101 X

100 4

236 4

321 4

A401 XxX

20 X

30 X

20 4

462 4

20 4

100

1000

126

30 X

ms XS

ms x

Tabanidae of Panama

mx ms XS mM XX

ws PS ria

PS PS PS PS PS PS

ms XS

mS PS

Sr PS MX

PPS PS PS PS

PS ms XS ms

ms PX

~*~ XS

mx PX

m~ X

xxx

MX MX

XX ms XS

ms PS

ms S

ms PS

14 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

MONTH No. oR AL OR Be ORR A ORB ED

Tabanus

albocireulus 1030 ee Fee RE RR ea Re we OPK

bigoti 32 Cae, Cue), Gana. &

colombensis 20 ae Sores Aiea, Soe, Gans’ Game,’ 4 X

commixtus 1491 DO i i Ree era eae Xx

defilippii 20 Fe er Pe ae Ue St

enanus 100 ee oe X

erebus 200 OR a ea XxX Xx

importunus 225 x Be Xo at, Gane &

lacajaensis 20 ee ae xX

nebulosus 149 DO Re OR I Ra XK KK eer RK

nereus 100 EC De AOR. x ee er

occidentalis

var. dorsovittatus 100+ De OR eR Oe » ae, Cea, Geman 4

occidentalis :

var. modestus 100+ mee XX Oe Oe OR Ro DES AR

nondescriptus 100+ X Xx X XxX Sa Nae 6

platyeerus 30 ee ee Oe X

praepilatus 29 > aa € Der ee ae X

praeteritus

ssp adiastolus 50 Gy ee Tee eae) ee 4 X X paw

pseudoculus 208 Pe RR XEN eK

punctipleura 20 pee ia alee a py, Gn, Wie ae aoe

pungens 959 Mee KR ay ee Oe Xx

stenocephalus 100+ Re ee Dw RE I pe a ome 4

surifer 100+ TR Re URE ae RG OE DO EG ee

unipunctatus 100+ Det Tae, Same 4 pCa, aay 4

unistriatus toe ee Re OR OR OR Ro EX Ka

vittiger

ssp guatemalenus 250 X DSR oie See Saar ie ate.

EXPLANATION OF TERMS

Since the terms used by various taxonomists for structures mentioned fre-

quently in descriptions are far from uniform, it may be useful to mention those

used here. Some of the more important of these structures are illustrated in

figure 1, p. 135.

Frons. The area between the eyes in female tabanids, lying above the sub-

callus and below the vertex. The frontal index is the vertical length of the

frons divided by its width at the base, below the callus.

The frontal callus is a bare and often swollen area at the base of the frons,

of various shapes, sometimes extended above into a ridge, which may form a

separate or connected median callus.

The eyes may be bare or pilose, bare when no short hairs are visible under

9x magnification, pilose when such hairs are obvious under low magnification. In

males a variable amount of the eye area may consist of more or less enlarged

facets. When obvious differences in size are apparent, the facets are said to we

well differentiated. Often these enlarged facets form a clearly defined patch in

the upper part of the eye with the line of demarcation between large and small

facets sharp; they are then said to be well demarcated. Eye color and pattern

G. B. Fairehild: Tabanidae of Panama 15

fades shortly after death, but at least the pattern may usually be revived in a

moist chamber or by the use of various techniques.

Vertex. The top of the head, often bearing a tubercle or bare area, fre-

quently with 3 ocelli or vestiges thereof.

Subeallus. The area, sometimes swollen or bare, lying below the frons, and

into the lower margin of which the antennae are inserted.

Frontoclypeus. The median area below the antennae. Its lower margin is

next to the proboscis. It, plus the anterior parts of the genae, are often spoken

of as the face.

Genae. The lateral areas below the eyes, bearing the beard.

Tentorial pits. More or less conical invaginations of the exoskeleton on each

side of the frontoclypeus.

The antennae are three-segmented, the first or basal segment is the scape,

the second the pedicel, and the third or terminal segment, is composed of no

more that 8 annuli, the basal 4 or more of which may be fused into a basal

plate, leaving a terminal style or flagellum of 2 or more annuli. The basal plate

may have a dorsal angle, tooth, or spine of variable length.

The proboscis is composed of paired slender mandibles and maxillae, and un-

paired labrum and hypopharynx, all more or less enclosed in a trough-like labium

bearing paired labella at its distal end. Normally the last two structures are the

only visible parts.

The palpi lie on either side of the proboscis and are two-segmented. The

terminal segment is the largest and most prominent and is the one whose con-

dition is usually referred to in descriptions.

The mesonotum (scutum) is the dorsal surface of the thorax, the scutellum a

posterior dorsal rounded or broadly triangular sclerite slightly overlapping the

abdomen. On each side are a pair of notopleural lobes at about the middle of

the mesonotum. The pleura are the lateral areas of the thorax.

The abdomen is composed of 9 or 10 segments, but only the first 6 or 7 are

normally visible, so that "last segment" generally refers to last visible segment

on a living or undissected fly. Tergites and sternites are the dorsal and ventral

surfaces of the abdominal segments.

The wing venation is very constant, but veins and cells are often referred to

in describing wing patterns. I prefer the terminology used by Curran (1934)

rather than the, to me, awkward terminology of the Comstock-Needham system

and its various modifications, especially in regard to the cells. The fork, or fork

of third vein is the junction of R4 and R5 of the Comstoeck-Needham system,

often bearing a proximal stub or appendix. The first posterior cell (5th R) may

be open, narrowed or coarctate towards the margin, or closed, sometimes some

distance from the wing margin, when it is said to be petiolate. The crossband

in descriptions of species of Chrysops is a dark band of variable extent extend-

ing from the costa towards or to the hind margin in the middle of the wing.

The apical spot is a dark area of very variable extent distal to the cross band

and always including at least part of the marginal cell. It is separated from the

crossband, when extensive, by the hyaline crescent.

The basicosta is a small seale-like selerite overlying the base of the costal

vein. It is said to be bare when it lacks setae similar to those on adjoining

costa, although fine pubescence is always present.

The vestiture of the body and legs consists of pollinosity, a fine colored

powder-like substance, and pilosity or short or long, variously colored hairs,

which are slender, sometimes ligulate, or strap-shaped and recumbent or erect.

Pruinosity refers to an appearance of the surface which changes its color de-

pending on the incidence of the light.

16 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

The legs show few outstanding characters, except for the presence or ab-

sence of paired apical spurs on the hind tibiae. The tibiae are sometimes

swollen or incrassate, and the hind pair may bear long hairs in a row, the hind

tibial fringe.

The colors of Tabanids are a combination of that of the vestiture and the

integument, which should be described separately, but seldom are. The pattern

of stripes, spots, ete. can be variously described, but in this work stripes are to

be understood as being parallel to the long axis of the body, bands as being

transverse or perpendicular to the long axis.

The above terminology is that traditionally used in taxonomic publications on

Tabanidae and that used by me in previous publications. It differs in a few

respects from that used in McAlpine et. al. (1981), but is retained here to main-

tain continuity with the previous work on the Panama fauna. The modern

terms, where different, are inserted above in parentheses.

GLOSSARY OF GENERAL TERMS LIKELY TO BE FOUND IN KEYS

apical - at or towards the tip or outer end of a part.

banded - markings at right angles to long axis of body or structure.

basal - opposite of apical.

bicolored - of 2 contrasting colors, as yellow and black.

ealli - plural of callus.

callosity, callus - a generally bare and shiny protuberance.

eonecolorous - of the same color as an adjoining part.

contiguous - in contact; touching each other.

diffuse - not clearly defined; with fuzzy or indefinite margins.

discrete - clearly defined; opposite of diffuse.

distal, distally - the end or direction away from the main body; opposite of

proximal.

dorsolateral - between the mid-line and the side margin on the upper surface.

flavidus - light yellow or light yellowish brown.

fulvus or fulvous - dark yellow or dark yellowish brown.

fuscus or fusecous - blackish.

hyaline - transparent; like glass or water; often modified by such words as yel-

lowish, smoky, grayish, etc.

inflated - swollen, as if blown up like a balloon or bubble.

infusecated - blackened or darkened.

integumental - having to do with the basal integument rather than the surface

pollinosity or pilosity.

petiolate - stalked; see under wing venation in explanation of terms.

pilose - covered with pile, or very short erect hairs, like velvet or a non-shag

carpet.

protuberant - sticking out, as a nose on a face.

proximal - opposite of distal.

pubescence - soft, short, reclining hairs.

setose - covered with longer hairs or bristles.

striped - markings parallel to long axis of body or structure.

tubercle - a small discrete wart-like protuberance.

unicolorous - of one color only.

vestiges - remains of a structure which no longer functions, as vestiges of

ocelli.

G. B. Fairchild: Tabanidae of Panama t%

REFERENCES AND SYNONYMIES.

I have tried to keep the latter to a bare minimum, including only those ci-

tations necessary to clarify the name used or to refer to an illustration of the

species. When possible I have included only citations referring to Panama

specimens. Full synonymy as well as a fairly complete list of references to

Neotropical Tabanidae will be found in the Catalogue of the Diptera of the

Americas South of the United States, Fascicle 28, Family Tabanidae, Museu de

Zoologia, Universidade de Sao Paulo (Fairchild 1971). I have abbreviated this ti-

tle in the references under the species to "Cat. S. Amer. Dipt. Fase. 28" in the

interest of brevity. All but a very few recently described Panama species are

listed in this Catalogue, with synonymy and distribution. The earlier references

to the species may be found in Kroeber's 1934 Catalogue, or in various early

papers by me on the Panama species, generally cited under each species, and to

be found in the bibliography at the end of this work. Listing all references to

each species would enormously increase the labor of preparation and greatly in-

crease the length and cost of publication, without a concomitant increase of

usefulness. The list of references at the end of this publication contains only

those publications specifically cited in the text, plus a complete listing of pub-

lications referring primarily to the Panama fauna. The single most useful publi-

cation for identifying most of the Panama species is that of R. C. Wilkerson

(1979), as it contains figures of many Panama species.

Key to Subfamilies, Tribes, Genera, and Subgenera

1 Hind tibiae with 2 apical spurs. Basicosta and subcostal vein without

macrotrichiae. Styles of male genitalia bifid, rounded, or pointed.

Caudal ends of female spermathecal ducts simple, without cup-like

EXPANSIONS ..ccccceeee Rhee G Res WRK 6 OAC TREATS 4 Win UR Res eK RAG hardin dnaeies 2

Hind tibiae without apical spurs. Basicosta bare or setose, subcostal

vein with at least macrotrichiae below. Styles of male genitalia

truncate. Spermathecal ducts with cup-like expansions at caudal

CGS sek Rae SaidwadaVnasereen PRs vee tease tees delaa cnt GA Tabaninae 7

2. Third antennal segment with 7 to 8 distinct annuli. Ninth tergite entire

IN DOIN SONGS. nanececidita dies LI ns ee itiiioune Pangoninae 3

Third antennal segment never with more than five distinct annuli, the

basal one much the largest. Ninth tergite divided in both sexes .....

nadie danacauvve lb engeun dies cadena danvssieedacdWeloweniat sebnntebeaueniiaans .ee Chrysopsinae 6

3. Eyes bare. Style of male genitalia bifid. Frons with ridge-like callus,

which may be bare or pollinose ......00 Pangoniini. Esenbeckia (p. 22)

Eyes pilose. Style of male genitalia simple, pointed. Frons flat, without

BNY° SOR OL CAEP auc onl a oe RAS PN ng Seionini 4

4. Third antennal segment with annuli bearing dorsal and ventral pro-

jections, bipectinate. First posterior cell petiolate, closed by a for-

ward. bending of ‘MI’ to Meet RGE! Mai cake ctaees Pityocera (p. 34)

Third antennal segment simple, subulate. First posterior cell open or

closed, but if closed, R5 bends backward to Meet M1 .eseccccccesecccees 9)

6.(2)

7.(1)

10.

12.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

First and fourth posterior cells Doth ClOS€d.......ccceccccescescees Scione (p. 31)

Fourth posterior cell always open, first posterior Closed OF ODEN ....ccccceeee

ci atasen gab cwebechpele vase wieadh GAB cdadaweh iGO oC REL Lap tues Fidena (p. 25)

First and second antennal segments elongate, both markedly longer than

third segment. Eye pattern of small irregular.dotss ..is.ioctscsscccscedccpecs

ban Sata ds « Daaniatesigatie Dee av eva We Sena ie sepa Sen teks ASA ce Silvius (Assipala) (p. 35)

Third antennal segment always as long or longer than second, usually

longer than either first or second segment. Eye pattern of dots and

bars; characteristic of (he SpeCieSwscsassdisveccmsacssieans Chrysops (p. 36)

Basicosta without strong setae or these less dense than on adjoining

eosta. Often with one or more of the following: vestiges of ocelli;

sclerotized labella of proboscis; long dorsal spine on third antennal

segments; bare and shiny or inflated subeallus, frontoclypeus or first

antennal segment; inflated tibiae; pictured wings. ....... Diachlorini 8

Basicosta with numerous strong setae, equal in size and density to those

on adjoining costa; or if setae sparse, then a tubercle at vertex ......

pene tes eeag eM aseas iaree tees aMtia ees eis s ok) dedas ae sded i sewddebootssie os secede Tabanini 26

Third antennal segment with a strong dorso-basal tooth or forward

pointing spine which often reaches to or beyond end of basal plate 9

Third antennal segment with at most an acute dorso-basal angle.......... 16

Eyes densely short pilose. Antennal tooth reaching beyond first annulus.

Proboscis at least twice length of long slender palpi, about equalling

head height, labella short, pollinose. Callus club-shaped, much nar-

rower than frons. Wings with diffuse discal dark patch ......c.cccccccces

eas « kaa ly Souk dS lesintheud duasebtulds wdcaceeucr ibis <Bdevae «ds ‘Dasychela (p. 74)

i accede wyncus es ee caaue coach ca gertacececew snap siete caress Re eeeeetGRU Pe edies sacwesecses. 10

Stout, generally hairy and bee-like species with usually inflated fore

tibiae, long hair fringes on at least hind tibiae, inflated palpi, short

stout antennae with dorsal tooth extending beyond first annulus, and

SMI SEIS OLISZSd ADC 11a, See sc ceb eves sstuielenss sedescsles stbesdedscwiscccdecccsvced 11

More slender species or without the above combination of characters...12

Abdomen not greenish, densely hirsute. Hind tibial fringe long; at least

fore tibiag dinPlaterh ss iscitnesethcacdcnspiendienaaesdies aud’, Stibasoma (p. 75)

Abdomen green or greenish, sparsely clothed with hairs. Hind tibial

fringe moderate; all tibiae slender ............. St. (Rhabdotylus) (p. 75)

Frontal callus reduced to a small ill-defined median bare patch or eclub-

shaped mark. Palpi inflated basally. Antennal tooth short, not

reaching end of basal plate. Pale unicolorous stout yellow or orange

Species with greenish body Color and UNMarKked WINGS. ceccccccccccccecccees

sill ead ieee VNR UT, JRA iia AUD ais va eunaats tevin sknwase Cryptotylus (p. 81)

Frontal callus well defined. Antennal tooth longer, slender, acute.

Palpi slender. Wings with dark MarkiNngS 2...crccccccscccccscscccccccccccosees 13

13.

14,

16.

17.(8)

18.

12,

G. B. Fairchild: Tabanidae of Panama 19

Stout, wholly black shiny species. Wing with a black crossband but dis-

eal cell with a round clear area. Dorsal antennal tooth long and

acute. Vestiges of 3 ocelli at vertex. Labella of proboscis not or

weakly selerotized:. :.c<clesctenced Ui cadbvanccencevvedss Gio Dicladocera (p. 75)

Not black and shiny, but if so wings without clear spot in diseal cell.

ANTENNAE LOOT: VALlaDIle). cao vsins<ovcéevehtbnebanaies cdabiigeabedteviewletascccsneceses 14

Frontal callus slender, ridge-like, narrower than frons. Labella largely

shiny sclerotized. Eyes unicolorous bright green in life, rarely bi-

eolored or with faint median line. Mesonotum unicolorous or weakly

Striped, not transversely banded ....... Catachlorops (Psalidia) (p. 70)

Frontal callus as wide as frons. Eyes banded or unicolorous blackish in

life. Mesonotum often transversely banded .........cscscscccsscccsscsceccsces 15

Slender black species, the wings with base and a broad diseal band black,

leaving area around ends of basal cells and apex beyond fork hyaline.

Fore tibiae bicolored, mid and hind pair wholly black pilose. Fourth

tergite with prominent white pilose dorsal triangle ..........cccccccccsscees

spldebis es URL dslhd pods auaeMhbaeeecde seseee. Dichelacera (Desmatochelacera) (p. 69)

Not entirely black, but if so then wing pattern not as above and all

tibiae extensively white. Abdomen and mesonotum usually trans-

versely banded, wing pattern a diagonal band, sometimes faint ........

Sue gee eK UO 2 eden bs ehds Sgn dilda canna swantys Nodameggne ssshvncrvedsvenialvesdaseevevcecs LO

Mesonotum dark with 2 short pale stripes anteriorly and sides and noto-

pleural lobes yellow. Wings yellowish hyaline, the outer 3rd beyond

fork faintly smoky. Subcallus with a median bare patch ......06 Livivead

fidildhi, alba wuts. scccccccccecccccccceseceseeee DicChelacera (Idiochelacera) (p. 69)

Mesonotum without longitudinal stripes, usually with at least a pale pi-

lose transverse prescutellar band. Wings various. Subcallus ecom-

PileRelDOLUNOGE: ns codecseunoshedakeasaunes . Dichelacera (Dichelacera) (p. 61)

Subeallus, and usually first antennal segment, greatly inflated and shiny.

Third antennal segment very long and slender with an obtuse dorso-

basal angle. Tibiae slender or slightly incrassate. Wings black or

partly so, the extreme apex always sharply hyaline, vein R2+3 bent

Sharply forward. Palpi moderately slender, pollinose. Frontoclypeus

DOLINGSS naveesuesccokeutiloes sb Ryde apappnekts daha teh ogethas Bolbodimyia (p. 56)

Without the above combination of Characters. iccccccccccecesccsosccecdcccsscecesees 18

Tibiae, especially first two pairs, greatly inflated. Subcallus, fronto-

clypeus and genae bare and shiny. Palpi shiny and flattened. Wings

basally black at least to ends of basal cells. Labella unsclerotized

sins enipesinnantanpaebnin kona Rupntecensde ieNeee aNeers ees annpaweusenmmecSemeees cubeesodeee cies 19

Tibiae not or but slightly inerassate, and without the above combination

Of .CHAPAGtErS . cevitamisees as ald bk ware eneaeneng’s abe ailign uke seaalea Risa beaten sanksannnteds 21

Large shiny bluish black species with basal black of wings filling half

discal cell. Basal plate of third antennal segment broad and flat,

over 3 times as long as style, its dorsal angle obtuse and at middle

ER aa a ak Rls Selasoma (p. 58)

Small species with slender antennal plate, its obscure dorsal angle near

DaSe.- SUlesOPODAPLIONBITY, LONGO x isnussasspnednssinnnraencnselensseanpace shone 20

20.

21.(18)

22.

23.

24.

25.

26.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Wings black only to ends of basal cells, the distal margin of the black

area vertical from costa to tip of anal cell. Abdomen bare and

STHAY, VIOIACCOUS ccdiscidsseeacves oud reese .cvosdtasdeaceedes Himantostylus (p. 53)

Wings black to beyond end of diseal cell, with a hyaline triangle in

fourth (M3) and fifth (Cul) posterior cells. Mesonotum, and often

abdomen, with metallic brassy or greenish seale-like hairs ..... Sesnennns

Ticeeak ecasel oie ites Cock Meee dee sued (aces eae eas ac kate Lepiselaga (p. 101)

Mesopleura shiny or pearly pollinose, in contrast to rest of pleura. La-

bella unselerotized. Fore tibiae nearly always moderately incras-

sate. Frontoclypeus nearly always shiny and inflated. Wings usually

WIth: GOEK ADIGAL DAEGH Foss. cldiil i ceideccdccssccsecosseds Diachlorus (p. 54)

Mesopleura not so, at least not contrasting with other pleural sclerites...

sdk Ahh RAd eine tba eased oad LAMET ks Ktke eaedieh hase <AdhcedoSehthes sap eodvavacapedertaseeensces ry

Frontal callus absent. Uniformly pale pilose insects with greenish in-

tegument, unbanded eyes, sclerotized labella and hyaline or spotted

WHS cvadascedl ccnaddaccecbas Rerieetss des cocdacddlvocdvucceesseats Chlorotabanus (p. 58)

POMC CATs DPOSCIIE cir iaivasies sisi cVeseseeedeceesesveeecee vo sccebeececesetscccccessesecess 23

Frontoclypeus and genae subshiny, produced, the face conical. Third an-

tennal segment subeylindriecal, without dorsal angle. Frontal callus

eclub-shaped, flat. Wings unusually long, entirely black except for

apical halves of basal CelIS ......sccrscocscccscccssoosee Hemichrysops (p. 55)

Frontoeclypeus and genae not produced, the latter very rarely shiny.

Calis Gn WINGS NOU BS QDOVE lil... ccsdscevavebecsandebdictscbeseesssessedeccsecs 24

Labella at least partly shiny sclerotized. Frons narrow, generally over 5

times as high as basal width, the callus small, narrower than frons.

Eyes unicolorous, unbanded. Third antennal segment with strong

OPA ARIS ais bies io sion cesneye ee acdessvencsstiavesssonsscets Phaeotabanus (p. 60)

Labella unseclerotized. Frons generally less than 4 times as high as basal

width. Eyes usually banded. Third antennal segment with weak dor-

BED Ae ec chspractercerdteircerecstcphcboveceerusdtskeyserdstavelsettesedscthcdessenccstteces 29

Frons excessively broad, less than twice as high as basal width. Frontal

eallus small, rounded, narrower than frons. Tentorial pits black and

shiny within. Style of third antennal segment with annuli partly

fused "SO"thal “OMY SAPS GiIStINGUISNADIe eee eis cis ee ead ce ste a ceesesccences

pkebetlbcnes séabecpterlaalaeouss co seientaaets Stenotabanus (Brachytabanus) (p. 52)

Frons narrower, the callus as wide as frons. Tentorial pits pollinose.

UPS SME US SATA: (TU Lar ei gerd ke dase secede ae seeheseretetecketeeterevecccsosoone 26

Frontal callus usually higher than wide, and with a prolongation above

widened into a more or less distinet median callus, usually sur-

rounded by a dark contrasting hair patch. Eyes generally purple

With 2 ereen “harids h ieasssuitileisees Stenotabanus (Stenotabanus) (p. 46)

Frontal callus usually wider than high, without median callus or con-

trasting hair pateh. Eyes generally purple with three green bands or

greenish with four dark bands. Mostly pale beach-inhabiting species

PT aveecdealenesne cave tebe NES Stenotabanus (Aegialomyia) (p. 50)

27.(7)

29.

30.

31.

32.(27)

G. B. Fairchild: Tabanidae of Panama 21

With a well marked tubercle at vertex and often clear vestiges of ocelli.

Eyes bare, frons narrow, callus club-shaped or ridge-like ........ seeken eo

Without a tubercle at vertex, though there may be a slightly raised shiny

or discolored spot. If a tubercle is rarely present, then eyes are

pilose, or frons broad, or Callus rounded ....sccccccceceee Snmange sive L devout 032

Ocellar vestiges nearly always present. Tibiae either all strongly bi-

colored or entirely black. Wings never patterned or spotted, some-

times tinted or costal cell yellowish, usually glass clear. Scutellum

and usually mesonotum with yellow or white vestiture, often con-

trasting with abdomen. Setae on basicosta as dense as on costa .....

Laxeadnigiegsebeat banaue cs seccscccccccccscececcsccscsscccesescsseees LEUCOtADANUS (p. 98)

Ocellar vestiges evident or not. Mid and hind tibiae never prominently

biecolored. Wings seldom glass clear, either tinted, spotted or with

dark pattern. Setae on basicosta nearly always reduced in density,

rarely ADSENt © c.ccccccccccccee Kies Leek A Ei tieintanahetthes Gok ROTO ce oe Seabseeeiel sacewenne

Wings clear, tinted, or the veins margined, or cross veins with prominent

clouds, often the apex more or less strongly darkened ...ccccccscccecceeeee

sth Ceosts sdacian. chee idebenditdan binds editin iad . Stypommisa (p. 92)

Wings with prominent dark pattern, which may be reduced to a small

area below stigma, but does not consist of clouds on cross-veins or a

prominent apical dark patch .....ccccccoee ‘vekiitaphoinerese iienwey TPE ey Sere 30

Wings with an irregular dark pattern of variable extent which always

leaves clear areas surrounding all cross-veins and fork of third vein.

Frons narrow to very narrow, 7 to 10 times as high as basal width.

Palpi slender. Eyes bronze in life, unbanded. Always rather slender,

long winged species, the wings nearly or quite equalling body length.

Males with large eye facets bare or rarely Short PilOS€ ...coccccscsecseoes

Seis aca «Us esas aS sesosseeeeeee Philipotabanus (Philipotabanus) (p. 83)

Wings ranging from almost wholly blackish to hyaline with a small dark

area below stigma, but cross-veins at end of discal cell and fork of

third vein not surrounded by clear fenestrae when included in dark

DAUICI Geist Gadsiesk bie iiEetamckuihecnt ties aha MRR 31

Slender species with frons 7 times as high as wide or narrower, the palpi

and antennae slender, the proboscis considerably longer than palpi,

with small labella. Eyes bright green in life ......... esbeen Raion ckiesuKetnis

inaeh Ganeeadielts pn csntdadhde nesneh Os tian diced the (Melasmatabanus) (o. 90)

Stouter species with broader frons not over 6 times as high as basal

width, the palpi inflated, the antennae broader, the proboscis hardly

longer than palpi, the labella large. Eyes green or brick red in life,

fading to greenish black. Males with large eye facets long pilose ..

<ssadasstbc senile RbiAsiiee Malkes Hs ebedatiiads -cahniiitieseste ee (Mimotabanua) (p. 89)

A small rounded tubercle at vertex. First antennal segment enlarged,

eap-like, wider than third segment. Eyes of female usually pilose,

of male densely so. Wings with all cross-veins prominently spotted

and first posterior cell (5th R) closed or Coarctate ...cccccccccccccccccecees

pivddondtbiibadeesines FO 8 mR a ON meen (p. 102)

Without the above combination of characters. Never with a tubercle at

vertex, sclerotized labella or bare basicosta .......00. Tabanus (p. 103)

Ze Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Genus Esenbeckia Rondani

Rondani 1963, Arch. Zool. Modena 3: 83. Philip, 1945, Rev. Brasil. Ent.,

2: 13-60, pp. 1-10, key. Wilkerson and Fairchild 1983, Journ. Nat.

Hist. 17: 519-567, figs 1-18, key.

In addition to the characters in the key, species of Esenbeckia do not have

produced conical fronto-clypeus, the palpi project at nearly a right angle to the

axis of proboscis and are usually fairly long and curved. The proboscis is seldom

over twice the height of the head, often less than head height, slender to quite

heavy, the labella ranging from slender to broad, and either partly or wholly

sclerotized. The antennae have the first two segments short, hardly longer than

wide, the third of 8 annuli, the first few much wider than long, the terminal

segments longer than wide, so that the whole segment tapers from a broad base

to a slender and attenuated apex. The eyes are always bare and unicolorous,

green, bronzy or blackish in life. The wings are clear, tinted, blackish or vari-

colored, and nearly always have the first posterior cell closed and a long ap-

pendix at fork of third vein. Legs are slender and relatively long. Three sub-

genera occur in Panama, but are not keyed separately.

Most species seem to be avidly haematophagous, and in Panama at least

each has a definite and fairly short flight season. Males of but two of our 5

Species are known.

Key to females of Esenbeckia

B. Proboscis thick and heavily selerotized, the labella curved, thickened and

entirely sclerotized. Thorax and abdomen blackish, the former with

whitish hairs on pleura, the latter with narrow white hair fringes on

BIE SEST STS OAT SISENIESS: . eaiwcissvarsnssgnscnsdenseretnorcees sdedieacdudecddececenevedds

Boece ab vend basta dab. (Proboscoides) ecuadorensis ssp. chagresensis (p. 23)

Peoutseis more slender, the labella not as above. Thorax and at least

first two abdominal segments not DIACK .......ccccccccsscssscscscccscscscscscees 2

Zs Palpi very short, hardly one fourth length of long proboscis. First three

sternites yellowish-horn color, the second and third each with a pair

of round sublateral brown spots, remainder of sternites dark brown .

Sisto ivansadeegn. te er Geinlicideededdbenlemareesesss ewes (Ricardoa) subguttata (p. 24)

Palpi at least half length ne proboscis. Abdomen unspotted beneath..... 3

3. Proboscis less than head height, the labella broad and only partly sclero-

tized. Legs black. Mesonotum and scutellum clothed with orange

pollinosity and hairs. Abdomen with first two segments translucent

horn-colored, remainder black. Palpi broad and square tipped .........

peat iaele Bieiade scccccccccscscccccsscccscscscscscseees (ESenDECKia) translucens (p. 24)

Proboscis at least head height, slender, the labella narrower. Legs

brown or ‘yellow? “ADdOmMeH NOG DIA CK oot cia eia eds coeenecsecees 4

G. B. Fairchild: Tabanidae of Panama 23

4. Stout species. Palpi with blunt apices. Abdomen bright jade green.

Thorax dark brown. Hind legs largely black haired .......cccccccccccccecees

sioie vp lchdecesbiedeced eile toed ecaestbaeaneisen Prieee bees caeds .»»- prasiniventris (p. 24)

Slender species. Palpi acutely pointed. Thorax and abdomen dull pale

yellowish brown, the first two abdominal tergites paler. All legs

pale; Waited Ho. ikea Eocene cee ess eee tke dab osornoi osornoi (p. 25)

Subgenus Proboscoides Philip.

Proboscoides Philip, 1943, J. New York Ent. Soe., 51 (2): 111.

Esenbeckia (Proboscoides), Fairchild, 1969, Arq. Zool. S. Paulo, 17(4):

201. Fairchild and Wilkerson 1981, Florida Ent. 64(1): 158-175, figs.

1-9.

This group is separated from typical Esenbeckia by having the labella of the

proboscis lengthened, entirely sclerotized and apparently lacking pseudotracheae.

Only one species occurs in Panama.

Esenbeckia (Proboscoides) ecuadorensis ssp. chagresensis Fairchild.

Esenbeckia chagresensis Fairchild, 1942, Ann. Ent. Soe. Amer. 35(2):

192-194, P1. 1, fig. 9, female, Panama.

Esenbeckia (Proboscoides) ecuadorensis chagresensis: Wilkerson 1979.

Cespedesia 8(31-32): 141-142. Fairchild and Wilkerson 1981, Fla.

Ent. 64(1): 165.

This species appears to be rare in Panama, as only a single specimen has

been taken since those included in the type series, from the Chagres river basin

and from Utevey near Pacora. The additional specimen is from Morti river,

Darien Prov., 29 July 1967. The subspecies seems confined to eastern Panama

and N.W. Colombia. It differs from the nominate form mainly in having black

hairs on mesonotum, rufous in equadorensis, and in a darker abdomen with only

the first 2 tergites partly dull yellowish, the first 3 nearly wholly yellow in

ecuadorensis. All Panama records are from June, July or August.

Subgenus Ricardoa Enderlein

Enderlein, 1922, Mitt. Zool. Mus. Berlin, 10(2): 340.

Esenbeckia (Ricardoa): Fairchild, 1969, Arq. Zool. S. Paulo, 17(4): 201.

This group differs from Esenbeckia in having a very slender proboscis with

small labella, and palpi usually not over half length of proboscis. Only a single

species is known from Panama, the group being mainly Mexican.

24 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Esenbeckia (Ricardoa) subguttata Fairchild

Esenbeckia subguttata Fairchild, 1964, J. Med. Ent. 1(2): 169, fig. 1.

Only the type specimen is known, taken at about 2000 ft. elevation at La

Zumbadora, Cerro Azul, Panama Prov. in March 1959. It is easily recognized by

the combination of long slender proboscis and very short palpi, hardly 1/4 length

of proboscis. It is a light brown insect, the abdomen shiny reddish brown, the

anterior tergites translucent, and with lateral round dark brown spots and me-

dian dark triangles on sternites 2 and 3.

Esenbeckia (Esenbeckia) translucens (Macquart)

Pangonia translucens Macquart, 1845, Mem. Soe. Sci. Lille (1844), p. 154,

Pi. TF ig.23-

Esenbeckia translucens: Fairchild, 1942, Ann. Ent. Soe. Amer., 35(2):

191-192, Pi, II, fig. 15, female, full references. Fairchild, 1971,

Cat. Dipt. S. Amer., 28, p. 9 synonymy. Wilkerson and Fairchild,

1983, Journ. Nat. Hist. 17: 555.

The eyes are greenish black in life. The second tergite is yellowish horn-

colored and bears a black mark in the middle, from a broad inverted triangle to

a thin streak, and the sides of this tergite are sometimes extensively dark; re-

mainder of abdomen black. The antennae are black, or dark reddish brown, and

the wings strongly infuseated.

The species is fairly common. Panama records are from the Atlantic coast

from Boeas del Toro to San Blas, and from Cerro Azul and El Valle de Anton. It

appears to prefer areas of heavy forest and high rainfall, and occurs up to about

2000 ft. elevation. It attacks man and horses eagerly, biting the latter primar-

ily on the belly. Time of flight is from May to September, with most records in

June and July.

The range is from Mexico to Brasil and eastern Peru.

Esenbeckia (Esenbeckia) prasiniventris (Macquart)

Pangonia prasiniventris Macquart, 1845, Mem. Soe. Sci. Lille (1844) p.

161. Pl. Ill, fig. 9.

Esenbeckia prasiniventris; Fairchild, 1942, Ann. Ent. Soe. Amer., 35(2):

197-198, Pl. II, Fig. 16, female, full references. Wilkerson and

Fairchild 1983 Journ. Nat. Hist. 17: 552.

The eyes are bright bluish green in life. Color of the abdomen varies from

blackish green in fed specimens to pale yellowish green. Size varies from 12 to

16mm. in wing length. Beard and pleural hair varies from white or grey to or-

ange. Males are paler, yellowish green, the eyes without enlarged facets and

the palpi short and porrect.

Common to abundant. Records are mainly from the Pacifie side of the

isthmus, but it has been taken in Boeas del Toro and on the Atlantic side of the

Canal Zone. A diurnal species with a preference for drier, open habitats at

lower elevations. Nearly all Panama specimens were taken in months of Dee. to

Mar., but scattered specimens in May, July, and August. Seldom taken in heavy

G. B. Fairchild: Tabanidae of Panama 25

forest or above 2000 ft. It attaeks larger domestic animals and man vigorously.

The species ranges from Guatemala to Brazil. It is possible that more that one

Species is involved here, but evidence is not yet conclusive.

Esenbeckia (Esenbeckia) osornoi osornoi Fairchild

Esenbeckia illota osornoi Fairchild 1942, Ann. Ent. Soe. Amer., 32(2):

196, P1. Il, fig, Il. Female, Panama.

Esenbeckia osornoi osornoi: Wilkerson and Fairchild 1983, J. Nat. Hist.

17: 547.

The eyes are yellowish bronze in life. The male is similar in color, the up-

per eye facets enlarged but not sharply demarcated from the small facets. The

male palpi are long, thread-like, about a third length of proboscis.

The species is fairly common. It occurs throughout the Canal Zone area, in

Darien Prov., San Blas, and Chiriqui Prov. on the coast. It is a ecrepuscular and

nocturnal species, often taken in mosquito stable traps and light traps, and ap-

pears to show a preference for localities near rivers and swamps. Time of

flight is from January through March, in the height of the dry season.

The species as a whole ranges from Costa Rica to southern Brasil, the sub-

species osornoi from Costa Rica to Colombia. I now feel that E. illota Will. is

a Central American species with a limited range and specifically distinct from

the complex of forms treated recently by Wilkerson and Fairchild as subspecies

of osornoi.

Genus Fidena Walker

Walker 1850, Ins. Saud., Dipt. 1: 8. Mackerras, 1955, Aust. J. Zool.,

3(3): 487-490, full references. Fairchild, 1971, Cat. S. Amer. Dipt.

Fase. 28, p. 16, synonymy. Wilkerson 1979, Cespedesia (Cali,

Colombia) 8(31-32): 143-156.

Members of this large and difficult genus are easily recognized in Panama

by the combination of pilose eyes, produced faces, long and slender proboscides

and open cell M3. All are species of areas of high rainfall and are seldom found

outside of heavy forest. Several are tree-top or forest canopy inhabitants.

Males of but one regional species are known.

Key to species

1. Legs prominently bicolored, the femora black, the tibiae and tarsi yel-

low. Wings black or heavily infuseated basally to ends of basal cells

sndleais sais «ERAT AND wa yh ata s dunce o LGUs Dilseshlis's kG ls s meSb de ighaGRE Shi hevccikecenncveuses o

Legs not bicolored, all black or yellowish. Wings rarely heavily infus-

cated basally, usually evenly tinted ...........cccsscsees iWin donheadh Leudteniniae

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Beard snow-white. Sides of mesonotum with a stripe of white hairs

from before suture to posterior margin. Abdomen shiny black, with

small tufts of silver-white hairs in middle of tergites 2 to 5, and at

Sides of tergites 2, 5, and 6. Sternites 2 to 4 with white lateral

hair tufts; 2 sometimes with white hind marginal band. Face par-

tially denudedsandeshiny iii... aivaesd si ibien doce le eu eriomeroides (p. 27)

Beard black or dark brown. Mesonotum without contrasting pale hairs.

Abdomen shiny black or deep brown with segments 4 or 5 to 7

clothed with pale straw yellow, rufous orange, brown or black hairs.

Tergite 2 with a pateh of white hairs at sides, and sternites 2 to 4

with small lateral white hair tufts. Rarely tergites 3 and 4 may

have small median white hair tufts .........ccccccssceees rhinophora (p. 29)

Legs pale yellowish brown. Beard, pleura and venter of abdomen pale

yellowish pilose, contrasting with dark dorsal SurfaCesS .......ccccsscsseees

a hiNlebn Saves tos bcs MISE DES a Leaanllsly HMR So tha Lidlevctes auribarba (p. A,

Legs dark brown to black. At least pleura dark PilOSeé.........scccccccscccscees

Beard brown to black. Abdomen largely black piloSe ...........scccccccceccceece 5)

Beard white. Abdomen extensively golden pilose ........sccccccsccccceccccccccccees 6

Large species, generally over 18 mm. in wing length. Frons narrow,

over 4.5 times as high as wide, convergent below. Antennae brown-

ish black. Face conically produced, brown, wholly grayish pollinose.

Wings uniformly deep yellowish brown tinted. Abdomen dark brown

to black, shiny, sparsely black pilose, with tufts of white hairs at

sides of tergites 2, 5 and 6 ..ccccccccece icunebedesbedevan sds flavipennis (p. 28)

Small species, wing length generally less than 12 mm. Frons broader,

less than 3 times as high as wide, parallel sided. Antennae bright

yellow. Face less produced, yellow, with extensive bare patches lat-

erally. Wings smoky hyaline, the costal cell blackish and end of

first basal cell darkened. Abdomen black, the second segment

Sparsely white haired above and below, forming a more or less dis-

tinct hind marginal band widened at sides above. Tergites 5-7

sparsely white haired and often with median white triangles on ter-

item: Ad. Aiccas sccetetin eosSedlonendnedgenptecnnes POE ans saeleid schildi (p. 30)

Face relatively short, shorter than frons and subeallus, largely shiny.

Antennae short and unusually broad, the basal annuli markedly wider

than long. Thorax black, generally with prominent white hair tufts

on each side of scutellum, rarely reduced to a few white hairs mixed

with black. Abdomen largely bright golden pilose, but with first

tergite black pilose, second with golden hairs forming a large median

triangle and lateral patches, third with only dorsolateral anterior

black patches, remainder wholly golden pilose. Wings yellowish

tinted, the basal cells and costal cell darker; first posterior cell

eogretate, rarely ClOSOC)) Guy sik ssitiinad ncasccoes siebeaganee trapidoi (p. 30)

Face relatively long, as long or longer than frons and subeallus; gray

pollinose above, shiny at sides. Antennae long and slender, the basal

annuli at least as long as wide. Thorax without white hair tufts be-

G. B. Fairchild: Tabanidae of Panama at

side scutellum. Abdomen bright golden pilose from third to last

segments only, with but a small patch of yellow hairs on posterior

lateral borders of second tergite. Wings evenly brownish yellow

tinted, the first posterior cell closed and petiolate .. howardi (p. 28)

Fidena auribarba (Enderlein)

Melpia auribarba Enderlein, 1925, Mitt. Zool. Mus. Berlin, 11 (2): 276,

female, Colombia.

Melpia auribarba var. albibarba Enderlein, 1925, Mitt. Zool. Mus. Berlin,

11(2): 276, female, Colombia.

Fidena auribarba and var. albibarba: Fairchild, 1958, Ann. Ent. Soc.

Amer., 51(6): 528-529, full references. 1971, Cat. S. Amer. Dipt.

Fase. 28, p. 18. Wilkerson 1979, Cespedesia 8(31-32): 145-146.

Melpia columbiensis Kroeber, 1930, Mitt. Zool. Mus. Hamburg, 44: 177,

female, Colombia.

Fidena columbiensis Kroeber, 1934, Rev. Ent., 4(2): 247. Fairchild, 1951,

Ann. Ent. Soe. Amer., 44(3): 442, Panama, full references.

The face is conically produced, mostly brown pollinose, but with a shiny

apex and subshiny lateral stripes. Antennae nearly black and quite long and

slender, much longer than palpi. Beard cream colored, the pleura more yellow-

ish. Legs yellowish brown. Wings uniformly dilute yellowish brown, first poste-

rior cell coarctate, rarely closed at margin, appendix short, usually absent. Ab-

domen black pilose above, with small median white hair tufts on tergites 2 to

0, rarely vestiges on 1 and 6. Extreme sides of all tergites, and whole venter

pale creamy white pilose.

All Panama material has come from the Atlantic side of the Isthmus or

from elevations above 2000 ft. Records are from Bocas del Toro, Chiriqui, and

Colon Provinces. The species attacks man readily and has been taken in tree

top collections as well as at ground level. All specimens seen have been taken

in September, October or November. )

I have studied the types of auribarba and var. albibarba on loan from Berlin

Museum through the courtesy of Dr. H. Schumann. They are indistinguishable

except for the color of the vestiture of beard and pleura, rich yellow orange in

auribarba, creamy white in var. albibarba. The latter appears to be the normal

form in Panama. The types and other Colombian specimens from Rio Raposo,

near Buenaventura, 13 March 1963, V. Lee coll. and Rio Yurumanqui in Paris

Museum are larger than most Panama specimens and with more produced faces.

The types of F. columbiensis in British Museum, were probably from the Choco

region of Colombia, next to Panama, and agree closely with Panama material.

Use of the name albibarba is precluded by another Fidena albibarba (Enderlein),

1925, p. 293, which, though later, takes priority over a variety, which has no

Standing under the International Rules.

Fidena eriomeroides (Lutz)

Epipsila eriomeroides Lutz, 1909, Zool. Jahrb. Suppl. 10, H. 4, p. 649,

fig. 27, S. Paulo, Brasil. Fairchild, 1961, Mem. Inst. Osw. Cruz,

§9(2): 217, Pid figh:6.

28 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Fidena eriomeroides: Fairchild, 1971, Cat. S. Amer. Dipt., Faces. 28, p.

19, synonymy. Wilkerson, 1979, Cespedesia 8(31-32): 147-148.

This species is easily separated from rhinophora, the only other regional

species with bicolored legs, by the white lateral hair stripes on mesonotum, usu-

ally prominent abdominal white triangles, and lack of contrasting pale dense

hairs on posterior segments of abdomen.

Records from Panama are as follows: Rio Mandinga, San Blas, 16, 17 and

19 May 1957, in tree tops, P. Galindo coll. 4 females; La Zumbadora, Cerro

Azul, Panama Prov., 12 June 1959, 1 female; Rio Paya Yellow Fever Station,

Rio Tuira, Dairan Prov., 2 July 1958, 1 female; Rio Tuira Y. F. Sta., Darien

Prov., R.P., 20 May 1958, in Shannon trap, 1 female. All localities are below

2500 ft. in areas of heavy rainfall and dense forest. The specimens range in

wing length from 11.5mm to 16 mm. The eyes are bright green in life.

I have seen the types of all the synonyms listed in 1971, as well as speci-

mens from Brazil (Para, S. Paulo), Colombia (Valle, Antioquia, Meta), Peru

(Tingo Maria), Ecuador (Napo) and Costa Rica (Puntarenas) and although some

variation is evident, it is not correlated geographically.

The male was deseribed by Wilkerson (loc. cit.) and another of this sex from

Monteverde, Puntarenas, C.R. agrees with his brief description.

Fidena flavipennis Kroeber

Fidena flavipennis Kroeber, 1931, Zool. Anz., 95(1-2): 24, fig. 8, male,

Venezuela. Wilkerson, 1979, Cespedesia 8(31-32): 148-149.

Fidena isthmiae Fairchild, 1941, Ann. Ent. Soe. Amer., 34(3): 642-643,

fig. 1, female, Panama.

The species is entirely black or dark brown, with small tufts of white hairs

before wing bases, on squamae, and on sides of second, fifth and sixth tergites,

occasionally on second sternite as well. This applies to all Panama specimens

seen, except that rarely the pre-alar white spot is faint. As pointed out by ©

Philip (1954) and in the original description of isthmiae, specimens from Mexico

may show more variation which Philip (1978) has decided warrants recognition at

the subspecies level. In Panama the species is occasional throughout the low-

lands up to 2500 ft. on both coasts, being the only species of the genus taken in

the Canal Zone area. It does not seem to be restricted to areas of heavy forest

and high rainfall, but has been taken in cleared areas and even in the suburbs of

Panama City. Records are at hand from the provinces of Chiriqui, Bocas del

Toro, Veraguas, Panama, San Blas, and Darien as well as from both coasts of

the Canal Zone. Specimens have been taken in every month from March to Oc-

tober. It seems not to be especially attracted to man, but has been taken biting

a large boa, and more recently 4 were taken biting a dead cayman (Crocodilia)

by C.O. Myers in Darien.

Fidena howardi Fairchild

Fidena howardi Fairchild 1941, Ann. Ent. Soc. Amer., 34(3): 640, fig. 5,

female, Panama; 1953, Ann. Ent. Soc. Amer., 45(2): 262, Pl. 1, fig.

G. B. Fairchild: Tabanidae of Panama 29

This searece species, in addition to the characters in the key, has the

scutellum reddish, white hairs on squamae, and often a small white spot before

wing bases, all lacking in trapidoi. Aside from the type material, taken at

about 6000 ft. in Chiriqui Prov., I have seen but 4 additional specimens, as fol-

lows: Almirante Yellow Fever Station. Bocas del Toro Prov., 1 Aug. 1951, taken

biting in camp, 1 female; Rio Changena, Bocas del Toro Prov., 2400 ft., 1 Sept.

1961, 1 female; Upper Rio Changuinola, Bocas del Toro Prov., 3000 ft., 15-25

Aug. 1960, R. Hartmann coll., 2 females. All these localities are in heavy rain

forest, the first being also the type locality of F. trapidoi. The types from

Chiriqui were taken in June, all others in August or September. A single speci-

men from Costa Rica, Paul Serre coll. 1922 is in the Paris Museum.

Fidena rhinophora (Bellardi)

Pangonia rhinophora Bellardi, 1858, Sagg. Ditt. Mess., 1. pp. 46-47, Tab.

1, fig. 1, female, Mexico.

Fidena rhinophora: Fairchild, 1953, Ann. Ent. Soc. Amer., 46(2): 262.

Wilkerson 1979, Cespedesia 8(31-32): 152-153.

Pangonia pyrausta Osten Sacken, 1886. Biol Centr. Amer., Dipt. 1, pp.

43-44, female, Panama.

Fidena pyrausta: Fairchild, 1941, Ann. Ent. Soc. Amer., 34(3): 644-645,

full references; 1951, Ann. Ent. Soe. Amer., 44(3): 441; 1956, Smiths.

Miseell. Colls., 131(3): 26, type seen.

Melpia auricauda Enderlein, 1925, Mitt. Mus. Berlin, 11: 276, female,

Colombia, Venezuela. Kroeber, 1930, Mitt. Zool. Mus. Hamburg, 44:

179, fig. 28.

Although not rare at suitable localities this species is rarely taken at eleva-

tions under 2000 ft. and always in areas of high rainfall. It attacks man readily,

and has been taken once biting pigs at 6 A.M. Records are available from sev-

eral localities in Chiriqui Prov. up to 6000 ft., in Bocas del Toro Prov., at El

Valle in Cocle Prov., and in the mountains of Darien Prov. from 1900 to 4800

ft. One of the specimens from Darien also was taken in early morning. Speci-

mens have been taken from March to September. I have seen specimens from

Mexico (Oaxaca), Colombia, Venezuela, Ecuador and Peru, and others have re-

ported it from Argentina.

There is a confusing amount of variation, both structural and tinctorial in

this species, but I have been unable to correlate these variations to define

clear-cut differences. A series of 15 specimens from Sta. Clara, Chiriqui aver-

age smaller than other material, with shorter, less produced faces. Four of

these have the abdomen entirely black pilose, except for small tufts of white

hairs at sides of tergite 2, while the remainder have the fifth to last segments

rufous orange pilose. All are somewhat rubbed, but several, including one of the

black pilose specimens, show vestiges of median white hair tufts on tergites 3

and/or 4. Four from Palo Santo, Chirqui, include 2 with the fourth to last seg-

ments densely orange pilose, one with these segments pale yellowish pilose, and

1 with only the fifth to last tergites orange pilose. The first 3 have long

snouts, the last shorter. One of the specimens with the fourth tergite orange

pilose is homotype of pyrausta. The short-snouted specimen and one of the

long-snouted ones bear vestiges of a median white pilose spot on tergite 3. Two

specimens from Rio Cana, Bocas del Toro have the fourth to last tergites or-

ange pilose but shorter snouts, while one from Rio Changena, Bocas del Toro,

30 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

has the same abdomen but a long snout. One specimen from El Volcan,

Chiriqui, has the hairs on fourth to last segment pale straw colored, as do two

from Taecarcuna, Darien. All three have long snouts. Three others from Tacar-

ecuna area also have long snouts, but only fifth to last segments with pale hairs,

and these deep orange rufous. One of these agrees with types of aurieauda, ex-

eept for a short appendix on fork of third vein and darker wing bases. The

specimen from Peru has pale yellowish hairs on fourth to last tergites and a

shorter snout, while the Ecuadorian specimen has only fifth to last segments

pale yellow pilose and a short snout. In general, the larger the specimen the

longer the snout.

It is possible that there are two closely similar sympatric species here, dif-

fering only in degree of prognathism, but with similar color variations. If so,

Statistical analysis of much longer series will be necessary to prove this. In any

ease, rhinophora, pyrausta and auricauda were all based on long-snouted speci-

mens. Fidena analis Fab. and F. basilaris Wied. also belong to this complex,

though they are both abundantly distinet, as a recent study of the types showed.

Fidena schildi (Hine)

Erephopsis schildi Hine, 1925, Oece. Pap. Mus. Zool. Univ. Michigan, No.

162, pp. 11-12. female, Costa Rica.

Fidena schildi: Fairchild, 1941, Ann. Ent. Soc. Amer., 34(3): 644, full

references; 1971, Cat. S. Amer. Dipt. Fase. 28 p. 23, synonymy.

Wilkerson 1979, Cespedesia 8(31-32):; 153-154, Colombia.

The characters in the key should serve to identify this species easily. It is

the smallest species of the genus in Panama. The eyes in life are brilliant pea-

cock blue. The species is fairly common in heavy forest in Bocas del Toro

province, attacking man at ground level, and once taken biting pigs before dawn.

Other reeords are: La Zumbadora, Cerro Azul, Panama Prov., 2000 ft., 13 Sept.

1956, 1 female, and Rio Tacareuna, Darien Prov., 1900 ft., 7 and 18 July, 1963,

2 females. Both Darien specimens were taken in the early morning, between 6

and 7:30 A.M. All our records are in the months from June to September. It

was known elsewhere only from Costa Rica, but I have recently seen 4 perfectly

typical examples labelled Charvein, Guyane Francaise, R. Benoist 1914, in the

Museum d'Histoire Naturelle, Paris, and Kroeber's types of the synonymous nigri-

eans in Halle are labelled Colombia and Brasil. Wilkerson (loc. cit.) reports it

from Antioquia, Colombia on the authority of C. Porter.

Fidena trapidoi Fairchild

Fidena trapidoi Fairchild 1953, Ann. Ent. Soe. Amer., 46(2): 260-262, Pl.

1, fig. 1, female, Panama.

Aside from the recognition characters given in the key, this species is en-

tirely deep black, the white beard and mesonotal tufts, and bright golden vesti-

ture of abdomen making it an unusually striking insect. The few additional

specimens taken since the species was described have all come from the same

locality,mostly from tree top collections, and all taken in May or June of 1951

and 1952. This locality is about 12 miles inland from Almirante, at the foot of

the cordillera at less than 1000 ft. elevation, in heavy rain forest. The arboreal

G. B. Fairchild: Tabanidae of Panama 31

habits and short flight season no doubt account for failure to collect it else-

where. The specimens were mostly taken attacking mosquito collectors on plat-

forms in the forest canopy, occasionally also at ground level. More recently, a

specimen labelled Finca La Selva, Puerto Viejo, Costa Riea, 3-VI-75, D. R. Perry

has been seen. It is the first collection of the species not from the type local-

ity and a new record for Costa Rica.

Genus Seione Walker

Walker 1850, Insect. Saundersiana, Vol. 1, P. 10.

The chief character separating this group from Fidena is the closed condi-

tion of cell M3. A number of species have quite hirsute faces and patterned

wings, conditions not found in Fidena, while a bare or partly bare frontoclypeus

is not known in Seione though frequent in Fidena. The center of distribution of

the genus appears to be the Andean highlands, and the bulk of the species are

mountain forms. A few species extend into Panama, one reaches Mexico, and

others occur in the highlands of Venezuela and the Guianas and in Southern

Brasil. A curious fact is the apparent absence of Seione aurulans Wied. in

Panama, as barely subspecifically distinct forms occur from southern Mexico to

Honduras and in Colombia. The nomenclature of the group is tangled, as the

Species are variable, numerous and separable with difficulty.

Key to species

1. Wings particolored, yellowish and grey, all crossveins with obscure dark

clouds. Thorax striped with yellow. Abdomen yellow to brown with

a middorsal row of yellow pilose triangles ....... maculipennis (p. 31)

Wings hyaline or tinted, not particolored and without clouds on cross

VEANS) Siebhss ies Te as iu cuhpscvaan daseushivenaaasessedes pee 4. MB is 2

2. Thorax with whitish stripes. Abdomen with tergites bearing median and

lateral posterior white hair tufts. Costal and basal cells yellowish

Ace ee bab idWdas cdhe Ua asia cdabecvaddedle Medien cabbunseash sete Caneel (a, 32)

Thorax without strongly contrasting stripes. Abdomen without contrast-

ing pale hair tufts ........sccee. Pa sakhacttggtedeesd bie HA he ha alee SUEUR “ae

3. Palpi broadly leaf-shaped, much shorter than third antennal segment.

Beard and pleura light brown, paler than mesonotum. Abdomen dark

brown, black pilose, with a median row of dark red pilose triangles,

venter paler, yellow pilose .......... edi uaed SRK Pise, ees ablusus (p. 34)

Palpi sickle-shaped, subequal to third antennal segment. Beard, pleura

and venter of abdomen black pilose. Abdomen with all tergites be-

yond first clothed with pale yellow to bright orange rufous hairs .....

‘sigh his th depen MRE da Ca scecscsscseseeee PUfEeSCENS (Dp. 33)

Scione maculipennis (Schiner)

Diclisa maculipennis Schiner, 1868, Reise Novara, Zool., II, Abt. 1, Vol.

B, Dipt., p. 102, Pl. Il, figs. 7, Ta, female, South America.

Fairchild, 1967, Pacific Ins., 9(2): 248, fig. 7.

32 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Seione maculipennis: Fairchild, 1942, Ann. Ent. Soc. Amer., 35(2): 189-

190, Pl. I, fig. 5, full references.

The species is variable in color and size, some specimens being quite pale

and washed out, others dark and boldly marked. The prominent hair tufts on

each side of the blackish scutellum may be pale straw colored, yellow, orange or

largely blackish. Legs vary from pale brown to nearly black. Beard may be

white, yellow or brownish. Wings may be brightly gray and yellow with promi-

nent dark clouds, or nearly uniform grayish with but a trace of yellow and

barely discernible clouds on crossveins. Profile of face and proportion of frons

are fairly uniform, but shape of palpi varies from broad and leaf-like to quite

slender with drawn out tip. ?

The species is common to abundant at suitable localities. Records are

available from the provinees of Chiriqui, Bocas del Toro, Veraguas, and Darien

at elevations from 1400 to 7000 ft. in areas of heavy rainfall. At higher eleva-

tions it seems to be the dominant Tabanid and attacks both man and domestic

animals avidly. Specimens have been taken in every month except May.

I have seen Schiner's types, and Panama specimens agree well with his de-

seription and poor figure. Although the species may be confused with other

Similar species, I have seen true maculipennis from Venezuela, Colombia and

Costa Rica.

Seione costaricana Szilady

Scione costaricana Szilady, 1926, Biol. Hung., 1(7): 29, female, Costa

Riea. Not Se. ecostaricana: Kroeb. 1930. Stett. Ent. Zeit., 91(2):

156.

Seione claripennis: Fairchild, 1942, Ann. Ent. Soe. Amer., 35(2): 188-189,

pl. 1, fig. 3, female, Panama, full reference. Kroeber, 1930, Stett.

Ent. Zeit., 91(2): 154-155, in part, Panama material only. Philip,

1954, Rev. Brasil. Ent., 2: 23, in key only. Not Se. claripennis Ric.

1900.

Specimens from Panama vary much in color of vestiture. Beard may be al-

most white, yellow, orange or practically black, and the hair tufts on each side

of scutellum show the same range of color, though the scutellum itself seems

always reddish. The abdomen may be light brown to nearly black, but is always

black pilose, with silvery white median hair tufts on tergites 1 to 5, on sides of

tergites 2, 4, and 5, and with a more or less complete white hind marginal band

on tergite 2. Venter of abdomen is brown to black with a white pilose hind

marginal fringe on second sternite. Legs are unicolorous, brown to black. Frons

and face are fairly uniform, but palpi vary in degree of attenuation of tips.

Wings are unspotted, hyaline, more or less yellow tinted basally and along fore

border.

This species is far less abundant and more restricted in range than mac-

ulipennis. All material seen, except the one from El Valle, Cocle Prov. listed in

1942, has come from the highlands of Chiriqui and neighboring Bocas del Toro

Provinees, at elevations over 2000 feet. The species has been taken in Jan.,

Aug., Sept., and Dec. and probably, like maculipennis, flies throughout the year.

So far as now known, the range is limited to Western Panama and Costa Rica.

Types of both ecostaricana Szil. and claripennis Ric. were in Budapest and

presumably destroyed. Study of types of strigata End. and brevipalpis End. on

G. B. Fairchild: Tabanidae of Panama 33

loan from Berlin Museum, from Ecuador and Bolivia respectively, indicates two

Similar but distinct species. Brevipalpis seems to agree best with the descrip-

tion of eclaripennis, with small black frontal patch, less protuberant face, and

dark scutellum. It was placed as a synonym of claripennis by Kroeber (1930),

and is certainly different from any Panama specimens. Se. strigata is very sim-

ilar to some Panama specimens, but differs in wider, more divergent frons, more

protuberant and whitish pollinose face, and in lacking white pilose band on sec-

ond tergite. A specimen from E. Peru agrees well with type of strigata struc-

turally and in lacking a white band on tergite 2, but has dark brown pollinose

face and yellow rather than white beard and scutellar hairs. The description of

costaricana compares the species with claripennis, but says that it has dark

spots around crossveins and no white fringes on hind borders of segments. In

the key in the same publication, however, the wings are given as without spots.

Szilady presumably had Ricardo's type before him for comparison. Kroeber

(1930) keys both claripennis and costaricana as having unspotted wings, but sep-

_arates them on basis of color of hair tufts at sides of scutellum, white for

claripennis, yellow for costaricana, but he had no Central American material of

the latter. It is also to be noted that Szilady in his key says of costaricana:

"Sides of scutum posteriorly with white hair tufts. Abdomen with median spots

of white hair." This agrees with some Panama material, but not with Kroeber's

remarks. Kroeber also lists material of eclaripennis from Panama. In view of

the loss of both Ricardo's and Szilady's types, certainty is probably impossible.

However, the fact that the only name based on Central American material is

costaricana, and that Panama material differs from the type of brevipalpis and

its probable senior synonym elaripennis and from other S. American specimens,

leads me to prefer this name for Panama material. It is possible that Szilady

had a weakly marked specimen of maculipennis as well as costaricana before

him, whieh would account for the ambiguities in his key and description, as the

species are quite similar except for the wings.

Scione rufeseens (Ricardo)

Erephrosis rufescens Ricardo, 1900, Ann. Mag. Nat. Hist., (7)6: 294,

Peru, Bolivia. Fairchild, 1956, Smiths. Miseell. Coll., 131(3): 27.

Type from Peru in Budapest presumably destroyed; paratype from

Bolivia in B.M. seen.

Rhinotriclista rufescens: Kroeber, 1930, Stett. Ent. Zeit., 91(2): 147 with

generosa End. as syn.

Seione rufescens: Kroeber, 1934, Rev. Ent., 4(2): 234. Wilkerson 1979,

Cespedesia 8(31-32): 173-174.

Rhinotriclista generosa Enderlein, 1925, Mitt. Mus. Berlin, 11: 302, Peru.

Fairchild, 1966, J. Med. Ent., 3(1): 13, fig. 20. Type studied.

Fidena fulvosericea Kroeber, 1931, Zool. Anz., 95: 26, fig. 11, female,

Colombia. Fairchild, 1956, Smiths. Miscell. Coll., 131(3): 17, type

in B.M. seen.

Scione aureopygia Fairchild, 1942, Ann. Ent. Soe. Amer., 35(2): 186-188,

Pl, 1, fig. 4, female, Panama.

As noted by Ricardo (1900), the species is quite un-Seione-like in general

appearance, which seems to have led to its repeated redescription. The greatly

produced face lacking long hairs, long curved palpi, unmarked wings and meso-

notum without prominent stripes resemble Fidena. The wings are peculiar in

34 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

having the membrane on both surfaces unusually glossy and finely wrinkled.

There is little variation in long series from Panama, except for the colors of

abdominal hairs, which range from pale straw yellow to deep rufous orange.

The species is very local, but may be abundant where found. Single speci-

mens are at hand from three localities in the lowlands of Bocas del Toro

Province, and 2 from Rio Mandinga, San Blas. There are also 2 from Porto

Bello, Colon Prov., 22 Apr. 1912, Buseck coll. in U.S.N.M. The species was ex-

eceedingly abundant at Tacarcuna Village, on the Rio Tacarecuna, Darien Prov.,

1900 ft., an area of heavy forest and high rainfall, and three were also taken at

Cerro Mali, 4800 ft. in the same area.

At the camp on Rio Tacarcuna I had the opportunity to observe the habits

of this species over a three-week period. The flies were active for only a very

short period just after dusk, from about 6:30 to 7:00 p.m., and to a lesser extent

at the corresponding period before dawn. They were attracted to man, and to

the warm part of a Shannon trap where it was heated by a lighted gasoline

lantern, but did not seem much affected by the light itself. Several specimens

when captured disgorged droplets of clear fluid, presumably nectar, as it was

sweet to the taste. None were captured at tree platforms, nor at other times

of the day. Seanty records, except for the long series from Darien, show the

Species to be on the wing from April to September. The species ranges from

Costa Rica to Bolivia.

Seione ablusus Fairchild

Scione ablusus Fairchild, 1964, J. Med. Ent., 1(2): 169-170, fig. 2.

The species is known from the type, taken biting man in heavy forest at

Cerro Campana, Panama Prov., 2800 ft., 2 July 1960, and another from El Valle

de Anton, 21 July 1963, R. Dressler coll. A specimen det. Se. rufipes Kroeb. by

Dr. J. Bequaert from Restrepo, Meta, Colombia agrees closely with Kroeber's

brief description and appears to be the most closely related species. It differs

from the present species in somewhat broader frons, red antennae, much less

produced face, broader and blunter palpi, much shorter and heavier proboscis,

and in lacking dark red median triangles on abdomen. The uniformly brownish

tinted wings, brown unmarked mesonotum and brown abdomen (the red pilose

median triangles are very inconspicuous) will, in addition to the characters in

the key, easily distinguish this species in Panama.

Genus Pityocera Giglio Tos

1896, Boll. Mus. Zool. Anat. Comp. R. Univ. Torino 11(224): 3, fig 1.

Fairchild, 1969, Arq. Zool. S. Paulo 17(4): 203-204.

Easily told from all other American Tabanidae by the bipectinate antennae.

The typical subgenus is monobasic and only the one species occurs in Panama.

G. B. Fairchild: Tabanidae of Panama 35

Pityocera (Pityocera) festae Gig. Tos.

Pityocera festae Giglio Tos, 1896, Boll. Mus. Zool. Anat. Comp. R. Univ.

Torino 11 (224): 3, fig. 1, female, Panama. Fairchild, 1942, Ann. Ent.

Soe. Amer., 35(2): 183-184, fig. 7, female, full reference. Wilkerson

1979. Cespedesia 8(31-32): 179-180.

This is a brown Fidena-like fly without conspicuous markings, but with

lightly tinted wings, and shiny bulbous face. The first posterior cell is closed

far from the margin. It is not uncommon in Panama from the Canal Zone east

into Darien, but the flight season is short, from July to Sept., with most speci-

mens taken in August. It is crepuscular and flies until after it is too dark to

see, as well as before dawn, but may be on the wing in heavily shaded areas

throughout the day. It is attracted to man, but does not bite readily. It has

been taken in horse-baited stable traps, Shannon traps, and biting horses, and

does not seem very limited in habitat requirements, as it has been taken in open

country, around town, and in heavy forest. The range is from the Canal Zone

area east and south to western Ecuador.

SUBFAMILY CHRYSOPSINAE

Tribe Chrysopsini

This tribe is the only representative of the subfamily ocurring in Panama,

and consists here of but 2 genera, Silvius (Assipala) and Chrysops. They are to

be distinguished from other Panamanian tabanids by generally small size, not

over 10mm in wing length, the wings with a dark pattern, the eyes in life either

speckled or with dark spots and stripes, the hind tibiae with spurs, the frons

broad, generally as wide or wider than high, with a frontal callus, and the ter-

minal antennal segment of a long basal part and a 4 annulate style. The scape

and pedicel are both generally long, together usually exceeding the terminal

segment.

Genus Silvius subgenus Assipala Philip

Assipala Philip, 1941, Canad. Ent., 73: 4, 9, in key only; 1941, Rev. Ent.

12(3): 470-473;1942 (1940), Rev. Chilena Hist. Nat., 44: 190-

194;1954, Rev. Brasil. Ent. 2: 58-59.

Silvius (Assipala): Mackerras, 1955, Austr. J. Zool., 3(4): 616, fig. 18.

This is a small group of slender long-winged flies resembling Chrysops in

many respects, but having the eyes irregularly speckled and with the third an-

tennal segment very short in relation to the first two. Only a single species oc-

curs in Panama, the group ranging thence north into southwestern U. S.

36 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Silvius (Assipala) tanycerus (Osten Sacken)

| Chrysops tanycerus Osten Sacken, 1886, Biol. Centr. Amer. Dipt., 1: 46-

47, female, Costa Rica. Kroeber, 1930, Zool. Anz. 90: 73-74, figs

11-12.

Assipala tanycerus: Fairchild, 1946, Ann. Ent. Soc. Amer., 39(4): 564,

Panama, full references.

The species is apparently confined to extreme western Panama, as specimens

have been seen only from El Volean, Chiriqui Prov., 15 Feb. 1943, T.H.G. Aitken

coll. and Potrerillos, Chiriqui Prov., 20 Jan. 1934, D.V. Brown ecoll. It has been

recorded from Costa Rica several times. The El Nolpan specimen was compared

and found to agree with the types in British Museum in 1953. It is a slender

brownish Chrysops-like fly with clouds around all ecrossveins and fork of third

vein as well as an irregular patch below stigma and along fore border at apex.

The first two antennal segments are very long, the third very short, less than

half length of either of the other two. It appears to have a short flight season,

since all specimens seen were taken in Jan. or Feb.

Genus Chrysops Meigen

Meigen 1803, Illiger's Mag. f. Insektenk., 11, p. 267. Kroeber, 1925-

1926, Konowia 4(3-4): 210-256, 319-375, 5 plates. Fairchild, 1942,

Proc. Ent. Soc. Washington, 44(1): 1-8, figs. 1-7. Philip, 1955, Rev.

Brasil Ent. 3: 47-128, keys. Mackerras, 1955, Aust. J. Zool., 3(4):

617-620, full generic references.

The genus is characterized by having generally bare facial callosities,

lengthened first two antennal segments, more or less sclerotized proboscis and

labella, eyes characteristically marked in life with purple spots and bands on a

bright green or golden ground, and wings usually with a dark ecrossband. All

Panama species are small, not attaining a wing length of 9mm. The group is

world-wide in distribution. In Panama, with certain exceptions, the species are

local and only occasionally abundant. They attack man readily, usually about

the head, and are more persistent in their efforts to bite than most local

Tabanidae. Thirteen species and 1 subspecies occur in Panama, discussed below

in alphabetical order.

Key to Species

1% Frontoclypeus with a median pollinose stripe. Wings with broad black

crossband, first basal cell about half black basally, apical spot drop

Shaped. Second abdominal tergite yellow with a thick black median

inverted V-shaped mark, and unconnected sublateral black spots ......

Wis aasinidacavnua dieeindeae eas Niinosivebd deep apahyeesdde Wiebdasnb seeders @hiriquensis (p.. 40)

PPORTOCIVOEUR GNLICRIY DAPS BAG BUN liiissenisersasneedeavonendiersescesecenaseensscas 2

2. Diseal cell largely hyaline, at least a central spot or streak hyaline...... 3

LEAL COU WOT: WANT MAO ACES J ilidssdhusnecesnehsbasieaduemnnsdebe ten culsnecnavds ends asuseuns 9

G. B. Fairehild: Tabanidae of Panama 37

Abdomen largely yellow, with 2 dorsolateral dark stripes from 2nd seg-

ment to 3rd, generally divided into 2 stripes on 3rd and succeeding

segments. Legs and frontal and facial callosities largely yellow.

Thorax yellow to brown, with yellow pollinose stripes. Wing pattern

yellowish! DrOWM: Weds PANE OA lee dadbl sli eed variegata (p. 45)

Abdomen with entirely black or dark brown integument. At least coxae

and femora largely black, as are frontal and facial callosities ....... 4

Abdomen with prominent grey pollinose transverse bands’ on posterior

margins of all tergites. Wing pattern of 2 dark bands crossing wing

at ends of basal cells and beyond distal end of diseal cell. Apical

spot a faint narrow infusecation along fore border. Apices of femora

and at least bases of tibiae yellow © ........cccccsecssscccscsnees alleni (p. 39)

Abdomen entirely black, at most slight gray pollinosity at sides of ter-

gite 2. Wing pattern fragmented, not as above. Legs brown to

edion NOL. DAT LIBOlOLEM. . acccccnkadoncenconnnsnaibuciaaianenduons reticulatus (p. 43)

Apical spot of wing a slender band or slightly widened, not extending

beyond apex and encroaching into second submarginal cell (4th R.)

OUP RE POREMO NG KAIS Ki) is Fiheeudis cse'd'eaieh advice dba cee 04 Gomelen dab ap sandsieinekssceccenss 6

Apical spot of wing extensive, filling most of wing apex and leaving but

a narrow and irregular hyaline crescent or series of spots between it

SAC CPOSSDONGE: wcsdeallidepecisaaiee din sbdve pwns MALLU capa Gkiinn div kek Caklidetédsineds 11

Fifth posterior cell (1IstCu) entirely infuseated. Fourth (3rdM) usually

with a small hyaline spot. Crossband broad and intensely black.

Thorax black, without dorsal stripes or lateral spots. Abdomen

black, sides of second tergite with small dull yellow triangles, and

with small dull yellow median triangles on tergites 2 to 4 — ..cccccccoeee

Y tepeed hs Ab MR cB ets een} leucospilus (p. 41)

Fifth posterior cell always with at least apex hyaline, often over half

hyaline. Fourth with or without a hyaline Spot .......c.ccccccccccscccccccces 7

Mesonotum and pleura blackish, without stripes or spots. Hyaline area

in fifth posterior cell clearly extending into fourth cell, but not

crossing it. Abdomen black with small dull yellow triangles laterally

on second segment and median yellow triangles on tergites 2 to 5 ...

Linn taiphbnpt ea ndad@inkikuandanulanesiads Web iehs i dee te melaena (p. 41)

Abdomen with a dorsolateral series of pale dashes, forming a pair of in-

terrupted stripes, as well as a row of pale median triangles on ter-

gites 2 to 5. Sides of tergites 1 to 3 or 4 yellowish. Abdomen be-

neath yellowish, with a broad even black longitudinal stripe. Cross-

band of wing with outer border irregular, fading posteriorly and

failing to reach hind margin in fourth posterior cell sealaratus(p.43)

Abdomen without dorsolateral stripes or even ventral stripe. Outer

margin of crossband straight or slightly concave, the crossband fill-

ing apex of fourth posterior Cell .......cceeoe. Bea a Nel hs il eeesedubans 9

10.

11(5).

13.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Second tergite black with yellow median hour glass-shaped mark ex-

tending longitudinal width of tergite, the sides broadly yellow. Third

to fifth tergites black with yellow hind borders, widened into narrow

median triangles which usually reach anterior border of segments.

Wings with broad dark crossband, without hyaline fenestra in first

submarginal cell (3rdR), and with hyaline area in posterior margin of

erossband confined to fifth POSterior Cel] .......cccccccccccccccccccccscccccees 10

Second tergite black, broadly yellow at sides, very rarely with a small

median yellow spot, generally without any median mark. Tergites 3

and 4, black, with equilateral yellow triangles whose apices may

reach anterior borders, tergite 5 with yellow hind border widened

into a low triangle in middle. Wings with narrower crossband, the

hyaline area in posterior margin of crossband large, filling all but

base of fifth posterior cell and entirely crossing fourth posterior cell

in middle. First submarginal cell with small hyaline streak at base .

sUETe op enue sberee re sabsvnrseveelelsciebosssebtsestsemetts cauroguttata (Dp. 39)

Apieal spot black, extending as a clearly defined narrow costal stripe of

even width to Wing APeX .......ccccsccsssseeseee Varians var. tardus (p. 44)

Apical spot dilute, ill defined, spread as a brownish shade over much of

wing apex ..... eodde eale ce Vateidedebegsbiadsscdeveshens varians ssp. vargus (p. 44)

Hind femora at least one-third basally black...........cccccccccccccccccccece nae 12

Hind femora red or yellow, at most extreme apex black...... jeiaucobececees 13

Abdomen black with second segment anteriorly narrowly, laterally

broadly yellow, and with a broad yellow median triangle which rarely

extends through the black to join the anterior yellow band. Tergites

3 to 5 or 6 with narrow yellow hind borders, widened into median

triangles of variable size on 3 and often 4. Pleura usually with a

spot of yellow pile, rarely lacking. Wings with crossband joined to

eloud on fork of third vein by a dark bridge above the vein,leaving a

small hyaline spot between cloud and Crossband ON VEIN ...ccccccccscceees

a boaeatcenucosesadeerassnerceciwedssnedeasdctecsscyenden sax sseccscesceeeeeeCalogaster (p. 40)

Abdomen wholly black, or rarely with the pattern of calogaster faintly

indicated in grayish pollinosity. Pleura always all black. Cloud on

fork of third vein never joined by a complete bridge to crossband,

rarely almost joined DY SPUPS .ccccccceseee Hledudcieeiaeluedcunetnes. soror (p. 43)

Cloud on fork of third vein joined to crossband by a broad spur of dark

color. Midtibiae black or blackish, contrasting with pale femora.

Abdomen with second tergite narrowly white to yellow anteriorly,

widened to full length of segments at sides and with a generally

broad, small median yellow triangle. Third and fourth tergites

rarely with faint se of small pale median triangles ...... meoneucless

lcweaterivedtvbedeess PAL doce ssisvac leet esse vorblsdeluiwederd Civckebebeshes MOMMIES AD. Ad)

Cloud on fork of third vein unconnected with crossband. Midtibiae at

most slightly brownish, not strongly contrasting with pale femora.

Abdomen as above, but pale triangle on second tergite usually taller

and narrower, occasionally joined to anterior yellow by a narrow

median yellow stripe. Third tergite usually with a dull yellowish

median streak, sometimes lacking, and fourth rarely with a minute

RIE SEPOBIC’ \.spacuveuredeeeeceden UE Rr re RTE IC EE ENS Fos seeee Mexicana (p. 42)

G. B. Fairehild: Tabanidae of Panama 39

Chrysops alleni Fairchild

Chrysops alleni Fairchild, 1939, Proc. Ent. Soc. Washington, 41(9): 257-

258, fig. 1, female, male Panama; 1942, Proc. Ent. Soc. Washing-

ton,44(1): 5. Philip, 1955, Rev. Brasil. Ent., 3: 49, 72, female, male,

in key. :

This is the smallest Chrysops in Panama, some specimens having a wing

length under 5 mm. The species seems confined to mangrove swamps on the

Pacific coast of the Isthmus, from Darien (Garachine) to Los Santos (Tonosi). It

has been taken in February, April, May, and September, hence probably flies

through the year. It may be very abundant at suitable localities and flies about

the collector's head, though it seems reluctant to bite. Males have been taken

resting on foliage in sunny spots along the edge of mangrove swamps, and in a

mosquito light trap.

Chrysops auroguttatus Kroeber

Chrysops auroguttatus Kroeber, 1930, Zool. Anz., 90(3-4): 71-72, figs. 6-

8, female, Colombia, Trinidad; Fairchild, 1946, Ann. Ent. Soc.

Amer., 39(4): 565, Panama, Colombia; Philip, 1955, Rev. Brasil Ent.,

3: 68, 80, 88, male, Panama. Wilkerson, Cespedesia 8(31-32): 182-

184, fig. 52.

Chrysops auroguttata var. pallidefemorata: Pechuman, 1937, Rev. Ent.,

7(2-3): 136, Panama. Not Kroeber 1930.

Chrysops incisa: Fairchild, 1942, Proc. Ent. Soe. Washington, 44(1): 3-4,

fig. 3, female, Panama. Not Macquart 1846.

All Panama material seen is auroguttatus. The closely similar pallide—

femoratus may be recognized by its broader frons with definitely transverse bar-

like callus, and by having the second and third antennal segments considerably

shorter than in auroguttatus. Pallidefemoratus usually has a yellow triangle on

second tergite and usually lacks the hyaline spot in base of second submarginal

cell (R2), while in auroguttatus these conditions are reversed. These color char-

acters and the color of the legs are not wholly reliable, while those of structure

have held in all material seen.

In Panama the species is quite abundant on the Atlantic side of the Canal

Zone in the Ft. Sherman area, and has been taken at a number of other locali-

ties from San Blas to Bocas del Toro, all at low elevations on the Atlantic

coast. It was not secured at our stations in the foothills of Boeas del Toro

Province. It has been taken in all months from January to October and probably

flies throughout the year. It attacks man persistently and enters horse-baited

mosquito traps and light traps.

Auroguttatus has been seen from Vera Cruz, Mexico (in Coll. L. L. Pechu-

man), Costa Rica, Panama, Colombia and Trinidad; pallidefemoratus from Yu-

catan, British Honduras, and Trinidad, a curiously discontinuous distribution.

40 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Chrysops calogaster Schiner

Chrysops ecalogaster Schiner, 1868, Reise Novara, Zool., Il, Abt. 1, Vol.

B. Dipt., p. 103, female, South America. Fairchild, 1942, Proc. Ent.

Soe. Washington, 44(1): 7, fig. 4, female, Panama, full references.

Philip, 1955, Rev. Brasil.Ent., 3: 115-116; Goodwin and Murdoch

1974, Ann. Ent. Soe. Amer., 67(1): 98 fig. 23. Wilkerson 1979, Ces-

pedesia 8(31-32): 184, fig. 46.

This species is separable from soror largely on the basis of the yellow

banded abdomen. About two-thirds of the available specimens from Panama have

a yellow pollinose patch on the mesopleura, while in the remainder this area is

dark grayish. The abdominal pattern is also quite variable. Two specimens have

the yellow triangle on second tergite joining the anterior yellow band of this

tergite in the middle, while in the others the triangle varies much in size. One

very dark specimen has the yellow bands on tergites 3 to 5 very narrow and only

evident in the middle. All have the crossband joined to the spot on fureation by

a slender dark streak above the vein.

In Panama the species is uncommon but widespread. It has been taken in

the lower mountains in Darien Province, on the Atlantic side of the Canal Zone,

in San Blas, and in Chiriqui Province at a low elevation. All localities are in

areas of relatively high rainfall. The species ranges from Costa Rica to

Venezuela and Peru, though it seems nowhere common. The specimens reported

by me (Fairchild 1961) from Brasil are not calogaster, as they have red femora

and lack the dark connection between crossband and spot on fureation. Goodwin

and Murdoch (1974) have deseribed and figured the pupa, found in a small slug-

gish stream among dead leaves.

Chrysops chiriquensis Fairchild

Chrysops chiriquensis Fairchild, 1939, Proc. Ent. Soe. Washington, 41(9):

259-260, figs. 2-3, female, Panama, Guatemala;1942, Proc. Ent. Soe.

Washington, 44, (1): 5. Bequaert and Renjifo Salecedo,1946, Psyche,

53(3-4): 58-59, male, Colombia. Philip, 1955, Rev. Brasil. Ent., 3: 52,

74, male,female, in key only. Wilkerson 1979, Cespedesia 8(31-32):

186-18,fig.49.

Chrysops subeaecutiens: Hine, 1925, Occ. Papers. Mus. Zool. Univ.

Michigan, No. 162, p. 20, Boquete, Chiriqui.(In part,not Bellardi).

This rare mountain species should be easily recognizable from the characters

given in the key. It is a large,robust, hairy species, with drop-shaped apical

spot and boldly marked abdomen. Philip (loc. cit.) has given the male charac-

ters in his key. In Panama the species has only been taken a few times, the

holotype and Hine's specimens at Boquete (in 1939) and another female, Palo

Santo, Chiriqui, 15 April 1964, both at elevations over 3000 feet. I have seen

other specimens from Costa Rica, Chiapas, Mexico and Guatemala, while Be-

quaert and Renjifo report a male from Colombia. The first abdominal segment

may be extensively pale in Mexican and Guatemalan specimens, though wholly

blaek in both those from Panama.

G. B. Fairchild: Tabanidae of Panama 41

Chrysops leucospilus Wiedemann

Chrysops leucospilus Wiedemann, 1828, Auss. Zweifl. Ins. 1: 202, female,

Brasil. Philip, 1955, Rev. Brasil. Ent., 3: 155. Fairchild, 1971, Cat.

S. Amer. Dipt.Fase. 28, p. 32, synonymy. Wilkerson 1979, Ces-

pedesia 8(31-32): 188-189, fig. 55.

The single Panama specimen of this species was taken on the Rio Tacarcuna,

Darien Prov., 1900 ft., 6 July 1963, in heavy forest. It differs from all other

specimens I have seen in entirely lacking any hyaline in fourth or fifth posterior

cells and in having the yellow patches at sides of second tergites smaller and

duller than usual. In South American material the extent of the hyaline spot in

fourth posterior cell varies considerably, being very minute in some specimens

from Villaviceneio, Colombia, and quite large, at least half width of cell in

specimens from Paraguay and southern Brasil. The record from Panama repre-

sents a large extension of range. The species was reported by Wilkerson (loc.

cit.) only from the eastern side of the Andes, and his specimens had very small

hyaline spots in fourth posterior cells.

Chrysops melaenus Hine

Chrysops melaena Hine, 1925, Occ. Pap. Mus. Zool. Univ. Michigan, No.

162, p. 17, female, Costa Rica, Venezuela. Fairchild. 1942, Proc.

Ent. Soe. Washington, 44(1): 3, fig. 2, female, Panama, full refer-

enees. Fairchild, 1953, Ann. Ent. Soe. Amer., 46(2): 259-260, male,

Panama. Philip, 1955, Rev. Brasil. Ent., 3: 68, 80, in key only.

Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer.,67(1): 100, fig.

24. Wilkerson, 1979, Cespedesia, 8(31-32): 189-191, fig. 48. |

In addition to the characters given in the key, the species has the frontal

and facial callosities largely or wholly yellow, the femora bicolored, black

basally, reddish yellow apically. It is the commonest and most widespread

species of the genus in Panama, occurring mainly in heavy forest, from sea-level

to 3000 ft. along the Pacific coast from Darien to Chiriqui provinees. There are

numerous reeords from the Canal Zone, a few from Colon and San Blas, but it

has not been taken in the Boeas del Toro area in spite of intensive collecting.

It is especially frequent at El Valle, and in forested areas in the Canal Zone.

We have a single record from Coiba Island, off the Pacific coast of Veraguas

Province, one of the few Tabanidae recorded from there. It attacks man readily

and is taken in horse-baited stable traps. The single recorded male was taken in

a light trap. We have taken it in every month save October, so that it probably

flies throughout the year. The range is from Costa Rica to Venezuela. Good-

win and Murdoch (1974) deseribed the pupae of both sexes.

It is probable that future collecting will show that melaenus is but a race of

C. latifasciatus Bell. The two are separated only by the extension of the hya-

line area in the fifth posterior cell into the fourth posterior cell in melaenus,

while it is confined to the fifth posterior cell in latifasciatus. Occasional

specimens of the latter show a small subhyaline spot in the cell. Latifasciatus

has been seen from Nicaragua, Honduras, Guatemala and the states of Chiapas

and Tabaseo, Mexico. The few Costa Rican specimens seen have been melaenus.

I have not seen the specimens from Nicaragua reported as melaenus by Woke

(1947); my two from that country are both latifasciatus.

42 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Chrysops mexicanus Kroeber

Chrysops calogaster var. mexicana Kroeber, 1925, Konowia, 4: 248, Pl.

II, IV, male, female, Mexico.

Chrysops mexicana: Fairchild, 1942 Proc. Ent. Soc. Washington, 44(1): 8,

fig. 6, Panama, in part; 1946, Ann. Ent. Soe. Amer., 39(4): 565-566.

Philip, 1955, Rev. Brasil. Ent., 3: 109-110, types seen. Wilkerson

(1979, Cespedesia,8(31-32): 191-192, fig. 53.

As suggested previously (Fairchild 1946), Panama material represents two

closely similar species, separated in the key. Fig. 6 in the 1942 publication

represents mexicanus, which averages larger and seems to have a different dis-

tribution. The bulk of our material has come from several localities in Bocas

del Toro province, with lesser numbers from the mountains in Veragus and Cocle

provinces, and 2 from the eastern part of Colon provinee. All specimens were

taken in areas of high rainfall and heavy forest, mostly between 1000 and 3000

ft. elevation. In localities where forest canopy collections were made it was

only once taken in the treetops.

Aside from Panama specimens, I have seen 2 from Chiapas, Mexico in Coll.

L. L. Pechuman, one of which was compared with the type by Dr. Philip, who

noted that it had more yellow on abdomen than the type. It is also more yellow

than Panama specimens, having the first abdominal segment extensively yellow

and the triangles on third and fourth tergites large and conspicuous. Dr. Philip

also compared a specimen from Almirante with Kroeber's types and noted that

the hyaline crescent was wider in the Panama specimen and that it laeked the

narrow dark spur along the vein from crossband to fork. Both these are variable

characters. I have also seen a third female, Rio Raposo, near Buenaventura,

Colombia, 25 Nov. 1963, taken at a tree platform 13 meters above ground level,

V. Lee leg., which agrees closely with Panama material. It was one of 11

specimens taken at that locality (Lee et al. 1969). The species thus ranges from

southern Mexico to Western Colombia, as reported by Wilkerson (1979).

Chrysops nexosus Fairchild

Chrysops nexosus Fairchild in Wilkerson 1979, Cespedesia 8(31-32): 192-

194, fig. 45.

Chrysops mexicana: Fairchild 1942, Proc. Ent. Soc. Washington, 44(1): 8,

fig. 7, in part; 1946, Ann. Ent. Soe. Amer., 39(4): 565-566, in part.

Not Kroeber 1925.

Holotype female, Santa Fe, Veraguas province, Panama, 9 Aug. 1950, taken

at platform in forest canopy 26 feet above ground level.

Paratypes, 4 females, same locality as holotype, taken 17, 26 July 5; 10

Aug. 1950, all at ground level; 3 females, Cerro Tute, vic. Sta. Fe, Veraguas

prov., 20 Mar. 1947;1 female, Almirante, Bocas del Toro Prov., 11 July 1957, in

Shannon trap at Yellow Fever Station; 1 female, Ft. Sherman,C.Z., 14 June

1947; 1 female,Rio Pequeni, Madden Lake, Colon Prov., 21 Aug. 1940; 1 female,

La Victoria, Cerro Azul, 2400 ft., Panama Prov., 23 Jan. 1947; 2 females, Cerro

Santa Rita,Colon Prov., 6 May 1952; 4 females, Rio Mandinga, Colon Prov., 23

G. B. Fairchild: Tabanidae of Panama 43

Sept. 1956 and 25, 27 Jan. 1957; 2 females, Pito, San Blas, 20 June 1943. Holo-

type and some paratypes to be deposited in F.S.C.A.

This species is to be separated from the other Panama species by the char-

acters given in the key. From C. rossi Philip and C. fairehildi Philip, it differs

in having all femora yellow on at least basal two-thirds. From rossi it differs

further in less infusecated basal cells, and from fairchildi in having yellow spots

on pleura and lacking yellow hind marginal bands on any tergites.

Chrysops reticulatus Wilkerson

Chrysops reticulatus Wilkerson 1979, Cespedesia 8(31-32): 195-197, figs

28, J-4,5, female, Panama, Colombia.

Chrysops soror: Lee, Fairchild and Baretto, 1969, Caldasia 10(49): 445.

Not Kroeber 1925.

An almost wholly black species indistinguishable from C. soror Kroeb. except

by the fragmented wing pattern, all cells in the area of the cross band with

hyaline centers, whereas in the apical wing area all cells have dark central spots

or streaks.

I at first thought the specimens I had seen were but aberrant specimens of

soror, but Dr. Wilkerson studied additional Colombian material and feels that

these specimens represent a distinct species. The only Panama specimen known

is the holotype, taken at Rio Mandinga, San Blas, 13-V-57.

Chrysops sealaratus Bellardi

Chrysops sealaratus Bellardi, 1859, Ditt. Mess. Pt. 1, p. 72, Pl. 2, fig.

19, female, Mexico. Fairchild, 1946, Ann. Ent. Soe. Amer., 34 (4):

964-565, Panama, full references; Philip, 1955, Rev. Brasil. Ent., 3:

68, 77, M, F, in key only.

This is the largest species of the genus occurring in Panama. It is easily

recognized by the irregular step-like outer margin of the crossband, which also

fails to reach the hind margin of the wing in full intensity, and by the three

rows of light spots on the abdomen. In Panama it has been taken on the coast

of Bocas del Toro Prov. around Almirante, and at Robalo, on the Chiriqui La-

goon. It does not occur in heavy forest inland, but seems confined to the vicin-

ity of coastal fresh water swamps. It bites man and horses and may be abun-

dant and annoying. Our records indicate captures from January to May and in

August, but it probably flies throughout the year. The range extends northward

on the Atlantic coast into southern Mexico. Recent collections in Boeas del

Toro Prov. included a number of males taken in U.V. light traps. This sex is

similar to the female, differing chiefly in more extensively infuscated basal cells

and in having sublateral dark abdominal stripes reduced to a slender dusky line.

Chrysops soror Kroeber

Chrysops soror Kroeber, 1925, Konowia, 4; 245, Pl. 2, female,

Venezuela. Fairchild, 1942, Proc. Ent. Soe. Washington, 44(1): 7-8,

Ae Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

fig. 5, Panama, Guatemala, full references; 1946 (1947), Ann. Ent.

Soe. America, 39(4): 565. Philip, 1955, Rev. Brasil. Ent., 3: 59, in

key.

Easily recognized by being the only all blaeck-bodied species in Panama, ex-

cept for the aberrant C. reticulatus Wilk. Occasional specimens show faint grey

middorsal triangles and lateral spots on second abdominal tergite.

In Panama the species occurs from Darien to Chiriqui, mostly in the moun-

tains over 2000 ft., but also at lower elevations on the Atlantic coast. None

have been taken in the Canal Zone, and in general it does not occur with ecalo-

gaster. Dates of capture include all months but April and September.

The species ranges from Guatemala to Venezuela and Colombia. I have also

seen material from British Honduras (Belize), Hells Teeth Creek, 6 Oct. 1955,

Galindo and Trapido, and Honduras, Lancetilla, 23 Nov. 1953, W. Hils.

Chrysops varians var. tardus Wiedemann

Chrysops tardus Wiedemann, 1828, Auss. Zweifl. Ins. 1: 577.

Chrysops varians var. tardus: Fairchild, 1971, Cat. S. Amer. Dipt., Faces

28, p. 34, synonymy. Wilkerson 1979, Cespedesia 8(31-32): 198-199,

fig. 51. )

The species most resembles C. melaena of the local fauna, differing in hav-

ing a large yellow triangle on tergite 2 of the abdomen which joins the yellow

base of the segment, in having all tergites with posterior borders yellow, and in

lacking hyaline in 4th posterior cell of wing. It has been taken but once in

Panama, at 1500 ft. on Cerro Pirre, Darien Prov., where 7 females were taken

in a Shannon trap from 27 Jan. to 6 Feb., 1961. The species and its variety

range south to Argentina, and it seems to be one of the commonest species of

the genus throughout much of its range. Typical C. varians is paler, with more

yellow on abdomen and often a reddish scutellum; it seems rare in the northern

parts of the range of the species.

Chrysops varians ssp. vargus n. ssp.

Differs from varians var. tardus only in the following respects. Apical spot

diffuse, more dilute than crossband and fading out posteriorly so that in some

specimens the whole wing apex is faintly tinged, leaving only a restricted com-

pletely hyaline stripe bordering the crossband. Yellow markings of abdomen on

the average less extensive, so that median yellow triangles on tergites 3 and 4

rarely extend to anterior margins of these tergites.

Holotype female, vic. Chitaria, Turrialba-Siquirres, Limon Prov., Costa Rica,

18 Feb. 1965, R. Dressler coll. Paratypes: 2 females, same locality and collec-

tor, 22 Feb. 1965; 1 female 10 mi. N.W. Pto. Jimenez, Costa Rica, 16 Aug.

1957; 2 females 5 mi. S.W. Sta. Clara de San Carlos, Alajuela Prov., Costa Rica,

400 ft. el. 19 Feb. 64, H. E. Evans coll.; 1 female Golfito, Costa Rica, 15 Aug.

1957, Arnold Menke ecoll.; 1 female Almirante, Bocas del Toro Prov., Panama, 2

Oct. 1951, in Shannon trap at ground level. The name is Latin for "vagabond" in

allusion to its distribution. The holotype will be deposited in F.S.C.A., 1

paratype each in collections L. L. Pechuman, C. B. Philip and G. B. F. 2

paratypes are in Los Angeles County Museum of Natural History.

G. B. Fairchild: Tabanidae of Panama 45

This appears to be an additional case of discontinuous distribution where the

range of a species is broken by the isthmian area of Panama. Typical varians

var. tardus is found only in extreme eastern Panama, the present subspecies in

extreme western Panama, and neighboring Costa Rica.

Chrysops variegatus (De Geer)

Tabanus variegata De Geer, 1776, Mem. pour Servir a l'Hist. des Ins., VI,

p. 230, P1.XXX, figs. 7, 8, female, Surinam.

Chrysops_ variegata: Bequaert, 1940, Rev. Ent., 11(1-2): 276-279.

Fairchild, 1942, Proe. Ent. Soe. Washington, 44(1): 4-5 fig. 1. Philip,

1955, Rev. Brasil. Ent., 3: 66, 85, female, male, in key.

The numerous references and synonyms of this common and widespread

species are listed by Bequaert (1940).

This species is quite variable in color. The hyaline crescent may be either

well marked or reduced to a diffuse paler area along outer border of crossband,

only the rounded spot in first submarginal cell being distinct. The abdominal

blaek dorsolateral stripes may be broad and intense, or narrow and faint, clearly

divided into stripes on each side at third tergite, or not until fourth. Outer

border of crossband may be straight, convex or slightly sinuous. Whether these

variations represent separate populations, as some have thought, or only fortu-

itous variations, will need careful and probably statistical analysis of large num-

bers to clarify.

In Panama the species occurs in small numbers throughout the lowlands of

the Pacific Coast and the Atlantic coast in the Canal Zone. It has not been

taken in areas of heavy forest, such as at the Almirante Yellow Fever Station,

or the forest stations in Darien, but seems to show a preference for low swampy

terrain along rivers, and coastal mangrove swamps. It bites man and horses

readily, and both sexes have been taken in mosquito light traps. Our records

are from all months of the year save March, so that it probably has no definite

flight season. The species ranges from southern Mexico and Cuba south through

the Antilles and Central America to northern Argentina and Paraguay.

SUBFAMILY TABANINAE

Tribe Diachlorini

The Diachlorini are a structurally diverse group and a dominant one in the

Neotropics. In addition to the characters given in the key, Diachlorini are often

mimetic, resembling Hymenoptera or other families of Diptera. Their wings are

often boldly patterned, ocelli or ocellar tubercles are frequently present, the an-

tennae are frequently modified, their labella often strongly sclerotized. This

plethora of structural and tinctorial specializations, combined with the retention

of other characters such as ocelli and bare basicostas has led to the belief that

they are the more primitive of the 2 tribes of Tabaninae ocurring in the

Neotropics. It has also led to the creation of many named segregates in con-

trast to the Tabanini. There are 59 genera and subgenera of Diachorini in the

Neotropics compared to only 5 of Tabanini. In Panama the corresponding num-

bers are 25 and 2.

46 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Genus Stenotabanus Lutz

Lutz 1913, Brazil Medico, 27(45): 6. Fairchild, 1942,Ann.Ent. Soc.

Amer., 35(3): 297-298, full references. |

The three subgenera as defined in the key are treated separately below. All

the species of the nominate subgenus are small to moderate in size, seldom ex-

eeeding 10 mm. in wing length and generally smaller. Colors are diverse, the

abdomen banded, spotted or striped, the wings generally clear, occasionally with

spots on crossveins or tinted, but never with a pronounced dark pattern. The

eyes are always banded, the frons usually fairly broad. Known males have the

upper eye facets greatly enlarged and sharply demarcated from the small facets.

All the species have short proboscides with soft, unsclerotized labella and short

inflated palpi.

_ Subgenus Stenotabanus Lutz

Key to females

1. Femora yellow. Abdomen pale brown with a broad yellowish white stripe

from base to apex. Wings slightly brown tinged. Eye purple with 4

NALrOW Lreen DAaNnds ...ccccccccceee Shes ctlcsvieuuessoces verses’ fulvistriatus (p. 47)

Femora black or brown. Abdomen without median pale Stripe ........sccees 2

Ze Legs entirely dark. Mesonotum with a pair of prominent dorsolateral

whitish stripes and the black scutellum bordered with white. Ter-

gites blackish, 2 to 6 with narrow pale hind borders, 1 and base of 2

bluish pruinose. Wings glass clear. Eyes purple with two narrow

PTeEeN DANS ..cecceccsseees indncavvasssupueavbauetee ocebuue epeaseaae minuscula (p. 48)

All tibiae basally pale.........scov osetcuepes Wheceesnses teeeucneey peiebaaeiy Padesre ks twesiseed

3. Subeallus inflated, bare and _ shiny. Mesonotum sparsely, scutellum

densely clothed with bright golden hairs. Abdomen with hind borders

of tergites 2 to 6 banded with bright golden hairs, the band narrow

on 2, broad on the remainder and broadest in the middle. Eye pur-

ple with 2 narrow green bands and the upper and lower borders

greenish purple ....... a bia oie s sais ee tains elec ». ealvitius (p. 47)

Subeallus pollinose, or if partly bare, then abdomen black and wings

deeply fumose ........ divabiesNeneboslusliaes eer a error rs iat aN ee one ahaha aces

4. Abdomen with prominent pale hind marginal bands on tergites 2 to 6.

Frontal callus about aS wide aS Nigh ....ccccccccccccscseccssoseees sitecetseieacces 3

Abdomen with faint hind marginal bands or none. Frontal callus clearly

higher than wide ......cc0.«. Pe pcnieenccas send iste beens Aah sib becanpestees np RON Ha 6

De Antennae wholly orange. Mesonotum unstriped, steel grey pollinose,

with numerous greenish shining secale-like hairs. Palpi blackish, at

least mainly black-haired. Eye reddish bronze with 2 green bands

Gers ekatuewecdes Coy Le ALR ee ie Pee estusk ees ...-+-incipiens (p. 48)

G. B. Fairehild: Tabanidae of Panama 47

Antennae with at least style black, basal plate usually apically dusky.

Mesonotum brownish pollinose, with a pair of broad golden-haired

dorsolateral stripes and scutellum prominently pale on posterior half.

Palpi pale yellow, wholly pale haired. Eye purple with 2 green

bands ...... lpenetcsaeeiediebiieesee. ssiinee oe tidgen saebaatcs weeucite minuscula var (p. 48).

6. Whole insect black, including palpi and venter of thorax and abdomen.

Mesonotum with seattered greenish seales. Antennae wholly dark

orange. Wings deeply fumose on apical half, darkest anteriorly. Eye

green with 3 purple bands ......scccses pevabacgsute Bendonise ...sordidatus (p. 49)

Venter of abdomen and thorax grey pollinose and white haired, meso-

notum without greenish seales, abdomen with narrow and faint pale

hind-marginal bands and small median pale hair tufts on all tergites.

Antennae with at least style black. Palpi pale basally, with at least

some white hairs. Wings nearly glass clear, faintly and unevenly

brown tinted. Eye purple with 2 very narrow green bands .....cccccsoee

ds RCLECIA aad ntBTE Raa Oath aE Lib da has jiicalia. scecscscsssceeeeee ODSCUPUS (PD. 49)

Stenotabanus (Stenotabanus) ecalvitius Fairchild

Stenotabanus ecalvitius Fairchild, 1942, Ann. Ent. Soec.Amer., 35(3): 306-

307, fig. 10, F, Panama.

This little species, only 7-8 mm. long, is easily recognizable by the combina-

tion of bare subcallus and densely golden-haired secutellum. Only 3 specimens

are known, as follows: Panama City, in automobile, 10 Jan. 1942; Rio Tan-

ganti,Camp 2, Rio Mandinga, San Blas, 24 Jan. 1957; upper Rio Pucro, Darien, in

Shannon trap, 15 Mar. 1958. The last two localities are in areas of heavy forest

and high rainfall. The most similar species appears to be Stenotabanus roxannae

for which Wilkerson (1979) erected the subgenus Stilbops. The present species

agrees quite well with roxannae in structure of frons, but differs in having the

abdominal tergites bordered behind with pale pollinosity and golden hairs, in

lacking any brown coloring in the wings, except for the stigma, and in having

white pollinose and pilose genae and frontoclypeus.

Stenotabanus (Stenotabanus) fulvistriatus (Hine)

Tabanus fulvistriatus Hine, 1912, Ohio Naturalist, 12(7): 515, female,

Mexico. Fairchild, 1942, Ann. Ent. Soc. Amer., 35(3): 303, fig. 11,

female, Panama; 1953, Ann. Ent. Soc. Amer., 46(2): 270, male, ref-

erences and synonymy.

Easily recognizable by the broad yellowish-white abdominal stripe, this

Species seems to reach its southern limits in western Panama. We have records

of only 4 specimens, 3 from Puerto Armuelles, Chiriqui Prov., 27 July 1929, L.

H. Dunn, coll. and one taken in a railroad car at Concepcion, Chiriqui Prov., 10

Aug. 1950. The species ranges north to Mexico.

_ Other closely similar species occur from eastern Peru south to Brasil and

Argentina, but there appears to be a large gap in distribution which includes

48 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

most of Panama and Colombia. The eyes in life are purple with 4 narrow green

bands. The male has the upper facets of the eye well demarcated from the

lower facets and the facets noticeably enlarged.

Stenotabanus (Stenotabanus) incipiens Walker

Tabanus incipiens Walker, 1860, Trans. Ent. Soe. London, (N.S.) 5: 275.

Amazon region. Fairchild, 1956, Smiths Miscell. Coll. 131(3): 19.

Type in London headless.

Stenotabanus (Stenotabanus) incipiens: Fairchild, 1971, Cat. S. Amer.

Dipt., 28, p. 46, synonymy. Wilkerson 1979, Cespedesia 8(31-32): 212-

214.

Tabanus maculifrons Hine, 1907, Ohio Naturalist, 8(2): 222, female,

Guatemala.

Stenotabanus maculifrons Fairchild, 1942, Ann. Ent. Soe. Amer., 35(3):

304, fig. 8, female, Panama,Colombia, references. Goodwin and Mur-

doch, 1974, Ann. Ent. Soec.Amer., 67(1): 110 fig. 32.

This species can be separated most quickly from minuscula by bicolored tib-

iae, and from the variety by blackish palpi, wholly yellow antennae and generally

black and white color. The eyes in life are reddish bronze with two green

stripes. What I believe are males of this species are similar to the females, and

distinguishable with difficulty from presumed males of minuscula and its variety.

They are placed here on the combination of wholly yellow antennae, dark fore

eoxae and basally whitish tibiae.

Incipiens has been taken at several localities in the Canal Zone and nearby

Panama and Colon provinces, and at Almirante, in Bocas del Toro prov., almost

entirely in light traps,Shannon traps, and in buildings. Some of the collecting

localities are in heavy forest, others in relatively dry and open areas, all at low

elevations. The species is probably on the wing throughout the year as records

are from Jan., Mar., July and Dec. The known range is from Guatemala to

Trinidad and Bolivia.

Goodwin and Murdoch (1974) succeeded in rearing a single female found near

an old log, and figured the pupa.

Stenotabanus (Stenotabanus) minusecula (Kroeber)

Pseudacanthocera (?) minusecula Kroeber, 1930, Zool. Anz., 90(3-4): 80-

81, fig. 21, Vulean Colima, Mexico. Fairchild, 1966,Stud. Ent., 9(1-

4): 354. Type studied.

Stenotabanus (Stenotabanus) minuseula: Fairchild, 1942, Ann. Ent. Soc.

Amer.,35(3): 305, fig. 7, Panama;

This species is clearly separable from ineipiens and the variety discussed

below by the wholly blackish legs, and from ineipiens in addition by largely

white palpi and striped thorax. The eye in life is dark reddish, with two green

transverse bands, seemingly narrower than in ineipiens. In Panama the species

has been taken at several localities in the Canal Zone, notably a long series

taken in a Malaise trap on Barro Colorado Id. by C.W. and H. E. Rettenmeyer,

in nearby Panama prov., and in Darien prov. up to 1500 ft. Most of the locali-

ties are in or near forest, in areas of moderate to high rainfall. All were se-

G. B. Fairchild: Tabanidae of Panama 49

cured in the dry season, from Jan. to Apr. in various years. Presumed males

are browner and lack the mesonotal stripes. Their tibiae are all dark, fore

eoxae dark but with pale pollen, and antennal styles blackish. The known range

is from Mexico to Panama.

What appears to be only a well marked variety of minuscula differs only in

having all tibiae extensively white, fore coxae pale in ground color, and thorax

and abdomen brown rather than black, the pale markings of thorax and abdomen

yellowish. It has only been taken with minuscula, at the same localities and

season. Since leg color is used as an important key character, minuscula ap-

pears twice in the key. Males of what appear to be this variety differ from

minuscula males only in having the tibiae pale basally, and fore coxae pale in

ground color. I ean find no structural distinctions in either sex, and some

specimens are intermediate in leg color, having reddish tibiae and black femora.

The type in Munich agreed well with a specimen from Chiapas, Mexico, with

wholly dark tibiae.

Stenotabanus (Stenotabanus) obseurus Kroeber

Stenotabanus obseurus Kroeber, 1930, Eneyel. Entom., Ser. B., II,

Diptera, V p. 124, Pl. 1, figs. 21; Pl. 2, figs. 5, 35; Pl. 3, fig 51.

Venezuela. Wilkerson, 1979, Cespedesia 8(31-32): 217-218.

Stenotabanus (Stenotabanus) constabulorum Fairchild, 1942, Ann. Ent.

Soe. Amer., 35(3): 304-305, fig. 6, female, Panama.

Aside from the characters in the key, this species is 8-9 mm. long, largely

black dorsally with seattered small metallic seales on thorax, the abdominal

bands very narrow, the mid and hind tibiae very largely white, the frons about 7

times as high as wide, considerably narrowed below, the callus higher than wide

and the median callus large and prominent. The vertex is extensively bare, with

prominent vestiges of ocelli.

A male from Rio Tacarcuna, Darien, 19 July 63, though slightly teneral,

agrees well with the female, except that the antennae are wholly yellow. The

eyes are holoptic, wider than thorax, the upper area of enlarged facets well dif-

ferentiated and demarcated from the small, occupying about two-thirds eye

area. The upper facets are dull bronzy, the lower red with a narrow green

streak. The female eye is reddish purple with two narrow green bands.

Obseurus has been taken rarely and at but 4 localities in Panama, on the

headwaters of Madden Lake on the Rio Boqueron and Rio Pequeni, Colon Prov.,

at Cerro Campana, Panama Prov., and at Rio Tacarecuna, Darien Prov. All are

areas of heavy forest and high rainfall. Specimens have been taken in March,

May, June and July.

The species has been taken also at Palmar, Dept. Puntarenas, Costa Rica,

from where it ranges south to Venezuela, and eastern Peru and Bolivia.

Stenotabanus (Stenotabanus) sordidatus Fairchild

Stenotabanus sordidatus Fairchild, 1958, Ann. Ent. Soc. Amer., 51(6):

925-526, fig. 6, female, Panama. Wilkerson, 1979, Cespedesia 8(31-

32): 218-219.

50 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

This species has the abdomen entirely black, the wings strongly blackish fu-

mose, and all tibiae extensively white. The eye in life is green with 3 reddish

purple bands. It has been taken in Bocas del Toro, Colon, Panama, San Blas,

and Darien provinces, always rarely and always in heavy forest below 2000 ft.

Our records are for Jan., Feb., March, May, June, and July. Perfectly fresh

Specimens have the subeallus thinly pollinose in the middle and at sides, not en-

tirely bare and shiny as shown in the original description. The species has so

far been taken outside Panama only in Antioquia, Colombia and at Golfito,

Puntarenas Prov., Costa Rica, the latter locality being a new record for Costa

Rica.

Subgenus Aegialomyia Philip

Aegialomyia Philip 1941, Canad. Ent. 73: 10, in key, T. psammophilus O.

S. type species.

Stenotabanus (Aegialomyia): Fairchild, 1942, Ann. Ent. Soc. Amer.,

35(3): 298-299. Philip, 1958, Amer. Mus. Novit., No. 1921, pp. 104,

key.

Flies of this subgenus are mostly small to medium sized insects, up to 15

mm. in length, generally with relatively broad frons and often pale or whitish in

general color. The eyes are generally purple with three green bands or greenish

with four dark bands. Most of the species are inhabitants of coastal environ-

ments, beaches or mangrove swamps. The main concentration is in the

Caribbean area, some sixteen species being known from the West Indies and ad-

jacent coasts from Florida to Venezuela. Some species will attempt to bite

man, while others seem to show little interest in him. Five species are known

from Panama. |

Key to females.

1; Abdomen without dorsolateral pale patches, grayish brown with hind and

lateral margins of all tergites whitish and a more or less complete

middorsal pale stripe. Frons 2.5 to 3 times as high as basal width.

Basal plate of third antennal segment as wide aS ONG ....ccccccccscecsees

walt lo tiie « deen Ns hunaes took RIDES uae stele de seal e vend sleaukech onnens changuinolae (p. 51)

Abdomen with dorsolateral pale patches. Basal plate frequently longer

LILAC" Wine svete begun eseeds des suues shicksenncdnome thas tan wekeleeeseureaeceeacebenee apadte ods 2

2 Wings with definite brown clouds on crossveins and fork of third vein.

Frons 3 times as high as basal width, narrowed below, the callus un-

usually protuberant. Palpi slender and curved, not markedly inflated

basally. Eyes purple with 2 green bands .......cccccoee pompholyx (p. 52)

Wings without clouds. Palpi markedly inflated ...........cecccccccccccccccsccescsees 3

os Frons broader, hardly twice as high as wide, parallel sided, the vertex

entirely bare and shiny. Antennal style longer than plate ........esseee

ai sulaiiaya c IEC natwuminnee Wer ve csuesdieeeet tec ha laceuee ath totes uenyeaias blantoni (p. 51)

Frons narrower, over twice as high as wide, the vertex not entirely bare.

APB a eap A TEP RANT COGAN Moy HA cant CLGLSAEEALAS AAG REENE ChELSERseeneesneee

G. B. Fairehild: Tabanidae of Panama 51

4, Frons about 2.2 times as high as wide. Antennal style subequal to basal

plate, the latter longer than wide. Median abdominal stripe nearly

parallel sided and continuous, the sublateral pale spots small and

GiSCrete ..ccceese IA, Cl Ae: HUA Adel. AV. OS seooeeee Littoreus (p. 51)

Frons about 2.8 times as high as wide. “Antennal style longer than plate,

the latter as wide as long. Median abdominal stripe is a series of

contiguous triangles, the sublateral spots large and diffuse ..........s00.

sites sswawsdensll Silda LavBAew iiea eee aA Weneepeitillensis (p52)

Stenotabanus (Aegialomyia) blantoni Fairchild

Stenotabanus (Aegialomyia) blantoni Fairchild, 1953, Ann. Ent. Soc.

Amer., 46(2): 269, P1. 1, fig. 5, female, Panama. Philip, 1958,

Amer. Mus. Novit. No. 1921, p. 3, in key.

Only the types are so far known with certainty, taken in a mosquito light

trap at Jaque, Darien prov. in July, 1952. This locality is on the Pacific coast

near the Colombian border. What may be the male of this species is repre-

sented by a specimen taken in a car in Panama City 9 Apr. 1957 by my col-

league Pedro Galindo. It is slightly larger than the female, the abdominal pat-

tern like the female though paler and less distinct, but it has a long appendix on

fork of third vein, longer than either St. (A.) paitillensis or St. (A.) littoreus, the

most similar regional species. The eyes are holoptic, bare, the upper facets

well differentiated and demarcated from the lower, occupying at least 2/3 of

eye area. There is a very small tubercle sunk between eyes at vertex. The up-

per facets are pale tan in life, the lower bronzy with one pale green stripe.

Stenotabanus (Aegialomyia) changuinolae Fairchild

Stenotabanus (Aegialomyia) changuinolae Fairchild, 1942, Ann. Ent. Soe.

Amer., 35(3): 300, fig. 5, female, Panama.

Stenotabanus (Aegialomyia) ananasi Fairchild, 1951, Ann. Ent. Soc.

Amer., 44(3): 454-455, fig. 6, female, Panama. 1971, Cat. S. Amer.

Dipt. Fase. 28, p. 48, synonymy.

This species, the largest of the group in Panama, reaches 13 mm. in length.

It is not uncommon on white sand beaches on the Atlantic coast from Bocas del

Toro to the Canal Zone and neighboring Colon prov. It bites man readily and

has also been taken in light traps. Our records are from Jan. to May in various

years. A presumed male, lacking abdomen, has the upper facets well differ-

entiated and demarcated, occupying over two-thirds eye area. There is no tu-

bercle at vertex. The above synonymy was established through the kindness of

Dr. L. L. Pechuman, who compared specimens of ananasi with the damaged type

of changuinolae.

Stenotabanus (Aegialomyia) littoreus (Hine)

Tabanus littoreus Hine, 1907, Ohio Naturalist, 8(2): 227-228, female,

male, Puerto Barrios, Guatemala.

52 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Stenotabanus (Aegialomyia) littoreus: Fairchild, 1953, Ann. Ent. Soe.

Amer., 46(2): 270, Pl. 1, fig. 6, Panama, references.

Only a few specimens have been taken in Panama, a female from near Pina,

Colon prov., biting man on the beach, Nov. 1950; 8 females, Ft. Sherman Res-

ervation, Canal Zone, 24 Dec. 1965, biting man on beach; and a male taken in a

light trap in Mojinga Swamp, C. Z. Jan. 1953. The latter locality is a tidal

Swamp along the lower Chagres river near the Atlantie coast., The female from

Pina was kindly compared with Hine's type by Dr. Philip. The species ranges

north to Belize, from where I have a long series taken at Corozal in May 1960

by John Strangways-Dixon. These latter specimens have the pale sublateral

patches more extensive than the Panama specimens, generally connected by a

narrow isthmus to the median pale stripe and to the pale hind borders. The

eyes in life of Panama specimens are reddish purple with 2 broad grayish green

stripes above and below the middle. The upper margin is also grayish green. Or

the eye may be described as grayish green with 3 reddish purple stripes.

Stenotabanus (Aegialomyia) paitillensis Fairchild

Stenotabanus (Aegialomyia) paitillensis Fairchild, 1942, Ann. Ent. Soc.

Amer., 35(3): 299, fig. 12, female, Panama; 1947, Op. cit., (1046),

39(4): 569; Philip, 1958, Amer. Mus. Novit., No. 1921, p. 3, in key.

Five specimens of this little species (9-10mm) have been taken in or near

mangrove swamps on the Pacific coast from Garachine, Darien prov., to

Aguadulee, Cocle prov. Two were taken attempting to bite, one in a light trap,

in Feb., Mareh and April. It is to be separated from related species by smaller

size as well as the characters in the key. The eyes in life have the upper and

lower margins purple, the green middle of the eye crossed by 2 narrow purple

stripes.

Stenotabanus (Aegialomyia) pompholyx Fairchild

Stenotabanus pompholyx Fairchild, 1953, Ann. Ent. Soe. Amer., 46(2):

273-274, Pl. 1, fig. 4, female, Panama.

Only the two type specimens are known, both taken in a light trap at Patino

Point, Darien Prov., July 1952. This locality is at the mouth of the Tuira river,

on the Pacific coast. The type is denuded, the paratype lacks the abdomen.

When describing this species, I had doubts as to its subgeneriec placement, but

feel now that it fits best in the present subgenus, in spite of having but 2 green

bands on eye. The wings are more clearly spotted on ecrossveins than others of

the subgenus in Panama.

Subgenus Brachytabanus Fairchild

Brachytabanus Fairchild 1942, Ann. Ent. Soc. Amer., 35(3): 300; 1971,

Cat. S. Amer. Dipt., Fase. 28, p. 50.

G. B. Fairchild: Tabanidae of Panama 53

The subgenus can be recognized by its small size, broad frons, and 3 annu-

late antennal style. Only one species occurs in Panama.

Stenotabanus (Brachytabanus) longipennis Kroeber

Stenotabanus longipennis Kroeber, 1930, Encye. Entom., Ser. B., Dipt. 5

(1929), p. 125, Pl. 1, fig. 23, Pi. 2, fig. 7, °37, PI. 3, fig.055, fe-

male, Venezuela.

Stenotabanus (Brachytabanus) longipennis: Fairchild, 1942, Ann. Ent.

Soc. Amer., 35(3): 300-301, fig. 4, female, Panama; 1964, J. Med.

Ent., 1(2): 175, male. Wilkerson 1979, Cespedesia 8(31-32): 219-220.

Aside from being the smallest species of Stenotabanus in Panama, less than

7 mm., this species is easily recognizable by the combination of very broad

frons, small round callus, black tentorial pits, 3-annulate style, and abdomen

with a broad yellowish middorsal stripe. The eye in life is glaucous green with

three narrow reddish purple bands and a reddish purple patch in the upper inner

corner, next the vertex. The male has much enlarged upper facets and two

round black velvety spots on subcallus. In Panama, females have been taken

mostly biting man or in a Shannon trap, the single male at light. It has been

taken from the Canal Zone east to Darien Prov., entirely on the Pacific coast,

and on the Pacifie coast of Costa Rica, generally near rivers or the coast.

Records indicate it flies at least from April to September, but it may be more

abundant and widespread, as its small size causes it to be easily overlooked. It

ranges from Costa Rica to Colombia and Venezuela.

Genus Himantostylus Lutz

Lutz 1913, Mem. Inst. Osw. Cruz, 5(2): 183-184. Fairchild, 1965, Psyche

72(3): 211-213, fig. 1. Panama.

The genus contains but one hitherto rare species, similar in appearance to

Lepiselaga and Selasoma. From the first it differs in lacking metallic scale-like

hairs, and in normal discal cell and less extensively black wings, from the last in

smaller size, less swollen tibiae, more slender and cylindrical third antennal

segment, and in having a prominent tubercle at vertex and banded eyes. The

wings are basally black to ends of basal cells in Himantostylus, to middle of dis-

eal cell in Selasoma.

Himantostylus intermedius Lutz

Himantostylus intermedius Lutz 1913, Mem. Inst. Osw. Cruz, 5(2): 183-

184, Pl. 13, fig. 22, male, Yahuarmayo, E. Peru. Stone, 1934, Rev.

Ent., 4(2): 191-192, female, Canamina, Bolivia. Wilkerson 1979. Ces-

pedesia 8(31-32): 223-224, fig. 58.

This species has been known only from the type male and the female de-

scribed by Stone, until found in Panama (Fairchild, 1966). In July 1963 I was

fortunate enough to secure a series of 59 females and 1 male on the upper Rio

Taecarcuna, Darien Prov., 1900 ft. The habits are rather peculiar. The flies

24 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

were concentrated about the shore of a semi-permanent forest rain pool. They

flew close to the ground and clustered on my boots, which were of black rubber

with canvas tops. They attempted to bite the rubber, but not the canvas, and

when I placed my hand on the rubber, they crawled freely over the fingers, but

without attempting to bite. None were taken in a Shannon trap placed within a

few feet of the pond, though one was taken on my boots in camp a quarter of a

mile away, and another seen crawling over a discarded ration can in camp. The

male was taken at light in camp. Tapirs had used the pool as a wallow and

were apparently common in the vicinity, while large frogs and basilisk lizards

were other possible sources of a blood meal.

These Panama specimens differ from the descriptions of Lutz and Stone and

from a specimen from Quincemil, Dept. Cuzco, Peru, 16-31 oct. 1962, 2450 ft.,

L. E. Pena coll. given me by L. L. Pechuman, in having the black on wing bases

covering the whole base of the wing basal of a line from costa at ends of basal

cells to tip of anal cell (ist A). The Peruvian specimens have only the basal

cells black, the remainder of the basal area only slightly dusky. The male has

the head notably wider than thorax, the upper area of enlarged facets well dif-

ferentiated and demarcated from the small facets, and occupying about two-

thirds of eye are. There is a small tubercle at vertex, with vestiges of 3 ocelli.

The basicosta is sharply pointed and without macrotrichiae in both sexes. The

Species may be more common and widespread than suspected, as its peculiar

habits render it quite inconspicuous. A recent specimen from Costa Rica,

Puntarenas prov., is like those from Panama, as is one from Rio Raposo, Bue-

naventura, Colombia. It is possible that specimens from West of the Andes, in-

cluding those from Panama and Costa Rica may prove subspecifically distinct

from the Amazonian nominate form.

Genus Diachlorus Osten Sacken

Osten Sacken 1876, Mem. Boston Soc. Nat. Hist., 2, Pt. 4(4): 47.

Fairchild 1942, Ann. Ent. Soc. Amer., 35(3): 295, full references;

1972, Fla. Ent. 55(4): 219-229, key. Wilkerson and Fairchild 1982,

Proc. Ent. Soe. Washington 84(3): 636-650, figs. 1-5, key.

Only two species of this chiefly South American genus have been taken in

Panama, separated below. Both attack man readily, as well as domestie ani-

mals. They are both small yellow insects with black and yellow striped

mesonota, vividly patterned eyes, and black-tipped wings.

Key to Species

Frons over 5 times as high as basal width. Thorax with prominent iso-

lated dark spots on each side anteriorly, and a broad median dark

stripe with hair-fine central yellow line ........ By sake wee. jobbinsi (p. 55)

Frons 4 times as high as basal width. Thorax without anterolateral dark

spots,and median dark stripe divided by a broad yellow line widened

posteriorly .......00. es ah daca venue pit palais Lebeteian ee tasaue curvipes (p. 55)

G. B. Fairehild: Tabanidae of Panama 55

Diachlorus curvipes (Fabricius)

Haematopota curvipes Fabricius, 1805, Syst. Antl., p. 107, South Amer-

ica.

Diachlorus curvipes: Fairchild, 1942, Ann. Ent. Soe. Amer., 35(3): 296-

297, fig. 3 full references; 1946, Op. Cit., 39(4): 567, male. Wilker-

son 1979, Cespedesia 8(31-32): 225.

The species occurs most abundantly in the lowlands of Darien Prov., but has

been taken in the Canal Zone and neighboring Panama Prov., and at Almirante,

in Bocas del Toro Prov. It appears to fly throughout the year, though seemingly

more abundant in the dry season. It has been taken in light traps, stable traps,

and biting man and horse. All specimens have come from near sea level, except

for two collections on the slopes of Cerro Pirre, Darien Prov., 1500 ft., and

there was a swamp at that locality. It has not been taken in heavy forest, but

seems to prefer open country near swamps or large rivers. The male is similar

to the female. The species ranges from Costa Rica south to Paraguay.

Diachlorus jobbinsi Fairchild

Diachlorus jobbinsi Fairchild, 1942, Ann. Ent. Soe. Amer., 35(3): 296,

female, Panama; 1946, op. cit., 39(4): 567, fig. 7; 1953, Proc. Ent.

Soc. Washington, 55(5): 241. Wilkerson 1979, Cespedesia 8(31-32):

326-227.

This was one of the dominant species of Tabanids at our Yellow Fever Sta-

tion near Almirante, Bocas del Toro Prov., over 3000 specimens being taken over

a period of 2 years, almost entirely attacking man at ground level in the forest.

Elsewhere in Panama, the species occurs at El Real, and Yape in the Rio Tuira

Valley, and at Cerro Pirre, Darien Prov. A few specimens, including males,

have also been taken in light traps and stable traps on the Atlantie side of the

Canal Zone. At Almirante, the species flies throughout the year, but all records

from elsewhere are for Jan., Feb., and Mar. The male is like the female in

color and structurally similar to the male of curvipes.

Specimens have been seen from Costa Rica, Colombia and Ecuador.

Genus Hemichrysops Kroeber

Kroeber 1930, Zool. Anz., 88(9-10): 226. Monotypic.

Hemichrysops fascipennis Kroeber

Hemichrysops fascipennis Kroeber 1930, Zool. Anz., 88(9-10); 237-238,

figs. 9-10, female, Colombia; Fairchild, 1964, J. Med. Ent., 1(2):

175-176, fig. 12, full references. Wilkerson 1979, Cespedesia (8(31-

32): 227-228, fig. 59.

This is a small, slender, long-winged insect, the wings black except for a

small hyaline area covering ends of basal cells, and axillary cell lightly infus-

cated. The body is wholly shining black, the frons narrow with slender callus,

26 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

the antennae slender and Diachlorus-like. Only two specimens have been taken

in Panama, at Rio Changena, 2400 ft., and Rio Uri, 3000 ft., Boeas del Toro

Prov., in heavy forest. The species is known from Colombia and Costa Rica as

well.

The male is like the female in color, except that the hyaline area in the

wings is smaller. The eyes are bare and an upper area of much enlarged and

well demarcated facets covers about half eye area. There is a small tubercle

sunk between the eyes at vertex. The antennae are dull yellowish, the style

dusky. Palpi dull yellowish, shorter than in female and porrect, and with long

sparse hairs. The abdomen also has long sparse pale hairs on sides of first 3

segments, and vestiges of a small white-haired triangle in middle of tergite 4.

The description is based on a specimen taken at light, Rio Raposo, Valle,

Colombia, 17 Aug., 1965, by Vernon Lee.

Genus Bolbodimyia Bigot

Bigot 1892, Wien. Ent. Zeits., 11: 162. Kroeber,1929, Encycl. Ent., B,

Dipt., 5: 109-112 and figs. Stone,1954, Ann. Ent. Soe. Amer., 47(2):

248-254, keys, synonymy. Wilkerson 1979, seguir entoaa 8(31-32): 228-

2359, key.

Species of this genus are, with two exceptions, largely black insects with

black wings having the extreme tip sharply hyaline. In addition to the charac-

ters in the key, the upper branch of third vein(R2+3) is bent sharply forward, so

that the first submarginal cell (3rd R) is somewhat coarectate. The third anten-

nal segment is very long and slender, with a blunt dorsal angle close to the

base. The eyes in life have the upper third purplish black, the lower 2/3 pur-

plish black with irregular green speckles. The species are apparently crepuscular

or nocturnal, though sometimes taken during the day in heavy shade. They seem

limited to heavy forest, and most of our specimens were taken in Shannon traps

or horse-baited stable traps. Members of the genus seem nowhere abundant,

perhaps due to secretive habits. Three species are known from Panama, keyed

below.

Key to females

ds Largely bright yellow species, the wing basally yellow, the abdomen with

broad median yellow stripe ADOVE .....scscoccscscscecees oes Zalindoi (p. 57)

Wing and dorsal surface of body wholly intensely black..........cccccsessseees 2

Be Hyaline of wing apex extending into first submarginal cell. Frons, sub-

callus and fronto-clypeus wholly orange. First antennal segment

orange, only slightly inflated. Mid and hind tarsi basally white .......

il ldo UAE raul dcardtbin she rhe tipantats, BaNeRN erythrocephala (p. 57)

Hyaline of wing apex confined to second submarginal cell (4th R). Ab-

domen and thorax beneath, all femora, as well as frons, subeallus

and frontoclypeus orange. First antennal segment black, greatly in-

flated. All tarsi dark .....cccccccccceee arewess OE MEN DEST Oele philipi (p. 57)

G. B. Fairchild: Tabanidae of Panama ST

Bolbodimyia erythrocephala (Bigot)

Atylotus erythrocephalus Bigot, 1892, Mem. Soc. Zool. France, 5: 668,

female, Panama.

Bolbodimyia erythrocephala: Fairchild, 1942, Ann. Ent. Soe. Amer.,

35(3): 294, full references. Stone, 1954, Ann. Ent. Soe. Amer., 47(2): |

252-253. Wilkerson 1979, Cespedesia, 8(31-32): 231-232.

In addition to the characters in the key, this species has the palpi, all legs

except tarsi, and entire thorax and abdomen black.

I have seen, besides the type, 13 specimens, from 6 localities in Panama,

Almirante and Rio Changena, Bocas del Toro Prov., Cerro Campana and Cerro

Jefe, Panama Prov., and Tacarecuna Yellow Fever Sta. and Cerro Mali, Darien

Prov. Al these localities are in heavy forest in areas of high rainfall, and all

except Almirante are at elevations between 2000 and 5000 ft. All were taken

between May and September, mostly in Shannon traps. The species is known

from Panama, Costa Rica and Colombia.

Bolbodimyia galindoi Fairchild

Bolbodimyia galindoi Fairchild, 1964, J. Med. Ent., 1(2): 172, fig. 5.

Wilkerson 1979, Cespedesia 8(31-32): 232-234.

Only the type and 2 other specimens of this aberrant species are known. It

is similar to philipi in degree of inflation of first antennal segment, and to

erythrocephala in extent of hyaline at wing tip, but differs from the other local

Species in color. The wing is yellow with a subapical black patch and a broad

band of grey extending along hind margin from R4 to base of anal cell. The

thorax is bright orange yellow with a median black streak, the abdomen yellow

with a pair of dorsolateral broad black stripes. The type was taken in a Shan-

non trap at the Tacarecuna Yellow Fever Station, Darien Prov., 30 Aug. 1958,

2200 ft., and the other known Panama specimen at Cerro Campana, Panama

Prov. 16 July 1966, 2000 ft., C. W. Myers coll. Wilkerson (loc. cit.) discusses the

differences, largely tinetorial, between the single known Colombian specimen

and the 2 from Panama.

Bolbodimyia philipi Stone

Bolbodimyia philipi Stone, 1954, Ann. Ent. Soc. Amer., 47(2): 253-254,

female, Guatemala, Costa Rica, Panama, Colombia. Fairchild, 1964,

J. Med. Ent., 1(2): 172, male described. Wilkerson 1979, Cespedesia

8(31-32): 230-235.

Bolbodimyia bicolor: Fairchild, 1951, Ann. Ent. Soe. Amer., 44(3): 452.

Not bicolor Bigot.

This species has been taken at Ft. Davis., C. Z., at Cerro Campana, and Las

Cumbres, Panama Prov., and on Barro Colorado Id., C. Z., in Dee., June and

July in various years. Three of the 10 specimens taken in Panama were males

taken at light, the females in Shannon traps and horse baited mosquito traps.

Wilkerson (loc. cit.) believes it very likely that this species is but a color vari-

58 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

ant of bicolor, or at most a subspecies, an opinion with which I concur. Mate-

rial is still very seanty in this genus, and a final decision on specific limits must

await its accumulation.

Genus Selasoma Macquart

Macquart 1838, Dipt. Exot, 1(2): 187. Fairchild, 1942, Ann. Ent. Soc.

Amer., 35(3): 289, full references; 1966, Psyche 72(3): 210-211(1965).

- This genus contains but one widespread species.

Selasoma tibiale (Fabricius)

Tabanus tibialis Fabricius, 1805, Syst. Antl., p. 102.

Selasoma tibiale: Fairchild, 1942, loc. cit. p. 290, fig. 1, full references.

Wilkerson 1979, Cespedesia 8(31-32): 235-236, fig. 60.

The species is easily recognizable by its stout build, swollen tibiae, shiny

blue-black abdomen, basally black wings, and third antennal segment with the

broad flat basal plate many times longer than the reduced style. The eye is

greenish black, without bands. In Panama the species is definitely nocturnal,

and has been taken attacking man and horses and- in horse-baited stable traps.

The flight season appears to be short; all our material was taken in December

or January, in various years. Specimens have been seen from three localities in

the Canal Zone, and from Rio Mandinga on the border between Colon and San

Blas, all at low elevation and near water. Range is from Oaxaca, Mexico, to

southern Brasil.

Genus Chlorotabanus Lutz

Lutz 1909, Inst. Osw. Cruz em Manguinhos, p.30. Fairchild, 1940, Rev.

Ent., 11(3): 714, full references. Philip and Fairchild, 1956, Ann. Ent.

Soe. Amer., 49(4): 313-324, keys. Fairchild 1969, Arq. Zool. S. Paulo

17(4): 208.

The genus was revised with keys, figures and synonymy by Philip and

Fairchild in 1956. The species are nearly unicolorous pale greenish insects of

erepuscular habits and without frontal callus, attacking domestic animals but

rarely man and often attracted to lights. Two species, keyed below, are known

from Panama.

Key to species.

Wings with distinct dark spots on all crossveins and at tips of longi-

tudinal veins. All tibiae black-haired at tips ........ mexicanus (p. 59)

Wings without black spots, glass-clear, only costal cell yellowish. Tibiae

whelly ‘pale hatred’ lh, Saties Waele eseels nb ONAL edi Ba ce inanis (p. 59)

G. B. Fairchild: Tabanidae of Panama 59

Chlorotabanus inanis (Fabricius)

Tabanus inanis Fabricius, 1794, Ent. Syst., 4: 368.

Chlorotabanus inanis: Fairchild, 1940, Rev. Ent. 11(3); 714-715, fig. 1,

full references. Philip and Fairchild, 1956, Ann. Ent. Soe. Amer.,

49(4): 316-317, fig. 1, synonymy. Goodwin and Murdoch, 1794, Ann.

Ent. Soe. Amer., 67(1): 102, figs, 13, 25. Wilkerson 1979, Cespedesia

8(31-32): 239.

This species is similar to mexieanus in habits and occurrence. It has been

taken in all months save April and October, in the Canal Zone, Panama, Colon,

San Blas and Darien Provinees, and probably occurs throughout the country at

low elevations. There are, however, no records for the Almirante area, where

mexicanus is common. The range is more extensive than that of mexicanus. I

have seen specimens from Mexico (Tabasco), Colombia, Venezuela, Trinidad and

Brazil as far south as Matto Grosso. Goodwin and Murdoch 1974) described the

larva and pupa, taken among decaying leaves in small pools in a nearly dried up

stream bed. Wilkerson (loc. cit.) did not find this species in western Colombia,

though it has been taken in Antioquia and Meta, and is a common species

throughout the Amazon Valley.

Chlorotabanus mexicanus (Linnaeus)

Tabanus mexicanus Linnaeus, 1767, Syst. Nat., 12th Ed., 1, pt. 2, p.

1000.

Chlorotabanus mexicanus: Fairchild, 1940, Rev. Ent., 11(3): 715-716, fig.

2, full references. Philip and Fairchild, 1956, Ann. Ent.Soc. Amer.,

49(4): 315-316, fig. If., synonymy. Goodwin and Murdoch, 1974,

Ann. Ent. Soe. Amer., 67(1): 102, fig. 13, 26. Wilkerson 1979, Ces-

pedesia 8(31-32): 240-241.

In Panama, the species has been taken in all months except October, in the

Canal Zone and in the provinees of Bocas del Toro, Panama, San Blas, and

Darien, always at elevations below 2000 feet. Males are frequently taken in

light traps, and females have been collected biting pigs, cattle, and horses after

dark, in Shannon traps operated with a light at night, and in horse-baited

mosquito traps. It appears to attack man rarely if at all. Both sexes have been

reared from field collected larvae on several occasions, the larvae among dead

leaves in slow flowing water or in aquatic vegetation. It occurs with C. inanis

in most localities, but the latter seems to be absent from the Almirante area in

Bocas del Toro, where mexiecanus is common. Neither species has been taken in

the forest canopy, and both seem rare or absent in areas of heavy forest, per-

haps because bait animals were not available. The species ranges from Mexico

to Ecuador and Surinam, but I have seen no authentic Brazilian specimens.

Goodwin and Murdoch (1974) deseribed the larva and pupa, taken in similar habi-

tats to those where inanis occurred. The greenish brown-mottled larvae of the

2 species are very similar.

60 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Genus Phaeotabanus Lutz

Lutz 1913, Brasil Medico, 27(45): 6. Barretto, 1950, Ann. Face.

Med.Univ. S. Paulo, 25: 26, full references. Medem, 1981, Ces-

pedesia 10(37-38): 123-146.

Tabanus (Phaeotabanus): Kroeber, 1934, Rev. Ent., 4(3): 304.

Three species of this small genus occur in Panama. All seem to be crepus-

cular or nocturnal and not strongly attracted to man. Although not referring to

any Panamanian species, Medem (op. cit.) observed that species of Phaeotabanus

were the dominant tabanids attacking caimans in eastern Colombia. He did not

report nocturnal activity.

Key to Species —

1; Dorsum of thorax and abdomen black. Whole ventral surface including

subeallus, face, antennae, palpi and legs bright orange. Wings deep

blaek with a sharply hyaline apex. Male similar but with mesonotum

and seutellum bright Oren@es,. .icoreadsansebobeusiscestaets phaeopterus (p. 61)

Whole inseet dull yellowish, abdomen slightly greenish in life. Wings

yellowish or grayish, with an unusually long appendix on fork of third

VEIN s MASS SHNTIAL EPS eiligsale co cbecellivews vented sdecccsasdeentbneiescesssesicenesenss 2

os Legs yellow. Abdomen yellowish, unicolorous..longiappendiculatus (p. 60)

Legs largely blackish. Abdomen with a broad yellowish middorsal stripe.

sib deni kak s Rebudisioins t4 deb Buda uh busattenss's had ceeded tone vba bivs esanWeaeiesw esau atopus (p. 60)

Phaeotabanus atopus (Fairchild)

Stenotabanus atopus Fairchild, 1953, Ann. Ent.Soec. Amer., 46(2): 271-

272, Pl. 1, fig. 3, female, Panama.

Phaeotabanus atopus: Philip and Fairchild, 1956, Ann. Ent. Soe. Amer.,

49(4); 314.

The type series was taken at our Yellow Fever Station inland from Almi-

rante, Bocas del Toro Prov., in a Shannon trap at ground level, in the months

from June to August, 1951. Aside from these specimens, only a few others have

been seen, from Palmar and Golfito, Dept. Puntarenas, Costa Rica. In addition

to the key characters, the species is darker throughout than longiappendiculatus,

with quite smoky wings, and the antennal plate is broader, more excavated, the

dorsal angle more acute.

Phaeotabanus longiappendiculatus (Viacquart)

Tabanus longiappendiculatus Macquart, 1855, Dipt. Exot., Suppl. 5, p. 32,

female, Honduras.

Cryptotylus luteoflavus Bell.: Fairchild, 1940, Rev. Ent., 11(3): 718-719,

fig. 3, female, Panama, references; 1946, Ann. Ent. Soc. Amer.,

39(4): 567, male.

G. B. Fairchild: Tabanidae of Panama 61

Phaeotabanus longiappendiculatus: Philip and Fairchild, 1956, Ann. Ent.

Soe. Amer., 49(4): 314-315, references and synonymy.

The species has been taken in some numbers along the Rio Chagres above

Gamboa, and occurs also in Panama Provinee and at Almirante, in Bocas del

Toro Prov. It is nocturnal and has been taken at light, in Shannon traps and

biting domestic animals. Most records are from the dry season months Dec. to

May, with isolated records in Nov. and June. From the limited data, it appears

that the species prefers lowland areas near swamps and large rivers. It ranges

north to Mexico and south to Venezuela. It somewhat resembles a species of

Chlorotabanus, but is less greenish, has a frontal callus, and very long appendix

at fork of third vein.

Phaeotabanus phaeopterus Fairchild

Phaeotabanus phaeopterus Fairchild, 1964, J. Med. Ent., 1(2): 176, figs.

8, 9. Wilkerson 1979, Cespedesia 8(31-32): 241-242.

In coloring this species resembles some species of Bolbodimyia, but the

larger size, lack of swollen subeallus and basal antennal segments, narrow frons

and broad antennal plate easily distinguish it. In Panama it is known only from

the types from Darien Prov., both sexes taken at light. Wilkerson (loc. cit.) re-

ports the species also from Western Colombia and some paratypes came from

Madre de Dios in eastern Peru, where more recent collectors have secured fur-

ther material in flight traps.

Genus Dichelacera Macquart

Maequart 1838, Mem. Soc. Roy. Sei. Agric. Arts. Lille, pt. 2, p. 116.

Fairchild and Philip, 1960, Stud. Ent. 3(1-4): 1-90. Pl. 1-X, key, syn-

onymy.

Three subgenera and twelve species occur in Panama, treated separately by

subgenus below. All the species appear to be forest inhabitants, some in the

canopy and others at ground level. All attack man readily and, so far as known,

all are diurnal, though males and occasionally females, are taken attracted to

light.

Subgenus Dichelacera

The ten Panamanian species of this subgenus are all slender species usually

with banded abdomens and wings with a diagonal dark fascia from apex to fifth

posterior cell, which may fill most of the wing beyond basal cells. All have at

least a single eye band and dark secutellums in the female. The species are dis-

cussed in alphabetical order for ease of reference.

T42)

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Key to Species

Males (unknown in crocata, hartmanni, regina)...........000 Bev seat id iskckessea 2

Females....... sid ieczsreiehats Pepe eee Ly rere are Ce ae eT ee Ere PsA be cecescnes 9

Upper eye facets greatly enlarged, sharply differentiated and demar-

cated from small facets. Inner border of wing fascia even or un-

even iveliu i cones hoe dese ch a Rustic DeWeese deeesculecs@bs dudel den vedbacduLpess eevee cudeey bbe 3

Upper eye facets slightly enlarged, not sharply differentiated and de-

mareated from small facets. Wing fascia faint and dilute, its inner

Margin nearly Straight ......ccccccccccces a dadajactoseeleeses bce uecsbesecuausemeuseveesis 7

Wing largely black, hyaline only along hind margin from apex to fifth

posterior cell and a band from base of discal cell to hind margin.

Legs black except for white mid tibiae and all tarsi. Abdomen and

thorax entirely black ..... Vide Dees kUGa WARS KS Ue erRSNedesesedais .melanosoma (p 66)

Wing with extensive hyaline or yellow areas. Abdomen always and tho-

rax usually with pale transverse Dands .......ccccccccccccscscscscccsccscccvcccces 4

Thorax wholly yellow haired, including scutellum, Wings with dark fas-

Cia dilute, its inner Margin EVEN ........ccccccccsccsscceees marginata (p. 65)

Thorax with scutellum and a band between wing bases black. Dark wing

fascia intense, its inner margin step-like OF SIMUOUS ........cscecccececeee 9)

Dark wing fascia with a clear area in first to third posterior cells and

its inner margin sharply step-like. Black thoracic band over twice

as wide as yellow prescutellar band. Yellow hairs only on third and

FOUPTH BOCOMI Al LACAILOS « .rcccevncescicnenednoncsdovsennevascetes fasciata (p. 64)

Dark wing fascia uniform, without light fenestra, its inner border sinu-

ous. Black thoracic band narrower, and yellow hairs on additional

LEP PIGES iiicl seriRdicoss verre ee sabiveusadcabeacondue Riv eh Wale ese nwadetecacees spe D Ate sices 6

Inner margin of dark fascia passing through fork of third vein. First ab-

dominal tergite wholly yellow haired, 3 to 5 with yellow hairs on

hind: Margins “ivsseceosss ie Gi. iaias. submarginata (p. 68)

Inner margin of dark fascia passing well proximal to fork of third vein.

First tergite yellow-haired, second to fourth with broad yellow-

haired median triangles ........scccccccscssecscese eats ets Ueda shasnierk rex (p. 67)

Thorax without preseutellar pale band, almost wholly dark haired. Ab-

domen largely dark haired, at most with sparse yellow hairs on hind

borders of tergites 1 to 4. Legs entirely dull blackish ..........ssecceceee

sietuedeioes Upilacss PAVE WENT eiduaiaPelakiacabacenvecs scapularis var. aquilus (p. 68)

Thorax with a prominent prescutellar pale Dand........cccccccccccce PebntPldae acne 8

Prescutellar pale band as wide as or narrower than inter-alar black band.

Abdominal tergites 2 to 5 predominantly black haired. Legs yellow

Tey Wg EASES, RRR te, a se SRA scapularis (p. 67)

Prescutellar pale band markedly wider than inter-alar black band. Ab-

dominal tergites predominantly yellow haired. Legs yellow except

for blaek hind tibiae and tarsi, and fore tarsi ....... ..princessa (p. 66)

9.(1)

10.

11.

12.

13.

14.(10)

15.

G. B. Fairehild: Tabanidae of Panama 63

Thorax and abdomen entirely black, except for narrow white-haired hind

marginal band on first tergite. Wings largely black, as in male. All

femora black, all tibiae over half white. Fronto-elypeus black, bare

and shiny ....... Wiis Rieck bh lina ahem aie melanosoma (p. 66)

Thorax and abdomen rarely unicolorous; at least basal cells always hya-

line; fronto-clypeus always POL]INOSE .....ccccccccecees sophie VR tiivddavedeeess 10

Fifth and following abdominal tergites wholly dark and dark haired...... 11

At least fifth tergite with pale hind border or tuft or triangle of pale

PUA esi, sag shen g'c PEN SWIM vies uh Sewn lelley LA does Vii ide h ean Me ssveeses 14

Dark wing fascia with markedly step-like inner border and a more or

less prominent hyaline patch in first to third posterior cells First

tergite and narrow hind borders of third and fourth pale yellowish-

white pilose, the remainder of abdomen black .......... -fasciata (p. 64)

Wing otherwise or second tergite with yellow haired hind border or tri-

GIGS: dbs vgeencsaintnneds oveaake pees Kiel seas oseapsvanedne sis:tipvie Cinque tds chanevonadecseyess 12

Proximal margin of dark fascia weakly sinuous to nearly straight, passing

through fork of third vein. Fascia filling wing evenly to margin, the

basal clear areas strongly yellow tinged. Frons rather narrow, the

callus nearly square. Yellow abdominal bands on tergites 2 to 4

even or slightly wider in Middle .......cssssccsessesssssrerees regina (p. 66)

Proximal margin of fascia strongly sinuous to step-like passing well

proximal to fork of third vein. Frons broader, the callus higher than

WAS: Sissies aoe Sea cated taken aks iRtds n salpnetnwacee eonbiiildanghclees Greil nee aan 13

Black interalar band very broad, the prescutellar yellow band narrow,

hardly one-third as wide. Outer parts of dark fascia dilute, some-

what as in faseiata. Yellow abdominal bands narrow and even ........

rere eee Tn sivnbisdnsins’s Vensenenedesiebeddsoesnsedsevbeedeseccdseresss Martmanni (p. 65)

Black interalar band generally narrower, rarely over twice as wide as

yellow prescutellar band. Dark fascia usually of even intensity.

Abdominal bands consisting of deep yellow triangles, that on the

second tergite smallest, those on third and fourth often extended

laterally to sides of tergites wisi n hae edie sacar Xe eels cnaeenbhaeet rex (p. 67)

Proximal margin of wing fascia more or less sinuous, passing through

fork of third vein. Fascia dark and of even intensity to wing mar-

gin, rarely with a more dilute area in first to third posterior cells.

Eyes green with a broad median band and upper third purple ...........

iso Dibasavenganendd vanes ucneenanneds pn dpniensliad Codd «Mahon nvaubiin viens vag sawed MATEUSa Wapee st LS

Proximal margin of fascia straight or somewhat dentate, generally pass-

ing proximal to fork. Fascia faint and dilute, strongest along proxi-

mal border. Eyes green with a narrow purple median stripe ........ 16

Proximal border of fascia smoothly sinuous, basal hyaline area strongly

yellow. Abdominal yellow bands very wide, occupying over two-

thirds width of their respective segments, widest in middle, that of

fifth tergite usually a broad triangle © ..........ccccscsesceees crocata (p. 64)

64 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Proximal border of fascia almost step-like, basal hyaline area slightly or

not at all yellow. Abdominal yellow bands even, not over half width

of segments, that on fifth tergite often reduced to a small pateh ...

bp sobRTaasessswcsatvevsssssesnsstbeaeies tues sccccccccccscssssceceeees SUDMarginata (p. 68)

16. Mesonotum and abdomen almost wholly dark haired, at most a few pale

hairs before secutellum, and on hind borders of tergites, the pale

bands of abdomen in integument only. Femora dark yellowish to

DISCKISh: Al. SO oe nieve Sei scapularis var. aquilus (p. 68)

Mesonotum and abdomen with prominent yellow-haired bands...........s««- 17

Iii. Preseutellar pale band narrower than interalar dark band. Abdominal

pale bands usually narrower than the black. Legs with fore and hind

femora dusky to pea black. Pale hairs generally whitish ...... asbawen

es cas stb eR OR RE BORA scapularis (p. 67)

Prescutellar pale band as “halide or widen than dark band., Abdominal

pale bands as wide or usually wider than dark bands. Femora all

yellow. Pale hairs generally rich golden or Drassy .......cccccccccecsccees 18

18. Frons wider, 2.5-3.1 times as high as basal width........... marginata (p. 65)

Frons narrower, 3.4-4 times as high as basal width......... princessa (p. 66)

Dichelaecera (Dichelacera) croeata Fairchild

Fairchild 1953, Ann. Ent. Soc. Amer., 46(2): 263-264, P1. 2, fig. 10, fe-

male, Panama. Fairchild and Philip, 1960, Studia Ent., 3(1-4): 27-28.

An intensely yellow species with tergites 1 to 5 broadly yellow pilose and

the wings deeply yellow tinted. Specimens have been taken only at our Yellow

Fever Station near Almirante, and on the Rio Changena, 2400 ft., both in Bocas

del Toro Prov. Both localities are heavy forest in areas of true rain forest.

The species is highly arboreal; only one of 45 specimens seen from Panama was

taken at ground level. It is known elsewhere from Costa Rica and Nicaragua, at

the latter locality probably also from tree top collections, although not so la-

belled. Specimens have been taken from April to December. The eye in life is

green with a broad median purple stripe and the upper margin broadly purple.

Dichelacera (Dichelacera) fasciata Walker

Dichelacera faseiata Walker 1850, Ins. Saunders. Dipt., 1: 68. Fairchild

and Philip, 1960, Stud. Ent., 3(104): 32-33, P1. 3, fig. 9, full refer-

ences and synonymy. Wilkerson 1979, Cespedesia 8(31-32): 251-253,

fig. 63.

Dichelacera analis Hine: Fairchild, 1940, Ann. Ent. Soe. Amer. 33(4):

696-697, fig 1.

This is the most abundant and widespread species of the genus in Panama.

It is readily recognized by the irregular step-like dark wing fascia, narrow pale

abdominal bands only on tergites 3 and 4, contrastingly whitish first tergite, and

paired dark spots on anterior border of mesonotum.

G. B. Fairchild: Tabanidae of Panama 65

The species attacks man and domestic animals readily at ground level in

forested areas below 2500 ft. It was taken once attacking a sloth in the forest

canopy. It has been taken almost wherever collections have been made, even in

fairly young second-grown forest, and is often sufficiently abundant to be a real

pest. The flight period includes the whole rainy season, from May through

November. Outside of Panama, the species ranges north to Nicaragua and south

to western Colombia and Ecuador.

Dichelacera (Dichelacera) hartmanni Fairchild and Philip

Dichelacera (Dichelacera) hartmanni Fairchild and Philip 1960, Studia

Ent., 3(104): 37-39, Pl. 3, fig. 8, Pl. 7, fig. 4, female, Panama.

This uncommon species is close to D. rex, but darker, the yellow markings

reduced in extent, the abdominal triangles often reduced to small spots or a

narrow pale pilose hind margin on tergites 2 to 4. The frons is wider than in

rex and the callus larger and more prominent. Specimens have been seen from

higher elevations in Chiriqui prov. (Sta. Clara, La Fortuna), Bocas del Toro prov.

(Rio Changena Yellow Fever Camp), Veraguas prov. (Sta. Fe); Cocle prov. (El

Cope) and Panama prov. (La Zumbadora, upper Rio Pacora), taken in the months

of April, June, Aug., and Sept., mostly in Shannon traps or U. V. light traps.

The male is like the female, but with wider pale abdominal bands. The eyes

have the upper facets well demarcated and differentiated from the small, occu-

pying about 1/2 total eye area, structurally like the male of D. rex, but consid-

erably darker.

Dichelacera (Dichelacera) marginata Macquart

Dichelacera marginata Macquart 1847, Dipt. Exot., Lille Mem., Suppl. 2:

30, female, Cayenne. Fairchild and Philip, 1960, Studia Ent., 3(1-4):

44-47, Pl. 2, fig. 7, Pl. 8, fig. 10, full references and synonymy.

Wilkerson 1979, Cespedesia 8(31-32): 253-254, fig. 79.

In spite of further study of additional material, I am still unable to separate

females of marginata, princessa, and scapularis with certainty, although the

males of at least the first two are quite distinct. I have now seen 2 males of

marginata from Almirante, Bocas del Toro Prov. taken at light in the forest

canopy, a finding which negates our previous statement (Fairchild and Philip,

1960, p. 55) that marginata does not occur west of the Canal Zone. Our previ-

ous records of prineessa from at least the Atlantie coast of Panama in Bocas

del Toro Provinee should thus probably be referred to marginata. Whether fe-

males from Chiriqui are marginata or princessa must await the finding of males.

As defined in the key, and with the exception of the doubtful Chiriqui mate-

rial, specimens have been seen from 18 localities in Darien, Panama, Colon, and

Boeas del Toro Provinces, and the Canal Zone. Most localities are in lowland

forest, a few up to 2400 ft. elevation. The species is taken in light traps,

horse-baited mosquito traps, and attracted to man, almost entirely at ground

level. Records are from May to October, although at Almirante it has also been

taken in January. The species ranges from Costa Rica to northern Brazil and

66 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

eastern Peru. Wilkerson (loc. cit.) records it only from extreme northern Choco

on the Pacifie coast of Colombia, but it is abundant on the eastern side of the

Andes and the Amazon basin.

Dichelaecera (Dichelacera) melanosoma Uine

Dichelacera melanosoma fine 1920, Ohio J. Sei. 20(8): 316, fig. 1, fe-

male, Costa Rica. Fairchild and Philip, 1960, Studia Ent. 3(1-4): 47-

48, Pl. 5, fig. 7, full references. Wilkerson 1979, Cespedesia 8(31-

32): 254-255, fig. 64.

The species is easily recognizable by the largely black wings and body with

largely white tibiae. It seems to be rare and local in Panama, only two speci-

mens having been taken, one at Progreso, Chirqui Prov., the other on the Rio

Paya, in Darien Prov. Both were taken at ground level in heavy lowland forest,

in July and August in different years. The species is known elsewhere from

Costa Riea and Colombia, the Interoceaniec Canal Survey having taken a series

at Teresita, Choco, in a Malaise trap. These Colombian examples and the one

from Rio Paya are darker than Costa Rican and Chiriqui specimens, the hyaline

area in fifth posterior cell and base of discal cell, and the hyaline outer margin

of wing noticeably narrower. This may be another case of discontinuous distri-

bution, the main isthmian area separating 2 populations which show signs of be-

coming morphologically distinct.

Dichelacera (Dichelacera) prineessa Fairchild and Philip

Dichelacera (Dichelacera) prineessa Fairchild and Philip 1960, Studia

Ent., 3(4): 53-55, Pl. 4, fig. 3, M, F, Costa Rica.

Whether this species is actually represented by females from western

Chirqui must await the finding of males, as pointed out under marginata. The

only material seen is 1 from Chorcha, Chiriqui and a considerable series from

Camp Pital in the Coto region near the Costa Rican border taken by Dunn in

1929. These average smaller than Costa Rican examples, and the frons seem

narrower than in marginata.

Dichelacera (Dichelacera) regina Fairchild

Fairchild 1940, Ann. Ent. Soc. Amer., 33(4): 699- ‘KOO we laid; fig ndifer

male, Panama. Fairchild and Philip, 1960, Studia Ent., 3(1- A): 59- 60,

full references. Wilkerson 1979, Cespedesia 8(31-32): 256-258, fig

65.

Regina is readily separated from submarginata by the characters in the key.

In addition, the pale bands on tergites 3 and 4 are generally paler and broader

than the band on tergite 2, and the basal clear area of the wing usually more

yellowish.

The species is taken both at ground level and in the canopy, and at Almi-

rante, where it is abundant, was secured in all months except Oct. and January.

Elsewhere it is a rainy season form. Within Panama, it has been taken only in

G. B. Fairchild: Tabanidae of Panama 67

forested areas of high rainfall, mostly at elevations between 1000 and 2500 ft.

It occurs with submarginata at several localities, but is most abundant in Bocas

del Toro and Veraguas provinces, where submarginata has not been taken. The

range is from Honduras to Ecuador. At a camp on Rio Tacarcuna, Darien, June-

July 1963, regina was the dominant Dichelacera in the forest canopy, sub-

marginata in the canopy and along the forest edge, fasciata at ground level

within the forest. All three attacked man readily within their habitats.

Dichelacera (Dichelacera) rex Fairchild

Dichelacera (Dichelacera) rex Fairchild 1951, Ann. Ent. Soc. Amer.,

44(3): 474-448, P1. 1, fig. 1, female, Panama. Fairchild and Philip,

1960, Studia Ent., 3(104): 60-61, male, full references.

The broad dark yellow abdominal triangles on second to fourth tergites char-

acterize this species. The dark fascia of wing is often less intense along hind

margin. The eye in life is green with a narrow purple median stripe and the up-

per margin tinged with purple.

All specimens, except two taken in a light trap in the forest canopy, have

been taken attacking man at ground level or in Shannon traps. The species has

been secured in small numbers only. It appears to be confined to areas of heavy

forest and high rainfall below 2500 ft. The species is similar to hartmanni, but

is paler, more yellow, with the abdominal triangles nearly reaching foreborder of

their respective segments and the frons narrower. Further material may show

the 2 species to be but ecological forms, as they have been taken at the same

locality only once. D. rex is known from Almirante, Bocas del Toro prov., Rio

Mandinga, San Blas prov., Cerro Azul, Panama prov., Taearcuna, Darien prov.

and Heredia prov. Costa Rica, localities mostly at lower elevations than those

listed for hartmanni.

Dichelacera (Dichelacera) scapularis Macquart

Dichelacera scapularis Macquart 1847, Dipt. Exot. Suppl. 2: 31,female,

Mexico. Fairchild and Philip, 1960, Studia Ent., 3(104): 61-64, P1. 1,

fig. 12, Pl. 10, fig. 8, full references.

There is much color variation in both sexes of this species. The typical

form has dark brown to black femora, a narrow, nearly white, prescutellar band,

and narrow whitish abdominal bands. Panama specimens agreeing with this form

have been taken only in the Canal Zone area and neighboring Panama Province,

in July and October to January. Occurring with this in the Canal Zone area is a

paler form with pale femora and generally broader and yellower thoracic and

abdominal markings. This pale form is also abundant around Almirante, in Bocas

del Toro Frov., where the typical form has not been secured. It has been taken

from May to September in Almirante, June to December in the Canal Zone

area.

The typical form was described from Mexico, and specimens from Vera Cruz

and Tabasco have been seen, as well as 1 from Nicaragua. The pale form oc-

curs also in Nicaragua and Honduras. |

Finally what appears to be but a highly melanistic form of scapularis is de-

scribed below.

68 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Dichelacera (Dichelacera) scapularis var. aquilus var.nov. Male and Female.

Differs from seapularis only in lacking all or nearly all pale hairs. Palest

specimens have yellowish though dark-haired femora, and small tufts of dull

yellowish hairs on notopleural lobes, before scutellum and the first tergite partly

yellow-haired. Darkest specimens have all body and leg hairs blackish. Of

three males seen, two have dark femora and large eye facets not sharply de-

mareated from small, the other has pale femora and upper eye facets larger and

sharply demarcated, in this respect intermediate between males of seapularis

and marginata. Of this dark form,, the following specimens have been studied.

Chilibre, Panama Prov., Aug. 1944, 2 females; Ft.Clayton, C. Z. 19 July 1954,

horse trap, 1 female; Albrook Field, C. Z., 24 July 1951, horse trap, 1 female;

Madden Airstrip, Panama Prov., 15 Nov. 1948, horse trap, 1 female; El Real,

Darien Prov., 11 July 1950, in forest canopy, 1 female; Rio Paya, Darien Prov.,

10 Sept. and Aug., 1958, 2 females; Ft. Kobbe, C. Z., 24 Aug. 1951, light trap, 1

male; Las Cumbres, Panama Prov., 4 Aug. 1958, on window of house, 1 male;

Las Palmas, Veraguas Prov., 16 July 1953, light trap, 1 male. The females from

Darien Prov. have pale femora, the male from Veraguas pale femora and demar-

eated eye facets. As suggested earlier (Fairchild and Philip 1960 p. 63) this

dark form and the pale legged form of seapularis may be segregates from a hy-

brid population derived from seapularis x marginata crosses. Although varietal

names have no standing in Zoological nomenclature, I have chosen to give this

well-marked variety a name in order to be able to key it and discuss it, but

there seems no point in designating conventional types. Named specimens will

be deposited in M.C.Z., F.S.C.A. and U.S.N.M.

Dichelacera (Dichelacera) submarginata Lutz

Dichelacera submarginata Lutz 1915, Mem. Inst. Oswaldo Cruz, 7(1): 86-

87, Pl. 20, fig. 8, female, Venezuela, Peru. Fairchild and Philip,

1960, Studia Ent., 3(1-4): 66-67, P1. 2, fig. 9, Pl. 7, fig. 3, full ref-

erences and synonymy. Wilkerson, 1979, Cespedesia 8(31-32): 258-

259 fig. 69.

This species is likely to be confused only with regina. In addition to the

yellow on tergite 5, submarginata averages smaller and has a broader frons.

In Panama this seems to be a lower montane species. With few exceptions,

Specimens have come from elevations between 100 and 5000 ft. in areas of high

rainfall in Darien, Panama, Colon, Cocle and Chiriqui provinces. The species

seems partly arboreal. At Rio Tacarecuna, Darien prov., submarginata was taken

abundantly at platforms in the forest canopy, and in the open along the river,

but seldom in the forest itself at ground level. At the same locality, D. regina

also occurred in the treetops, though in far fewer numbers, while D. fasciata

was abundant at ground level within the forest. The season of flight probably

includes the whole rainy season, but our records show collections only from

March to September.

The range outside Panama is from Costa Rica to Venezuela and eastern

Peru. Wilkerson (loc. cit.) records the species only from northern and eastern

Colombia.

G. B. Fairchild: Tabanidae of Panama 69

Subgenus Idiochelacera Fairchild

Dichelacera (Idiochelacera) Fairchild 1969, Arg. Zool. S. Paulo 17(4):

210.

Recent collections using a U. V. light trap at La Fortuna, Chiriqui Prov., by

Henk Wolda have shown the unexpected presence of a species of this subgenus.

Dichelacera (Idiochelacera) subeallosa Fairchild and Philip.

Dichelacera subcallosa Fairchild and Philip 1960, Studia Ent. 3(1-4): 83-

84, pl IV fig. 8, M., F., Santander Colombia.

Dichelacera (Idiochelacera) subeallosa: Wilkerson, 1979, Cespedesia 8(31-

32): 261-263, fig. 67. Colombia.

A short series of both sexes was taken at La Fortuna, Chiriqui Prov. in U.V.

light traps by Dr. Henk Wolda of the Smithsonian Tropical Research Institute, as

follows: 1 female, 28; IX-76; 1female,16-22IV-77; 1 male, 23-29-IV-77; 2 males

7-13-V-77; 1 female, 14-20-V-77; 6 females, 21-27-V-77; 1 female, 28-V to 3-VI-

77; 1 female, 4-10-VI-77; 1 male, 6-12-VIII-71; 1 male, 3 females, 13-19-VIII-77;

1 female, 10-16-IX-77; 1 male, 1 female, 29-VIII to 4-IX-78; 1 female, 16-22-V-

79. The specimens were collected in Kahl solution, and I was not able to make

very attractive pinned specimens of them, though they are quite recognizable.

These Panama specimens agree with those from western Colombia discussed by

Wilkerson in having the antennal plate strongly infuseated and the mid and hind

femora more or less prominently infusecated on their apical 1/4 to 1/3. Long se-

ries of well preserved specimens may enable definition of a subspecies on each

side of the Andes. My specimen from Peru is from Huanuco and is like the

types from Santander, Colombia, though the apical darkening of femora is hardly

evident. The only specimen seen from Ecuador is from the coast and like the

western Colombian specimens cited by Wilkerson.

The males are quite unlike the females in appearance, being largely yellow

and yellow pilose, the thorax unmarked, the abdomen yellow and yellow pilose,

except for, at most, black pilose hind margins on tergites 2 and 3, the last

wholly black pilose except for a median yellow pilose triangle. When denuded,

as most of my 8 specimens are, the black markings do not show, as they are due

only to pilosity, and the specimens appear entirely yellow. All femora and

ecoxae and mid tibiae are yellow, the hind tibiae and tarsi dusky and black pi-

lose, the fore tibiae white basally, black apically and tarsi black. The male eye

has the upper facets well-differentiated and demarcated, the area of enlarged

facets occupying over 1/2 the eye area. The area of large facets is densely

Short pilose and there is a small tubercle sunk between the eyes at vertex.

There is no evidence of the bare patch on subecallus so prominent in the female

and the dorsal antennal tooth in shorter than in the female.

Subgenus Desmatochelacera Fairchild

Desmatochelacera Fairchild 1969, Arg. Zool. S. Paulo, 17(4): 210

70 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Dichelacera (Desmatochelacera) transposita Walker

Dichelacera transposita Walker, 1854, List Dipt. Ins. Brit. Mus., 5: 151,

female, West Coast of America.

Catachlorops transposita: Fairchild, 1940, Ann. Ent. Soe. Amer., 33(4):

690, fig. 8, full references. Wilkerson 1979, Cespedesia 8(31-32):

263-265, fig. 66.

Only a single species occurs in Panama. It differs from Subgenus Dichela-

cera in having unicolorous greenish black eyes, black wings with a broad hyaline

band crossing ends of basal cells and whole wing apex hyaline. Body is black,

scutellum reddish and abdomen with a white haired triangle on tergite 4. Legs

blaek, only fore tibiae basally white.

All Panama specimens have been taken east of the Canal Zone on the At-

lantie side of the continental divide, from the Rio Pequeni and Rio Boqueron at

the head of Madden Lake, to Pito, near the Colombian border. All localities are

in heavy forest in areas of high rainfall. Only eleven specimens have been

taken, in Feb., Mar., Apr. and September.

The species ranges south through Colombia into Ecuador, always west of the

Andes. The type in B. M. is labelled Ecuador, and I have seen specimens from

Cachabe, as well as Colombian examples. Barretto (1957) records a specimen

from Mt. Roraima, Br. Guiana, but I have not seen this specimen and it may not

be conspecific with transposita.

Genus Catachlorops Lutz

Lutz 1909, Inst. Osw. Cruz. em Manguinhos, p. 29. Fairchild, 1940, Ann.

Ent. Soe. Amer., 33(4): 689, full references. Barretto, 1946, An.

Fac. Med. Univ. S. Paulo, 22: 151-183; Fairchild, 1969, Arq. Zool. S.

Paulo, 17(4): 210.

Only 1 of the several subgenera of Catachlorops is represented in Panama,

the bulk of the species of the genus being Brazilian.

Subgenus iia Enderlein

Psalidia Enderlein 1922, Mitt. Zool. Mus. Berlin 10: 344.

Four species and 1 well-marked variety occur in Panama, separated by the

subjoined key. All have narrow frons and slender ridge-like callus, very slender

palpi, unbanded eyes, and characteristic wing patterns. Males of 3 species are

known and easily associated with their respective females.

Key to females of Psalidia

Ly Dorsal spine of third antennal segment short, seldom reaching end of

basal plate. First posterior cell of wing broadly open. Black or

grayish species with a white triangle on fourth tergite. Wings hya-

line with an irregular blackish band beyond end of discal cell, the

WINE MOCK HUANG J ccacvicscnoananciabemnsy berehinaranees. umbratus (p. 73)

G. B. Fairchild: Tabanidae of Panama | 71

Dorsal spine longer, extending beyond end of basal plate. First posterior

cell closed and petiolate, rarely narrowly open. Dark brown to yel-

lowish species, with abdomen unmarked or a faint median stripe.

Wings. Otherwise & as sgices ss sdevadauassaadenssvuqusnieswasiss yuslebaGecweckpee de chutes coun chee 2

Wings sharply patterned, with a clearly defined hyaline band extending

across wing from costa to hind margin between end of discal cell

and ends of basal cells. Basal cells and distal band dark brown, the

latter with large oval clear fenestra in marginal and submarginal

cells, and most of apical Cell hyaline .........ccccccccccescoes scurrus (p. 73)

Wings diffusely patterned, without a sharp median band.........cccccccscsscses 3

Legs entirely deep black and black pilose, except for narrowly yellow

knees and paler coxae. Body pilosity brown ventrally and laterally,

black dorsally, without a median dorsal abdominal stripe. Wings as

12): ES secu ids Ssinundencamnutensvash <anapsultsciahanasiceatt ences fortunensis (p. 72)

Legs yellow to red except darker apices of foretibiae and all tarsi more

or less apically infuseated. Pilosity of legs and body yellow to cop-

pery red, the abdomen always with a more or less distinet broad

dersormedian.stripe. of Daler. DLLOSILY 4 caisdespadasesnpncisrse-onssndbecdentnsdinnsss 4

Costa, basal cells, discal cell and bases of marginal and submarginal

cells yellowish to lightly smoky, a contrasting black spot at end of

stigma, and fenestra in marginal and submarginal cells large and dis-

tinct. Body vestiture yellowish. Eyes generally uniformly bright

ey Ia ao OR RED ok OE De AS fulmineus var. ocellatus (p. 71)

Whole wing more or less deep brown, with hyaline or subhyaline fenestra

in discal cell, below stigma, and in apices of marginal and sub-

marginal cells. Body color generally deep rufous. Eyes usually

faintly bicolored, darker in upper half ......scccccssesees fulmineus (p. 71)

Catachlorops (Psalidia) fulmineus (Hine)

Tabanus fulmineus Hine, 1920, Ohio J. Sci., 20(6): 186. Female, Gatun,

Canal Zone.

Dichelacera (Psalidia) fulminea: Fairchild, 1942, Ann. Ent. Soc. Amer.,

35(4): 472.

Psalidia ocellata Enderlein, 1925, Mitt. Zool. Mus. Berlin, 11(2): 393, fe-

male Muzo, Colombia. Fairchild, 1940, Ann. Ent. Soe. Amer., 33(4):

693, fig. 9, full references.

Dichelacera (Psalidia) fulminea form ocellata: Fairchild, 1961, Rev.

Biol. Trop. 9(1): 27.

Tabanus festivus Hine, 1920, Ohio J. Sci., 20(6): 187, female, Gatun,

Canal Zone. Dunn. 1934, Psyche, 41(3): 174. Not T. festivus Wied.

1828.

Bellardia fureata Bigot, 1892, Mem. Soc. Zool. France, 5: 631, female.

Patrie inconnue. Not Dichelacera (Psalidia) fureata (Wied.) 1828.

Catachlorops (Psalidia) fulmineus: Fairchild 1971, Cat. Dipt. Americas

S. of United States, 28, Tabanidae p. 71. Wilkerson 1979, Cespedesia

8(31-32): 270-272, figs. 73, 74.

12 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

The two forms of this species separated in the key are indistinguishable

structurally, and not all specimens can be placed with certainty on color char-

acters. I have seen the types of all the names referred here; fulmineus is the

rufous form with dark wings, all the other names refer to light specimens for

which ocellatus End. appears to be the only available name.

In Panama both forms occur in the same localities and seem to range

throughout the country in heavy forest at altitudes below 2500 ft. Both were

fairly abundant around Almirante in Bocas del Toro Prov. and at Sta. Fe in the

mountains of Veraguas Prov. Elsewhere the form ocellatus seems the more

abundant, though fulmineus has been secured at eight localities, including the

Canal Zone and Darien Prov. Both forms are also on the wing at the same

time. With the exception of three oecellatus in January and two in September,

all specimens were taken in the months from May to August.

Males of each form have been seen, 1 of ocellatus, 5 of fulmineus. They

differ only as to color, in the same way as the females. Notes on the color of

the eyes in life, available for three fulmineus and five ocellatus, indicate that

the eyes are bicolored in fulmineus, either bicolored or uniformly green in ocel-

latus. In the bicolored eye, the upper two fifths is darker, sharply separated

from the lower lighter green.

Aside from Panama material, I have seen specimens of f. oecellatus from

Vera Cruz and Tabasco, Mexico; Peten, Guatemala; Lancetilla, Honduras; Blue-

fields, Nicaragua; Limon Prov., Costa Rica; Muzo and Buenaventura, Colombia.

Fulmineus seems rarer, but I have specimens from Lancetilla, Honduras and

Costa Rica. Wilkerson (loc. cit.) took the ocellatus form in western Colombia

(Cauca, Guapi), but reported the fulmineus form only from Choco and Antioquia.

Catachlorops (Psalidia) fortunensis Fairchild n. sp.

A large dark species with black legs and dark brown antennae, bare sub-

callus, closed first posterior cell and wing pattern like C. (P.) ocellata.

Female. Length 18mm., of wing 15.5 mm. Eyes bare, color and pattern un-

recorded. Frontal index 10.4, divergence index 1.7. Frons dark reddish brown

pollinose, the callus dark brown, nearly black. Subcallus largely bare, somewhat

inflated, rugose, yellowish brown, the pollinose margins yellow brown, somewhat

paler than frons. Frontoclypeus and genae concolorous, the sparse beard dark

brown. Vertex with a slender bare stripe continuing down behind the head. A

dense fringe of stout black bristles forming a postocular band just below eye

margin. Seape and pedicel dark brown, thinly pollinose, densely black setose.

Basal plate dark. reddish brown, dorsal spine and style blackish. Palpi dark

brown, densely black haired. Proboscis thick, dark brown, labella black, largely

shiny and sclerotized.

Mesonotum with reddish brown integument, dark brown pollinose and with a

slender dark median stripe, wholly black pilose. Seutellum concolorous. Pleura

slightly paler, its pollinosity grayish brown, with long brown hairs. Coxae con-

eolorous with pleura. Femora, tibiae and tarsi all entirely black and black

haired, except for the yellow femoro-tibial junction. Wings not distinguishable

from those of ocellata. Basicosta blackish without macrotrichiae. Halteres dark

orange.

Abdomen with mahogany red integument, subshiny, dorsally entirely black or

very dark brown pilose, ventrally with paler integument and pale pollinosity with

slightly paler hairs on hind margins of sternites.

G. B. Fairehild: Tabanidae of Panama 73

Holotype female, Chiriqui Prov., Panama, Fortuna, 1000 meters elev. 12-23-

VIII-1976, H. Wolda coll. To be deposited in F. 8. C. A. The specimen lacks

the left mid leg and the terminal annulus of left antenna.

It is with some hesitation that this single specimen is deseribed, but its very

distinctive black legs and dark color separate it readily from the 2 other Pana-

Manian species, ocellata and fulmineus. The frons is also narrower than any

other specimens measured.

A male taken in a U. V. light trap at Fortuna on 27-1977 by Dr. H. Wolda

may be the male of this species. It is, however, more reddish than the type,

with red legs and orange pilose thorax and abdomen. The wings, though ecrum-

pled (the specimen was preserved in alcohol for 5 years) appear to have the

same pattern as the female, with the substigmatie spot emphasized, as in ocel-

lata. It is, however, very much darker than the single available male of that

Species, both in body and wings. In body color it is closer to fulmineus males,

but the wing lacks the extensive dark infuseation of that species.

Catachlorops (Psalidia) seurrus Fairchild

Catachlorops (Psalidia) seurrus Fairchild 1958, Ann. Ent. Soc. Amer.,

51(6): 526-527, figs. 8, 18, female, Panama; 1971, Cat. Dipt. S.

Amer., Fase. 28: 71 synonymy.

This species is rare in Panama, though probably widespread, as records are

available from Darien and Panama provinees and the Canal Zone. All were

taken during the dry season from Jan. to March and on the dryer Pacific side of

the Isthmus, so the species is separated both ecologically and seasonally from

fulmineus. It may be more common than indieated by the records, since little

collecting has been done during its flight season. The species occurs in

Guatemala and Mexico, and most probably in intervening areas, though no

records are available. The eye is bronzy in life, without markings. There seem

no records from south of Panama.

Catachlorops (Psalidia) umbratus (Hine)

Tabanus umbratus Hine, 1920, Ohio, J. Sci., 20(6): 187, female, Costa

Rica.

Catachlorops umbratus: Kroeber, 1934, Rev. Ent. 4(2): 274.

Dichelacera (Catachlorops) umbratus: Fairchild, 1951, Ann. Ent. Soc.

Amer., 44(3): 448-450, fig. 2, Panama, full references;

This species is highly arboreal. Of 32 specimens now available, 27 were

taken attracted to human collectors in the forest canopy, two were taken at

ground level and three lack information. The bulk of our specimens therefore

have been collected where canopy collections were made, and distribution may

be wider than records indicate. The species has been taken east of the Canal

Zone, along the continental divide at about 2000 ft., on the Atlantic side of the

same range (Cerro Azul), and at a few other localities in the same general area.

We also have a short series from the upper Rio Tuira, in Darien Province, and

from Chorecha, Chiriqui Prov. Most specimens were taken in February, with

lesser numbers in December, January and March, in various years. The eyes in

life are bright emerald green, unicolorous.

74 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

The male is similar in all non-sexual characters to the female. The eyes

are bare with an upper area of greatly enlarged facets clearly demarcated from

the small facets and occupying about two-thirds of the eye area. There is a

small tubercle in the notch at vertex, without vestiges of ocelli. The palpi are

clubbed and bluntly faleate Plesiotype male, Las Cumbres, Panama Prov., 23

Feb. 1959, G. B. Fairchild coll., taken on window of house.

Due possibly to its highly arboreal habits, the species is known so far only

from Costa Rica and Panama, though Wilkerson (1979) gives an unconfirmed re-

port of the species' occurrence in Antioquia, Colombia.

Genus Dasychela Enderlein

Enderlein 1922, Mitt. Zool. Mus. Berlin, 10(2): 345. Fairchild, 1969, Arq.

Zool. S. Paulo, 17(4): 212.

Subgenus Dasychela End.

The genus is a small one of a few specialized species localized in north-

western South America. The species have unicolorous pubescent or rarely bare

eyes, long proboscis, long dorsal tooth on third antennal segment and a dark

wing pattern which excludes the crossveins at apex of basal cells. Only a single

species occurs in Panama.

Dasychela (Dasychela) badia (Kroeber)

Dieladoeera badia Kroeber, 1931, Rev. Ent., 1(14): 402, fig. 3, female,

Panama. Fairchild, 1940, Ann. Ent. Soc.Amer., 33(4): 689, fig. 7.

Stypochela badia: Fairchild, 1958, Ann. Ent.Soc.Amer., 51(6): 518, Pl. 2,

fig. 14.

Tabanus venenatus: Hine 1917, Trans. Amer.Ent. Soc., 43(765): 295,

Costa Rica. Not Osten Sacken 1886.

The species is easily recognized in our fauna by the uniform dark brown

color, the abdomen with faintly paler broad mid-dorsal pilose triangles on ter-

gites 3 and 4, and the wings with a diffuse broad brown band which leaves the

crossveins and apex hyaline. The dorsal tooth on third antennal segment is very

long and with a recurved tip, reaching well beyond end of basal plate. The pro-

boseis is unusually long, over twice length of palpi, brown pollinose with small

compact labella. The pilose eyes are dark purplish green in life, unbanded.

Panama material is all from areas of heavy forest and high rainfall, in

Darien, Colon, and Bocas del Toro provinees, and in the mountains at about 2000

feet in Panama, Cocle and Veraguas provinces. The species has usually been

taken from January through March, in the dry season, but a long series from

Cope, Cocle prov. was secured in Aug. 1977 by R. B. and L. S. Kimsey. At the

right place and time, it may be abundant and annoying, as it attacks man read-

ily.

I have seen the Costa Rican specimen on which Hine's misdetermined record

was based. Hine later, in his MS notes on a visit to the British Museum, saw

the types of venenatus, and two specimens of the present species and realized

his error. In his collection when I studied it in 1940, this Costa Rican specimen

G. B. Fairchild: Tabanidae of Panama 75

was labelled Dichelacera peruviana Bigot, comp. with type, and it is indeed

closely related. It is possible that further collecting in Colombia and Ecuador

will show that badia is but a northern race of the earlier described peruviana.

The species ranges from Costa Rica into eastern Panama and probably extreme

northwestern Colombia, though I have no records from the latter country.

Genus Diecladoeera Lutz

Lutz 1913, Brazil Medico, 27(45): 5. Fairchild, 1969, Arg. Zool. S.

Paulo, 17(4): 217.

Dicladocera nigrocoerulea (Rondani)

Tabanus nigrocoeruleus Rondani, 1850, Nouv. Ann. Rend. Accad. Sci. 1st.

Soe. Agric. Bologna (3)2: 191.

Dicladocera nigrocoerulea: Fairchild, 1972. Cat. S. Amer. Dipt. Fase. 28,

p. 74, synonymy. Wilkerson, 1979, Cespedesia 8(31-32): 297-298, fig.

94.

The single species of this genus occurring in Panama is a large stout shiny

black insect, with the wings basally black to beyond end of discal cell. There

are clear fenestrae in ends of basal cells and middle of diseal cell, and a small

median tuft of white hairs on tergite 4. It differs from species of Stibasoma by

having some setae on basicosta, soft pollinose labella on proboscis and slender

tibiae.

Three females were taken at Cana, in the mountains of Darien Prov. be-

tween 28 July and 11 Aug., 1965. The species ranges thenee to Venezuela and

Ecuador. It is surprising that only one of 16 species of Dicladocera listed by

Wilkerson from western Colombia reaches Panama, and that one only in extreme

eastern Panama.

Genus Stibasoma Schiner

Schiner 1867, Verh. z. b. Ges. Wien, 17: 310. Fairchild, 1940, Ann. Ent.

Soc. Amer., 33(4): 684, full references; 1953, Ann. Ent. Soc. Amer.,

46(2): 267, key. Goodwin and Murdoch, 1974, Ann. Ent. Soc. Amer.,

67(1): 112-114. |

Rhabdotylus Lutz, 1909, Inst. Osw. Cruz em Manguinhos, p. 29.

Fairchild, 1942, Ann. Ent.Soc. Amer., 35(4): 472. Barretto, 1951,

Arq. Mus. Nac. Rio de Janeiro, 42: 72-73. Fairchild, 1950, Psyche,

57(4): 125.

Flies of this genus are generally thick-set hairy bee-like species, with bare

unicolorous or bicolored eyes. In Panama most of the species seem predomi-

nantly arboreal in habits, and unless collections are made in the forest canopy,

they are rarely taken. The group may not be a natural one, as one or more of

the combination of characters used to define it may be absent or poorly devel-

oped in any given species. Thus the tibiae are sometimes slender and do not

always bear especially long hair fringes. The frontal callus is usually flat and

nearly as wide as frons, always extended above in a narrow or broad ridge to

76 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

the vertex. Many species have the frons extensively bare, especially at vertex.

Most species have the wings colored or patterned, seldom entirely hyaline. The

antennae seem always short, seldom equalling the usually inflated palpi in

length. |

Rhabdotylus Lutz is retained in a subgeneric sense for a small group of

greenish bodied species which have the head characters of Stibasoma but are

less hairy, and lack inflated tibiae. The known males resemble their females,

(except chionostigma) but have bare holoptie eyes, with a large and well demar-

eated area of enlarged facets in the upper two-thirds of the eye. All of the

Species discussed here, and no others, were reported from western Colombia by

Wilkerson (1979), who also figured the wings of 3 of the species.

Key to species

P Tibiae not inflated. Wings with only costal cell tinted, veins at most

lightly brown-margined. Abdomen grass green to bluish green in

life, fading to yellowish, sparsely black haired except for obscure

pale-haired median triangles on second to fifth tergites ..........sessese

SRR OP Be TaD Se SOE Te Ute 5 anima ts Ve Rl GE (Rhabdotylus) venenata (p. 80)

Tibiae inflated. Wings black, yellow or patterned. Abdomen not green 2

2. Wings entirely black except for dilute apex and hind margin. First two

abdominal tergites densely sulfur yellow-haired, remaining tergites

varying from almost wholly blaeck-haired to wholly orange rufous.

Subecallus denuded and Shiny .....ccccscscsosccscvcscsccceees panamensis (p. 79)

Wings not entirely black. Abdomen Otherwise .........scccccccscrsscscccscccesces 3

3. Wines with: PPORINENE DICK WNAPRINGS s0s8.2. 0k idelcaeete ces décsdancacccsedgevcccocusceses 4.

Wings yellowish tinted or vaguely dusky, without prominent black mark-

PUR States antes oes otan ree hee tucat odes antec cdbedaclcsts oPeskbecsleaccsesndaticosvcgeccvecces. 3)

4, Wings with a prominent subapical black area in outer halves of marginal

and first submarginal cells, the anterior half of wing otherwise

bright yellow. Abdomen black, the first tergite with a short bright

yellow hind marginal band or transverse patch, the second with a

complete, narrower yellow DaNnd: | 22... ccscsesececsccscoees apicimacula (p. 77)

Wings largely black or brown, but with a prominent yellow hyaline patch

from costa to and including discal cell, and apex beyond tip of R2+3

hyaline. Abdomen black or brown above, but with all sternites with

broad yellow-haired hind Margins ........scccccccsescees chionostigma (p. 77)

Ds Large species, over 16 mm. Wings deep black at extreme base, remain-

der yellowish brown tinted basally, the apex and hind margin broadly

grayish hyaline. Thorax entirely deep velvety black. Abdomen with

first tergite black, second tergite black or usually partly or wholly

rufous orange-haired, remainder all rufous orange. Second to last

tergites with sulfur yellow-haired hind marginal bands of variable

WED "Oy a tRNA Oe BA. GRO eo ee flaviventre (p. 78)

Smaller species, less than 14 mm. Wings yellowish hyaline to base, the

outer margin and apex broadly dusky grayish. Thorax ranging from

blaek to orange rufous-haired, with a white or orange hair tuft be-

G. B. Fairehild: Tabanidae of Panama T7

fore wing bases and on each side of secutellum. Abdomen ranging

from nearly entirely black to partly orange rufous, always with at

least traces of yellow hind marginal bands on tergites 2 and 3, usu-

ally all tergites prominently banded .......04. seocseeeeeeftlvohirtum (p. 79)

Stibasoma apicimacula Fairchild

Stibasoma apicimacula Fairchild 1940, Ann. Ent. Soc. Amer., 33(4): 686-

688, Pl. 1, Fig. 6, female, Panama; 1951, Ann. Ent. Soe. Amer.,

44(3): 451-452; 1953, Proce. Ent. Soe. Washington, 55(5): 241.

This species is the least bee-like of the Panama species. In addition to the

characters in the key, the insect is black including antennae and palpi. Legs

black, including tarsi, with white hairs at bases of all tibiae, hind coxae, and

base of hind femora dorsally. Thorax usually with small white hair tufts before

and below wing bases. Abdomen with yellow hairs in middle of hind margin of

first tergite and a complete narrow hind marginal band on seeond. All sternites

with narrow pale-haired hind margins. Subcallus bare and shiny in middle.

The bulk of our material has come from Yellow Fever camps along the up-

per reaches of the Rio Mandinga, between Colon and San Blas. It also occurs at

Almirante, Bocas del Toro, at El Valle, Cocle Prov., and at Sta. Fe, Veraguas

Prov., the last two localities in mountain forest above 2000 feet. Of the

twenty-eight specimens bearing data, twenty-six were taken at platforms in the

forest canopy. Panama records run from May to September. Specimens from

Rio Raposo, near Buenaventura, Colombia, were taken in Jan., May, and June

1964. They were also taken at platforms in the lesa canopy. A single speci-

men is also known from Costa Rica.

Stibasoma chionostigma (Osten Sacken)

Tabanus chionostigma Osten Sacken, 1886, Biol. Centr. Amer.,Dipt. 1:

04, Pl. 1, fig. 11, female, Guatemala. Knab, 1913, Proc. U. S. Nat.

Mus. 46, No. 2033, p. 411.

Stibasoma chionostigma: Fairchild, 1940, Ann. Ent. Soe. Amer., 33(4):

685, fig. 5, Panama, full references; 1964, J. Med. Ent., 1(2): 171.

1971; Cat.S.Amer.Dipt. Fase. 28, p. 75 synonymy.

Stibasoma pachycephalum Bigot, 1892, Mem. Soc. Zool. France, 5: 636,

Mexico. Fairchild, 1956, Smiths, miscell. Colls., 131(3): 25, Types

seen.

Stibasoma flavistigma Hine, 1912, Ohio Nat., 7(7): 516, female, Mexico.

Stibasoma bifenestrata Philip, 1966, Ann. Ent. Soe. Amer. 59(2): 526,

figs. 5, 12, female,Nicaragua.

The species is easily recognized by the black and yellow patterned wings.

The subeallus and face are silvery pruinose, rarely the former bare in middle,

the beard and palpi white haired. Thorax is mainly black, but with silvery white

hair tufts above fore coxae, before and beneath wings bases, and on each side of

scutellum. Legs are black, black-haired except for silvery hair patches on bases

of all tibiae. Abdomen black haired above, apparently with pale hairs on last

one or two tergites, beneath all sternites with wide yellow-haired hind borders.

78 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

This is the rarest Panama Stibasoma, as only a few specimens are known,

from Chiriqui, Canal Zone, and Cerro Azul, Panama Prov., as previously re-

ported (Fairchild 1940). Additional specimens are 1 female, France Field, C. Z.,

12 May 1964 in horse baited mosquito trap; 1 female without data, probably

Canal Zone, and 1 male, Barro Colorado Id., C. Z., 25 June 1967, taken at U. V.

light, R. G. Beard coll. It is probably arboreal, like its congeners, but no infor-

mation is available. This species ranges form Mexico to Colombia.

The male is quite different in appearance, the wing having a broad yellow

band from costa to hind margin covering the basal and most of anal and auxil-

lary cells. The abdomen is yellow in ground color, with tergites 1 and 3 to 6

black-haired, 2 yellow-haired. Mesonotum chocolate brown, the scutellum red-

dish. Legs as in female. Eyes with upper facets much enlarged, covering fully

2/3 of eye area, brown and sharply demarcated from the small facets. Anten-

nae as in female, but more slender. Face and cheeks silvery pollinose, white-

haired. Palpi oval, porrect, blackish. Proboseis with fully selerotized labella. _

The species seems quite variable in color characters, some being wholly

black in body color, others quite reddish. The subeallus may be largely bare and

shiny, or thinly silvery pollinose, the former condition possibly due to wear.

Stibasoma flaviventre (Macquart)

Tabanus flaviventris Macquart, 1848, Dipt. Exot., Supp. 3, p. 11, Pl. 1,

fig 4, female, Rio Negro. |

Stibasoma flaviventre: Kroeber, 1934. Rev.Ent., 4(2): 263, references.

Fairchild, 1956, Smiths. Miscell. Colls., 131(3): 17, synonymy, type

seen. Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1): 112,

figs, 21, 33.

Stibasoma stilbium Fairchild, 1953, Ann. Ent. Soe. Amer., 46(2): 264-267,

Pl. 2, fig. 11 Panama; 1953, Proc. Ent. Soc. Washington, 55(5): 241.

In addition to the characters in the key, the legs are wholly black, including

tarsi, black-haired except for patches of silver white hairs at bases of all tibiae

on outer surface. Subeallus silvery white pruinose, as is face to a lesser degree.

Halters all black. Sternites with yellow hind marginal bands. As with most

species of Stibasoma, color is variable, the abdomen varying from mainly yellow-

haired to mainly copper-haired.

In Panama the species is rare and probably highly arboreal. Our four speci-

mens from Almirante,. Boeas del Toro Prov., were all collected in the canopy.

Three additional specimens, from Tacareuna and Rio Tuira, Darien Prov., and La

Zumbadora, Cerro Azul, Panama Prov., though not so labelled, were probably

taken in the canopy, as tree platform collecting was underway at all three lo-

ealities. We also have 7 females from Lancetilla, Tela, Honduras, all taken in

the treetops, and 2 from Teapa, Tabasco, Mexico, probably also from the forest

eanopy. Records inelude all months from March to September. The male is like

the female in color. The eyes are bare, holoptic, with a large area of sharply

demarcated large facets entirely surrounded above by a strip of small facets.

There is no visible tubercle at vertex. The eye of the female in life is bi-

eolored, black above, greenish black below, the two colors sharply separated.

The species ranges from Mexico at least to the northern Amazon basin. Speci-

mens have been seen from Guatemala, Surinam and Colombia in addition to

those mentioned above, while a subspecies occurs on Trinidad.

G. B. Fairchild: Tabanidae of Panama 79

Goodwin and Murdoch (1974) collected 10 larvae from between the leaf

bases of arboreal Bromeliaceae, from which they reared a single male. The lar-

vae are nearly black, the pupae with long horn-like thoracic spiracles.

Stibasoma fulvohirtum (Wiedemann)

Tabanus fulvohirtus Wiedemann, 1828, Auss. zweifl. Ins. 1: 155, female,

Brazil.

Stibasoma fulvohirtum: Fairchild, 1940, Ann. Ent. Soc. Amer., 33(4):

685, fig. 4, Panama, full references; 1951, Ann. Ent. Soc. Amer.,

44(3): 451; 1971, Cat. S.Amer. Dipt., Fase. 28, p. 76, synonymy.

Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1): 114, figs

21, 34.

In addition to the characters in the key, the subeallus and face are silvery

pruinose, palpi black or partly silver-haired. Femora black, tibiae bicolored,

first and last pair white at base, mid pair mostly white, tarsi yellowish, white-

haired. Eyes unicolorous, reddish green in life.

This is the most abundant and widespread species of the genus in Panama.

it has been taken at 11 localities from Darien to Chiriqui, most abundantly in

lowland forest on both coasts and appears to fly throughout the year. It is pre-

dominantly arboreal, and is attracted readily to man. Goodwin and Murdoch

reared over 20 adults from larvae collected in terrestrial Bromeliaceae at 6 lo-

ealities from Bocas del Toro to Darien.

I have seen the type of compactus Walk. in British Museum, and types of

fulvohirtum Wied. in Berlin, Frankfort and Vienna. All are paler than any

Panama specimens, perhaps due to fading with age. The species is reported

from Costa Rica to Brazil. Panama material varies much in color of vesiture,

some having largely golden-haired thorax and prominently banded abdomens, to

almost totally black specimens.

Stibasoma panamensis Curran

Stibasoma theotaenia panamensis Curran, 1934, Fam. Gen. N. Amer.

Dipt., p. 153, fig. 23, head, no description. Fairchild, 1951, Ann.

Ent. Soe.Amer., 44(3): 451, fig. 3.

Stibasoma theotaenia var. panamensis, Fairchild, 1940. Ann. Ent. Soc.

Amer., 33(4): 685-686, Panama, full references.

Stibasoma panamensis, Fairchild 1953, Ann. Ent. Soc. Amer., 46(20): 267,

key.

Easily recognizable among Panama species by the black wings and bright

yellow basal 2 segments of abdomen. The eyes in life are bicolored, black

above, paler grayish, below, with a sharp line of demarcation. Legs are black

except for rufous tarsi and patches of white hair at bases of at least hind

tibiae. Knob of halters ivory white, stem black. Subcallus somewhat inflated,

bare and shiny. Size is variable, from a wing length of 10mm. to 14mm.

The species is fairly common in the forest canopy at Almirante in Bocas del

Toro Prov., and on the Rio Mandinga, which forms the border between Colon

and San Blas. It has been taken a few times on the Atlantic side of the Canal

Zone (Barro Colorado Id. and Ft. Sherman) and in nearby Panama province

80 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

(Arraijan, Chorrera, Cerro Azul). We have also seen 4 females from Lancetilla,

Tela, Honduras, July, Aug. 1953, Jan. 1954, and 4 females Curiche, Choco,

Colombia, April, May, and Oct. 1967. Of the 24 specimens on which data are

available, 16 were taken at platforms in the forest canopy. It appears to be on

the wing throughout the year. Abdominal colors appear to vary with geography.

All those from Honduras have tergites 3-6 pale orange and orange pilose, as do

those from Boeas del Toro, and 1 from Araijan, Panama prov. The Mandinga,

Colon Prov. specimens, have tergite 3 largely black-haired, the remainder dark

coppery, more or less mixed with black. A specimen from near Pacora, Panama

prov., has these tergites all coppery red-haired, while those from Darien Prov.

and Choco, Colombia have tergites 3 to 5 entirely black-haired. These latter

specimens much resemble Stibasoma theotaenia Wied. in color, but are struc-

turally quite distinct, so that panamensis cannot be a subspecies of theotaenia.

Stibasoma (Rhabdotylus) venenatus (Osten Sacken)

Tabanus venenatus Osten Sacken, 1886, Biol. Centr. Americana, Dipt.,

24, female, Guatemala, Panama. |

Rhabdotylus venenatus: Stone, 1944, Bol. Ent. Venez. 3(3): 130,

Venezuela. Fairchild, 1956, Smiths. Miscel. Colls., 131(3): 31, types

seen. Philip, 1960, Proce. Calif. Acad. Sci., Ser. 4, 31(3): 87, desig-

nates lectotype.

Stibasoma (Rhabdotylus) venenata: Fairchild, 1971, Cat. Dipt. S.Amer.,

Fase. 28, p. 77. Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer.,

67(1): 114, figs. 21, 35.

Gymnochela (Amphichlorops) venenatus: Kroeber, 1932, Rev. Ent., 2(1):

92-93, fig. 32, male, female, Venezuela, Ecuador, Bolivia,

Guatemala, Panama, Costa Rica.

Amphichlorops venenata: Kroeber, 1934, Rev. Ent., 4(2): 271.

The three species belonging to this group, planiventris Wied., viridiventris

Macq., and venenata O.S., are difficult to separate, but I have now seen the

types of all three, and about 25 additional specimens. Planiventris Wied. has

the narrowest frons, the basal callus is small, rather flat, and surmounted above

by two slender depressions or pits which define the beginning of the slender

median ridge reaching toward the vertex. The antennae are short and stubby.

The abdomen, in undenuded specimens, usually has the second and third tergites

wholly yellow-haired and always with yellow-haired lateral margins, the fourth

to sixth tergites wholly black-haired, the seventh again pale haired. The wings

are yellowish tinted, with more or less strongly brownish or yellowish margin of

the veins. The femora are largely or wholly black-haired.

Viridiventris has a broader frons, the frontal callus rounder, more protu-

berant, and merging almost imperceptibly into the broader more flattened and

less defined central ridge. It lacks the pair of pits above the callus. The an-

tennae are less stubby than in planiventris. The abdomen has extreme sides of

all tergites with some pale hairs, the second and third tergites may have narrow

pale-haired hind borders as well. The wings are somewhat fumose in the basal

cells and below stigma, but lack the brown or yellow vein margins usual in

planiventris. The femora are consistently black haired.

Venenata is similar to viridiventris in the frons and callus, though generally

with still broader frons, but with both callus and central ridge better defined,

the former usually clearly separated from eye margins, and in one specimen,

G. B. Fairchild: Tabanidae of Panama 81

indications of the pits found in planiventris. The antennae are definitely longer

and more slender, more deeply bifid, than either of the other two species. All

but one of my specimens are badly denuded. The one good specimen, from

Panama, has sides of tergites 1 to 5 pale-haired, the last 3 blaeck-haired, the

dorsal surface largely black-haired, but with small flat white pilose triangles on

tergites 2 to 5. What remains of the vestiture on the other examples corre-

sponds with this pattern. The wings are evenly slightly grayish, the veins faintly

to strongly brown margined, as in planiventris. The femora are consistently pale

haired.

Males of planiventris and viridiventris have been studied. They are very

similar in all respects. The antennae of viridiventris are considerably more

Slender, less chunky. Femora of both are largely pale haired, and abdomens of

both entirely yellow haired. Males of venenata have not been seen.

Planiventris and viridiventris appear confined to S.E. Brasil, where they ap-

parently occur together.

Philip (1960) suggested that Central American specimens might belong to

two species, both represented in Osten Sackens type series. He felt that they

could be separated on color of the vestiture of palpi, femora, and pleura, orange

in the Panama specimen selected as lectotype, whitish in the Guatemala speci-

men. Two of my Panama specimens, from Rio Changena, Bocas del Toro Prov.,

2400 ft., 12 Aug. and 8 Sept., 1961, and another in U. S. N. M. from Caracas,

Venezuela, Sept. 1939 agree with the lectotype in these characters, and in hav-

ing more heavily brown margined veins. A third specimen, from Balzapambana,

Ecuador is like these two, but with vestiture but slightly yellow tinged. The

three other Panama specimens, from the Cerro Azul area, in Panama province

east of the Canal Zone, at altitudes of about 2000 feet, 6 Feb. 1953, 12 Feb.

1955 and 24 Sept. 1956, all have whitish hair on beard, pleura, and femora, but

black haired palpi. The wings are lighter, the veins with barely discernible

brown margin. A specimen from Carillo, Costa Riea, May 1903, C. F. Under-

wood, and another in U. S. N. M. from San Carlos, Miranda, Venezuela, are like

these last three Panama specimens. There is also a specimen from El Volean,

Chiriqui, Feb. 1936, in A. M. N. H., but I took no notes on color of vestiture.

My series also varies much in size, the smallest having a wing length of 11 mm.,

the largest 16 mm. Longer series of better preserved specimens will be needed

to settle this matter, but in view of the notable tendency of species of Stiba-

soma to show color variation, I feel it is more likely that at least viridiventris

and venenata will eventually prove but races of a single species.

In Panama the species has been taken only above 2000 ft. in areas of high

rainfall. The three Panama localities are also notable for the abundance of epi-

phytie and terrestrial Bromeliaceae. Goodwin and Murdoch (1974) succeeded in

rearing 2 of 25 larvae collected between the leaf bases of arboreal Bromeliaceae

at Cerro Azul, Panama Prov., and Cana, Darien Prov. These yielded a male and

a female, both small in size. The male has a yellowish abdomen and orange

hairs on pleura, and the wings are badly broken. The female is quite teneral,

also with orange haired pleura. The abdomen of the male is almost entirely de-

nuded; that of the female shrunken and discolored. In spite of differences in

appearance, I think these are probably venenata.

Genus Cryptotylus Lutz

Lutz 1909 Inst. Osw. Cruz em Manguinhos, p. 29; 1913, Brazil Medico,

45: 5. Fairchild, 1969, Arq. Zool. S. Paulo, 17(4): 212. Philip and

82 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Fairchild, 1956, Ann. Ent. Soe. Amer., 49(4): 313-324, figs. (as sub-

genus of Chlorotabanus). Wilkerson 1979 Cespedesia 8(31-32): 314-

316.

The species of this group resemble Chlorotabanus in unicolorous greenish or

orange coloration, but a small frontal callus is present and the basal plate of

antenna has a strong dorsal angle or sharp spine. Like Chlorotabanus they are

nocturnal. Two species oceur in Panama, separated as in the key below.

Key to species

Basal plate of third antennal segment broad, angled below, the dorsal

tooth not longer than its width at base. Vestiture orange to dark

rufous in females, pale yellow in males. Fork of third vein rarely

with A sherteappenditc tic arr ies Wiese unicolor (p. 82)

Basal plate slender, not angled below, the dorsal tooth longer than its

width at base, slender. Vestiture generally paler, yellow in both

sexes. Fork of third vein nearly always with a short to long ap-

OLS |b Ct eee ee Oa >, sie ele A oeseawccas eudtcieipebectns chloroticus (p. 82)

Cryptotylus unicolor (Wiedemann)

Tabanus unicolor Wiedemann, 1828, Auss. Zweifl. Ins. 1: 141.

' Cryptotylus unicolor: Fairchild, 1940, Rev. Ent., 11(3): 719-720, fig. 4,

full references. Wilkerson, 1979, Cespedesia 8(31-32): 315-316, fig.

34N.

Chlorotabanus (Cryptotylus) unicolor: Philip and Fairchild, 1956, Ann.

Ent. Soe. Amer., 49(4): 318-320, synonymy.

In addition to the characters in the key, this species is usually larger and

more rufous in color than chlorotiecus. Males of both species of Cryptotylus

have small tubereles between the eyes at vertex, lacking in Chlorotabanus. In

Panama the species is not uncommon along the Chagres river and about Gatun

lake. It has also been taken in Darien and Cocle provinces, always at low ele-

vations. Dunn (1934) reported rearing this species from larvae collected in

water lettuce plants (Pistia stratiotes). Both sexes have been taken in light

traps, and females in horse-baited stable traps and biting horses and cattle; it

does not appear to be attracted to man. The species probably occurs throughout

the eastern half of Panama, ranging thence southward to Bolivia and Brazil.

Records include the months from Dec. to Aug. Wilkerson saw specimens only

from northern Choco and eastern Colombia.

Cryptotylus chloroticus Philip and Fairchild.

Cryptotylus chloroticus Philip and Fairchild, 1956, Ann. Ent. Soec.Amer.,

49(4): 320-321, fig. 7. Wilkerson, 1979, Cespedesia 8(31-32): 314-315,

fig. 34M.

Cryptotylus limonus: Fairchild, 1940, Rev. Ent., 11(3): 720-722, fig. 5;

1942, Ann. Ent. Soe. Amer. 35(1): 88-90. Not Townsend 1897.

G. B. Fairchild: Tabanidae of Panama 83

In Panama, this species is quite abundant about Gatun Lake in the Canal

Zone and nearby Panama, and has been taken also at Aguadulee in Cocle

Provinee, and at Almirante, Bocas del Toro Prov. Unlike uniecolor it has a short

flight period, May through August. It is taken in light traps, horse-baited stable

traps, and biting cattle and horses, but not man. It probably occurs throughout

the lowlands of Panama in swampy areas. The species ranges southward at least

to Sao Paulo, Brazil, but records are seanty. Wilkerson (op. cit.) did not take

this in western Colombia.

Genus Philipotabanus Fairchild

Fairchild 1942, Ann. Ent. Soe. Amer., 35(4): 453 (Tabanus subgenus);

1964 J. Med. Ent., 1(2): 179.

Hemichrysops, Fairchild 1961, Rev. Biol. Trop. 9(1): 29. Not Kroeber

1930. Wilkerson, 1979, Cespedesia 8(31-32): 316-332.

This genus ineludes slender species of small to medium size with ridge-like

to drop-shaped callus, generally sparsely setose basicosta, pollinose labella, uni-

ecolorous eyes and wings with a dark pattern. The species of the three subgen-

era are treated separately here.

Key to Subgenera

1. Wings with an irregular dark pattern of variable extent which always

leaves clear areas surrounding all cross-veins and fork of third vein.

Frons narrow to very narrow 7 to 10x as high as basal width. Palpi

slender. Eyes bronze in life. Always rather slender long-winged

Species; wings nearly as long as or equalling body length ...........ceeeee

wins hagSiaetes odie staeites of dtucted, cetipae chat Nila ahaa tat. ue Philipotabanus (p. 83)

Wings ranging from almost wholly blackish to hyaline with a small dark

area below stigma, but cross veins and fork of 3rd vein not sur-

rounded by clear fenestra when included in dark pattern. .........cseeee 2

2: Slender species with frons 7x as high as wide or narrower, the palpi and

antennae slender, proboscis considerably longer than palpi and with

small labella. Eyes bright green in life ........ Melasmatabanus (p. 90)

Stouter species with broader frons not over 6x as high as wide, palpi in-

flated, antennae broader, proboscis hardly longer than palpi, the la-

bella large. Eyes green or brick red in life, fading to greenish black

iskigrasadincsnasacnetiaveeAchmaass ait) ssescccccscccsccscscseees Mimotabanus (p. 89)

Subgenus Philipotabanus

The nine species of this group known from Panama are keyed below. All

the species are primarily forest inhabitants and all attack man to some extent.

Most of the species are at least partly crepuscular. P. magnifieus is at times

sufficiently abundant and aggressive to constitute a pest. P. nigrinubilus has

been reared from larvae taken under the bark of rotten logs. The subgenus

ranges from southern Mexico to Bolivia and the northern Amazon region. The

species are all very similar in appearance and are difficult to separate in a key.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Key to Philipotabanus (Philipotabanus)

Basal cells of wing entirely hyaline. Dark wing markings reduced to a

small patch below stigma. First and second abdominal segments

pale horn-color, remainder darker ........cecsccseesecssceces grassator (p. 86)

Basal cells dark on at least basal third. Wing pattern more extensive... 2

Entirely black-bodied species with black legs, the abdomen with more or

less white hair on sides and posterior borders of some segments.

Wines Neavily OIBOh TA SNS BUR dvccsessrsoncacssccvesssasiesses suteusd

Dark to pale brown species with brown legs. The wings heavily or

LISI ARE PROD A AR ialieas TAO, as. th ORE, 4

Frons exceedingly narrow, over 10 times as high as basal width, callus

black, a narrow raised line. Black wing marking extends from near

ends of basal cells to beyond fork of third vein, leaving very small

hyaline fenestrae around end of discal cell and fork of third vein. A

small middorsal triangle of white hair on fourth tergite. Antennae

DPSS TY “LE Bien Li xnan Sach R wa ivdia adhe anand nigrinubilus (p. 87)

Frons broader, about 7 times as high as basal width, callus broader,

spindle or club-shaped. Black wing markings less extensive, not

reaching ends of basal cells and with large fenestrae about end of

discal cell and fork of third vein. Apex and hind border of wing

brownish, contrasting with hyaline fenestrae. Margins of all abdomi-

nal segments more extensively white-haired ........cccccees medius (p. 87)

Apex of wing markedly infuscated. Small species generally with wing

length less than 9 mm. Legs blackish, darker than pleura. Scutel-

lum concolorous or paler than MeESONOTUM ...sccccccees magnificus (p. 86)

Larger species, generally with wing length over 10 mm. Legs yellow to

DPOWN,. TOC ier Wee «bee TP ak We. ca sida an We Leave voces 5)

Mesonotum prominently striped. Seutellum subshiny, conspicuously

darker than mesonotum. First posterior cell (5th R) with a more or

less well defined longitudinal pale streak. Dark marking in second

submarginal cell (4th R) usually unconnected with dark pattern, the

latter generally rather streaky and broken. Abdomen light to dark

brown with a more or less well defined darker median integumental

stripe, and ill defined paler-haired triangles on tergites 3 and 4 ......

ssc sda OORELMARHEDN ha URL eh shea ETAT AAT TOUR MeV alee tc RGL Gals US ebrius (p. 85)

No dark spot in second submarginal cell (4th R), wing pattern reduced,

pale brown. Whole insect light brown, mesonotum faintly striped,

seutellum concolorous or paler. Antenna unusually broad, with long

dorsal tooth. Frons 7-8 times basal width .........scceccssss elviae (p. 86)

A dark spot at base of second submarginal cell. Antennae more slender,

the dorsal tooth no more than an acute angle .......c.ccccccccccsccscscveces 7

G. B. Fairchild: Tabanidae of Panama 85

ts Frons broader 7 to 9 times as high as basal width. Wing pattern black,

basal cells over half black, the diseal band broader and intense. At

least mid and hind tibiae brown or blackish ........eee0.- Seis RELRMES Lede enes

HO inal aaa COS pallidetinetus (p. 88)

Frons very narrow, 9 to 11 times as high as basal width. Wing pattern

pale brownish, the basal cells about half hyaline, the diseal band

narrow. |: Mid: and: hind ees: yellow: easccak si pcan ieainw eddies UTIL obs eceens 8

8. Abdomen with a row of small middorsal white-haired triangles. Hyaline

fenestra around end of discal cell completely surrounded by brown ..

TSA eRaa ind ean aber ala ead pterographicus (p. 88)

Abdomen with an ill-defined broad golden-haired triangle on fourth ter-

gite. Hyaline fenestra more or leSS Open DeNINd .........cccccccccccccccccece

Philipotabanus (Philipotabanus) chrysothrix Fairchild

Tabanus (Philipotabanus) chrysothrix Fairchild, 1943 (1942), Ann. Ent.

Soe. Amer., 35(4): 459-460, P1. 2, fig. 17, female, Panama.

This species is not separable structurally from pterographicus Fchld. but av-

erages slightly larger and more rufous in tone. The wing is like magnificus,

with dark spikes along posterior veins, but it lacks the apical darkening and the

pattern is more reduced, the spot in base of 4th R being small, faint and occa-

sionally absent. All but a few specimens have come from the mountains just

east of the Canal Zone in Panama prov. (Cerro Azul, Cerro Jefe) the others

from El Valle de Anton, Cocle prov., Rio Pequeni, Colon prov. and several other

localities in Darien prov., all localities of high rainfall at elevations of 1000 ft.

or more. All material was taken in March, April or May, at the end of the dry

season, mostly in Shannon traps or light traps, so that the species is probably

ecrepuscular or nocturnal. Neither seasonal nor geographic range overlaps that of

pterographicus. The species is so far not known outside of Panama.

Philipotabanus (Philipotabanus) ebrius (Osten Sacken)

Tabanus ebrius Osten Sacken, 1886, Biol. Centr. Amer., Dipt., 1: 49, Pl,

fig. 8, female. Irazu, Costa Rica and Chiriqui, Panama.

Tabanus (Philipotabanus) ebrius: Fairchild, 1943 (1942) Ann. Ent. Soc.

Amer., 35(4): 455-456, Pl. 2, fig. 14, female, male; 1971, Cat. S.

Amer. Dipt.Fase. 28, p. 78, synonymy.

This is a very variable species, ranging in body color from light yellowish

brown to nearly black. As interpreted here, the scutellum is always blackish,

and the wing pattern usually has a pale streak through the dark pattern in first

posterior cell, sometimes faint. The frontal index ranges from 6 to 8.25, aver-

aging 7.2. The types of ebrius are pale, with wings as figured by me.

The species occurs in Panama only in the mountains of Chiriqui and Bocas

del Toro provinees, mostly at elevations from 1500 to at least 6000 ft. It ap-

pears to fly from April to September, and attacks man and horses. It is at least

partly crepuscular or even nocturnal. The species is known elsewhere only from

Costa Rica.

86 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Philipotabanus (Philipotabanus) elviae (Fairchild)

Tabanus (Philipotabanus) elviae Fairchild, 1943 (1942), Ann. Ent. Soc.

Amer., 35(4): 460, Pl. 2, fig. 12, female. Buena Vista, Chiriqui

Prov., Panama.

The large size, reduced wing pattern and accentuated dorsal antennal tooth

serve for recognition. Aside from the types from Chiriqui, the species has been

taken only a few times, for a total of 5 specimens, near Almirante June and

July, and Rio Changena, Sept., both Boeas del Toro prov. Nothing is known of

its habits, except that the specimens were attracted to man. There are 2 fe-

males labelled Costa Rica, de Lafon, 1884, in Museum d'Histoire Naturelle,

Paris, and a long series was taken on the Osa Peninsula, Puntarenas, Costa Rica

in Feb. and Mar. (Hogue and Fairchild 1974).

Philipotabanus (Philipotabanus) grassator Fairchild

Tabanus (Philipotabanus) grassator Fairchild, 1953, Ann. Ent. Soc. Amer.,

46(2): 278-279, P1. 2, fig. 9, female, Almirante, Panama.

Philipotabanus_ stigmaticalis: Fairehild, 1958, Ann. Ent.Soe. Amer.,

51(6): 520. Not Kroeber 1931.

This little species is seemingly rare. I have seen but 3 Panama specimens, 2

from our Yellow Fever Station near Almirante, Bocas del Toro prov., taken in

the treetops 30 July 1952 and 30 May 1960, and another also taken in treetops,

along the Calderas - Chiriquicito trail, Bocas del Toro, in Oct. - Nov. 1955. I

synonymized this species with T. (Philipotabanus) stigmaticalis Kroeber in 1958

on the basis of the description of the latter and a specimen from British Guiana.

Acquisition of additional material from Colombia, Peru and Brasil reveals that 2

distinct species exist. Philipotabanus grassator is represented by the Panama

specimens above and a single female from Colombia, Dept. Valle, Centro Hydro-

elect. Anchicaya, 4 Aug. 1977, M. A.Tidwell coll. This specimen was received

too late to be included in Wilkerson's publication (1979). Of P. stigmaticalis

Kroeber, I now have the British Guiana specimen mentioned in 1958, 5 females

from the vicinity of Manaus, Amazonas, Brasil, all taken in an arboreal flight

trap in July or Aug. 1981, and 1 female from Peru, Madre de Dios, Rio Tam-

bopata Reserve, 20-31-X-1982, R. Wilkerson coll. P. grassator is paler, the

secutellum and first 2 abdominal segments yellowish brown, the legs, except fore

femora yellowish, the wing stigma pale brown, the basal cells without basal in-

fuseation. P. stigmatiecalis is darker, the scutellum blackish except in 1 speci-

men, the abdomen nearly black, the anterior 2 or 4 tergites bluish pruinose, the

legs all entirely black, the wing stigma black, the basal cells 1/2 to 2/3 basally

weakly infuseated. The wings of the Peruvian example are slightly darker.

Philipotabanus (Philipotabanus) magnifieus (Kroeber)

Tabanus (Phaeotabanus) formosus Kroeber, 1930, Zool. Anz. 86: 299, fig.

21, female, Ecuador. Not T. formosus Wlk, 1848.

G. B. Fairchild: Tabanidae of Panama 87

Tabanus (Phaeotabanus) magnificus Kroeber, 1934, Rev. Ent. 4(3): 305.

Ecuador, Costa Rica, Colombia. Nom. nov. pro. T. (Ph) formosus

Kroeb. 1930 nee Walker 1848.

Tabanus (Philipotabanus) magnificus: Fairchild, 1943, Ann. Ent. Soc.

Amer., 35(4): 456-458, Pl. 2, fig. 13, (1942) female, Panama.

Fairchild, 1953, Ann. Ent. Soc. Amer., 46(2): 278, male.

Philipotabanus (Philipotabanus) magnificus: Fairchild, 1971, Cat.

Dipt.Amer. S. of U. S., 28: 79. Wilkerson, 1979, Cespedesia 8(31-32):

327-329, fig. 103.

This is the smallest species of the genus, and the most abundant in Panama.

It occurs in forested areas from sea level to 5000 ft. throughout the country,

and although abundant at ground level is also taken frequently in the forest

canopy. It bites man readily and although taken at all hours, is primarily ecre-

puscular, having been taken until 9 p.m. and in light traps. It ranges from

Guatemala (Peten) south to Ecuador. The wing pattern varies slightly in extent

and intensity, but the species seems less variable than others of the genus. It

flies primarily in the rainy season from April to Dec., but seems most abundant

in May and June. The examples from east of the Andes in Colombia do not

seem to differ from specimens from Dept. Valle on the Pacifie coast.

Philipotabanus (Philipotabanus) medius (Kroeber)

Tabanus (Phaeotabanus) intermedius Kroeber, 1930, Zool. Anz. 90: 81,

female. Suiza de Turrialba, Costa Rica. Not T. intermedius Wlk.

1848, Egger 1859.

Tabanus (Phaeotabanus) medius Kroeber, 1934, Rev. Ent., 4(3): 305, nom.

nov. pro T. intermedius Kroeb. 1930 nec Walker 1848.

Philipotabanus (Philipotabanus) medius: Hogue and Fairchild 1974, Rev.

Biol. Trop. 22(1): 22.

The type was in Budapest and presumably destroyed. I confused the species

with pallidetinctus, (Fairchild 1943) but it is distinet; my figures and records

under medius really refer to pallidetinetus. It is separable from pallidetinetus in

being much blacker, with black scutellum and legs. The wing is almost the

same, the basal cells slightly less infuscated, the hyaline spot at end of discal

cell generally larger, and apex and hind border definitely smoky. The frontal

index is from 5.5 to 7.75. Specimens have been taken only in the mountains of

Western Panama, and adjoining Costa Rica as follows: Sta. Fe, Veraguas, May

1950 (1); Posada Guttierrez, Bocas del Toro, 2900 ft., 4 Apr. 1963 (9); Bocas del

Toro Prov., 5000 ft., Aug. 1963 (1); Palo Santo, Chiriqui, 25 Aug. 1950 (1);Sta.

Clara, Chiriqui, Aug. 1951 (2) and Nov. 1958 (3); Boquete, Chiriqui Aug. 1944

(3). The species is variable, some specimens being quite brownish with less

sharp wing pattern, and separable with difficulty from dark extremes of ebrius.

Philipotabanus (Philipotabanus) nigrinubilus Fairchild

Tabanus (Philipotabanus) nigrinubilus Fairchild, 1952, Ann. Ent. Soc.

Amer., 46(2): 279-280. P1. 2, fig. 8, female.

88 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Philipotabanus nigrinubilus Fairchild, 1964, J. Med. Ent. 1(2): 176-177.

Male. Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1):

110, fig. 31. Wilkerson, 1979, Cespedesia 8(31-32): 329-331, fig. 102.

This species is the blackest of the Panama species, easily recognized by the

very extensive dark wing patch and small clear spot at end of discal cell. It has

been taken near Almirante, in Boeas del Toro province, at Sta. Fe in the moun-

tains of Veraguas prov., in Colon prov. on the upper Rio Mandinga, in Panama

prov. at Cerro Azul, and in Darien prov., on the Rio Tacareuna. All these lo-

ealities are in areas of heavy forest and high rainfall, mostly at elevations of

1000-3000 ft. Specimens were taken attacking man at ground level or in Shan-

non traps, and in the months from April to October. Additional specimens were

reared from larvae at Rio Changena, Bocas del Toro prov. and Ft. Sherman, C.

Z. taken under bark of fallen or felled dead trees. (Murdoch & Goodwin 1974).

It is known elsewhere only from a short series taken at Lancetilla, Tela, Hon-

duras, July-Oect., 1953, and 4 females taken at Rio Raposo, Buenaventura,

Colombia. Wilkerson (Op. cit.) reeords the species from the departments of

Choeo, Valle and Cauca in Colombia.

Philipotabanus (Philipotabanus) pallidetinctus (Kroeber)

Tabanus (Phaeotabanus) pallidetinctus Kroeber, 1930. Zool. Anz., 86: 297,

fig. 19, female, Chiriquicito, Panama.

Tabanus (Philipotabanus) medius: Fairchild, 1943 (1942), Ann. Ent.

Soc.Amer., 35(4): 458-459, Pl. 2, fig. 16, female. Not T. medius

Kroeber 1934.

Tabanus (Philipotabanus) caliginosus: Fairchild, 1953, Ann. Ent.

Soc.Amer., 46(2): 276-277, male. Not T. ealiginosus Bellardi 1859.

Study of type of pallidetinetus in B. M. by myself and types of ecaliginosus

Bell. by Philip in Turin indicates that the latter is the prior name for alteri-

pennis Wlk. 1860, a Mexican species, and not the same as Panama material. Ph.

medius Kroeber, misidentified by me in 1943, is treated elsewhere here. The

present species is separated from medius by the contrastingly pale scutellum and

usually more brownish yellow color as well as characters in the key. It is

widespread in the lowlands of Panama up to 2000 ft., from Darien to Bocas del

Toro and Chiriqui. It has been taken from April to September, with a few

Specimen at Almirante in January. It attacks man and has been taken in horse-

baited stable traps and in light traps, and is probably mainly crepuscular, as

specimens have been taken after dark. The range ineludes Costa Rica and I

have seen a specimen from Ecuador which is probably but a variant. Dark

Specimens resemble medius, but have shorter wings, the distance from apex to

fork of third vein being 1/4 wing length, while in medius this distance is more

than 1/4 wing length. pe

Philipotabanus (Philipotabanus) pterographicus (Fairchild)

Tabanus (Philipotabanus) pterographicus Fairchild, 1943 (1942), Ann. Ent.

Soe.Amer., 35(4): 459, Pl. 2, fig. 15, female, Panama. Wilkerson,

1979, Cespedesia 8(31-32): 331-332, fig. 100.

G. B. Fairehild: Tabanidae of Panama 89

This species resembles large examples of Ph. magnifieus Kroeb., differing in

pale legs and lack of apical wing infuscation, as well as more reduced wing pat-

tern, whieh laeks the spike-like darkening extending toward the hind border

along veins RS to Cul. It averages slightly smaller than Ph. chrysothrix, is

More grayish, and with more extensive wing pattern, the spot in base of cell 4th

R being well developed. Specimens have been taken in Bocas del Toro and

Panama provinees, and in several localities on the Pacific side of the Canal

Zone all at low elevations, mostly in Shannon traps and horse-baited stable

traps. It has been taken once in the forest canopy and once biting a horse at

night. All but 2 specimens were taken in the months from Oct. to Feb. Three

females from 3 localities in coastal Ecuador are the only non-panamanian speci-

mens I have studied, though Wilkerson (Op. cit.) reports it from Choco,

Colombia.

Subgenus Mimotabanus Fairchild

Mimotabanus Fairchild 1964, J. Med. Ent., 1(2): 179; 1975, Ann. Ent.

Soc. Amer., 68(4): 689-694, 2 plates, key.

The 3 species of this group known from Panama are keyed below. The

group ranges from Guatemala to eastern Peru.

‘) Mesonotum prominently striped. Wings hyaline with a small dark shade

below stigma. Abdomen bluish pruinose with a row of middorsal

White “tvign@lese .iiluisdcbetalistuchedsstecld, ens eae. Wedbape plenus (p. 90)

Mesonotum not or faintly striped. Wings with more extensive black

markings. Abdomen black with at most a large white patch on

fourth tergite + IED UR Leer AES eel RM al MN DENY OE Ae” SRR, UP ERENT CR, RR PT, 8 COTO eseeee0 2

2 Seutellum and adjoining part of mesonotum with bright golden hairs.

Wing with basal cells entirely hyaline, dark patch filling whole discal

area from ends of basal cells to slightly beyond fork of third vein

and from costa to hind margin. Fourth tergite with a broad trape-

zoid of yellowish white hairs covering segment except for extreme

CER ORCA BONG canakguravebiahetiatebin ehieietiio eh inauratus (p. 89)

Scutellum black and black haired. Wings with basal cells 2/3 infuscated,

the dark diseal patch not extending distally beyond end of diseal cell

and not reaching hind margin. Pale hairs on fourth tergite forming

a rounded triangle 1/3 width of segment .........00. phalaropygus (p. 90)

Philipotabanus (Mimotabanus) inauratus (Fairchild)

Tabanus (Philipotabanus) inauratus Fairchild, 1947 (1946), Ann. Ent. Soc.

Amer., 39(4): 573-574, P1. 1, fig. 3, female, Panama.

The species is known from the provinces of Bocas del Toro, Veraguas,

Panama and Darien, and the Canal Zone, all at low to moderate elevations in

areas of high rainfall. All specimens were taken in Feb. or Mar. save one in

July. Two of the 6 female specimens were definitely taken in the forest canopy

and 2 others probably were also, all attracted to human bait. The species is

known elsewhere only from 1 female from Alajuela, Costa Rica. A recently

90 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

collected male, (Barro Colorado ld., C. Z. 3 Mar. 1976, at U.V. light, Silberglied

and Aiello colls.) is easily associated with the female on color and pattern. The

eyes are holoptic, the upper enlarged eye facets well differentiated and demar-

eated from the small facets, occupying about half eye area in a triangular patch

surrounded by small facets. The enlarged facets are shortly but densely pilose,

and there is a small tubercle between the eyes at vertex.

Philipotabanus (Mimotabanus) plenus (Hine)

Tabanus plenus Hine, 1907, Ohio Nat., 8: 255, female. Izabal

Guatemala.

Stenotabanus (Stenotabanus) plenus: Fairchild, 1942, Ann. Ent. Soc.

Amer., 35(3): 306, fig. 15, female. Panama. |

Philipotabanus (Mimotabanus) plenus: Fairchild, 1975. Ann. Ent. Soe.

Amer., 68(4): 694, P1. II fig. 5, revision of subgenus.

The species bears a striking resemblance to a large Sarecophagid fly, with

reddish bronze eyes in life. The male resembles the female, but the thoracic

markings less contrasting. The upper eye facets are enlarged and sharply de-

mareated from the small facets, occupy 2/3 or more of eye area, and are

densely but short pilose. The small facets are bare and extend in a narrow strip

to vertex, which bears a small tubercle sunk between the eyes. In Panama the

species is scarce but widespread ranging from Cocle prov. (El Valle de Anton) to

Darien, and from sea level to 2000 ft. It has been taken in light traps and

Shannon traps, and appears to have a limited flight season, Jan. to Apr. in the

dry season. Elsewhere it is known from Guatemala and Costa Rica.

Philipotabanus (Mimotabanus) phalaropygus Fairchild

Philipotabanus (Mimotabanus) phalaropygus Fairchild 1964, J. Med. Ent.,

1(2): 177, fig. 11, female, Panama; 1975, Ann. Ent. Soc. Amer. 68(4):

190, Panama to E. Peru. Wilkerson, 1979, Cespedesia 8(31-32): 322-

333, fig. 107, male, Colombia.

The 2 types were taken in Feb. 1958 in the forest canopy attracted to

mosquito eolleetors, Rio Tuira, Darien prov., while 2 further specimens are from

Cerro Quia and Rio Mono, both in Darien. Three specimens from Quincemil,

Cuzeo, Peru, 2450 ft., Aug. -Sept. 1962, L. Pena coll. seem to be the same. All

Panama specimens were taken Feb. or March and at least 2 of them in the for-

est canopy attracted to human bait. Wilkerson (Op. cit.) deseribes the male,

which has pilose eyes, and notes that hisesingle female from Colombia differed

slightly from both Panamanian and eastern Peruvian specimens.

Subgenus Melasmatabanus Fairchild

Melasmatabanus Fairchild 1964, J. Med. Ent., 1(2): 179. Wilkerson 1979,

Cespedesia 8(31-32): 318-322.

Only two species of this group, keyed below, occur in Panama; both are

rare, and little is known of their habits. The group ranges southwards to Matto

G. B. Fairchild: Tabanidae of Panama 91

Grosso, Brasil. Both Panama species are largely black insects with a solid black

wing pattern, long proboscis, slender palpi, yellow antennae and wholly black

legs. Wilkerson (Op. cit.) discusses and figures both Panamanian species.

Key to Species

Wing wholly black except for apical third of basal cells and apex beyond

PEROT ARG VIIA: dssscecvnenabhicmansneedasamencdensimnassncndd fascipennis (p. 91)

Wing with a black patch below stigma which ineludes whole discal cell

but fails to reach hind margin or fork of third vein. Basal cells

about half dark basally and axillary cell lightly infuscated .......csecee

Ne a MTL ei pk Le Ae ee Lee criton (p. 91)

Philipotabanus (Melasmatabanus) criton (Kroeber)

Phaeotabanus columbianus Kroeber, 1931, Stett. Ent. Zeit.,92: 90-91.

Female, Colombia. Not Enderlein 1925.

Tabanus (Phaeotabanus) criton Kroeber, 1934, Rev. Ent., 4(3): 304, nom.

nov. Fairchild, 1975, Proce. Ent. Soe. Washington 77(2): 260. Type

seen. |

Tabanus (Philipotabanus) keenani Fairchild, 1947 (1946) Ann. Ent. Soe.

Amer., 39(4): 574, fig. 2, female, Panama. 1953; Ann.

Ent.Soe.Amer., 46(2): 278, male. Colombia.

In addition to the type of keenani from Colon prov., the species has been

taken in the Canal Zone, at Cerro Azul in Panama prov. and at Rio Tacareuna

in Darien prov., for a total of 8 specimens. These were taken in the months

from July to November, two of them in light traps, one each in a house and a

horse-baited stable trap, the rest attacking man at ground level. The type of

criton, now in Warsaw, was recently seen on loan (Fairchild 1975). The male is

like the female in color and pattern, with a large area of clearly demarcated

and differentiated upper eye facets which are bare. There is no tubercle at

vertex. Specimens from Choco and Santander, Colombia and Pichincha Prov.,

Ecuador have also been seen.

Added to the characters in the key, both basal cells are about half infus-

cated basally, the mesonotum & scutellum are dark brown, unmarked, the ab-

domen black with a poorly marked white-haired triangle on fourth tergite. The

legs, palpi and proboscis are dark brown to black, but the antennae bright

orange yellow.

Philipotabanus (Melasmatabanus) fascipennis (Macquart)

Tabanus fascipennis Macquart, 1845, Dipt. Exot. Suppl. 1: 35, Pl. 4, fig.

1,female. "Nouvelle Grenade" (Colombia).

Tabanus (Philipotabanus) fascipennis: Fairchild, (1942) 1943, Ann. Ent.

Soc. Amer., 35(4): 462, fig. 19, female. Panama.

' Philipotabanus fascipennis: Fairchild, 1958, Ann. Ent. Soe. Amer., 51(6):

529, full references. |

92 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

The two examples seen from Panama came from the mountains east of the

Canal Zone (La Zumbadora, Cerro Azul, Panama prov., 4 Feb. 1957 and 17 Dec.

1965) taken by mosquito collectors in the forest canopy. I have seen other ma-

terial from Venezuela and Muzo, Colombia. In fresh specimens the eyes are

bright green, unbanded, the thorax brownish with faint coppery hair stripes, a

small white-haired triangle on fourth tergite, and the black wing areas with

metallic blue reflections. Wilkerson (1979) discusses Colombian material, de-

scribes the male, which has pilose eyes, and figures the wing.

Genus Stypommisa Enderlein

Enderlein 1923, Deuts. Ent. Zeits., p. 545; 1925, Mitt. Mus. Zool. Berlin,

11(2): 369. Kroeber, 1934, Rev. Ent., 4(2): 276, references.

Stictotabanus Lutz 1914, nomen nudum. _ Fairchild, 1961, Mem. Inst.

Osw. Cruz, 59(2): 244.

Stenotabanus: Fairchild, 1942, Ann. Ent.Soe. Amer., 35(3): 297, in part.

This genus is separated from Stenotabanus by having a few setae on the ba-

siecosta, the eyes with a single dark stripe, bicolored, or unicolorous, and the

frontal callus nearly always narrower than frons and generally prolonged upwards

in a narrow ridge. The included species are generally small flies, seldom over

12 mm. in wing length, usually with spotted or tinted wings and narrow frons.

They are structurally quite similar to Leucotabanus, but have much fewer setae

on basicosta and a different style of coloration. They seem to be close to what

would be expected of a link between primitive Diachlorini and primitive Taban-

ini, and could be placed in either group. The sparse setae on basicosta and well

developed tubercle at vertex suggest Diachlorini, the other characters are Taba-

nine or shared by both groups. The group is not very homogeneous in appear-

ance, and may well prove to be polyphyletie on further study.

The Panama species are all inhabitants of heavy forest in areas of high rain-

fall, two are quite strictly arboreal and one of these is strongly mimetic. All

attack man readily.

Key to females

di Wings glass clear. Frons less than 3 times as high as basal width, the

eallus broadly drop-shaped. All tibiae bicolored, basally pale. Ab-

domen dark brown with a series of contiguous middorsal triangles

and posterior margins Of all tergites grey ....csccccsccrseees serena (p. 97)

Wings tinted or cross veins elouded or spotted. Frons over 4 times as

PASTS ARR WCE VIG hy le a Rese daily VOL Red RRR Ese C ep a a ueGe deccceececeteesesene 2

ai Pork of, third vein Without AMpOnGiwns ines ccsvaiiioikide oeeiiva¥ ds ARE he adden veees 3

Fork of third vein always with an appendix which is rarely shorter than.

ENE. \AGACSHTVEIMOSORITIGHE SERS GRea sei lee C Raa eae oN FATIG Risee een 9)

3. Legs entirely black, black-haired. Mesonotum prominently bluish white

striped. Abdomen with at least first 2 tergites bluish pruinose, and

tergites 2 to 6 with small middorsal triangles and posterior lateral

GIG Wie NAPS inns tsacusiviedbeuscanecsbenbnarmnernescee maruccii (p. 96)

G. B. Fairehild: Tabanidae of Panama 93

Legs brown, at least in part, the tibiae at least partly pale-haired.

Mesonotum at most weakly Striped .......ccccccccccccccccces endebd sh Dihbeud toed cone 4

4, Brownish species, the mesonotum weakly striped, the abdomen without

prominent bluish pruinosity, but with pale hind margins and large

pale-haired median triangles on all tergites ...... vans jaculator (p. 95)

Black species, the mesonotum unstriped, the abdomen black, weakly

bluish pruinose anteriorly, but without median triangles or pale pos-

tOPIGP DONOS sssssssiassitievariccormines PIE Sere ey hypographa (p. 95)

5.(3') Wings with large prominent dark clouds on all crossveins and narrow pale

pilose bands on hind margins of all abdominal tergites ........cccccesces 6

Wings at most with small faint clouds on fork and distal end of discal

eell ae ee ei a a es SSCOSSHSHSHSHSSHHFHHSOHSSHHHHSHHHHHSHHHHHHHHEHEHHHHHHHHHHHHHHSEHHEHHHHEHE 7

6. Brown species, the thorax and abdomen nearly concolorous, the scutel-

lum with at least apex reddish. Second abdominal tergite with a

large but diffuse blackish integumental median spot. Frons slightly

broader (index 4.75). Abdominal pale pilose hind marginal bands

prominent, underlain by slightly paler broad integumental bands ......

ban baeS Ua EU TEIGUS Js aks sees sT5) sg tone OOURUd Doe w REE RIAA sgh eb oee lerida (p. 96)

Mesonotum and scutellum black in ground color, the paler brown ab-

domen contrasting. Second abdominal tergite without dark median

spot. Frons narrow (index 5.97). Abdominal pale pilose bands very

narrow, often not evident in worn specimens, not underlain by paler

inte THe cilia oma i changena (p. 94)

fi Wings with small to moderate clouds on crossveins and fork; marginal

and submarginal cells strongly brown tinted. Abdomen largely black-

haired, at most with small median pale-haired triangles. Smaller

species, generally less than 11 mm. in wing length ..........cessee. bachee

sh ik vee AE WEE NS OUR aad Ung Lbs Os lb aes ase pequeniensis (p. 97)

Wings without, or with very small and faint clouds on fork and

crossveins. Whole wing yellowish, sometimes brownish in marginal

and submarginal cells. Abdomen with broad yellow haired bands on

all tergites. Larger species, generally over 12 mm. in wing length

stint thls. Gah AS Ae GhawaG, BA et eaptiroptera (p. 93)

Stypommisa captiroptera (Kroeber)

Tabanus (Macrocormus) captiropterus Kroeber, 1930, Zool. Anz., 78(1-2):

10, fig. 8, female, Venezuela.

Stenotabanus xenium Fairchild, 1947, Ann. Ent. Soc. Amer., (1946) 39(4):

968, Pl. 1, fig. 5, female, Panama.

Stypommisa captiroptera: Fairchild,1971, Cat. S. Amer. Dipt. Fase. 28,

p. 81, synonymy. |

This species is generally larger than pequeniensis, as well as differing as de-

tailed under that species. It is neither so abundant nor widespread in Panama as

pequeniensis, occurring in Bocas del Toro Prov., (Nigua Creek, near Almirante

and our Yellow Fever Station), at Rio Pequeni on Madden Lake, Rio Mandinga,

Colon Prov., and Rio Canita, in the Rio Bayano valley. All localities are at low

94 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

elevations in areas of high rainfall. Several specimens were taken at night, and

most material was from Shannon traps. There appears to be no special flight

- season, aS specimens have been taken in Jan., May, June, July and Aug. The

species ranges from Nicaragua south to the mouth of the Amazon and eastern

Peru. |

Stypommisa changena Fairchild n. sp.

A moderate sized dull brown species with wings both spotted and the veins

brown-margined, and the abdominal tergites with obscure pale hind margins or

small median pilose triangles.

Female. This species is best described by comparison with St. lerida. It is

a more slender species and the overall coloration is darker. The secutellum is

dark blackish brown, (with at least the apex usually red in lerida). The frons is

slightly narrower in changena, the maximum, minimum, median and mean indices

of eight specimens being 6.25, 5.30, 5.78 and 5.97, while for seven specimens of

lerida the corresponding figures are 5.3, 4.5, 4.85, and 4.75. The indices of di-

vergence (frontal width at vertex divided by frontal width at base) are not sig-

nificantly different, ranging from 1.1 to 1.5 in both species. The wings of

changena have the appendix usually longer than the subtending vein segment due

to shortness of the latter, while in lerida the vein segment is often longer than

the appendix. The wings in changena have a noticeable anteroapical brown

tinting and all the veins are narrowly brown margined. In lerida the wing is

lightly and evenly tinted without noticeable darkening apically and the veins not

obviously brown margined. Lerida has a prominent but diffuse median blackish

integumental spot on abdominal tergite 2, wholly lacking in changena, where the

abdomen is uniformly dark brown. Changena has but faint traces of pale hair

fringes on the tergites, while these are prominent in lerida. The two species

have different times of flight, February, March and April for lerida, May to

November for changena. The eyes of both species are green with a single pur-

plish transverse stripe.

Male. Color and wing pattern as in the female. Eyes with enlarged upper

facets bare, well differentiated from small facets, but without a sharp line of

demarcation, the larger facets occupying about 2/3 to 1/2 total eye area. Small

facets extend in a narrow band to vertex behind and there is a small tubercle

sunk to eye level at vertex with vestige of at least the anterior ocellus. The

occiput bears a sparse fringe of long forwardly directed dark hairs.

Holotype female, Panama, Rio Changena, Bocas del Toro prov., 2400 ft.

elev., 26 Sept. 1961.

Allotype male, Panama, Chiriqui..prov., La Fortuna, 10-VI-1977, U.V. light

trap, H. Wolda coll.

Paratypes, 4 males, 67 females, as follows: Panama: 2 females, same data as

holotype; 3 males, 9 females, same locality and collector as _ allotype,

IV,V,VI,VII,VIlI-1977, U.V. light trap; 42 females, Yellow Fever Camp, same lo-

eality as holotype, human bait and Shannon trap, VIII-[X-1961; 3 females, Rio

Claro-Rio Changena, Bocas del Toro prov., 910 m elev., 20-21-V-1966, C. Myers

eoll.; 3 females Rio Uri, Changuinola, Bocas del Toro prov., VIII-IX-1968, R.

Hartmann coll; 1 female, Boquete, Chiriqui prov., no date or collector; 1 fe-

male, Robalo trail, Bocas del Toro prov., 6500 ft. elev., 6-VIII-47; 1 female

Calderas-Chiriquicito trail, Chiriqui prov., 600-1400 ft. elev., in treetops, 26-X

to 5-XI-55. Costa Rica: 1 female Cartago prov., Hacienda de Moravia de

G. B. Fairchild: Tabanidae of Panama 95

Chirripo, 7-10-VII-1964, McDiarmid coll.; 1 male, 2 females, Puntarenas prov.,

Monteverde area, 6-14-VI-73 and 21-V-76; 1 female, Alajuela prov., Chomogo

area, 1620 m. elev., 13-VI-73; 1 female, Carillo, no date, C. F. Underwood coll.

In addition, I have seen 1 female in C.A.S. from Heredia, Costa Rica, 19-20-VII-

(5, Fisher coll., and 27 females in Univ. California, Berkeley from Monteverde

area, Puntarenas prov., Costa Rica. :

Holotype, allotype and some paratypes to be deposited in F.S.C.A., other

paratypes in L.A.C.M., M.C.Z., U.C. Berkeley, and collections of L.L. Pechuman,

and the author.

Stypommisa hypographa (Kroeber)

Stypommia hypographa Kroeber, 1930, Zool. Anz., 86: 252-253, female,

Venezuela, Bolivia, Peru.

Stypommisa hypographa: Fairchild, 1967, Stud. Ent. 9(1-4): 359, fig.

32(1966). Philip, 1969, Acta Zool. Lilloana, Tucuman, 22: 124(1967).

Two specimens of what seems to be a form of this species were taken by R.

L. Dressler at Cerro Jefe, Panama Prov., 28 April and 8 May 1968. They differ

from my notes on the type in having the wings less strongly spotted, lacking the

clouds at tip of anal cell and tips of median veins. The tibiae have a few sil-

very hairs at base on all legs, most abundant on fore pair which are obscurely

bicolored. One specimen has the third antennal plate quite dusky, hardly con-

trasting with style, in the other it is dark orange, though not so brightly con-

trasting with black style as in ssp. neofurva Philip. Two specimens of the latter

from Bolivia, Mapiri, S. Carlos, 800 m. 9 Jan., 03 have wings intermediate be-

tween the Panama specimens and my figure of the type of hypographa. The

legs are browner in these, with but traces of silvery hairs on the fore tibiae

only in one. Kroeber describes the antennal plate as black, though the

Venezuelan type lacked antennae, so that his statement must refer to Bolivian

and/or Peruvian specimens, as may the remainder of his description, at least in

part. I believe that color of antennal plate is variable, and that the Bolivian

and Peruvian specimens discussed by Kroeber and Philip and my Panamanian

specimens are but variants of a single species. Whether they are specifically or

subspecifically distinct from Venezuelan material cannot be decided on present

evidence.

These Panama specimens closely resemble St. maruccii Fcehld. but differ in

lacking mesonotal grey stripes, in having a wholly black abdomen without mid-

dorsal pale triangles and with only a trace of bluish pruinosity on first tergite,

the wings more heavily fumose in marginal and submarginal cells, and the frons

slightly narrower with more slender tapered callus. Though both maruccii and

hypographa have been taken in the Cerro Jefe-Cerro Azul area, there seems to

be a seasonal separation at least in that locality, with maruceii occurring in

Sept. - Dec., hypograrha in April - May.

Stypommisa jaculator (Fairchild)

Stenotabanus jaculator Fairchild, 1942, Ann. Ent. Soe. Amer., 35(3): 309,

fig. 9, female, Panama; 1951, Op. cit., 44(3): 452-453, male.

Stypommisa jaculator: Barretto, 1957, Rev. Brasil, Malar., 8(1): 82

(1956), Br. Guiana.

96 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

This is a rather inconspicuous little fly, dusty cocoa-colored, the weakly

, Spotted wings with the fore borders tinted brownish. In well preserved exam-

ples, there is a series of broad sparsely white-haired triangles arising from the

sparsely white-haired hind borders of the abdominal tergites. The eyes are

bronzy green in life, with a narrow and often inconspicuous dark median trans-

verse line. The male is similar, the upper eye facets markedly enlarged but not

demarcated from the small facets by an abrupt change in facet size.

The species is abundant at about 2000 ft. on Cerro Azul, Panama Prov.,

where it was taken at human bait in the forest canopy and in a light trap.

Elsewhere it has been taken in the canopy and at ground level at several locali-

ties in the Canal Zone in stable traps and light traps, in Darien Prov. from near

sea level to 4000 ft., at Cerro Campana, Panama Prov., 3000 ft., and at El

Valle, Cocle Prov. The flight season is short. March to May, with one June

record. I have seen the specimen recorded by Barretto from British Guiana, and

more recently the species was taken at Teresita and Curiche river in the Dept.

Choco, Colombia by personnel of the Office of Interoceanie Canal Studies. The

species is separable with difficulty from St. pruniecolor (Lutz), from Brazil, which

has the antennae wholly orange yellow, while jaculator seems always to have at

least the terminal annulus of style black, generally the style all black.

Stypommisa lerida (Fairchild)

Stenotabanus lerida Fairchild, 1942, Ann. Ent. Soe. Amer., 35(3): 307-

308, Pl. 1, fig. 14. Female, Panama; 1947, Ann. Ent. Soc. Amer.,

(1946), 39(4): 568.

Lerida differs from changena as detailed in the key and in the description of

the latter, but seems to be a much rarer species. All specimens seen (23 fe-

males) have been taken on the Pacifie slope of Volean Bauru in Chiriqui

province above Boquete, at Cerro Punta and at Cerro Pando. Time of flight

seems short, as all were collected in Feb., Mareh and April in various years.

Conversely, most specimens of changena were taken in the rainy season from

May to Oct., and except for an undated specimen from Boquete, and a long se-

ri¢s from Fortuna, on the northern side of the Volean in Boeas del Toro

province. The specimens reported as lerida by mOSeP and Fairchild (1974) from

Costa Rica are changena on further study.

Stypommisa maruccii (Fairchild)

Stenotabanus maruccii Fairchild,“1947, Ann. Ent. Soe. Amer., 39(4): 568,

fig. 8 (1946) female, San Blas, Panama; 1958, Ann. Ent. Soe. Amer.,

51(6): 529, synonymy.

Stenotabanus frondivelia Fairchild, 1951, Ann. Ent. Soc. Amer., 44(3):

453-454, fig. 5, female, Chiriqui, Panama; 1953, Proc. Ent. Soe.

Wash., 55(5): 241.

This little species bears a close resemblance to a Sarecophagid fly, the eyes

in life being bright brick red, with a narrow transverse green stripe, fading to

green with a dark stripe shortly after death. The thorax bears prominent pale

stripes and the abdomen is extensively bluish-grey pruinose, with a median row

G. B. Fairchild: Tabanidae of Panama 97

of small pale-haired triangles. The wings bear small clouds on crossveins and

fork, and are lightly tinted along the apical half of anterior margin.

In Panama, the species is found in areas of high rainfall and heavy forest up

to elevations of at least 3000 ft., from Darien to Chiriqui and Bocas del Toro

Provinees. It is highly arboreal, being seldom taken at ground level, and appears

to be on the wing throughout the year. Practically all our material has been

taken with human bait. The range includes Nicaragua (Villa Somoza, 7 July

1953, in forest canopy) and Colombia (Rio Raposo, nr. Buenaventura, cited in

Lee, Fairchild and Barreto, 1969).

Stypommisa pequeniensis (Fairchild)

Stenotabanus pequeniensis Fairchild, 1942, Ann. Ent. Soc.Amer.,35(3):

308-309, fig. 13, female, Panama.

Stenotabanus pequiensis: Philip, (lapsus) 1960, Proc. Calif. Acad. Sci.,

Ser. 4, 31(3): 77, Peru.

Pequeniensis is quite variable in intensity of coloring and in proportions of

frons, the latter varying from 6 to 7 times as high as basal width. Certain pale

specimens, especially if the abdomen is denuded, are difficult to separate with

certainty from captiroptera. Pequeniensis generally has distinct though small

spots on crossveins and fork, and the apical wing infuseation is heavier and

ceases rather abruptly at the level of the tip of the stigma. The antennal style,

and sometimes the apex of the plate, are nearly always blackish. In cap-

tiroptera fork and crossveins rarely have perceptible clouds, the apical wing in-

fuseation is lighter, browner, and is not abruptly limited basally. The antennal

style is usually all yellow, or with only the apical one or two annuli blackish.

Pequeniensis usually has the pleura white or gray pilose, yellow in eaptiroptera.

Panama material has all come from heavily forested areas of high rainfall,

either on the Atlantic coast or from the mountains at elevations up to 5000 ft.

Records are from Darien, Panama, Colon and Boeas del Toro provinces. Most

specimens were taken in May, lesser numbers from June to September. The

species attacks man readily at ground level and is taken in Shannon traps. The

eyes are bright green in life, unbanded.

The male is colored like the female, the head wider than thorax with the

upper enlarged eye facets occupying about two-thirds of the eye area, strongly

differentiated and sharply demarcated from the small facets. In life the upper

facets are bronze, the lower green with purple reflections. There is a small tu-

berecle at vertex between eyes.

The species ranges from Costa Rica south to Colombia, Ecuador and eastern

Peru.

Stypommisa serena (Kroeber)

Tabanus serenus Kroeber, 1931, Stett. Ent. Zeit., 92: 304, female,

Venezuela. Type in Vienna seen.

Stenotabanus vapidus Fairchild, 1964, J. Med.Ent., 1(2): 172-174, fig. 10,

female, Panama.

This species should be recognizable from the characters in the key. it is

placed in this genus with some hesitation, as the frons is unusually broad and

98 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

the wings and color pattern of abdomen rather different from other species.

- The eallus is large and rounded, the upper extension thread-like, while the an-

tennal plate is broad with a strong dorsal angle. Aside from the type from

Venezuela, the species has been taken at Rio Tuira in Darien Prov., and from

Muzo, Colombia, and Tingo Maria, Peru. All localities are in the areas of high

rainfall, and all except the Rio Tuira specimen from altitudes of over 1500 feet.

Genus Leuecotabanus Lutz

Lutz 1913, Brasil Medico, 27, No. 45, p. 487. Fairchild, 1941, Ann. Ent.

Soc. Amer., 34(3): 629-631, references, key; 1953, Ann. Ent.

Soc.Amer., 46(2): 274-276, key; in press, Myia vol. 3.

This genus contains 17 deseribed species, of which 5 have been taken in

Panama. All the Panama species are basically black with the thorax or at least

the scutellum nearly always with contrasting white or yellow hairs, and the

wings clear or slightly tinted. The frons is generally rather narrow with a club-

Shaped or ridge-like callus and a definite tubercle at vertex. The basal plate of

third antennal segment is moderately broad, with a fairly strong dorsal angle.

Three of the species tend to have the abdomen more or less acutely pointed,

and all have bare unicolorous greenish-black eyes in life. The known males have

the upper eye facets greatly enlarged, clearly demarcated from the small facets,

the head wider than thorax, and a tubercle at vertex.

Key to females

4. Legs entirely black. Abdomen with at most a narrow band of white

hairs on hind margin of fourth tergite and sides of second. Costal

cell strongly brown tinted. Subeallus centrally bare and shiny ........ 2

Legs with all tibiae at least one third basally white. Abdomen with at

least second and fourth tergites with white hair. Costal cell not

darker than rest of wing. Subcallus pollimose .........ccccccccsccscsccscsees 3

2. -Mesonotum and scutellum entirely clothed with bright yellow pollen and

hairs. Abdomen above entirely black .......sccsceceee flavinotum (p. 100)

Mesonotum predominantly black haired, seutellum with black or dark

yellow hairs. Extreme sides of second and entire hind border of

fourth tergites with White HIPS 'ls....s.0ccbsisdeccbveceseccess aurarius (p. 99)

3. Antennae yellowish brown, the style much longer than basal plate.

Whole insect, with exception of notopleural lobes, blackish in ground

color, grey pollinose. All abdominal tergites usually with some

white hairs, more extensive on second and fourth tergites .........scse0.

dca siidue pu nenVibnvs toe cadeseh heeiiUudsur ashe asshieuinsd teeadsrechebass nigriventris (p. 99)

Antennae black or very dark brown. At least scutellum pale in ground

color, pollinosity of thorax and abdomen brown to black .......csccsess 4

4. Second tergite with a median and lateral white hair patches, usually

separated, sometimes narrowly joined. Fourth tergite more than two

thirds white, the band widest in middle and sides. Remaining ter-

gites entirely dark. Antennae black, strongly excavated dorsally, the

G. B. Fairchild: Tabanidae of Panama 99

basal angle prominent, the basal plate shorter or equal to style.

Frons about 4.5 times as high as basal width, the callus much nar-

rower, drop-shaped, brown or yellow. Palpi yellow or brown, black

or partly white-haired. Abdomen not pointed at tip ...ccccccccccccccccsees

Lpadevinnanessvoassuwshidounblaed pfiiieed iaktsseecia lide. ebamnerak is. 10)

Second tergite narrowly white-haired behind, widened in middle and

sides. Fourth tergite at most half white. Third, fifth and sixth ter-

gites often with narrow white hind borders or median white trian-

gles. Antennae less strongly excavated, the basal plate longer than

style Frons about 5.5 times as high as basal width, the callus pro-

portionally wider, black. Palpi black and black haired. Abdomen

usually sharply Pointed.” ..sesocsdscdsccsseccsesecscscessiooss .. exaestuans (p. 100)

Leucotabanus aurarius Fairchild

Leucotabanus aurarius Fairchild 1953, Ann. Ent. Soc. Amer., 46(2): 274,

Pl. 2, fig. 12, female, Panama.

Aside from the Holotype, only 3 other specimens have been taken, at Almi-

rante, Bocas Del Toro prov., one 11 July 1960 and two 7 Aug. 1963. The 1960

specimen differs from the others in entirely lacking yellow hairs on scutellum

and sides of mesonotum, the whole insect being black except for the white-

haired sides of tergite 2, hind margin of 4, and white haired sternites 2 and 4.

The type was taken in a Shannon trap, and the others probably also, although

not so labelled.

Leucotabanus nigriventris Kroeber

Leucotabanus nigriventris Kroeber 1931, Stett. Ent. Zeit., 92: 92, fe-

male, Chiapas, Mexico. Fairchild, 1941, Ann. Ent. Soe. amer., 34(3):

637, fig. 8, references and synonymy; 1975, Proc. Ent. Soe. Washing-

ton, 77(2): 260.

A single badly denuded specimen from Nigua Creek, Almirante, Bocas del

Toro Prov., July, 1962, in light trap, agrees with a specimen from Tabasco,

Mexico, determined as this species and with the types on loan from Warsaw.

Nigriventris is very similar in structure to L. ambiguus Stone from Arizona, but

differs in more evenly ridge-like callus, blunter palpi, pale antennae, and black

unicolorous ground color of mesonotum. My figure of this species (1941) was

drawn from a Guatemalan specimen determined by Hine as albiseutellatus Macq.,

a misdetermination, as noted by Stone (1938) when he described the Arizona

specimens as ambiguus. The palpi of the Tabasco and Panama specimens are

more slender, but still blunt tipped. Though almost entirely denuded of its

pilosity, the Panama specimen is deep black in ground color, entirely steel grey

pollinose.

100 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Leucotabanus canithorax Fairchild

Leucotabanus canithorax Fairchild 1941, Ann. Ent. Soe. Amer., 34(3):

634-636, fig. 5, female, Panama; 1953, op. cit., 46(2): 275, in key

only; 1956, Smiths. Miscell. Colls., 131(3): 10.

The species is rare in Panama, having been taken only in Darien prov., Rio

Tuira, 24 Feb. 1958, and at Moja Pollo on the Rio Chagres above Gamboa, Jan.

- May, 1940-1941. Specimens were taken biting domestic animals or in a Shannon

trap. I have seen females from British Honduras (Belize) and British Guiana

(Guyana), and a male from Trinidad. The latter is badly erushed, but resembles

the female, except that the white bands an abdomen are broader, that on second

tergite covering the whole segment. The figures for eanithorax and leuconotum

in my paper (1941) are mislabelled. Fig. 4 is leuconotum, fig. 5 eanithorax, the

reverse of the numbers on the plate.

Although originally proposed as a replacement name for T. albicans Macq.

1845, not Macq. 1835 and 1838, the name was accompanied by a description of

Specimens, and, is thus not invalidated by my subsequent (1956) discovery that

albicans Macq. 1845=exaestuans Linn. 1767.

Leucotabanus exaestuans (Linnaeus)

Tabanus exaestuans Linnaeus, 1767, Syst. Nat. Ed. 12, 1, pt. 2, p. 1000,

Amer. Merid. Philip, 1952, Ann. Ent. soc. Amer., 45: 312, type seen.

Leucotabanus exaestuans: Philip, 1960, Proc. Calif. Acad. Sei., Ser. 4,

31(3): 91, Peru. Goodwin and Murdoch, 1974, Ann. Ent. Soc.Amer.,

67(1). 108, figs. 14, 28.

Leucotabanus leucaspis (Wied.): Fairchild, 1941, Ann. Ent. Soc.Amer.,

34(3): 632-634, fig. 7, female, Panama, full references and syn-

onymy; 1942, Op. cit., 35(1): 88, seasonal distribution; 1953, Op. cit.,

45(2): 275, in key.

The species occurs throughout Panama at lower elevations, and seems to fly

throughout the year. It rarely attacks man but bites cattle and horses readily,

and is thus frequently taken in animal-baited stable traps. These have not been

used much outside the Canal Zone area, so that the species is probably more

abundant than our records indicate. Males have been taken in light traps, and

the species reared from larvae taken with those of L. flavinotum (Goodwin and

Murdoch 1974). The range appears to cover the whole Neotropics, from Mexico

to Argentina, but not in the West Indies.

The color varies considerably. Some specimens have the mesonotum almost

wholly white-haired, others with the disk dark and white hairs only on the mar-

gins and scutellum. The abdomen always has white bands on second and fourth

tergites, usually on fifth and sixth, and occasionally some white on third. The

pale hairs of thorax are generally white, less often with a more or less pro-

nounced yellowish tinge.

Leucotabanus flavinotum (Kroeber)

Tabanus flavinotum Kroeber, 1934, Rev. Ent. 4(3): 309, new name for

Tabanus nigriflavus Kroeber, 1931, nee. Kroeber 1930.

G. B. Fairchild: Tabanidae of Panama 101

Leucotabanus flavinotum: Fairchild, 1941, Ann. Ent. soc. Amer., 34(3):

632, fig. 1, female, Panama, references; 1956, Smiths. Miscell.

Colls. 131(3): 16, type seen; Goodwin and Murdoch, 1974, Ann. Ent.

Soe.Amer., 67(1): 108 figs. 15, 29.

This vivid little species hardly ever exceeds 10 mm. in length. The combina-

tion of bright lemon yellow thorax and black abdomen is unmistakable. In much

denuded specimens, the thorax is pale yellowish grey. The species seldom at-

tacks man, but enters stable traps and Shannon traps and attacks horses and

eattle readily. It has been taken in small numbers in Darien, Panama, Colon

and Bocas del Toro Provinees, and on both coasts of the Canal Zone, rarely at

elevations over 1000 ft. A considerable series of both sexes was reared from

larvae taken from mixed earth and rotten wood in the base of a hollow tree

containing bats on the rio Cocoli, Canal Zone. The larvae were of much the

Same size and all pupated and yielded adults within a few days of 1 May, 1956.

Most of the captured specimens were also taken in May, with seattered speci-

mens in March, April, June and July. The male differs only in sexual characters

and more slender antennae. The species is known from Costa Rica to Ama-

zonas, Brazil. Goodwin and Murdoch (1974) described the larva and pupa, finding

them in situations similar to those reported above, nearly always associated with

rotten wood. It is noteworthy that Burger (1977) found larvae of L. ambiguus

Stone in rot holes in trees in Arizona, while larvae of the other Nearctic

species, L. annulatus (Say), are always found associated with rotten wood or in

rot holes in living trees.

Genus Lepiselaga Macquart

Macquart 1838, Dipt. Exot., 1(1): 153. Kroeber, 1929, Encyel. Ent. Ser.

B, 2, Dipt. 5: 136. Fairchild, 1966, Psyche, 72(3): 210-217, fig. 5

(1965), key. Cosearon, 1968, Rev. Soc.Ent. Argent., 30(1-4): 51-59,

figs.

But one species of this small genus occurs in Panama, the widespread L.

erassipes; the other species occur in Colombia, southern Brasil and Argentina.

Lepiselaga crassipes (Fabricius)

Haematopota crassipes Fabricius, 1805, Syst. Antl., p. 108, no sex, South

America.

Lepiselaga crassipes: Fairchild, 1940 Psyche 47(1): 8-13, figs. 1-5, early

stages; 1942, Ann. Ent. Soc. Amer.,35(3): 291, fig. 2, full references.

Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1): 106, figs.

22, 29. Wilkerson, 1979, Cespedesia 8(31-32): 351-353, figs 113, 114.

In addition to the characters in the key, the species has a wholly black and

shiny face and shiny subeallus, flattened shiny palpi, all tibiae black and in-

flated, all tarsi white. The black parts of the wing have several small round

hyaline fenestra, and the base of M3,forming the posterior border of the discal

cell, is bent strongly forward,so that the discal cell is constricted in the middle.

In Panama the species is common to abundant in the vicinity of

swamps.lakes and large rivers. the larvae live in floating aquatic vegetation,

such as water lettuce and water hyacinth. the adults attack man persistently

102 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

especially about the legs, and may be severe pests. They are known locally as

"Congo". In the period shortly after the filling of Gatun Lake, large areas of the

lake became covered with water lettuce, and this insect became exceedingly

abundant. As conditions stabilized, the water lettuce disappeared except for

small patches, and the Congo fly is now only occasionally troublesome. Goodwin

and Murdoch (1974) have re-described the larva and pupa.

The range includes most of the Neotropics, from Southern Mexico and the

Greater Antilles,south to Paraguay and northern Argentina. Specimens have

been taken from aircraft landing in Miami, Florida from various Neotropical

countries. |

SUBFAMILY TABANINAE

Tribe Tabanini

Flies belonging to this tribe, in addition to the characters given in the key,

are generally without pictured wings or inflated legs or antennal segments.

They always lack sclerotized labella or vestiges of ocelli, and only a few species

have long dorsal spines dorsally on the antennae. All but one of those ocurring

in Panama belong in the Genus Tabanus, an enormous world-wide group of possi-

bly polyphyletie origin.

Genus Poeciloderas Lutz

Lutz 1921, Bol. Inst. Osw. Cruz, 1(1): 15. Fairchild, 1969, Arq. Zool. S.

Paulo, 17(4): 220; 1971, Cat. Dipt. S. Amer., 28: 85, synonymy.

This genus is characterized by having a small tubercle at vertex, eyes gen-

erally pilose in female, densely so in male, inflated antennal scapes, and wings

with first posterior cell closed or strongly coarctate. Most of the species are

south temperate or Andean, but the single Panamanian species ranges from

Mexico to Argentina. It has been included in the key to Tabanus, as it may

easily be mistaken for a species of that genus.

Poeciloderas quadripunctatus (Fabricius)

Tabanus quadripunctatus Fabricius, 1805, Syst. Antliat., p. 99.

Tabanus (Hybomitra) quadripunctatus: Fairchild, 1942, 35(4): 452-453,

fig. 8 (full references).

Poeciloderas quadripunctatus: Fairchild, 1971 Cat. S. Am. Dipt., Fase.

82,p. 86, synonymy. Goodwin and Murdoch, 1974, Ann. Ent.

Soc.Amer., 67(1): 116 fig. 36. Wilkerson, 1979, Cespedesia 8(31-32):

356-358, fig. 115. Cosearon and Fairchild, 1976, Physis, Sec. C. 35

(91): 294, fig. 1

This species is easily recognized by the combination of spotted wings, white

median figure on the second abdominal tergite and pilose eyes. It is uncommon

but widespread, apparently throughout Panama. The eyes of the female in life

are purplish black with 2 green transverse bands, while in the male the upper

G. B. Fairehild: Tabanidae of Panama 103

green band is covered by the area of slightly enlarged facets except at its outer

end. Females have been taken in horse-baited mosquito stable traps and Shan-

non traps, males hovering and at light. The variety amabilinis Philip, with ex-

tensively white second tergite seems to be less common than the type, but has

been taken in the same localities. Males have been taken hovering in the early

morning a few feet above ground level. It appears to fly throughout the year.

Goodwin and Murdoch (1974) described a male pupa from Chiriqui province.

Genus Tabanus Linnaeus

Linnaeus 1758, Syst. Nat. Ed. 10. 1: 601. Fairchild, 1971 Cat. S. Am.

Dipt. Fase. 28 p. 87, synonymy.

This genus includes 38 Panamanian taxa, and is hence the largest genus oc-

curring in the country. Efforts to split the group into subgenera have not met

with wide acceptance (Fairchild 1969) and subgenera are not used here. The key

is frankly artificial and based on easily seen characters. Difficulty may arise in

determining members of a group of small similar appearing species found in

mangrove swamps whose abdomens are faintly striped in well preserved speci-

mens, but apparently unicolorous when worn or denuded, as they often are. The

species are nondescriptus (couplet 29) and nereus and rhizophorae (couplet 40).

Specimens running to any of these species should be checked with the text and

with Fairchild (1973). Tabanus pungens apparently contradicts the definition of

the genus given in 1969 (Fairchild 1. ¢.) having a well-marked tubercle at ver-

tex. The eyes in both sexes are bare, however, and there seems no point in

erecting a separate category for it at this time. Males of about 2/3 of the

species are known, but since most of them will key out easily with the females,

they are not keyed separately. Notes on those that are known are given under

each species. The species are treated alphabetically for ease of reference.

Key to Females

ii Eyes of female generally with some short pilosity, those of male densely

pilose. Vertex with a round or oval low tubercle. Wings with first

posterior cell strongly coarctate or closed at or before wing margin,

and with all crossveins with strong dark clouds. First antennal seg-

ment strongly produced dorsally, forming a hood over the second

SCELMENt ..ecccccees suse bubainis tenis Poeciloderas quadripunctatus (p. 102)

Without the above combination of characters (If a tubercle at vertex is

present but wings unspotted see couplet 28) .......ccccccecee WUE sanicnadediende 2

2. A prominent black velvety pilose spot covering base of scutellum and

adjoining border of mesonotum, — bordered with paler hairs and

pollinosity ........ eT TT TC ee eT ee eon hisissdsn de 0

Without such a prominent spot though comatuene with a small velvety

Spot on posterior border Of MESONOTUM ....ccccccccsccccccccccscccccccccccccccoes 9

Xn With a long appendix at fork of third vein......... SUD ahs s SLRs Atenseeeite ses 4

Without an appendix at fork of third vein........ dedi x sie kiedia <abds eines cn venddons v4

104

7(3).

9(2).

10.

Li.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

First posterior cell always closed and petiolate. All crossveins and fork

with small, often faint, clouds. Eyes with 2 bands and lower margin

green, the bands as wide as the dark interval between them ...ccccccee

. LL OR LON. A La oculus (p. 118)

First posterior cell rarely closed, never petiolate and crossveins and fork

WH ENO T TOUS Peace aR cde eines Seance Peli dapipecdoiedevsewosducdbebscetuceiewescdcst 9)

Eyes with 2 narrow green bands, much narrower than the dark interval

between them, the lower margin sometimes narrowly green. Mid

abdominal pale triangles equilateral or higher than wide, the poste-

rior borders of tergites without pale hairs ......... pseudoculus (p. 122)

Eyes with 3 broad green bands, the upper 2 sometimes confluent at

outer ends, the lower margin of eye always dark. Mid abdominal

triangles broader than high, the posterior borders of tergites paler

SPP ROHN OE. Gee vi see Lean NTO UL i Se cacsebe rece evens soeeTUbsceueccsccesscee 6

Black species with femora largely black ........cssescesees albocirculus (p. 109)

Reddish species with femora largely pale ......... albocirculus var. (p. 109)

Abdomen unicolorous, dark brown, wholly black-haired. Frontal callus a

slender ridge, hardly widened below. Large species with dark brown

wings and almost wholly black legs .....ccsccscsscsceccsees defilippii (p. 112)

Abdomen with at least vestiges of pale-haired mid-dorsal triangles.

Frontal callus definitely widened DelOW ......ccccccccccccccccscesccccccccscccccs 8

Large stout species. Antennal plate orange, not unusually slender. All

femora orange. Abdomen dark reddish brown, blaeck-haired, with

small and often faint orange-haired median triangles. Eyes in life

Wiss Rees) Se GL eR PRR AL as polyphemus (p. 119)

Smaller slender species. Antennal plate black, unusually long and slen-

der. At least fore femora blackish, generally darker than the oth-

ers. Abdomen yellowish brown, brown haired, with faint slightly

paler-haired middorsal triangles which are sometimes contiguous,

forming a stripe. Eyes in life unicolorous, DrONZy ....ccccccccsscccccecscess

sti iS wed eee ei HORA occvccccccccccsceceeeeeeUipunctatus (p. 128)

With a more or less prominent black velvety spot on posterior margin of

MESONOTUM © ceocccccccccee PARaa theo ead Ci Sebwhaeeberevek une Obed Nees neeTiWdecssececceeeese 10

WED ted SU cE aa cs aed atks h ENE oe ea hdc bc cc cecsnceseecs 14

Abdomen without noticeable pattern, essentially UNICO]OLOUS ........ecseeeee 11

Abdomen with a pattern due to contrasting vestiture or underlying color

BE RGA ed aa rei rated eae ran cob cans ied cues daba ENSUE ab aNa Nabors ds dheceeoesnenenes 12

Abdomen pale green or straw colored, wholly yellow-haired. Mesonotum

gray, largely pale-haired. Wings glass-clear, or the veins slightly

brown-margined. Basal plate of third antennal segment with a mod-

erate acute dorsal angle .........00. devsoculsedvessucece olivaceiventris (p. 118)

Abdomen nearly black, bluish pruinose, black haired. Mesonotum dark

reddish brown, sparsely orange haired. Wings strongly brown tinted,

the veins broadly brown margined. Basal plate with a long dorsal

BG) daaveruvadsasddtevnuaonas ues bins eeoen) vesnberenavnge punctipleura (p. 123)

12.

13.

14(9).

15.

16.

17.

18.

19.

G. B. Fairchild: Tabanidae of Panama 105

Fork of third vein with a long appendix. Abdomen unicolorous brown

with a middorsal row of white-haired triangles. Slender species with

velvety black spot: Qiten TAIN . .scsccoccescrsscsentnccdecssoens rubripes (p. 125)

No appendix at fork. Abdomen otherwise. Velvety black spot prominent..

pcarer sss Maabeeterebedessdteldudesebactse whS bu SPAR R GH eso avi bnndscdnddeubapeeter chases enavenuass 13

Palpi unusually short and inflated. Abdomen yellowish brown in ground

eolor, often with an indistinct dark median stripe. Vestiture con-

sisting of median black-haired triangles and dark lateral margins, be-

tween which is a pair of broad dorsolateral orange-haired stripes or

series of more or less connected spots. Wings generally with a small

dark area below stigma and clouds around basal crossveins and fork

Of AMINE VEIN" .csccshauaetsetndse seed vals Fue EPH ete AES importunus (p. 113)

Palpi inflated basally, but with drawn-out slender apex. Abdomen as

above, though generally more reddish, but with a middorsal row of

pale-haired triangles superimposed on the dark-haired median stripe.

Wings with clouds around crossveins and fork, but without the sub-

stiPimatic dark Death, ..:cstisaccsussionoces Adsbiatabehaetvunt tes nebulosus (p. 115)

Large, wholly black species with deeply fumose to black WINGS .......0.0. mm

Otherwise colored or small species........ heed epnvip gens nda s exhaueye ts Phos sabhased oo 16

Frontal callus slender, ridge-like. Basal plate of third antennal segment

with a low but acute dorsal angle. Wings dark DrOWN ......ccccccccccccces

ish spskn ss nasil co AAACN CR es ek bia aoaab cs capdnkan ee cganatee morbosus ebeneus (p. 115)

Frontal callus oval below, extending in a ridge above. Basal plate with

a long slender curved dorsal spine. Wings deep black ......ccccccccccccces

EE GSS eet URN Wann ee EESU FO A, LOL AMMNO ROR NORM a a (p. 113)

Large black species with white striped mesonotum, white-haired scutel-

lum, abdomen with large median white patch on tergite 4 and

smaller white spots on tergites 1,2,5, and 6. Wings with all

ecrossveins and fork with large dark clouds, and base and apex in-

PBL EG. cussneetuns idedhani oon vtr we eeehcane ia RE RIPE eta xenorhynchus (p. 129)

Not as above, the wings not heavily spotted ............ Seed aeh capneiecsaheus cake “ae

Large brown species with reddish or orange brown abdomens without

conspicuous pattern and very narrow frons with slender ridge-like

COTE ines sstph beancoacien ss bilbhh bs ous aN MeaA LENO cabs Gleteny sa0h nuk SANS OkamaNee) exeaaee veeeNn 18

Smaller species, or the abdomen patterned or callus not ridge-like....... 20

Antennal plate with long dorsal spine reaching nearly to end of plate.

Abdomen dark reddish, black-haired, with posterior margins of ter-

gites and small middorsal triangles dark orange-haired. Beard and

cheeks grey. Wing lightly infuseated, generally brownish along veins,

fork of third vein with a short appendix or none ..macquarti (p. 114)

Antennal plate with at most a strong dorsal tooth......... Skaihnegesbeneessaness 19

Face, cheeks and beard brown. Frontal callus brown. Antennae orange,

unusually long and slender with the dorsal tooth close to base of

plate. Wings evenly brown tinted, with long appendix on fork of

third vein. Abdomen red, coppery-haired on tergites 1-4 and middle

of 5, remainder dark and black-haired ......seceeee scoooeeee SUPIfer (p. 126)

106

20(17).

21.

22.

23.

24(20)

25.

26.

27.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Face, cheeks and beard pale gray. Frontal callus black. Antennae dark

brown to black, broader. Wings brown tinted, with a short appendix

or none. Abdomen orange red on tergites 1 to 4, orange-haired, ter-

gites 5-6 abruptly black and black-haired ........cscssseeees bigoti (p. 110)

subeallus Dare aid sSniny, at 1EGSt I Darl .cieciicescosecesccensesevescevvebeseveeces 21

Subeallus pollinose unless denuded Dy WEA ........ccccccccccccccccccccccccccoccccecs 24

Abdomen black or dark brown with a single prominent white middorsal

stripe covering tergites 1-6. Frons over 6 times as high as basal

width. Wings somewhat smoky along fore border and ADeX .cccsccccoeee

Dr ols nav cloccccsiverEacses wes JetIere Ue seri te Uae cask tdi eeascens unistriatus (p. 128)

ode With dorsolateral StriPeS OF SPOTS ..... ccccccsccsccseccccevsentsstecsceesens 22

Subeallus with a pair of crescent-shaped bare areas above antennal

bases. Basal plate of third antennal segment broad, the plate sub-

equal in length to style. Abdomen markedly pointed. Middorsal ab-

dominal stripe a series of slender contiguous triangles, lateral stripes

BO ANC COMUINUOUS <coiceabe cccceicrcnncctavecsespectcecasauencses enanus (p. 113)

Subeallus wholly bare except for a narrow strip of pollinosity separating

it from frontal callus. Abdomen not pointed .........cccccccccccccccccccccs 23

Notopleural lobes lighter in color than adjacent mesonotum. Abdomen

black or dark brown, the middorsal stripe a prominent row of white

eontiguous triangles, more prominent than the white dorsolateral

stripes which rarely extend beyond tergite 4. Third antennal seg-

ment orange to brown, the style always black ...... platyeerus (p. 119)

Notopleural lobes concolorous with mesonotum. Abdomen dull yellowish

brown, the middorsal stripe a slender dull yellowish line, not more

prominent than the dorsolateral stripes. Antennae usually wholly or-

ange, the style seldom contrastingly black ...ccccccscccee aniptus (p. 110)

Abdomen dull yellowish with a single broad yellowish middorsal stripe

covering tergites 1 to 6. Wings with veins strongly brown margined

Frontal callus drop-shaped. Eyes in life green, UNDanded .....cccccccceee

aoa aca erm ar endo punto anas ene rixator (p. 125)

PC, Gs oro neces ce cr recestrancavsalscsevadcccsuvesneucccastuscccsecdecsas 29

Abdomen black with all tergites narrowly white on posterior margins,

the margins widened into low triangles. Small black stout species

WECIY FIV ELLIVS (WTS Oo ee esiu reece ces ceveseecerns quinquepunctatus (p. 124)

PUGOINEN OLN OL WISE cicecccccesccnccceccctencscctcclradeccacececccecdsauschetessseccosccecaceesss 26

Slender brown species with long brown-tinted wings, unusually long black

third antennal segment, and brown abdomen with a row of faint

pale-haired mid-dorsal triangles .........cccccccccscsseees lacajaensis (p. 114)

Without the above Combination Of Characters...cccccccccccccccccccsccccsssccccccees 27

Abdomen with well-marked median and dorsolateral longitudinal stripes,

sometimes composed of triangles or oblique patches, but at least the

MECIAN STrIPe CONTINUOUS seredicicaccccccecccecececdcccnsccseeceesnsncenassccnpesscess 28

28.

29.

30.

31.

32.

33.

G. B. Fairchild: Tabanidae of Panama 107

Abdomen otherwise marked. (If abdomen with very faint stripes, see

couplet 39) ....ccces arene povikin vc Pewtteh vobdasasthucbimuntesvaaeanl VUES» SAREE IEE Sr 36

Vertex with a small, discrete raised tubercle. rons less than 4 times

as high as basal width, the callus as wide as high, yellow. Abdomen

with a narrow even or irregular pale median stripe overlying a broad

black integumental stripe. Dorsolateral stripes of broad, pale

oblique contiguous patches. All femora pale, wings glass-clear, the

fourth posterior Gell broadly OPen .....cccccccssscscscsscesecs pungens (p. 123)

Vertex without tubercle, at most a Small Dare ALea ......ccccccccccscecccccscecs 29

Eyes unicolorous, bronzy to blackish green in life. Abdomen yellowish

brown with 3 rather inconspicuous yellow-haired stripes, the mid-

stripe of narrow contiguous triangles, the dorsolaterals of slightly

oblique patches. Wings with all veins brown-margined, sometimes

faint. Legs pale brown, the fore tibiae very obscurely bicolored .....

dis dddesinbici ns diana eeamentiel ania sovcscscseseeees MONGESCriptus (p. 116)

Eyes banded in life. If wing veins brown-margined, then fore tibiae

Strongly DiCOlOred ...cesevees veoneaiiueandwes iia Wieecnipaeeens' Lan onpenaneeeae sca)

Abdominal mid-stripe of a series of narrow connected triangles ........... 31

Abdominal mid-stripe of even width throughout, rarely slightly irregular .

Sisbicas sibsckesens Wi dsasen nisagiteee oweanaacrte'e doa leahecgnewtin Gu Meme ehads Ais si caeiere see

All femora entirely black or dark brown, the tibiae paler brown, the fore

pair very obscurely bicolored. Wings glass clear. Abdomen nearly

black, the dorsolateral stripes even ....... cosspeatessaeesessete GUN (Ds.422)

Mid and hind femora largely pale, only somewhat infusecated basally, the

fore pair blackish and fore tibiae sharply bicolored. Wings with

small faint clouds on crossveins and fork of third vein. Abdomen

reddish brown with a small black integumental triangle on middle of

tergites 1 and 2. Dorsolateral stripes markedly step-like, of oblique

AC CNS. wiccicasnedsen save hey asks stiatacansabeereidenelcs enteass commixtus (p. 111)

Frons broader, less than 4 times as high as basal width, the basal callus

round or square, yellow to red. All femora pale; fore tibiae clearly

bicolored. Wings glass clear. Abdominal stripes broad and even,

yellowish. Scutellum red, contrasting with blackish mesonotum ......

sia aces asbeidindisiasbeet Meine brcuiuek tia dasibutasabivedensbes vittiger guatemalanus (p. 127)

Without the above combination of Characters........cccccccceccees sisnsececameaneus 33

Frons about 4 times as high as basal width, parallel sided, the callus

usually dull yellowish. Mid and hind femora pale, fore femora black,

fore tibiae conspicuously bicolored. Wings glass clear. Abdominal

Stripes broad, chalky white, the dorsolaterals slightly uneven.

Scutellum black, concolorous with mesonotum ....colombensis (p. 110)

Frons narrower, 4.5 to 6 times as high as basal width, markedly conver-

gent below. Wings always slightly to moderately brownish tinted,

especially costal cell and tips of first 2 veins ...........000 aditswevcesss 34

108

34.

35.

36(27)

37.

38.

39.

40.

Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Mid and hind femora entirely pale, rarely slightly dusky at extreme base.

Secutellum reddish, at least at apex. Abdomen yellow to brown,

without conspicuous black patch at base either dorsally or ventrally.

Abdominal stripes generally broad and even, yellow to nearly white

re trai rk occidentalis var. dorsovittatus (p. 117)

Mid and hind femora at least 1/4 black at base. Scutellum black........ 30

Abdomen yellow to brown, the stripes yellow pilose as are sternites. A

conspicuous black integumental median patch on first 2 sternites and

usually a black triangle on first 2 tergites ...... stenocephalus (p. 125)

Abdomen black to dark brown, the median stripe white, narrow, the dor-

solateral pale stripes usually faint, reduced or absent. Sternites

with dark integument but white pilose and PollinoSe ........sccccccccccccees

basisedesibaluddssaddccaccadeoeteeratte cavvenseses occidentalis var. modestus (p. 117)

Abdomen with conspicuous dorsolateral rows of pale haired spots, and a

smaller median row of pale triangles ...........ccccccccccsccccsccccceces sasnate 37

FAGGOTIC CUMOL WISE ii lisedii cp Pil bia ls cables thacddaneneWass¥ os cacegdasaidessPaedabeddisscecsens 38

Large stout species, black or brown, mesonotum striped, abdomen with

median row of small slender pale triangles and dorsolateral rows of

small round diserete pale spots. Wings glass clear. Eye purple with

2 NAFPOW SP ee: DANES ssiascdnesashasstcasdccesacasiardidsescdias’ pruinosus (p. 122)

Slender grey species with dorsolateral oblique pale-haired patches on ab-

domen, wings grayish. Eyes greenish bronze with single narrow dark

BEVIS - aedasi ser thVednndvahstddncvetecneis cca pate sdaeeeases Hiitis shes praepilatus (p. 120)

Hind femora basally at least 1/3 black, remaining femora and coxae all

or largely black. Abdomen orange brown, with a black integumental

inverted triangle on first 2 tergites, sometimes also median black

streaks on one or more of succeeding tergites. Vestiture black ex-

eept for a dorsolateral row of orange haired round or oval patches,

sometimes contiguous and forming broad stripes. Median stripe,

when evident, faint, slender, yellow-haired. Frons about 5 times as

THEN BS DESAY WIL: sudhircitvesssnbssvasscadeevsceosteataesvecs praeteritus (p. 120)

PUY PEMOPE BUTIP SEY DA Sistas ckcecdtietacs ces dee lidccbvetecsds betes scaccessdseaesepeecoescente 39

Abdomen with a black inverted integumental triangle on first 2 tergites,

as in praeteritus, but pale median pilose stripe more evident and

dorsolateral pale pilose patches more slender and paler .........cecscccees

js apesASESVsGaLAdU bo agua hSadbELAd RAAELAERTNESIRAOAAAAKS praeteritus adiastolus (p. 121)

Abdomen without middorsal integumental black triangles and _ frons

PIE ORISD GERORsAUTVSESVE GA SEGENERSELLSRONLEUOLDASAK CREEL DSR SEREART hapakUO Os s06bgenaceasenssedes 40

Abdomen dull yellowish brown, contrasting with steel gray thorax, with 3

very faint and inconspicuous pale-haired stripes. Wings hyaline.

Frons broad, about 2.5 times as high as basal width, the callus black

veh Van ahdeOGEAd UUUESbOA Cd erL EAE PESRIA RE SINGasEba TaKueER Kes bauauateioes rhizophorae (p. 124)

Abdomen dull yellowish brown, slightly paler than the dull brownish tho-

rax, with 3 equally faint paler stripes. Wings with all veins narrowly

brown-margined. Frons 3.5 to 4 times as high as basal width, the

ALIS VOLOW cassinsanpinnnassns seashabecd benbsd sh lebesbhvaseasineebaions nereus (p. 116)

G. B. Fairchild: Tabanidae of Panama 109

Tabanus albocireulus Hine

Tabanus albocirculus Hine 1907, Ohio Nat., 8(4): 227, female, Tucurrique,

Costa Rica. Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28 p. 88,

Synonymy. Goodwin and Murdoch, 1974 Ann. Ent. Soe. Amer., 67(1):

128, Figs. 16,37. Wilkerson, 1979, Cespedesia 8(31-32): 363-365.

Tabanus (Bellardia) albocireculus: Fairchild, 1942, Psyche, 49(1-2): 11-12,

figs. 4, a, b, Panama. Fairchild, 1946, Ann. Ent. Soe. Amer., 39(4):

279-576, Almirante and Robalo, Bocas del Toro, Panama.

Tabanus (Lophotabanus) albocireulus: Fairchild, 1953, Proc. Ent. Soc.

Washington, 55(5): 241. |

This species is exceedingly variable in size and color but can be separated

from related species in Panama by the eye pattern, which consists of 3 broad

green transverse bands on a purple ground color. The typical form, which is

nearly black in body color with almost wholly black femora, occurs chiefly in

areas of heavy rainfall, such as Bocas del Toro province where it is the only

form taken, Darien (Cerro Pirre) and the Atlantic coast generally, with a few

records from the Pacific coast in the Canal Zone. The pale form is smaller,

with light reddish ground color and femora almost wholly reddish yellow, and is

dominant along the Pacific coast from Darien to Chiriqui provinees, though

specimens have also been taken on the Atlantie coast in the Canal Zone area.

Intermediate forms, with darker ground color and femora basally blackened

are dominant in San Blas and Darien provinces, but oceur on both coasts in the

Canal Zone, though not, apparently in Bocas province.

The few males collected have all been light or intermediate in color, and

are easily associated with the female. The eyes have the area of large facets

small and triangular, less than half eye area, and the facets not greatly en-

larged. There is a tubercle at vertex between the eyes, reaching eye level.

The palpi are oval, inflated, porrect, and the body more hairy than the female,

with the abdominal mid-dorsal triangles smaller and less distinet. All males

seen have been taken in mosquito light traps.

The species is one of the commonest taken in the lowlands and in open ar-

eas, though it occurs also in heavy forest, at ground level. It attacks horses,

but not man and is taken abundantly in horse-baited mosquito stable traps,

Malaise traps and Shannon traps. Both sexes have been taken in light traps,

though females are taken during the day as well. The flight season appears to

extend throughout the year, though there are more records for the rainy season

from May to December. Males were taken in April (1) and September (4). The

range is from Nicaragua to Colombia west of the Andes. Goodwin and Murdoch

(1974) reared one of 3 larvae collected. It yielded a male of the light form,

from Santiago, Panama.

T. albocirculus is very similar to T. antareticus Linn. (=xipe Kroeber) but

the latter has the male eye with strongly differentiated large facets, the wings

usually with faint clouds around cross veins and fork of third vein, more intense

than the general wing infuscation, and the frons of the females is slightly

broader, on the average. Antarcticus has so far been seen only from S. America

east of the Andes, and the two species may prove to be only subspecifically dis-

tinct. Eye patterns of both are the same.

110 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Tabanus aniptus Fairchild.

Tabanus aniptus Fairchild, 1976, Stud. Ent. 19(1-4): 241-244, fig. 1 fe-

male. Colombia, Panama. Wilkerson, 1979, Cespedesia 8(31-32):

365-367, male. Colombia.

Tabanus trivittatus, Lee Fairchild and Barreto, 1969, Caldasia 10(49):

455. Not Fabricius 1805.

Only a single specimen of this recently described species has been seen from

Panama, taken at Jaque, Darien Prov., 7 April. 1967 by the noted herpetologist

Charles W. Myers. It differs from its nearest Panama relative, T. platycerus

Fehld. in having dark concolorous notopleural lobes and the stripes of the ab-

domen yellowish and rather inconspicuous. The species is apparently abundant in

Colombia along the Pacific coast, extending to Ecuador.

Tabanus bigoti Bellardi

Tabanus apicalis Macquart, 1847, Dipt. Exot. Suppl. 2: 20. Mexico. Not

Wiedemann 1828.

Tabanus bigoti Bellardi, 1859, Sagg. Ditt. Messico, 1: 58. New name for

T. apicalis Macq. Fairchild, 1942, Ann. Ent. Soe. Amer., 35(4): 443,

Pl. 1, fig. 4. in part, form B; 1971, Cat. S. Amer. Dipt. Fase. 28 p.

89, synonymy. |

This species ean be separated from the similar and more abundant T. surifer

Fehld. by black antennae with shorter dorsal tooth, black and more slender

frontal callus, grey pollinosity of face, cheeks and thorax, and lack of a long

appendix at fork of third vein. The eye is bright bluish green in life.

In Panama the species is uncommon. Most material was taken at our Yel-

low Fever station S. W. of Almirante, Bocas del Toro, in an area of heavy rain

forest. Scattered specimens were taken at Mandinga, San Blas, Cerro Azul east

of Panama City and near Gamboa, C. Z., in Shannon traps, in a tent, or flying

about the collector. The season of flight is short, as of 32 specimens available

with data, 28 were taken in May, 1 in April, 2 in June and 1 in July. Specimens

from Wauchope, Limon, Costa Rica were taken on 3 June and a series of 6 from

Lagos Montebello, Chiapas, Mexico, R. L. Dressler coll. were taken in May. The

species ranges from Mexico to eastern Colombia.

Tabanus colombensis Macquart

Tabanus colombensis Macquart 1846, Dipt. exot., Suppl. 1, p. 37, Pl, f,

fig. 2. Fairchild, 1956, Smiths, Miscell. Col., 131(3): 13; 1971, Cat.

S. Amer., Dipt. Fase. 28, p. 90. Synonymy. Goodwin and Murdoch,

1974, Ann. Ent. Soc. Amer., 67(1): 120, fig. 38. Fairchild, 1983, Ent.

Soe. Amer. Mise. Publ. No. 57, p. 19. |

Tabanus truquii Bellardi, 1859, Sagg. Ditt. Messicana 1: 64, male, Mex-

ico. (New synonymy).

Tabanus amplifrons Kroeber, 1933, Rev. Ent., 3(3): 354. Fairchild, 1942,

Ann. Ent. Soe. Amer., 35(2): 178-179, Pl. 2, fig. 17. Philip, 1942,

Psyche. 49(1-2): 23.

_ G. B. Fairchild: Tabanidae of Panama | 111

This species can be separated from others of the group with 3 striped ab-

domens by the combination of broad, even, chalky white stripes, wholly pale

femora in the female, glass clear wings and parallel sided frons. The male has

the upper facets but slightly differentiated and densely hairy, while the femora

are largely black. The eyes of the female are purple with two broad green

bands, the area above the upper band greenish purple. In Panama it is not a

common species. A year's collecting at Mojo Pollo on the Chagres river yielded

39 specimens, and 1 male 3 females were taken at Sta. Fe, Darien in April and

May 1967. I recorded a few others from the Canal Zone in 1942, and have since

seen a male and a female taken in a light trap at Sta. Clara, Cocle prov. and a

female taken near Gatun, C. Z. in a horse baited mosquito trap. It apparently

does not attack man, and may be more abundant in drier areas of Panama where

little collecting with bait animals has been done. Our records include the

months from February to September, so it probably flies throughout the year.

The range is extensive, from Texas to northern Brasil. Goodwin and Murdoch

(1974) collected over 65 immature specimens from which 50 adults emerged,

mostly from moist soil near water on the Pacific side of the isthmus. The lar-

vae seem indistinguishable from other species of the lineola group, but the pupa

is figured.

Tabanus commixtus Walker

Tabanus commixtus Walker 1860, Trans. Ent. Soe. London, 5: 273. Mex-

ico. Fairchild, 1956, Smithsonian Miscell. Colls. 131(3); 1983, Ent.

Soc. Amer. Mise. Pub. No. 57, p. 19, figs.

Tabanus maya Bequaert, 1932 (1931), J. N. Y. Ent. Soc., 39: 546, fig. 2,

female, Yucatan, Mexico. Fairchild, 1942, Ann. Soe. Amer.,35(2):

176, Pl. 2, fig. 19, male, female, Panama.

Tabanus truquii: Philip, 1965, Ann. Ent. Soe. Amer., 58(6): 876.

Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28, p. 103. Goodwin and

Murdoch, 1974, Ann. Ent. Soc. Amer., 67(1): 126, fig. 42, pupa. Not

truguli Bell.

This species is most readily separated from the other local species with 3

abdominal stripes by having the mid-stripe a series of narrow connected trian-

gles and the lateral stripes of somewhat oblique dashes. The wings have a faint

though unusually distinct cloud on fork of third vein, the frons is parallel sided,

and there is usually a dark integumental patch beneath scutellum extending onto

the second abdominal tergite. The male has the abdominal pattern faint and the

large eye facets bare and slightly enlarged, in a triangular patch surrounded by

small facets. Both sexes have the mid and hind femora basally black. )

The species is widespread at low elevations, but not abundant. All records

are from horse-baited stable traps, Malaise traps, taken biting animals or in

light traps. It appears to fly throughout the year, but with a definite peak of

abundance from April to June. Most records are from the Canal Zone area, due

to concentration of horse-baited traps, where it is almost equally abundant on

both coasts. Other records are from the Pacifie coast with a single record from

Sasardi, San Blas on the Caribbean coast. It was not taken during several years

intensive collecting in Bocas del Toro prov., and appears to favor open country.

The range is from Mexico to Trinidad.

112 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Goodwin and Murdoch (1974) reared 3 specimens from the Pacific side of the

Isthmus. These were the only specimens seen and only the pupa was described.

They were taken in habitats similar to those where colombensis larvae occurred.

Tabanus defilippii Bellardi

Tabanus defilippii Bellardi 1859, Sagg. Ditt. Messicana, 1: 57-58. Female,

Mexico.

Tabanus (Bellardia) defilippii: Fairchild, 1942, Psyche, 49(1-2): 10-11,

figs. 2, a, b. Panama, full references; 1971, Cat. S. Amer. Dipt.

Fase. 28 b. 91, synonymy.

This large blackish brown species with prominent pale-cireled black spot on_

seutellum is likely to be confused only with T. polyphemus Fehld. The latter is

paler in color, with red-brown abdomen usually bearing small yellow-haired tri-

angles, orange legs and antennae and golden-haired palpi.

Defilippii is known in Panama mainly from heavy forest in Bocas del Toro

provinee, where it is highly arboreal, over 90% of specimens taken at our Yel-

low Fever station near Almirante were taken in the forest canopy attracted to

humans. The few other specimens were taken in a Shannon trap or biting a

horse. We have also 1 female from Sta. Fe, Veraguas also taken in the forest

canopy, and there is another female in M. C. Z. from Buena Vista, Chiriqui,

both areas of high rainfall and heavy forest. The eyes in life are unicolorous

bronzy green.

Our records show the species flying from March to September in Panama,

with most records in April, May and June. The range is from western Panama

to southern Mexico.

Tabanus dunni Fairchild

Tabanus dunni Fairchild 1942, Ann. Ent. Soc. Amer., 35(2): 166-167, P1.

2, fig. 18; 1983, Ent. Soc. Amer. Mise. Publ. No. 57, p. 21, figs.

This rare species resembles T. commixtus Walk. in the markings of the ab-

domen, but is black in ground color and with a broader frons. The femora are

dark grey to black, the wings glass clear. The eyes in life bear 2 rather narrow

green stripes on a purple ground. The mesonotum in fresh specimens is also

quite markedly striped, unlike other members of the group in Panama. The

species was described from 5 females from Miraflores and Ancon in the Canal

Zone, taken in Jan. 1930 by L. H. Dunn. Since then only 3 females have been

secured, 1, Old Panama, bred from larva 11 Aug. 1942, the larva collected in

tidal mangrove swamp but pupal shell lost; 1, vic. Nata, Cocle prov., 20 May

1946, and 1, Sta. Fe, Darien prov., O. I. C. S. 1591, 26 May 1967. The species

seems confined to the Pacific coast of Panama and may have specialized habits.

In 1942 (Ann. Ent. Soe. Amer. 35(4): 472) I recorded a specimen from Nicaragua,

but this later proved to be T. pruinosus Big., as I noted in 1951 (Ann. Ent. Soc.

Amer., 44(3): 460)] The two are somewhat similar, but pruinosus lacks the ab-

dominal stripes. The Nicaraguan specimen was badly denuded.

G. B. Fairchild: Tabanidae of Panama 113

Tabanus enanus Fairchild

Tabanus enanus Fairchild 1942, Ann. Ent. Soe.Amer., 35(2): 157-158, P1.

1, fig. 8; 1976, Stud. Ent. 19(1-4): 239, 348, fig. 5. Panama.

The holotype of this species was an unusually small specimen, as later col-

lections have shown the size to be from 10-12 mm. The subeallus generally

shows a narrow transverse bare area just above the antennal bases, the abdomen

has a narrow median stripe of narrow connected triangles and broader lateral

Stripes of straight disconnected pale dashes, while the hind margins of the ter-

gites are narrowly pale margined. The markedly narrowed and pointed abdomen

is very characteristic. The eyes (revived) have 3 green bands, the upper 2 con-

nected or nearly so at their outer ends, similar to unistriatus and trivittatus.

The male is unknown.

We have taken a few specimens in horse baited mosquito traps on the Pa-

cifie side of the Canal Zone in Feb. and March 1953, and a single specimen at

Anton, Cocle Prov. in Jan. 1964. The Interoceanic Canal Survey took a long

series of over 100 specimens in a Malaise trap at Sta. Fe, Darien Prov. from

Jan. to April 1967, so that the species appears to fly only at the height of the

dry season. All the localities are close to the coast on the Pacifie side of

Panama. The species is not known elsewhere.

Tabanus erebus Osten Sacken

Tabanus erebus Osten Sacken 1886, Biol. Centr. Amer., Dipt., 1, p. 50.

Female, Nicaragua, Panama. Fairchild, 1942, (1943). Ann. Ent. Soc.

Amer., 35(4): 443, fig. 1. full references. Goodwin and Murdoch,

1974, Ann. Ent. Soc. Amer., 67(1): 128, Fig. 20.

One of the largest Tabanidae in Panama, this species is easily recognized by

its entirely black color, including the wings. Only the halters are ivory white.

the abdomen is bluish pruinose, as in the N. American T. atratus, which it

greatly resembles. The rare T. morbosus ebeneus is the same color, though with

paler wings, but has a narrower frons and slender antennae without a strong dor-

sal spine. The eyes are black in life, without pattern and the male is still

unknown.

In Panama the species is widespread but limited to forested areas of heavy

rainfall, ascending in the mountains to 5,000 ft. It bites man occasionally, but

prefers larger animals. Hunters say that its abundance indicates the presence of

tapir in the vicinity. Most of the material was taken in Shannon traps. The

flight season appears to be rather short, as with the exception of 1 specimen

taken in December, all of nearly 200 specimens were taken from April to

September, over half in May. The range is from Honduras to Panama.

Goodwin and Murdoch (1974) figure a very large larva taken at Ft. Kobbe,

Canal Zone, which they failed to rear, but felt was probably T. erebus, as this is

the only Panamanian species of sufficient size likely to be found in this area.

Tabanus importunus Wiedemann

Tabanus importunus Wiedemann 1828, Auss. Zweifl. Inskt., 1, p. 127, fe-

male, Brasil. Fairchild, 1943, Ann. Ent. Soe. Amer., 35(4): 444-445,

114 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

fig. 2, references; 1971, Cat. Dipt. S Amer. Fase. 28, p. 95, syn-

onymy. 1984, Contr. Amer. Ent. Inst., 21(3): 25, Map 1. Wilkerson,

1979, Cespedesia 8(31-32): 376-377 Cosearon, 1979, Obr. Cent. Mus.

La Plata, 6: 264, fig. 4.

This large brown species is likely to be confused only with T. nebulosus,

from which it is easily separated by the unusually short stout palpi and absence

of pale-haired middorsal abdominal triangles. The eyes in life are unbanded,

pale glaucous green. Males of importunus and nebulosus are difficult to sepa-

rate, though the latter usually have middorsal pale triangles and a more exten-

sive and distinet pateh of black pilosity on the mid posterior border of meso-

notum.

All specimens seen from Panama have come from the Canal Zone area, and

all but a very few from horse-baited mosquito stable traps or biting cattle and

horses. Specimens have been taken in every month save Feb., June and Dec.,

though there seem to be peaks of abundance in May and August. it is probably

more widespread and abundant than records show, but since it does not appar-

ently occur in forested areas and is attracted only to large animals, our col-

lecting methods outside the Canal Zone did not reveal its presence. The range

is extensive, from Panama to Paraguay; it apparently does not extend into Costa

Rica or farther north.

Tabanus lacajaensis Kroeber

Tabanus lacajaensis Kroeber 1931, Stett. Ent. Zeit., 92(1-2): 303, Costa

Rica. Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28 p. 95, syn-

onymy.

Tabanus umbraticolus Fairchild, 1943, Ann. Ent. Soc. Amer., 35(4): 450-

451, Fig. 7 (1942), female, Panama.

This is a slender brown fly with unusually long brown-tinted wings, narrow

frons, and long, slender black third antennal segment. It closely resembles T.

unipunctatus Big., but lacks a black scutellar spot. The eye is unbanded, dull

bronzy in life. Records are all from the mountains of Chiriqui and Bocas del

Toro provinces, those from Cerro Pando, Bocas prov. being larger and darker

than those from Boquete. The only available male is like the female from

Cerro Pando, in being very dark, the eyes bare, holoptic, the upper eye facets

not differentiated or enlarged, and with a small bristly tubercle sunk between

the eyes at vertex. The species has been taken from March to May at Boquete,

with a single record for October. It was described from near San Jose, Costa

Rica, and I have seen material from the same area. It has not been reported

from elsewhere. If the single male mentioned above is properly associated, then

a further difference from unipunctatus occurs, since in the latter, the upper eye

facets are enlarged and well demarcated and differentiated from the small

facets.

Tabanus macquarti Schiner

Tabanus macquarti Schiner 1868, Reise Novara, Dipt., p. 89, female,

Colombia. Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28, p. 96, syn-

onymy. Wilkerson, 1979, Cespedesia 8(31-32): 377-379.

G. B. Fairchild: Tabanidae of Panama 115

Tabanus (Chelotabanus) macquarti: Fairchild, 1964, J. Med. Ent., 1(2):

181-183, fig 6, references. Panama, Colombia.

This species is similar in appearance to T. bigoti and T. surifer, but the dor-

sal antennal tooth is very long and spine-like, reaching to or beyond end of basal

plate. The antennae and legs are largely black, the wing brownish, especially

along fore border, and there is frequently a short appendix on fork of third vein

on one or both wings.

In Panama the species has only been taken in wet montane forest in Darien

province, on Cerro Pirre and Cerro Quia, and on the upper Rio Mcno, all locali-

ties close to the Colombian border. Some were taken attacking man at ground

level, others in Shannon or Anderson traps, from Jan. to March, 1961 and 1969.

Flight season is not necessarily limited to these months, as little collecting has

been done in this area and only at that time of year. The range is from Costa

Rica to Ecuador and Venezuela, apparently mainly in lower montane rain forest.

Tabanus morbosus ebeneus Philip

Tabanus ebeneus Philip, 1966, Ann. Ent. Soc. Amer., 59(3): 525-526, fig.

4. Female, Guatemala, Panama.

Tabanus morbosus Stone, 1938, U.S. Dept. Agric. Miscell. Pub. No. 305,

pp. 89-90, fig. 36. Female, Arizona. |

Tabanus morbosus ebeneus: Fairchild, 1971, Cat. S. Amer. Dipt. Fasc.

28, p. 96. Burger 1976, Southwestern Naturalist, 21(1): 117-121.

Only a single specimen, a paratype from Chiriquicito, Bocas del Toro prov.,

May 1917, is known from Panama. The holotype and another paratype were

taken in Guatemala. The species differs from erebus O. S., the only similar

local species, in more slender callus and narrower frons, paler wings, and more

slender antennae without a strong dorsal spine. Nothing is known of the habits

of ssp. ebeneus; Burger (loc. cit.) collected and reared larvae of morbosus from

several localities in Arizona, but found the adults very rare.

Tabanus nebulosus De Geer

Tabanus nebulosus De Geer 1776, Mem. Serv. Hist. Ins., 6: 227, Pl. 30,

fig. 2, female, Surinam. Fairchild, 1971, Cat. S. Amer. Dipt., fase

28, p. 96, synonymy; 1984, Contr. Amer. Ent. Inst. 21(3): 27, Map 3.

Goodwin and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1): 130, Fig.

39.

Tabanus ferrifer Walker, 1850, Ins. Saunders., 1, Dipt., p. 30, Barbados.

Fairchild, 1943 (1942), Ann. Ent. Soe. Amer., 35(4): 444, fig. 3, fe-

male, Panama.

The earlier references are listed by Fairchild (1943) under ferrifer; the ex-

tensive synonymy in the Cat. S. Amer. Dipt. This is a large brown horse fly

with parallel-sided frons, ridge-like callus and a median row of small yellow-

haired triangles and sublateral rounded spots on abdomen. The eyes in life are

yellowish green with reddish purple highlights, without pattern. The male is

similar in color and the eyes, though not much enlarged, have the area of en-

larged facets well differentiated and demarcated from the small. T. importunus

116 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

is the most similar species, but differs in very short inflated palpi, in lacking

the mid-dorsal pale-haired triangles on abdomen, and in usually having at least a

small area of infuscation on the wing beneath the stigma. Both species have a

patch of black hairs before the scutellum.

In Panama nebulosus is common and widespread in the more open habitats at

lower elevations. It bites primarily horses and cattle, not man, and may be a

serious pest at times. Most of the available records are from horse-baited

mosquito traps in the Canal Zone area, as little collecting from bait animals has

been done elsewhere. Males are frequently taken in light traps. The species

flies throughout the year, though it appears to be most abundant from March to

May and from August to October. The species ranges from British Honduras to

southern Brasil.

Goodwin and Murdoch (1974) describe and figure the female pupa, one of 2

they reared from a total of 3 specimens collected, apparently as pupae, as the

larva is not mentioned.

Tabanus nereus Fairchild

Tabanus nereus Fairchild 1943, Ann. Ent. Soe. Amer., (1942) 35(4): 446,

fig. 9, female, Panama; 1958, Ann. Ent. Soc. Amer., 51(6): 530;

1973, Proce. Ent. Soe. Washington, 73(3): 322-323, fig. 3. Wilkerson,

1979, Cespedesia 8(31-32): 379-380. Colombia. Fairchild, 1983, Ent.

Soe. Amer. Mise. Publ. No. 57, p. 24, fig.

This small inconspicuous yellowish-brown species has the eyes in life green

with a narrow midstripe and the lower margin broadly purple. The abdominal

markings consist of 3 faint stripes. It much resembles small specimens of non-

descriptus, but in addition to the patterned eyes has a broader frons, nearly

round yellow callus, and more slender palpi. Head characters and eye pattern

suggest relationship with T. guatemalanus.

In addition to the specimens previously listed, from the Canal Zone, Darien

and Herrera provinees, it has been taken in horse baited stable traps at Ft.

Kobbe, C. Z., and a long series was taken at Curiche, Dept. Choco, Colombia, in

a Malaise trap by personnel of the O.I.C.S. survey. There is also a single

specimen from Guayaquil, Ecuador, May-June 1913, C. T. Brues coll. in M.C.Z.

Collections show this to be also a coastal species and flying throughout the

rainy season from May to Dec.

Tabanus nondescriptus Fairchild

Tabanus nondescriptus Fairchild 1973, Proc. Ent. Soe. Washington, 75(3):

323-326, fig 4.

Tabanus rixator: Fairchild, 1958, Ann. Ent. Soe. Amer., 51(6): 530, in

part, not Fairchild 1943.

A small dull brownish species with 3 inconspicuous pale stripes on abdomen,

wings with all veins narrowly brown margined, frontal callus brown, higher than

wide, and eyes unpatterned.

The species is exceedingly close to nereus Fehld., but differs in having un-

banded eyes, a higher, more rectangular callus, differently shaped antennae,

more prominently striped abdomen and larger average size. The means and me-

G. B. Fairehild: Tabanidae of Panama 117

dians for wing lengths of available specimens of nereus are both 9.5 mm., and

for nondescriptus 11.13 and 11.25 mm., for those from Sta. Fe, Darien,and 11.0

and 10.5 for those from all other localities. It differs from rixator in generally

larger size, reddish scutellum, long oval or oblong callus, and in lacking a broad

prominent yellow stripe on abdomen. |

The species occurs near or in mangrove swamps along the Pacific coast from

Darien prov. to the Canal Zone area. Specimens have been taken flying about

the collector, in horse-baited mosquito stable traps and most abundantly, in

Malaise traps. It appears to fly throughout the year, with perhaps a peak of

abundance from Feb. through May. It has not yet been collected outside of

Panama. :

Tabanus occidentalis var. dorsovittatus Macquart

Tabanus dorsovittatus Macquart, 1855, Dipt. Exot., Suppl. 5. p. 50.

Tabanus dorsiger var. dorsovittatus: Fairchild, 1971, Cat. S. Amer.,

Dipt. Fase. 28, p. 92, synonymy. Goodwin and Murdoch, 1974 Ann.

Ent. Soe. Amer., 67(1): 124 fig. 17.

Tabanus lineola var. carneus: Fairchild, 1942, Ann. Ent. Soc.Amer.,

35(2): 173-174, Pl. 2, figs 21-25. Synonymy.

Tabanus occidentalis var dorsovittatus: Fairchild, 1983, Ent. Soc. Amer.

Mise. Publ. No. 57, p. 26.

This is the most abundant horse-fly in Panama, and the one most likely to

be taken by the casual collector, as it is found primarily in open country and

cleared land, and is a serious pest of cattle and horses. It varies considerably

both in size and coloring. Females with wing lengths of 7 mm. to 11 mm. have

been seen, while the abdominal stripes may be chalky white to distinctly yel-

lowish. The mid and hind femora are always entirely pale. Males have been

taken hovering in open areas, and are abundant in light trap catches, like it's

close relative lineola in N. America. The extent and size of the upper eye

facets is also variable, but they are always clearly demarcated from the small

facets. The species occurs throughout Panama up to elevations of at least 3000

ft., but is apparently absent from heavy primary forest. The various forms of

occidentalis range from North Argentina to Mexico.

This and the 2 following forms are treated here as in my recent revision of

the lineola group (Fairchild 1983), though this may not be the final word. Nei-

ther a statistical study nor examination of the female genitalia has been done.

Goodwin and Murdoch (1974) found this one of the commonest species they

collected as larvae, and they reared over 100 to the adult stage. They figured

the larva and stated that the pupa was not separable from that of T. ecolom-

bensis Viacq. Larvae were widespread in damp soil near water, mostly on the

Pacific side of the isthmus. The other forms of the species were unfortunately

not secured.

Tabanus occidentalis var. modestus Wiedemann

Tabanus modestus Wiedemann, 1828, Auss. Zweifl. Ins., 1: 146.

Tabanus lineola var. plangens Walker: Fairchild, 1942, Ann.Ent.

Soc.Amer., 35(2): 174.

118 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Tabanus dorsiger ssp. modestus: Fairchild, 1971, Cat. Dipt. S. Amer. 28:

92.

Tabanus occidentalis var. modestus: Fairchild, 1983, Ent. Soc. Amer.

Mise. Pub. No. 57, p. 26.

This form resembles stenocephalus in structure and eye color, but is much

darker, the abdomen nearly entirely black with a narrower white middorsal

Stripe and the mid and hind femora over two-thirds black basally. The only

males I have seen resemble those of stenocephalus with the abdomen dark red-

dish brown and the stripes yellow; they differ however in having the upper eye

facets well demarcated and differentiated, being in this respect like occidentalis

var. dorsovittatus. The 2 males studied were collected in Antioquia, Colombia

together with females, and both sexes of dorsovittatus.

The form appears to range from Honduras to southern Brazil and eastern

Peru. Brazilian examples show every intergrade with dorsovittatus, while

Panama specimens can nearly always be separated by the black seutellum and

largely black hind femora.

In Panama, modestus has been taken on both sides of the isthmus in the

Canal Zone area, but shows a marked preference for primary forest in areas of

high rainfall, being common at our Yellow Fever station near Almirante, and at

our Rio Taecarcuna camp in Darien, where neither dorsovittatus nor steno-

cephalus were taken. Most of the available specimens were taken in Shannon

traps.

Tabanus oculus Walker

Tabanus oculus Walker 1848, List Dipt.Brit. Mus., 1: 157. Female, Hon-

duras. Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28 p. 98, syn-

onymy.

Tabanus (Bellardia) oculus: Fairchild, 1942, Psyche 49(1-2): 12-13. figs.

3 a, b. Female, Panama, full references.

This species can be separated from the very similar T. pseudoculus Fehld. by

having the first posterior cell of the wing always closed and petiolate and hav-

ing distinct though pale brown clouds around ecross veins at ends of diseal cell

and at fork of third vein. The frons is also slightly narrower and the eye pat-

tern with 2 or 3 fairly wide green bands and the lower margin green.

Only a single specimen has been taken in Panama, and that was collected

many years ago by L. H. Dunn in extreme western Chiriqui province, close to

the Costa Rican border. This specimen was taken together with T. albocirculus

and T. pseudoculus attacking a mule at Camp Pital, and all were reported by

Dunn (1934a) as T. albocireulus. The species ranges north to southern Mexico.

Tabanus olivaceiventris Macquart

Tabanus olivaceiventris Macquart 1847, Dipt. Exot. Suppl. 2: 18. Male

(Female), Para, Brasil. Fairchild, 1964, J. Med. Ent. 1(2): 184-185.

Male, female, Panama, references; 1971, Cat. S. Amer. Dipt. Fase

28, p. 98, synonymy; 1984, Contr. Amer. Ent. Inst. 21(3): 30, fig. 17,

Map 8. Wilkerson, 1979, Cespedesia, 8(21-22): 380-382.

G. B. Fairehild: Tabanidae of Panama 119

In life this species has an unmarked yellowish green abdomen which fades to

dirty yellow in older preserved specimens. The thorax is steel grey with a small

contrasting spot of black hairs just in front of scutellum. The wings are gener-

ally glass clear, occasionally brownish tinted along the veins. The frons is par-

allel sided, narrow, with a slender ridge-like callus. The eyes in life are pale

glaucous green, unbanded. The male is similar in color, but with abdomen

densely white-haired, upper eye facets only moderately enlarged but well demar-

eated from small facets, and with a very small tubercle sunk between eyes at

vertex. This species appears to belong to a group including T. importunus, T.

nebulosus and T. punctipleura, all of which have a black spot on hind border of

mesonotum and unicolorous glaucus green eyes. The species are superficially

quite different in color, but frons and frontal callus are quite similar in all.

Only a single specimen has been taken in Panama, at Cerro Pirre, Darien

Prov., 5 Feb. 1961. The locality is at 1500 ft. in wet montane forest. The

specimen was taken in a Shannon trap. The species ranges southwards across

northern South America at least to the mouth of the Amazon.

Tabanus platycerus Fairchild

Tabanus platycerus Fairchild 1976, Stud.Ent., 19(1-4): 242, 251-253, fig.

9. Female, Panama, Costa Rica.

Tabanus (Neotabanus) fumatipennis: Fairchild, 1942, Ann. Ent. Soc.

Amer., 35(2): 162, P1. 1, fig. 3. Not Kroeber 1933, misident.

Although not abundant, this small species appears to occur throughout

Panama at low elevations. Most specimens were taken in horse-baited mosquito

stable traps, in Malaise traps, or from the screens of a railroad track car run-

ning through banana plantations. The male has not yet been taken. Specimens

look like small T. occidentalis var. modestus, but the frons is much narrower

and the subcallus bare and shiny. In the Canal Zone area and Darien records

are all from April to June, though in Almirante it was taken into October. It

appears to range north to Nicaragua, but has not so far been taken south of

Panama.

Tabanus polyphemus Fairchild

Tabanus (Lophotabanus) polyphemus Fairchild, 1958, Ann. Ent. Soc.

Amer., 51(6): 527-528. Female, Panama.

Tabanus (Lophotabanus) fumomarginatus (Hine): Fairchild, 1951, Ann.

Ent. Soe. Amer., 44(3): 455-456, figs. 7, 7a, 7b. Female, Panama.

Not Hine 1920, misident.

This species is quite similar in general appearance to T. defilippii, but is

paler, with reddish orange legs, a middorsal row of pale haired triangles on

abdomen, and a broader frons with better developed basal callus. The third an-

tennal segment is orange, with but a moderate dorsal angle. The eyes in fresh

specimens are bicolored, the lower 2/3 green, the upper third bronzy, the line

between the two colors sharp.

In Panama the species has only been taken in areas of heavy forest in the

eastern half of the country, from Cerro Azul and several localities in Darien

Provinee. All specimens were taken April or May, in Shannon or Malaise traps,

120 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

and it does not appear to be common. Elsewhere it is known from Honduras,

Guatemala and the Mexican states of Tabasco and Vera Cruz, and Colombia. A

series of 10 specimens were taken attracted to man in the forest canopy at

Lancetilla, Honduras, so the species may be predominantly arboreal. This com-

bined with an apparently short flight season, may account for the few specimens

taken in Panama, where arboreal catches may not have been made in localities

where it occurs at the right time of year.

Polyphemus is very similar to T. fumomarginatus Hine, differing in slightly

better defined frontal callus, broader third antennal segment, and smaller, or-

ange-haired median triangles on abdomen. It is quite possible that the two may

be only subspecifically distinct. Wilkerson (1979) did not find the species in

western Colombia.

Tabanus praepilatus Fairchild

Tabanus praepilatus Fairchild 1943, Ann. Ent. Soe. Amer., (1942) 35(4):

445-446, fig. 11, Panama; 1953, Ann. Ent. Soc. Amer., 51(6): 530,

male; 1973, Proc. Ent.Soc. Washington, 75(3): 320, fig. 2. Goodwin

and Murdoch, 1974, Ann. Ent. Soe. Amer., 67(1): 130, figs. 18, 40.

A slender grayish brown species with clear wings and 3 rows of pale spots

on the abdomen. It somewhat resembles T. pungens Wied. but is usually larger,

lacks the broad black middorsal abdominal stripe, and has quite different anten-

nae, with style markedly longer than basal plate. The eyes are dull greenish

with, usually, a slender median dark stripe which fails to reach lateral eye

margin. All specimens seen have come from the vicinity of the Pacific coast,

from Darien to Los Santos provinces, generally in or near mangrove swamps.

The male and 1 female were taken in a light trap. The male resembles the fe-

male in color and pattern, the bare eyes holoptic, with upper enlarged facets

well-differentiated and demarcated from small facets, covering about 3/4 of eye

area. There is a small tubercle sunk between the eyes at vertex. Specimens

have been taken in Jan., Feb., May and August, so that it probably flies

throughout the year. The species has not been reported from elsewhere.

_ Goodwin and Murdoch (1942) took 3 larvae from white sandy soil at Guarare,

Los Santos prov., Panama. A male was reared, and they were able to describe

and figure both larva and pupa.

Tabanus praeteritus Fairchild

Tabanus praeteritus Fairchild 1947, Ann. Ent. soe. Amer., (1946) 39(4):

979, 572, fig. 6, male, female, Panama; 1958, Op. cit., 51(6): 529-

530. (in part). Wilkerson,1979, Cespedesia 8(31-32): 382-383.

A blackish-brown species with small round yellowish-brown callus extended

above in a slender line. Wings grayish hyaline, no appendix on fork of third

vein, thorax unstriped, blackish, as is scutellum; abdomen dull yellowish-brown

with a black integumental triangle on first and second tergites, sometimes ex-

tended to form a continuous or broken black streak on succeeding tergites.

Dorsolateral pale hair patches usually present on tergites 1 to 5, and often also

a faint pale middorsal line, though both markings easily lost in denuded speci-

mens. Fore tibiae obscurely bicolored, mid and hind tibiae dull yellowish-brown.

G. B. Fairehild: Tabanidae of Panama 121

Fore femora blackish, mid and hind femora all dull yellowish or basally more or

less blackish. The eyes are yellowish or greenish bronzy, unbanded.

The male has the upper eye facets but slightly enlarged and poorly demar-

cated from the small facets. it is otherwise similar to the female and easily

associated with it. Only a single male, from Parrita, Puntarenas Prov., Costa

Riea, has been seen. It is in the Los Angeles County Museum.

There appear to be two forms of this species, separable on small color char-

acters and separated geographically. The typical form is slightly darker, has at

least bases of mid and hind femora infuscated and the oval yellowish-haired dor-

solateral spots on abdomen quite prominent. This form seems to be limited to

the Pacific coast, where it occurs from Puntarenas prov., Costa Rica to Rio

Raposo, south of Buenaventura, Colombia. In Panama, in addition to the type

from Chepo. Panama prov., and paratypes from Ancon C. Z. and El Real Darien,

I have a short series from Sta. Fe, Darien, taken in a Malaise trap in Jan., Feb.,

April and May 1967, by personnel of the O.I.C.S. The other form, with wholly

reddish-yellow mid and hind femora and less prominent dorsolateral hair patches

is deseribed below.

Tabanus praeteritus adiastolus Fairchild n. ssp.

Tabanus praeteritus Fairchild 1958 Ann. Ent. Soe. Amer. 51(6): 529-530.

Not Fairchild 1947.

A dull blackish brown insect differing from the nominate subspecies largely

in color, as follows: fore femora and coxae reddish brown, at most slightly

darker than tibiae (in praeteritus both are black, contrasting with bicolored

tibiae); mid and hind femora wholly yellowish brown (in praeteritus basally about

1/2 blackish); abdominal vestiture largely black, but with a tuft of pale yellowish

hairs in middle of first tergite at apex of scutellum and in well preserved

specimens a narrow middorsal pale pilose stripe (in praeteritus, there is no such

tuft nor middorsal pale stripe, the tergites bearing fairly prominent orange

pilose dorsolateral patches on at least tergites 2 to 4, often also on 1 and 5,

though reduced in size. The callus in the present subspecies is usually yellow or

brown, tapering upward gradually into the median callus, while in the nominate

form the callus is more often black, abruptly joined to a slender thread-like

median callus

Holotype female, Panama Canal Zone, Ft. Gulick, 28-V-1953, horse baited

Stable trap. To be deposited in Florida State Collection of Arthropods.

Paratypes, all females: 1, same locality as holotype, 28 Jan 1954, light trap;

Canal Zone: 24, Fance Field, Jan to May 1953, horse traps, all somewhat

rubbed; 13, Navy tank farm, Gatun, Galeta Point road, horse traps, Jan to June,

1953; 2, Coco Solito, horse trap, 1-VII-53; 1, Ft. Sherman, horse trap 13 Nov.

1951; 1, Ft. Davis, horse trap 19 Jan 1953; 1, Ft. Randolph, horse trap 11-VI-63;

Bocas del Toro prov., 2, Almirante, biting horse, 14-VIII-63; Colon prov.,1 ,

Porto Bello, Shannon trap, Oct. 63, 1, Mandinga, ?Shannon trap, 30-V-64; San

Blas prov., 1, Rio Mandinga, 10-V-57, 1, Rio Tangandi Yellow Fever Camp,

Shannon trap at dusk, 23-V-57, 1, Cuadi River 14-IV-67, 1 Sasardi, 14-IX-67,

O.1.C.S. 3914. Specimens range from 10 to 14 mm in length. The name is from

Greek, adiastolos, meaning not separated, confused, as its distinction from prae-

teritus was only recently recognized.

The two taxa are both coastal, praeteritus known from localities on or near

the Pacific coasts of Colombia, Panama and Costa Rica, while ssp. adiastolus

122 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

seems confined to the Caribbean coast of Panama from San Blas province, to

Almirante in Bocas del Toro prov. The bulk of our material was taken in horse

baited mosquito stable traps on army reservations on the Atlantie side of the

Isthmus in the former Canal Zone. A few were also taken in Shannon traps and

light traps or biting a tethered horse. The eyes in life are unbanded, dull

greenish or yellowish bronze. The 3 specimens from Atlantic coast localities

listed in the original description of praeteritus are not now before me, but were

no doubt the present subspecies.

Tabanus pruinosus Bigot

Tabanus pruinosus Bigot 1892, Mem.Soc. Zool. Franee 5: 683, male,

Mexieo. Philip, 1950, Ann. Ent. Soe.Amer., 43(1): 116, fig 2C.

Fairchild, 1951, Ann. Ent.Soec.Amer., 44(3): 459-460, female, Panama;

1971, Cat. Dept. S.Amer., Fase. 28,. p. 99. synonymy. Burger 1974,

Proc. Ent. Soc. Washington 76: 2): 106.

Hybostraba albovillosa Kroeber, 1931, Stett. Ent.Zeit., 92: 92-93, male,

Costa Rica. Fairchild, 1975, Proc. Ent. Soe. Washington, 77(2): 261.

Only 2 specimens have been seen from Panama, the female specimen men-

tioned by me in 1951 and a single male from Cerro Punta, Chiriqui, 5500 ft., at

light, L. J. Bottimer ecoll., in C.N.C. Both localities are in the mountains of

Chiriqui prov., not far from the Costa Rican border. This represents the far-

thest south record of the species, which ranges northward to southern Arizona.

It is a blackish brown species with striped mesonotum, wholly clear wings, and

abdomen with small median gray triangles and a row of sublateral small round

gray spots on each side of the abdomen. The eyes are dark purple with 2 nar-

row green bands. The male is strikingly different, the abdomen nearly white

due to long dense white hairs, and with the eyes densely short pilose on the area

of enlarged facets. The male type of H. albovillosus Kroeber was studied on

loan from Warsaw (Fairchild 1975) and agrees closely with this Chiriqui male.

Specimens from Mexico and Arizona differ slightly from the Panama specimens

in having more slender antennae, more prominent middorsal pale triangles on ab-

domen, and with male eyes essentially bare. If these differences hold up when

adequate series become available, Kroeber's name is available for the Panama

population.

Tabanus pseudoculus Fairchild

Tabanus (Bellardia) pseudoculus Fairchild, 1942, Psyche, 49(1-2): 13-14,

figs. 1 a, bh Female, Panama. Wilkerson, 1979, Cespedesia 8(31-32):

383-384.

Tabanus albocirculus (Hine): Dunn, 1934, Psyehe 4(3): 173-174, in part,

not Hine, misident.

Although very similar to T. oeulus, with which it has probably been con-

fused, it differs from that species in always having the first posterior cell of

the wing (4th R) open, though coaretate. The eyes in life are purplish bronze

with two very narrow bands and the lower margin green, while the wings are en-

tirely without clouds on the crossveins and fork. The male is like the female in

eolor, the large eye facets pale tan, well differentiated and demarcated from

G. B. Fairchild: Tabanidae of Panama 123

the small and occupying 1/2 to 2/3 of eye area. The small facets are purple

with a single narrow green band.

The speeies bites eattle and horses, and is taken in Shannon traps. It is

recorded from all parts of Panama at low elevations, mostly in forested regions,

though not by any means confined to a forest habitat. It is not a rare species,

but is never a dominant one. Specimens have been taken in all months of the

year. The range is from Guatemala to northern Colombia, Venezuela and

Trinidad, though there are few records north of Panama.

Tabanus punctipleura Hine

Tabanus punctipleura Hine 1920, Ohio J. Sci., 20(8): 314-315, female,

Costa Rica. Fairchild, 1946. Ann. Ent. Soe. Amer., 39(4): 569-570,

female, figs. 4, a, b. Panama, full references; 1984, Contr. Amer.

Ent. Inst. 21(3): 33-34, Map 4.

Tabanus defilippi: Kroeber, (sic) 1929, Zool. Anz., 83(1-4): 126-127 fig.

8. Not defilippii Bellardi, misident.

This is one of the largest species of Tabanidae in Panama and can be recog-

nized readily by the dark brown, grayish pruinose, unicolorous abdomen, brownish

wings and strong pateh of dense erect black pile on the prescutellum which is

bounded laterally by prominent white hair tufts. The pleura also bear a tuft of

black hairs below the wing bases. It is likely to be confused only with T.

defilippii, as Kroeber did, but the long tooth on antenna, and lack of black pile

on scutellum will easily separate punctipleura. The eye in life is uniformly light

yellowish green. I have seen a male from Costa Rica, P. Serre eoll., in Paris

Mus. The upper eye facets are but slightly enlarged and poorly demarcated

from small facets, occupying less than half eye area. There is a small wedge-

shaped tubercle between eyes at vertex. The palpi are brown, porrect, with a

small nipple at apex. The antennae are more slender and lack the ventral angle

found in female, but color and general appearance is the same.

In Panama the species has only been taken in the vicinity of Almirante, Bo-

eas del Toro prov., biting a horse and in a Shannon trap. It is not common

there, though specimens were taken from April to Nov. It was not secured at

our Yellow Fever Camp during two years of intensive collecting in earlier years,

but only at localities along the railroad in low or cut-over areas. It appears to

have a very restricted distribution, as all records are from the Atlantic coast of

Costa Rica or adjoining northwestern Panama.

Tabanus pungens Wiedemann

Tabanus pungens Wiedemann 1828, Auss. Zweifl. Ins. 1: 175. Fairchild,

1971, Cat. S. A. Dipt. Fase. 28, p. 100. (synonymy). Goodwin and

Murdoch, 1974 Ann. Ent. Soe. Amer., 67(1): 124, figs 19, 41. Wilker-

son, 1979, Cespedesia 8(31-32): 394-386. Coscaron 1979, Obra Cent.

Mus. de la Plata, 6: 270, figs.

Tabanus angustivitta: Fairchild, 1942, Ann. Ent. Soec.Amer., 35(2): 165,

Pl. 1, fig. 7. 1942 op. cit. 35(1): 88. Seasonal distribution.

This species is easily recognized in the Panama fauna by the presence of a

discrete tubercle at vertex, and an abdominal pattern which consists of a mid-

124 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

dorsal row of narrow connected pale triangles on a broad black middorsal stripe.

The sides of the abdomen are pale, reddish yellow, with a row of prominent

oblique pale-haired patehes. The eyes in life are purple with two transverse

green bands. The male has the eye facets differentiated into two sizes, the

large facets in a central triangular patch, occupying about 1/2 the eye area, and

there is a prominent tubercle at vertex.

The species is fairly abundant at low elevations in the Canal Zone and gen-

erally throughout the country in open areas. It attacks horses and cattle, but

not man, and is a dominant species in horse-baited stable traps. Males are fre-

quently taken in light traps. In Panama it occurs throughout the year, but is

definitely more abundant in the dry season, from December to May. It appears

to range from Texas to Argentina, but is absent from Chile and the West Indies.

Goodwin and Murdoch reported the larvae as abundant. They reared over

100. Larvae were found in the same habitats as dorsovittatus and colombensis,

but on both sides of the Isthmus.

Tabanus quinquepunctatus Hine

Tabanus quinquipunctatus Hine 1925, Occ. Papers Mus. Zool. Univ.

Michigan No. 162, p. 33. Nom. nov. pro T. quinquemaculatus Hine

1907. Fairchild, 1958. Ann. Ent. Soe. Amer., 51(6): 529, fig. 3;

1971, Cat. Dipt. S. Amer. Fase. 28, p. 100, synonymy.

Tabanus quinquemaculatus Hine, 1907, Ohio Nat., 8(2): 224. Not Hine

1904.

This is a small black species with a prominent row of small middorsal white

triangles on abdomen and the tergites white-margined behind, black femora but

reddish tibiae which are at least partly white-haired, and grayish hyaline wings.

All material seen from Panama was taken in light traps at elevations of over

1000 ft. in Chiriqui or Cocle provinces, as listed by me in 1958 (loc. cit.). I

have also seen 3 females in U.S.N.M. from Potrerillos, Chiriqui prov., D.V.

Brown coll. Specimens were taken in December, January and March. the

species ranges north to Guatemala, but seems nowhere common, perhaps because

of its apparently nocturnal habits.

Tabanus rhizophorae Fairchild

Tabanus rhizophorae Fairchild 1943, Ann. Ent. Soc. Amer. (1942) 35(4):

449-450, fig. 6, male, female, Panama; 1973, Proc. Ent. Soe. Wash-

ington, 75(3): 319-320, fig 1.

An inconspicuous small species with steel grey thorax, yellowish-brown ab-

domen with traces of median and dorsolateral pale-haired stripes on well pre-

served examples. The frons is quite broad, the eyes green or greenish black in

life, without pattern. The male is similar, though more yellowish and with more

obviously striped abdomen. The eyes have enlarged facets occupying about 1/2

eye area, but not well demarcated from the small facets. The type series was

taken sweeping salt marsh grass on the intertidal mud flats near Old Panama, a

locality now much modified by the growth of the city. It has also been netted

flying about the collector in a few other mangrove coastal habitats along the

Pacific coast in Panama prov. A single defective specimen lacking antennae

G. B. Fairehild: Tabanidae of Panama 125

from Coiba Id., off the coast of Chiriqui prov. may be the same, though the

frons is slightly narrower. All material was taken in the period Jan. to July,

with most specimens from Feb. to April. The species is not known outside

Panama.

Tabanus rixator Fairchild

Tabanus rixator Fairchild 1943, Ann. Ent. Soe. Amer., (1942)(4): 448, fig.

10, female, Panama; 1958 Ann. Ent. Soe. Amer., 51(6): 530, in part;

1973, Proc. Ent. Soc. Washington, 75(3): 321-322, fig. 5.

This little species is distinguished from nereus and nondescriptus by having

the frontal callus broadly clavate or pear-shaped, and a prominent broad yellow-

haired stripe the full length of the abdomen. Only the type and the 2 first

specimens mentioned in 1958, plus 1 female taken in a mangrove swamp in the

outskirts of Panama City, 3 Dec. 1964, are certainly this species. The

paratypes are not now before me, and may be in part nondescriptus. From the

scanty records, the species seems confined to coastal mangrove swamps on the

Pacific coast of Panama and flies during the rainy season, from July to Decem-

ber. The eyes in life are bronzy green, unbanded. It has not, as yet, been

taken outside Panama.

Tabanus rubripes Macquart

Tabanus rubripes Macquart 1838, Dipt. Exot., 1(1): 134, female, Cayenne.

Fairchild, 1964, J. Med. Ent., 1(2): 184-185, fig. 7, female, Paraguay,

Brasil, Colombia, Panama; 1971, Cat. S. Amer. Dipt., Fase. 28, p.

101, synonymy. Wilkerson, 1979, Cespedesia 8(31-32): 386.

Only a single specimen of this species has been taken in Panama, at Cerro

Azul, Panama Prov., 6 May 1961. It somewhat resembles in size and general

appearance T. pseudoculus Fehld., but lacks the prominent black hair spot on

scutellum and has a narrower frons and callus. Both species have an appendix

on fork of third vein and narrowed first posterior cell. The eye in life has two

bands and the extreme lower margin green, the upper band narrower than the

lower, the bands seemingly wider than in pseudoculus. Although Philip (1960)

States that the eyes of the synonymous T. lophus Phil. have 2 narrow purple

bands on a greenish ground, I believe this is due to a reversal in color some-

times seen when the pattern is revived. The male, according to Philip, has the

enlarged eye facets well demarcated from the small and occupying the upper 2/3

of eye, with a small tubercle sunken between eyes at vertex. South American

specimens have been taken in Shannon traps and attacking man. The species

ranges south to Paraguay east of the Andes, but was not taken on the Western

side of the Andes in Colombia during intensive collecting near Buenaventura, by

Lee et. al. (1969) or Wilkerson (op. cit.).

Tabanus stenocephalus Hine

Tabanus stenocephalus Hine 1906, Ohio, Naturalist 7(2): 27. Fairchild

1983, Ent. Soc. Amer. Mise. Pub. No. 57, p. 28, figs.

126 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Tabanus lineola var. stenocephalus: Fairchild 1942, Ann. Ent. Soc.

Amer. 35(2): 175, Pl. 2, fig. 26.

Tabanus dorsiger ssp. stenocephalus: Fairchild, 1971, Cat. Dipt. S.

Amer. 28: 92.

This species is readily separated from occidentalis var. dorsovittatus by

having the bases of at least hind femora contrastingly black, an integumental

spot or patch of black both above and below at base of second abdominal seg-

ment, and scutellum wholly black. In addition, the abdominal markings are usu-

ally darker yellow while the upper facets of the male eye are not differentiated

or demarcated from the small facets. The eye of the female in life generally

lacks the short upper dark band. Stenocephalus is not so abundant as the forms

of occidentalis and in Panama is almost confined to areas of heavy rainfall on

the Atlantic side of the isthmus and in Darien province. Though it was taken

frequently in the banana areas of Bocas del Toro Prov., intensive collecting in

primary forest at our Yellow Fever Station a few miles away did not yield

Specimens. The bulk of our material came from horse-baited stable traps, biting

horses, or in buildings or Shannon traps.

The species ranges from Guatemala to eastern Peru and Surinam, but I have

seen no typical examples from south of the Amazon basin.

Tabanus surifer Fairchild

Tabanus (Chelotabanus) surifer Fairchild, 1964, J.. Med. Ent. 1(2): 183-

184. M. F. Panama, Colombia. Wilkerson, 1979, Cespedesia 8(31-32):

386-389.

Tabanus thiemeana surifer Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28,

pp. 102-103, synonymy. !

Tabanus bigoti Bellardi: Fairchild, 1942. Ann. Ent. Soe. Amer., 35(4):

442. figs. 5, 5a, 5b. Female, in part, var. A.

This species differs from T. thiemeanus End. only in having the basal plate

of antenna largely or wholly yellow orange, and in having the dorsal spine much

shorter, often no more than an erect tooth. From bigoti and macquarti, the 2

related species in Panama, it differs in brown frontal callus, brown frontal and

facial pollinosity and yellowish brown tinted wings with a long appendix on fork

of third vein. The abdomen is usually black only on sixth and seventh segments,

not sharply black from fifth on as in bigoti.

The eyes of both sexes are bronzy in life, without pattern. The males,

taken only attracted to light, have a small area of large facets in the middle of

the eye, poorly differentiated and demarcated from the small facets. There is a

small tubercle sunk between the eyes at vertex.

T. surifer is the most abundant of this group in Panama, occurring through-

out the country in lower montane forest or areas of heavy rainfall, primarily

along the Caribbean coast and mountains on the Pacifie side. It has been taken

biting man, horses and pigs, but generally is taken in Shannon and Malaise traps

at ground level, often being the commonest species in its preferred habitat.

Records indicate it flies from January to September, though apparently more

abundant from April to August, during the first half of the rainy season.

Recent material of both surifer and thiemeanus taken in the Dept. of Valle,

Colombia at Anchicaya Dam, 10 Aug. 1973, by Wilkerson and Young, show that

both forms may fly together and surifer thus cannot be a subspecies of

G. B. Fairchild: Tabanidae of Panama 127

thiemeanus as I had listed it in the Catalogue (Fairchild 1971). Males of surifer

also differ from males of what I believe are thiemeanus and macquarti Schin. by

having an area of differentiated large facets in the upper half of the eye. This

area is not sharply differentiated from the small facets surrounding it, but very

evident. Males of thiemeanus and macquarti wholly lack enlarged eye facets.

The group to which surifer belongs is dominant in the andean region of Colombia

and Ecuador from where several additional nominal taxa have been described,

but specific limits await a careful analysis of long series from a variety of lo-

ealities.

Tabanus vittiger guatemalanus Hine

Tabanus guatemalanus Hine, 1906, Ohio Nat., 7: 21-24. Male, female,

Guatemala.

Tabanus vittiger guatemalanus: Fairchild, 1942, Ann. Ent. Soc. Amer.,

35(2): 180, P1. 1, fig. 16. Female, Panama; 1983, Ent. Soc. Amer.

Mise. Pub. No. 57, p. 30, figs.

Tabanus vittiger caymanicus Fairchild, 1942. Ann. Ent. Soc. Amer.,

35(2): 180-181. Male, female, Cayman Islands; Cuba.

Tabanus truquii: Bequaert, 1940, Rev. Ent. 11(1-2): 352-357, fig. 30. Not

Bellardi 1859. Cayman Islands; Puerto Rico.

Tabanus bellardii Szilady, 1926. Biol. Hung. 1(7): 23, Pl. 4, fig. 15. Fe-

male, Cuba.

Tabanus vittiger bellardii: Pechuman, 1957, Ent. News, 68(5): 118.

Florida.

Tabanus subsimilis guatemalanus: Philip, 1965, Ann. Ent. Soc. Amer.,

58(6): 877. Fairchild, 1971, Cat. S. Amer. Dipt. Fase. 28, p. 102.

The female of this species can generally be separated from occidentalis and

its forms by the markedly broader frons, and generally broader abdominal

stripes. The male has the upper facets greatly enlarged and short pilose. Both

sexes have wholly yellow to red mid and hind femora. From colombensis Macq.

It ean be separated by slightly broader and more convergent frons and yellowish

vestiture in the female, and by the enlarged facets and wholly pale femora in

the male. Colombensis has the abdominal stripes chalky white, and the male

eye without markedly enlarged facets, though they are also pilose. The eye pat-

tern of the female consists of 2 purple bands on a green ground, one across the

middle of the eye, the other covering the lower fifth of the eye.

In Panama the species is not abundant, though it appears to occur outside of

the forest at low elevations along the coast throughout much of the country.

Nearly all specimens seen were taken in horse-baited mosquito traps, the few

males in light traps, and most of the records are from the vicinity of the Canal

Zone, where such traps have been chiefly used. In this area, the species seems

about equally abundant on both sides of the isthmus. The species appears to fly

throughout the year, though limited catches suggest it may be more abundant in

April and May. The present subspecies occurs from Florida to Puerto Rico, and

from southern Mexico to the Guianas, Trinidad, and northern Brazil. It seems to

be strictly coastal in habits.

128 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

Tabanus unistriatus Hine

Tabanus unistriatus Hine 1906, Ohio Nat., 7(2): 28. Fairchild, 1942, Ann.

Ent. Soe. Amer., 35(2): 162, P1. 1, fig. 2 (References); 1976, Studia

Ent., 19(1-4): 240, 242, 260, 261, fig. 15. Wilkerson, 1979, Ces-

pedesia 8(31-32): 391-393.

The black abdomen with single narrow median pale stripe, wholly bare sub-

callus, and small size enable easy recognition of this species. The eyes are red-

dish purple with 3 blue-green bands, the bands about the same width as the in-

tervals between them, while the two upper bands are joined, or separated by a

narrow isthmus, at their outer ends. The male has the abdomen reddish brown

on first 3 segments, often with faint indications of sublateral stripes. The head

is wider than thorax, the upper eye facets greatly enlarged, occupying fully half

the eye area and well demarcated from the small facets. There is no tubercle

at vertex. Males were taken in a light trap.

In Panama the species seems to occur only in areas of high rainfall in the

western half of the Republic. There are records from Chiriqui prov. near the

Costa Rican border, Bocas del Toro prov. and from Sta. Fe in the mountains of

Veraguas prov. At our Yellow Fever station near Almirante, Bocas del Toro

prov., where continuous collections were made for 2 years, this was the second

most abundant species, 1349 specimens being taken attracted to man and in a

Shannon trap. Its peak abundance came from May to August though some were

taken in every month. It was seldom taken after dark except for a few males,

and but 2 specimens were secured in tree-top collections. This collecting sta-

tion was in heavy primary forest, as were the other localities where the species

has been taken. It seems to be rare or absent in the banana plantations and

open areas in the same general region, and only a few specimens were taken at

a jungle camp on the Rio Changena at 2400 ft. only a few miles away. The

Species ranges from Guatemala to Ecuador, west of the Andes. |

Tabanus unipunctatus (Bigot)

Atylotus unipunctatus Bigot, 1892. Mem. Soe. Zool. France, 5: 663. Fe-

male, Amer. meridional.

Tabanus unipunctatus: Kertesz, 1900, Cat. Tab. Orb. Terr. Univ. p. 76.

Fairchild 1971, Cat. S. Amer. Dipt. Fase. 28, pp. 103-104, synonymy.

Wilkerson, 1979, Cespedesia 8(31-32): 390-391.

Tabanus fumomarginatus: Dunn, 1934, Psyche 41(3): 174, female,

Panama. Not Hine 1920, misident.

Tabanus piraticus Fairchild, 1942, Psyche, 49(102): 15-16, figs. 7, a, b.

Female, Panama; 1953, Proc. Ent. Soe. Washington 55(5): 241.

This species can be readily separated from related species in Panama by the

unusually slender black or dark brown third antennal segment, often heavily

brown tinted wings and almost unicolorous dark yellowish brown abdomen, with

but a faint dull yellowish middorsal stripe, or series of vague triangles. The

legs, including fore tibiae are nearly uniformly dark brown. The eyes in life are

dark greenish black, without bands. Badly rubbed specimens may be confused

with T. lacajaensis Kroeber, the differences discussed under that species.

The male is similar to the female in color, the enlarged eye facets well de-

mareated from the small and occupying about half the eye area, the eyes uni-

G. B. Fairchild: Tabanidae of Panama 129

formly dark bronzy. The palpi are brown, densely dark-haired, inflated and

Slightly pointed.

The species is limited to areas of heavy forest, primarily where rainfall is

abundant, but occurs throughout Panama where these conditions exist at low to

moderate elevations. It is attracted to man, but there seem no records of its

actually biting, though it may well do so. It has been taken biting mules at

night and in Shannon traps. In the forest it flies mainly at ground level, though

it is occasionally attracted to men working on platforms in the forest canopy.

The 2 males, 1 from Almirante, Bocas del Toro prov. and the other from Rio

Tacareuna, Darien, were both taken at light, one in December, the other in

July. All our records are from May to December, during the rainy season, and

it is often abundant in suitable localities. The range is from Chiapas, Mexico to

Buenaventura, Colombia. The Mexican specimens are considerably paler and

smaller than those from further south.

Tabanus xenorhynchus Fairchild

Tabanus xenorhynchus Fairchild 1947, Ann. Ent. Soc. Amer., 39(4): 572-

573, Fig. 1, Male, Female, Panama, Costa Rica, Guatemala; 1984,

Contr. Amer. Ent. Inst. 21(3): 35-36, Map 7.

This large strikingly marked species is easily recognized by the white

secutellum and white spots on tergites 2, 4, and 5 combined with a wing with

strong large spots on all cross veins and fork of third vein, plus strong infus-

cation on base and apex. It has been taken sparingly in Chiriqui, Canal Zone,

San Blas and Darien, the females frequently in the forest canopy, the males at-

tracted to light. It probably flies throughout the year, as females have been

taken in Mareh, May and August, males in April, July and December. The

known range is from Panama to Guatemala. The structurally similar T. basilaris

Krob. and T. subviolaceus Fehld. occur in Brazil.

130 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

REFERENCES

BARRETTO, M. P. 1957. Tabanidas da Guiana Inglesa, com as descricoes de

seis novas especies da fauna guiano-amazonica (Dipt. Tabanidae). Rev.

Brazil. Malariol. 8(1): 75-89 (1956).

BEQUAERT, J. 1940. The Tabanidae of the Antilles Rev. Ent. 11(1-2): 253-269.

BURGER, J. F. 1976. Notes on Tabanus morbosus Stone in Arizona, with a de-

scription of the male. Southwestern Nat. 21(1): 117-121.

BURGER, J. F. 1977. The biosystematics of immature Arizona Tabanidae.

Trans. Amer. Ent. Soc. 103: 145-258.

BURNETT, A. M. and Kirby L. HAYS 1974. Some influences of meteorological

factors on flight activity of female horse flies. (Diptera: Tabanidae).

Environ. Ent. 3(3): 515-519.

COSCARON, S. 1979. Notas sobre Tabanidos Argentinos XV El Genero Tabanus

Linnaeus. Obra Centenario Museo de la Plata, 6: 251-278, figs. 1-7.

COSCARON, S. and G. B. FAIRCHILD 1976. El Genero Poeciloderas Lutz en

Argentina Physis (Buenos Aires) Sect. C, 35(91): 293-301. 2 figs.

CURRAN, C. H. 1934. The families and genera of North American Diptera.

Privately printed by the Ballou Press, New York, N.Y.

DUNN, L. H. 1934a. Entomological investigations in the Chiriqui region of

Panama. Psyche 41(3): 166-183.

DUNN, L. H. 1934b. Notes on the water lettuce, Pistia stratiotes L., as a nurs-

ery of insect life. Ecology 15(3): 329-331.

FAIRCHILD, G. B. 1941. Notes on Tabanidae from Panama IV. The genus Leu-

cotabanus Lutz. Ann. Ent. Soc.Amer. 34(3): 629-638.

FAIRCHILD, G. B. 1942. The seasonal distribution of some Tabanidae in

Panama Ann. Ent. Soe. Amer. 35(1): 85-91. 7

FAIRCHILD, G. B. 1943. Notes on Tabanidae from Panama X the Genus

Tabanus Linn. and resume of the Tabanidae of Panama. Ann. Ent. Soc.

Amer. 35(4): 441-474 (1942).

FAIRCHILD, G. B. 1946. Additional notes on the Tabanidae of Panama. Ann.

Ent. Soe. Amer. 39(4): 564-575.

FAIRCHILD, G. B. 1953. Arboreal Tabanidae in Panama. Proc. Ent. Soc.

Washington 55(5): 239-243.

FAIRCHILD, G. B. 1956. Synonymiecal notes on neotropical flies of the family

Tabanidae. Smithsonian Miscell. Colls. 13(3): 1-38.

FAIRCHILD, G. B. 1961. Insecta Amapaensia. Diptera: Tabanidae. Studia Ent.

4(1-4): 433-448.

FAIRCHILD, G. B. 1966. Notes on Neotropical Tabanidae VI A new species of

Lepiselaga with remarks on related genera. Psyche 72(3): 210-217(1965).

FAIRCHILD, G. B. 1969. Notes on neotropical Tabanidae XII Classification and

distribution with keys to genera and subgenera. Arq. Zool. S. Paulo, 17(4):

199-255.

FAIRCHILD, G. B. 1971. A catalogue of the Diptera of the Americas south of

the United States. Fase. 28, family Tabanidae. Mus. Zool. Univ. S. Paulo, 1-

163.

FAIRCHILD, G. B. 1973. Notes on neotropical Tabanidae XIV Two new species

of Tabanus from Panama and Colombia. Proc. Ent. Soe. Wash. 75(3): 319-

329.

G. B. Fairchild: Tabanidae of Panama 131

FAIRCHILD, G. B. 1975. Notes on Neotropical Tabanidae XV. Some species

described by O. Kroeber, formerly in the Stettin Museum. Proc. Ent. Soc.

Washington 77(2): 258-265.

FAIRCHILD,“ Ge: Be DO7ds Notes of Neotropical Tabanidae XVII. Genus

Philipotabanus Fehld. Subgenus Mimotabanus Fehld. Ann. Ent. Soc. Amer.

68(4): 689-694, 2 plates.

FAIRCHILD, G. B. 1976. Notes on Neotropical Tabanidae XVI. The Tabanus

trivittatus complex. Studia Ent. 19(1-4): 237-261, 15 figs.

FAIRCHILD, G. B. 1983. Notes on Neotropical Tabanidae XIX. The Tabanus

lineola complex. Miscell. Publ Ent. Soe. Amer. No. 57, pp. 1-51, pls. I-VI.

FAIRCHILD, G. B. 1984. Notes on Neotropical Tabanidae XX The larger species

of Tabanus of eastern S. America. Cont. Amer. Ent. Inst. 21(3): 1-50, figs 1-

29, maps 1-12.

FAIRCHILD, G. B. Notes on Neotropical Tabanidae XVIII The genus Leuco-

tabanus Lutz. Myia 3: in press.

FAIRCHILD, G. B. and T. H. G. AITKEN 1960. Additions to the Tabanidae of

Trinidad, B.W.I. Ann. Ent. Soc. Amer. 53(1): 1-8.

FAIRCHILD, G. B. and R. C. WILKERSON 1981. New species of Esenbeckia

(Proboscoides) with a key to the subgenus. Florida Ent. 64(1): 158-175, figs

1-9.

GOODWIN, J. T. and W. P. MURDOCH 1974. A study of some immature

Neotropical Tabanidae Ann. Ent. Soc. Amer. 67(1): 85-133.

HOGUE, C. L. and G. B. FAIRCHILD 1974. A revised check list of the Taban-

idae of Costa Rica. Rev. Biol. Trop. 22(1): 11-27.

KROEBER, O. 1930. Die Sudamerikanischen Arten der Gattung Scione WIk.

(=Rhinotriclista End.). Stett. Ent. Zeit. 91(2): 140-174.

LEE, V., G. B. FAIRCHILD and Pablo BARRETO 1969. Artropodos hematofagos

del Rio Raposo, Valle, Colombia III. Tabanidae. Caldasia 10(49); 441-458.

MEDEM, F. 1981. Horse flies (Diptera: Tabanidae) as ectoparasites on caimans

(Crocodylia: Alligatoridae) in eastern Colombia. Cespedesia 10(37-38): 123-

147. 2 figs. |

PHILIP, C. B. 1954. New North American Tabanidae VIII. Notes on and keys to

the genera and species of Pangoniinae exclusive of Chrysops. Rev. Brazil.

Ent., S. Paulo 2: 13-60.

PHILIP, C. B. 1960. Further records of Neotropical Tabanidae mostly from

Peru. Proc. Calif. Acad. Sei. 31(3): 69-102.

PHILIP, C. B. 1965. The identity and relationships of Tabanus (Neotabanus) vit-

tiger and notes on two cases of teratology in Tabanidae. Ann. Ent. Soc.

Amer. 58(6): 876-880.

PHILIP C. B., in Stone, Sabrosky, Wirth, Foote and Coulson, 1965. A catalogue

of the Diptera of America north of Mexico. U.S. Dept. Agriculture Handbook

No. 276.

PHILIP, C. B. 1978. New North American Tabanidae XXIV. Further comments

on certain Pangoniinae in Mexico, with special reference to Esenbeckia.

Proc. Calif. Acad. Sei. Ser. 4, 41(4): 345-356.

RICHARDSON, C. G. and B. H. WILSON 1969. Daily flight activity of male

Tabanus lineola var. hinellus Philip in the estuarine area of Louisiana. J.

Med. Ent. 6(3): 276-277.

STONE, A. 1938. The horse flies of the subfamily Tabanidae of the Nearctic

region. U.S. Dept. Agric. Miseell. Pub. No. 305, pp. 1-172.

WILKERSON, R. C. 1979. Horse flies (Dipt. Tabanidae) of the Colombian de-

partments of Choco, Valle and Cauca. Cespedesia (Cali, Colombia) 8(31-32):

87-435. figs. 1-119. ;

132 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

WILKERSON, R. C. and G. B. FAIRCHILD 1982. Five new species of Diachlorus

(Diptera: Tabanidae) from South America with a revised key to species and

new locality records. Proc. Ent. Soe. Washington 84(3): 636-650, figs. 105.

WILKERSON, R. C. and G. B. FAIRCHILD 1983. A review of the South Ameri-

ean species of Esenbeckia subgenus Esenbeckia (Dipt. Tabanidae). Journ. Nat.

Hist. 1983, 17: 519-567. figs. 1-18.

WILLIAMS, P. 1971. Some records of Tabanidae from British Honduras (Belize)

J. Med. Ent. 8(1): 98-109.

WOKE, P. A. Arthropods of sanitary importance in the Republic of Nicaragua.

Amer. J. Trop. Med. 27(3): 357-375.

EXPLANATION OF FIGURE 1

Fig 1A-J General horsefly morphology.

Fig. 1A. Head, Fidena auribarba.

Fig. 1B. Head, Chrysops variegata.

Fig. 1C. Head, side view, Dasychela badia. 7

Fig. 1D. Apex of hind tibia of Fidena flavipennis showing paired tibial spurs.

Fig. 1E. Base of wing of Fidena flavipennis showing bare basicosta and setose

base of costal vein.

Fig. 1F. Base of wing of Tabanus nebulosus showing setose basicosta and base

of costal vein.

Fig. 1G. Wing of Scione costaricana with the cells labeled according to Curran.

Note the closed and petiolate first and fourth posterior cells and anal

cell.

Fig. 1H. Wing of Tabanus pseudoculus with the veins named according to

Comstock-Needham. Note the appendix of fork of third vein and

ecoarctate first posterior cell.

Fig. ll. Side view of thorax of Poeciloderas quadripunctatus.

Fig. lJ. Dorsal view of thorax and first 2 abdominal segments of Tabanus

nebulosus.

G. B. Fairchild: Tabanidae of Panama 133

Ocelli

Subcallus Frontal callus

any

—_——~

a. PAE Ra oe

ge SSS SA

BM Ooo ea: hg A oo Salto

pe ee aoe

AZ" FASE

I rc

ae Ce

po ae

So Si LS Se

Vertex sy ee eee

Frontal callus

Antenna

Gena

Costal cell Subcostal Stigma

Mesocatepisternum

| Mesoanepimeron

Pronotal lobe

2nd Submarginal

1st Posterior

2nd Posterior

3rd Posterior Notopleural lobe

4th Posterior

5th Posterior

Transverse

suture

Mesonotum

Costa

Subcosta Wing base

Saas = es ee eer oon

Ew cdi ( rs

Appendix e

Longitudinal

Fork suture

M 4 Rs

Scutellum

wa oo Haltere

Abdominal tergite |

Cu, M3

Cu2 + 1st A

Abdominal tergite Il

22, no. 3, 1986

Contrib. Amer. Ent. Inst., vol

134

MAP OF PANAMA

(Courtesy Margo Duncan)

"eo ee

eS had

SI9IJOW

NV300 OldIOWd

VAS Nvaagalavo

G. B. Fairchild: Tabanidae of Panama 135

TAXONOMIC INDEX

Names thought to be valid are in boldface

page numbers of

references to Panama species are in boldface. ; ee

ablusus, Scione 8, 31, 34

adiastolus, ssp of praeteritus,

Tabanus 14, 108, 121

Aegialomyia 20, 50

albibarba, var of auribarba,

Fidena 27

albicans, Tabanus 100

albisecutellatus, Leucotabanus 99

albocirculus, Tabanus 10, 14,

104, 109, 118, 122

albovillosa, Hybostraba 122

alleni, Chrysops 8, 12, 37, 39

alteripennis, Philipotabanus 88

amabilinis, Poeciloderas var 103

ambiguus, Leucotabanus 99, 100

amplifrons, Tabanus 11, 110

analis, Dichelacera 11, 64

analis, Fidena 30

ananasi, Stenotabanus 51

angustivitta, Tabanus 11, 123

aniptus, Tabanus 106, 110

annulatus, Leucotabanus 101

antarcticus, Tabanus 109

apicalis, Tabanus 110

apicimacula, Stibasoma 6, 9,

12,40, 11

aquilus, var of scapularis,

Dichelacera 62, 64, 68

Assipala 18, 35

atopus, Phaeotabanus 60

atratus, Tabanus 113

aurarius, Leucotabanus 98, 99

aureopygia, Seione 33

auribarba var albibarba,

Fidena 27

auribarba, Fidena 8, 12, 26, 27

auricauda, Fidena 29, 30

auroguttata var pallidefemorata,

Chrysops 39

auroguttatus, Chrysops 8, 12,

38, 39

aurulans, Scione 31

badia, Dasychela 9, 12, 74

basilaris, Fidena 30

bellardii, Tabanus 127

bicolor, Bolbodimyia 57

bifenestrata, Stibasoma 77

bigoti, Tabanus 10, 14, 106,

110, 115, 126

blantoni, Stenotabanus 50, 51

Bolbodimyia 19, 56, 61

Brachytabanus 20, 52

brevipalpis, Seione 32, 33

ealiginosus, Tabanus 88

ecalogaster var mexicana,

Chrysops 42

ealogaster, Chrysops 8, 12, 38,

40, 44

ealvitius, Stenotabanus 46, 47

eanithorax, Leucotabanus 10,

99, 100

captiroptera, Stypommisa 13,

earneus, var of lineola,

Tabanus 117

Catachlorops 19, 70

chagresensis, Esenbeckia 11

chagresensis, ssp of ecuadorensis,

Esenbeckia 11, 22, 23

changena, Stypommisa 93, 94, 96

changuinolae, Stenotabanus 9,

13, 50, 51

chionostigmaa, Stibasoma 9, 76, 77

chiriquensis, Chrysops 8, 36, 40

Chlorotabanus 20, 58, 82

chloroticus, Cryptotylus 7, 9,

Ll, £43 O2

Chrysops 5, 6, 7, 11, 18, 36

Chrysopsinae 17, 35

Chrysopsini 395

ehrysothrix, Philipotabanus 7,

13, 85, 89

claripennis, Scione 32, 33

colombensis, Tabanus 10, 11,

14, 107, 110, 117, 124, 127

columbianus, Phaeotabanus 91

ecolumbiensis, Melpia 27

commixtus, Tabanus 10, 11, 14,

107, 111, 112

compactus, Stibasoma 78

eonstabulorum, Stenotabanus 49

eostaricana, Seione 8, 12, 31, 32

crassipes, Lepiselaga 10, 13, 101

criton, Philipotabanus 7, 9, 91

eas) Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

crocata, Dichelacera 6, 9, 12,

63, 64

Cryptotylus 18, 81

curvipes, Diachlorus 9, 13,

94, 55

Dasychela 18, 74

defilippii, Tabanus 6, 10, 14,

104, 112, 119, 123

Desmatochelacera 19, 69

Diachlorini 18, 45

Diachlorus 5, 20, 54

Dichelacera 19, 61

Dicladocera 19, 75

dorsiger ssp modestus,

Tabanus 118

dorsiger, Tabanus 117

dorsovittatus, Tabanus 124

dorsovittatus, ssp of occidentalis,

Tabanus 10, 117

dunni, Tabanus 107, 112

ebeneus, ssp of morbosus,

Tabanus 105,113, 115

ebrius, Philipotabanus 7, 9,

13, 85

ecuadorensis chagresensis,

Esenbeckia 8, 11, 23

elviae, Philipotabanus 9, 84, 86

enanus, Tabanus 10, 14, 106, 113

erebus, Tabanus 10, 14, 105, 113

eriomeroides, Fidena 6, 8, 26, 27

erythrocephala, Bolbodimyia 7,

26, 57

Esenbeckia 4, 5, 10, 11, 17, 22

exaestuans, Leucotabanus 10,

11, 13, 99, 100

fairchildi, Chrysops 43

fasciata, Dichelacera 9, 11,

12, 62, 63, 64, 67, 68

fascipennis, Hemichrysops 55

fascipennis, Philipotabanus 6,

10, 91

ferrifer, Tabanus 11, 115

festae, Pityocera 7, 8, 11, 35

festivus, Tabanus 71

Fidena 11, 18, 25, 31

flavinotum, Leucotabanus 13,

98, 100

flavipennis, Fidena 8, 12, 26, 28

flavistigma, Stibasoma 77

flaviventre, Stibasoma 6, 9,

76, 78

formosus, Tabanus 86

fortunensis, Catachlorops 71, 72

frondicolus, Stenotabanus 96

fulminea var ocellatus,

Catachlorops 11, 71

fulmineus, Catachlorops 9, 11,

Zi, ta, (3

fulvistriatus, Stenotabanus 47

fulvohirtum, Stibasoma 6, 9,

1 ty Te

fulvosericea, Fidena 33

fumatipennis, Tabanus 11, 119

fumomarginatus, Tabanus 119,

120, 128

fureata, Bellardia 71

galindoi, Bolbodimyia 56, 57

generosa, Rhinotriclista 33

grassator, Philipotabanus 6,

9, 84, 86

guatemalanus, ssp of subsimilis,

Tabanus 127

guatemalanus, ssp of vittiger,

Tabanus 10, 14, 107, 116, 127

hartmanni, Dichelacera 63, 65, 67

Hemichrysops 20, 55, 83

Himantostylus 20, 53

howardi, Fidena 8, 27, 28

hypographa, Stypommisa 93, 95

Idiochelacera 19, 69

illota osornoi, Esenbeckia 25

illota, Esenbeckia 11

importunus, Tabanus 10, 14,

105, 113, 115, 119

inanis, Chlorotabanus 7, 9, 13,

08, 59

Inauratus, Philipotabanus 6,

9, 89

incipiens, Stenotabanus 13,

46, 48

incisa, Chrysops 39

intermedius, Himantostylus 13,

53, 87

intermedius, Tabanus 87

isthmiae, Fidena 28

jaculator, Stypommisa 6, 13,

93, 95

jobbinsi, Diachlorus 9, 13,

04, 55

keenani, Tabanus 91

lacajaensis, Tabanus 10, 14,

106, 114, 128

latifasciatus, Chrysops 41

Lepiselaga 20, 53, 101

lerida, Stypommisa 13, 93, 94, 96

leucaspis, Leucotabanus 11, 100

G. B. Fairchild: Tabanidae of Panama 137

leuconotum, Leucotabanus 100

leucospilus, Chrysops 37, 41

Leucotabanus 21, 92, 98

limonus, Cryptotylus 11, 82

lineola var carneus, Tabanus

LU, 157

lineola var plangens, Tabanus 117

lineola var stenocephalus,

Tabanus 11, 125

lineola, Tabanus 7, 117

littoreus, Stenotabanus 9, 51

longiappendiculatus, Phaeotabanus

7, 9, 11, 13, 60

longipennis, Stenotabanus 9, 53

lophus, Tabanus 125

luteoflavus, Cryptotylus 11, 60

macquarti, Tabanus 10, 105,

114, 126

maculifrons, Stenotabanus 48

maculipennis, Scione 8, 12,

31, 33

magnificus, Philipotabanus 6,

(1, 9, 13, 83, 84, 86, 89

marginata, Dichelacera 9, 12,

62, 64, 65, 66, 68

maruccli, Stypommisa 6, 10,

13, 92, 95, 96

maya, Tabanus 11, 111

medius, Philipotabanus 9, 13,

84, 87

medius, Tabanus 11, 88

melaenus, Chrysops 8, 12, 37,

41, 44

melanosoma, Dichelacera 62,

63, 66

Melasmatabanus 21, 83, 90

mexicanus, Chlorotabanus 7, 9,

13, 58, 59

mexicanus, Chrysops 12, 38, 42

Mimotabanus 21, 83, 89

minuscula, Stenotabanus 13, 46,

47, 48

modestus, var of occidentalis,

Tabanus 10, 117, 118

morbosus ebeneus, Tabanus 105,

113, 195

nebulosus, Tabanus 10, 11, 14,

105, 114, 115, 119

neofurva, Stypommisa 95

nereus, Tabanus 10, 14, 103,

108, 116, 125

nexosus, Chrysops 12, 38, 42

nigricans, Fidena 30

nigriflavus, Tabanus 100

nigrinubilus, Philipotabanus 9,

13, 83, 84, 87

nigriventris, Leucotabanus 98, 99

nigrocoerulea, Dicladocera 75

nondescriptus, Tabanus 10, 14,

103, 107, 116, 125

obscurus, Stenotabanus 8, 47, 49

occidentalis dorsovittatus,

Tabanus 10

occidentalis modestus, Tabanus 10

occidentalis var dorsovittatus,

Tabanus 11, 14, 108, 117, 118,

126,127

occidentalis var modestus,

Tabanus 14, 108, 117, 119, 127

ocellata, Catachlorops 12, 71, 72, 73

ocellata, ssp of fulminea,

Catachlorops 9

oculus, Tabanus 10, 104, 118, 122 ©

olivaceiventris, Tabanus 104, 118

osornoi osornoi, Esenbeckia 11, 23,

osornoi, Esenbeckia 7, 8, 11

pachycephalum, Stibasoma 77

paitillensis, Stenotabanus 9,

51, 32

pallidefemorata, as var of

auroguttata, Chrysops 39

pallidetinetus, Philipotabanus

ts Oy1116:13):85, 87; 88

panamensis, Stibasoma 6, 9,

12, ‘76y:%9

panamensis, as ssp theotaenia,

Stibasoma 79, 80

Pangoniini 17

Pangoninae 17

pequeniensis, Stypommisa 13,

pequiensis, Stenotabanus 97

peruviana, Dichelacera 75

phaeopterus, Phaeotabanus 7,

60, 61

Phaeotabanus 20, 60

phalaropygus, Philipotabanus 6,

9, 89, 90

philipi, Bolbodimyia 9, 56, 57

Philipotabanus 21, 83, 84

piraticus, Tabanus 128

Pityocera 11, 17, 34

plangens, as var of lineola,

Tabanus 117

planiventris, Stibasoma 80, 81

138 Contrib. Amer. Ent. Inst., vol. 22, no. 3, 1986

platyeerus, Tabanus 10, 11, 14,

106, 110, 119

plenus, Philipotabanus 11, 89, 90

Poeciloderas 21, 102

polyphemus, Tabanus 6, 10, 104,

112, 119

pompholyx, Stenotabanus 50, 52

praepilatus, Tabanus 14, 108, 120

praeteritus adiastolus, Tabanus 14,

108, 121

praeteritus, Tabanus 10, 108,

120, 121

prasiniventris, Esenbeckia 8, 11,

23, 24

prinecessa, Dichelacera 62, 64,

65, 66

Proboscoides 23

pruinosus, Tabanus 108, 112, 122

prunicolor, Stypommisa 96

Psalidia 19, 70

pseudoculus, Tabanus 10, 14,

104, 118, 122, 125

pterographicus, Philipotabanus 7,

9, 13, 85, 88

punctipleura, Tabanus 10, 14,

104, 119, 123

pungens, Tabanus 10, 11, 14, 103,

107, 120, 123

pyrausta, Fidena 29, 30

quadripunctatus, Poeciloderas 10,

11,23, 102, 103

quinquemaculatus, Tabanus 124

quinquepunctatus, Tabanus 7,

106, 124

regina, Dichelacera 6, 9, 12,

63, 66, 67, 68

reticulatus, Chrysops 37, 43, 44

rex, Dichelacera 9, 12, 62, 63,

65, 67

Rhabdotylus 18, 75, 76

rhinophora Fidena 7, 8, 12, 26, 29,

30,

rhizophorae, Tabanus 10, 103,

108, 124

Ricardoa 22, 23

rixator, Tabanus 106, 116, 125

rossi, Chrysops 43

roxannae, Stenotabanus 47

rubripes, Tabanus 105, 125

rufescens, Scione 7, 8, 12, 31, 33

rufipes, Scione 34

scalaratus, Chrysops 8, 12, 37, 43

scapularis var aquilus, Dichelacera

62, 64, 68

scapularis, Dichelacera 9, 12,

62, 64, 65, 67

schildi, Fidena 7, 8, 12, 26, 30

Seione 11, 18, 31

Scionini 17

scurrus, Catachlorops 71, 73

Selasoma 19, 53, 58

serena, Stypommisa 92, 97

Silvius 18, 35

sordidatus, Stenotabanus 47, 49

soror, Chrysops 8, 12, 38, 40, 43

stenocephalus, Tabanus 10, 11,

14, 108, 118, 125, 126

Stenotabanus (Aegialomyia) 5

Stenotabanus 5, 20, 46, 92

Stibasoma 6, 18, 75

Stietotabanus 92

stigmaticalis, Philipotabanus 86

stilbium, Stibasoma 78

Stilbops 47

strigata, Scione 32, 33

Stypommisa 21, 92

subeaecutiens, Chrysops 40

subeallosa, Dichelacera 13, 69

subguttata, Esenbeckia 22, 24

submarginata, Dichelacera 6, 9,

13, 62, 64, 66, 67, 68

subsimilis guatemalanus, Tabanus

127

surifer, Tabanus 10, 14, 105,

110, 115, 126

surifer, as ssp of thiemeana,

Tabanus 126

Tabaninae 17, 45, 102

Tabanini 18, 102

Tabanus 5, 7, 21,102, 103

tanycerus, Silvius 36

tardus, Chrysops 44

tardus, var of varians,

Chrysops 38, 44

theotaenia panamensis, Stibasoma

79, 80

thiemeana surifer, Tabanus 126

thiemeanus, Tabanus 126, 127

tibiale, Selasoma 7, 9, 58

translucens, Esenbeckia 7, 8,

11, 22, 24

transposita, Dichelacera 9, 70

trapidoi, Fidena 6, 8, 12, 26,

29, 30

trivittatus, Tabanus 110

G. B. Fairchild: Tabanidae of Panama

truquiil, Tabanus 110, 111, 127

umbraticolus, Tabanus 114

umbratus, Catachlorops 6, 9,

12, %0,.138

unicolor, Cryptotylus 7, 9, 12,

82, 83 |

unipunctatus, Tabanus 7, 10,

14, 104, 112, 128

unistriatus, Tabanus 10, 14,

106, 113, 128

vapidus, Stenotabanus 97

vargus, ssp of varians,

Chrysops 38, 44

varians vargus, Chrysops

38, 44

varians var tardus, Chrysops

38, 44, 45

varians, Chrysops 44

variegatus, Chrysops 8, 12,

37, 45

venenata, Stibasoma 74, 76, 80

viridiventris, Stibasoma 80,81

vittiger caymanicus, Tabanus 127

vittiger guatemalanus, Tabanus

10, 14, 107, 116, 127

xenium, Stenotabanus 93

xenorhynchus, Tabanus 10, 6,

105, 129

xipe, Tabanus 109

139

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Contributions

of the

American Entomological Institute

Volume 22, Number 4, 1985

AN ATLAS OF WING PHOTOGRAPHS,

WITH A SUMMARY OF THE NUMERICAL CHARACTERS

OF THE NEARCTIC SPECIES OF CULICOIDES

(DIPTERA: CERATOPOGONIDAE)

W.W. Wirth, A.L. Dyce and B.V. Peterson

with Photography by I. Roper

AN ATLAS OF WING PHOTOGRAPHS, WITH A SUMMARY OF THE NUMERICAL

CHARACTERS OF THE NEARCTIC SPECIES OF CULICOIDES

(DIPTERA: CERATOPOGONIDAE)

W.W. Wirth!, A.L. Dyce2, and B.V. Peterson!

with Photography by I. Roper?

ABSTRACT

An atlas of 150 wing photographs of females of the described Nearctic species of

Culicoides is presented. There is no one taxonomic key now available that will serve for

the identification of all Nearctic species of Culicoides. This atlas will help in large part

to fill this void because the specific differences in wing patterns are, with few exceptions,

easy to distinguish and are useful field characters. The atlas will guide the user to a small

group of related species that can then be distinguished by one of the regional keys. A

table of numerical (morphometric) characters is also presented for the species. These

characters are useful in confirming an identification made on the basis of the wing

patterns shown. The 144 valid species (plus 4 subspecies) listed are distributed in ll

subgenera and 8 subgenerically unplaced species groups.

CONTENTS

JSST CICS: COIN aA Re tN a Oo aI Os rma ae esa Pace oT et a ene ee 2

TS AS IPTG AN Ch setae ce tai eee tay Bs roteid vals tesa wig ic eg Ue ated a En San Le Sg T Same aa seis Gala a ae bids long euiboeveusnena 2

Oma uecMeaehO cu (nell Gccdesacrenctacic merece cca wel Cu era cercieaae ula carekioaecculae nen Wahi som sinaculce ares uatecmmas 3

RAG NOG Saree noe ca ei am he. nan Whe kiiai rare Wom aceauakt. af tM an eMC abs ulate talents 4

NCAA OMS CINE AES eriadu cis Coat ois sa sea tens sateen ae chs wntisloee Hemea ere Wine Slvlnaielninseleinv ate Wale iaiclnn wae celoaennaemeweaune’ 4

TablevOraNGimericdn’ CHALaClOIS. - cee ook «ose seadcdecsere wens coeisudcadueUenasduedena sce cehonensescwseeestiene te 6

Palla Noa) fancy ea Fea Le eal reese ah cer a tema ees ec Mc ed sl a ea ciaiatarc Uieea RN Ws cious Bb atalalins d cit wise dalblad aber s eae eames 10

LEER AS WIRY eta 0 SF Eat a em PRS ca Nar eI pray Ren Sa Res LES a ee ae dE 40

Re De Geter stata ste eta get e pemietis otine ae dtues Cae atenmad cris clbraiNarg digesta, dure costa Halk Sidle-ccd-craclaaa-cielaaeie en okoaw ean mecnak 45

INTRODUCTION

Background

The biting midges of the genus Culicoides Latreille are notorious bloodsucking pests

of man and animals throughout the world. Because of their small size and crepuscular

habits they are not frequently collected and consequently are poorly known. In recent

years their incrimination as vectors of human and animal diseases has focused increasing

attention to their collection and study, and, at the same time, has caused a growing need

for manuals and keys for their identification. At present, there is no one taxonomic key

that will serve for the determination of all the Nearctic species, or even those of the

western states and provinces. The first and last authors of this paper are presently

l. Systematic Entomology Laboratory, IIBIII, Agricultural Research Service, USDA, c/o

National Museum of Natural History, NHB-168, Washington, D.C. 20560, U.S.A.

2. Division of Tropical Animal Science, Entomology Research Station, CSIRO, 55

Hastings Road, Warrawee, New South Wales, 2074, Australia.

3. Division of Animal Health, CSIRO, McMaster Laboratory, Private Bag No. 1, P.O.

Glebe, New South Wales, 2037, Australia.

Wirth, Dyce and Peterson: Nearctic Culicoides 5

working on a manual to the Culicoides of North America, but it will be some time before

the manual can be completed. In the meantime, we believe that this atlas of wing

photographs will help in large part to fill the void because the specific differences in wing

patterns are, with few exceptions, easy to distinguish and thus are useful in the field. In

most cases, this photographic atlas will guide the user to a small group of related species

that can then be distinguished by using one of the taxonomic keys listed below.

Microscopic slide preparations must be made of specimens in order to view and measure

the characters necessary for identification. Table | presents the numerical characters of

each species that may be useful in confirming an identification made on the basis of the

wing patterns or by use of supplementary keys.

It is not within the scope or purpose of this guide to present a comprehensive review

of the economic importance, morphology, biology, systematics, or distribution of the

Nearctic species. It is intended that the forthcoming manual will serve such a purpose.

One or more of the following references should be consulted for general information,

details of morphology, and guides to species of certain limited areas. $

General reviews of the morphology, biology, and classification of the family

Ceratopogonidae have been published by Edwards (1926), Wirth (1952), and Downes and

Wirth (1981). Keys to the genera of Ceratopogonidae were presented by Macfie (1940),

Johannsen (1943), and Wirth et al. (1974, 1977). Good reviews of the biology and economic

importance of the bloodsucking species of Culicoides were given by Downes (1958b),

Kettle (1962, 1965, 1969), and Blanton and Wirth (1979). Atchley et al. (1981) published a

complete world-wide bibliography of the family Ceratopogonidae with a keyword index.

Fox (1955) published a catalog of the Culicoides species of the Americas, and Wirth (1965)

cataloged the North American Culicoides. Arnaud (1956) listed the species of the world.

Jones (196la) published a comprehensive review of the larval habitats of the North

American species, and later (196lb) described the pupae of a number of these species.

Khalaf (1954) and Vargas (1960, 1973) published keys and descriptions to the subgenera of

North American Culicoides.

For lists, reviews, and keys to the Culicoides species occurring in various states or

other limited areas, the following publications are available: Malloch (1915) gave an early

review of the species occurring in Illinois, and Hoffman (1925) reviewed the species known

in North America at that time. Foote and Pratt (1954) reviewed the species occurring in

the eastern United States, while Wirth and Bottimer (1956) made a population study of the

species of the Edwards Plateau in Texas. Khalaf (1957) made a light trap survey of the

species of Oklahoma, and, in 1969, reported on the seasonal incidence of species in the

gulf coast of Louisiana and Mississippi. Downes (1958a) discussed the species occurring in

Canada. In 1965, Jamnback published a comprehensive report on the taxonomy, biology,

and immature stages of the species of New York, that is useful for the entire northeastern

region. Atchley (1967) made a taxonomic study of the species of New Mexico that is

generally useful for the southwest, and Jorgensen (1969) treated the species occurring in

southeastern Washington. More limited state reports were published by Childers and

Wingo (1968) for Missouri, and by Gazeau and Messersmith (1970) for Maryland. Battle and

Turner (1971) reviewed the Virginia species, provided a catalog and listed the known

distributions for the species of the eastern U.S.A. The most comprehensive and

up-to-date revision was published by Blanton and Wirth (1979) who provided a review of

the morphology, biology and classification of the North American Culicoides, and

presented a key to the species of Florida.

Format and Treatment tig

For each of the 144 valid species and 4 subspecies listed in the atlas, the following

information is provided: the current name of the species, its author(s), the original

citation including the year of publication and the page number on which the species is

named and described, and the state or province from which the species was first

described. Where the species was validated in a genus other than Culicoides the name of

the genus in which it originally was placed is noted. The general distribution for each

4 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

species is given at the state and province level, and this is followed by the actual locality

and collector of the specimen illustrated. In two instances, several photographs are

provided for species, viz variipennis (Coquillett) (3 photographs), and crepuscularis

Malloch (5 photographs), that are represented by several subspecies, or of species that are

somewhat variable and of uncertain status or consist of unresolved species complexes.

In the index are listed all the species names attributed to Culicoides in the literature

on the Nearctic fauna. Thirteen specific names are listed in the index that do not appear

in the atlas proper as valid species. The disposition of these names is as follows:

l. borinqueni Fox: not Nearctic.

2. canithorax Root and Hoffman: junior synonym of hollensis (Melander

and Brues).

. dickei Jones: junior synonym of furensoides Williams.

. dovei Hall: junior synonym of furens (Poey).

. hornae Foote and Pratt: junior synonym of travisi Vargas.

. inamollae Fox and Hoffman: junior synonym of insignis Lutz.

. khalafi Beck: junior synonym of debilipalpis lutz.

. minutissimus (Zetterstedt): not Nearctic.

. montanus Wirth and Blanton: junior synonym of neomontanus Wirth.

. pifanoi Ortiz: not Nearctic.

pseudopiliferus Wirth and Hubert: junior synonym of alexanderi Wirth

and Hubert.

. simulans Root and Hoffman: junior synonym of travisi Vargas.

. weesei Khalaf: junior synonym of stonei James.

—“OwoOOwWNKDUBW

— —

W BRO

Methods

The measurements and ratios of the female characters given in Table | are derived as

follows: The wing length is measured from the basal arculus to the tip of the wing; the

costal ratio is the length of the costa measured from the basal arculus to the tip of the

second radial cell divided by the wing length. The antennal ratio is the combined lengths

of the distal 5 segments divided by the combined lengths of the preceeding 8 segments of

the flagellum; the antennal sensillar pattern refers to the presence of sensilla coeloconica

that occur distally on some or all of segments 3 to 15. The palpal ratio is the greatest

length of the third palpal segment divided by its greatest width; the proboscis/head (P/H)

ratio is obtained by dividing the length of the proboscis from the torma to the tip of the

labrum-epipharynx by the distance from the interocular hair socket to the torma. Figures

of these morphological characters are available in Blanton and Wirth (1979), and Downes

and Wirth (1981).

The wing photographs were prepared from slide mounted specimens using a Wild

MKaz2 camera, a 2.5 x 3.5 in. film holder and a MKa5 Photoautomat exposure meter in

conjunction with a Wild M20 photomicroscope equipped with a 10x wide angle ocular and

3x, 6x and 10x objectives with a matching lowpower bright field condensor. The lighting

system consisted of a Wild Universal lamp-housing having a 12V, 1OOW quartz-iodine light

source controlled by a Wild regulating transformer. Natural and/or heat filters were used

with some specimens. Kodalith Ortho film, type 3, Kodagraph liquid developer D-76 and

Ilford P.Q. Universal developer gave good results. Exposure time was critical and was

determined by trial and error in accordance with the thickness and color of the mounting

medium in which the wings were embedded. Exposures always were made at 12V. The

development time was 2-3 minutes at 68°F according to instructions with the film.

Negatives were printed on Contrast Grade 0 or 1 glossy R.C. Kodak or Ilford paper.

Acknowledgments

We are greatly indebted to Robert H. Jones and Wayne L. Kramer of the

Arthropod-borne Animal Disease Research Laboratory, Agricultural Research Service,

U.5.D.A., Denver, Colorado, and to William L. Grogan, Jr. of Salisbury State College,

Salisbury, Maryland, for reviewing the manuscript and for supplying information on the

Wirth, Dyce and Peterson: Nearctic Culicoides 5

distribution of the species. We are grateful to Steve Nakahara, Systematic Entomology

Laboratory, IIBIII, Agricultural Research Service, U.S.D.A., Beltsville, Maryland, for

helpful comments on the manuscript. We also thank Henry K. Townes of the American

Entomological Institute, Gainesville, Florida, for helpful suggestions in setting up the

format of this publication. The senior author is grateful to Harold A. Denmark and

Howard V. Weems, Jr. of the Florida Department of Agriculture and Consumer Services,

Division of Plant Industry, Gainesville, Florida, for providing research facilities at the

Florida State Collection of Arthropods.

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Table 1

Systematic Arrangement and Numerical Characters of North American Culicoides

Species

cockerellil

freeborni

frohnei

gregsoni

lahontan

neofagineus

neomontanus

saltonensis

sierrensis

sommermanae

sordidellus

tristriatulus

canadensis

neopulicaris

yukonensis

insignis

venustus

alachua

chiopterus

hirtulus

juddi

obsoletus

pechumani

pusillus

sanquisuga

arizonensis

butleri

byersi

cacticola

copiosus

hinmani

insolatus

jamaicensis

Wing

length

(mm)

IDRIS oe Seco ate lo weet hol Picea ere

Rotnin ODWOu~yonue®

WNOWUW O77" WO ~

NO UIW

eof >

0.96

1.10

0.89

1.16

1.02

0.73

1.19

0.97

Costal Antennal

Ratio

Antennal

Sensillar

Pattern

Subgenus Culicoides

cockerellii Species Group

1.03

0.77

1.09

ish2

0.98

0.87

0.94

L053

1.19

E22

pulicaris Species Group

0.62

0.61

G559

OCooooogoom

Un OQ und uo

Owwyonr- 00

1.10

0.96

1.20

1.10

Subgenus Avaritia

— i os —

Mae, ia

Subgenus Drymodesmyia

ge ae

— — * WN

Now O

wo VM

NNWNHO

iB a 9, 11-15

Ne)

Cae

—

Oo :

(6-7), 11-15

Palpal

Ratio

Sr ghee eee

NNNONNOR-UNN A

WN MM hh

oaMa™~~j)~Con

ee a a On

NOWON™~ NW

P/H

Ratio

OOUIWOR DOH

UWNTIONOOS

Species

jonesi

loughnani

ryckmani

sitiens

torridus

baueri

bergi

blantoni

defoliarti

edeni

erikae

footei

haematopotus

inyoensis

daedalus

luglani

pampoikilus

arboricola

beckae

californiensis

cochisensis

flukei

guttipennis

oklahomensis

ousairani

pecosensis

villosipennis

alexanderi

bickleyi

cavaticus

chewaclae

denticulatus

doeringae

downesi

franclemonti

husseyi

jamnbacki

kirbyi

lophortygis

parapiliferus

Wirth, Dyce and Peterson: Nearctic Culicoides

Wing Costal Antennal Antennal

Length Ratio Ratio Sensory

(mm) Pattern

bi5 O52 LoS 3-15

eZ] 0.55 1.17 3-15

: 05> 1.19 3,(11-12),13-15

A? 0.53 Lb 3-15

22 0.51 E2t 3-9,11-15

Subgenus Diphaomyia

1.03 0.56 Be2 3,7-10

1.20 0.57 1.19 3,(4),5-10

1.02 0.56 et 3,8-10

1.16 0.54 E27 3,10-15

1.12 0.58 1.78 3,10-15

1.25 0.54 fea 3,8,10-15

0.90 0.61 1.14 3-10

1.15 0.60 1.70 3-15

1.24 0.56 @:99 3-10

Subgenus Unplaced, daedalus Species Group

1.01 0.62 1.34 3,11-15

i22 0.33 Re 2) 3-10

ey 0.54 os 5,9; 1,7, t1-15

Subgenus Unplaced, guttipennis Species Group

1.19 0.60 1.48 3,5,7,9, 11-15

1.15 0.61 eye) 3-15

1.28 0.52 EE? 3~9,]1-15

1.14 0.52 eB) 3-9, 11-15

0279 0.57 1.02 3,11-15

[52 0.60 1.54 3,5,7,9, 11-15

1.06 0.55 ES) 5,9, 7,7 lelS

LZ 0.57 1.00 3-1]

L209 0.57 1.15 3,5,7-9, 1-15

1.62 0.61 1.64 3,5,/7,9, 1-15

Subgenus Unplaced, piliferus Species Group

ae 0.59 1.08 3,5,7,9,11,13-15

1.07 0.60 1.20 3,13-15

1.60 0.56 1.00 3-15

0°95 0.61 1.36 5,547,9,10-15

1.4] 0.62 1.13 3,11-15

1.24 0.58 1.63 3,967, 9,11,13-15

1.58 0.61 1.67 J vii arlelS

L19 0.62 1.16 3,11-15

1.20 0.58 1.42 3,5,7,9, 11-15

1.14 0.60 1.23 be yy see wl

0.90 0.57 1.31 5,53 fe tellld-tS

1.16 0.58 1.50 3,5,7,9, 11-15

1.15 0.59 1.22 555A 13-15

Palpal

Ratio

acl al gh nag

CB) RY (GS) dS bes

Nay mee awa SY he Ia

O-o ONAWNW Oo

ee ee a RSS

ERUMNOMUA UD

ee tS FO FS 9 IS OPO" IY

UoDWnpwWnHoatnee—-NWe

Species

piliferus

riqgsi

scanloni

snowl

testudinalis

unicolor

utowana

calexicanus

davisi

hawsli

leechi

novamexicanus

oregonensis

palmerae

utahensis

wirthi

debilipalpis

eadsi

paraensis

reevesi

torreyae

barbosai

furens

furensoides

stellifer

gigas

variipennis s.str.

ssp. albertensis

ssp. australis

ssp. occidentalis

ssp. sonorensis

alaskensis

bermudensis

crepuscularis

hollensis

knowltoni

mississippiensis

sphagqnumensis

wisconsinensis

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Wing

Length

(mm)

el2

0.90

1.04

1.10

Subgenus Unplaced, palmerae Species Group

SU sie Oe eoel toc vote teh a sm ton

NESS UN AWWW

ba SP diss (00) Cy bad tio} dss

0 Oy

ae)

NO f&NMWHW

ONOWMOO

Costal Antennal Antennal

Ratio Ratio Sensillar

Pattern

0.62 1.87 3,5,/7,9-15

0.61 1.14 3,5,7,9,(1l),13-15

0.62 [ay 3,5,7,9-15

0.61 1.20 3,5,7,9, 11-15

0.60 1.24 3,9,7,9)13-15

0.56 1.08 3,13-15

0.58 els 3, 11-15

3,5,7,9, 1113-15

3-15

3,5,7,9, 11-15

3-15

3-5,7,9,1,13-15

3-15

a.l5

3-15

Ooooooogeoe

DOA nnonn7inan

~— WH YWYHOOAn Sf

co eee Nome sera. Scge Monae Core” Ree Sam

VINNNMOARHRHO

SIOMDWN wm wn

Subgenus Haematomyidium

0.65 0.83 3,8-10

0.57 0.78 3,8-10

G59 0.77 3,8-10

0.56 0.65 3,8-10

0.59 G97 3,8-10

0.58 0.93 3,7-10

0.58 1.28 3,7-10

0.53 0.80 3,8-10

0.59 0.97 3,8-10

0.56 0.82 3,8-10

0.63 0.94 3,8-10

3,7-10

0.55 0.88 3,(4-7),8-10

0.55 3,(7),8-10

0.52 0.81 3,(6), 7-10

Subgenus Beltranmyia

0.60 1.48 3-15

0.60 1.00 3,13-14

0.56 1.40 3-15

0.61 1.16 3, 11-15

0.58 1.48 3-15

0.58 1.30 3,U1-15

0.61 ee) 3-15

0.57 1.10 3,(4,5,7,8),1l,

13-14

Palpal

Ratio

NNNNNNN

™ BWM DD PD

NNNN NNNNN NNNNNNNNN

—~aIW & NP CNT ae ae (GOT IND NOW HANH SPD

NNNNWN

MPIWofS s+)

NNWNYUNNN

™“FNMOUOONNYWN

P/H

Ratio

Wirth, Dyce and Peterson: Nearctic Culicoides 9

Species Wing Costal Antennal Antennal Palpal P/H

Length Ratio Ratio Sensillar Ratio Ratio

(mm) Pattern

Subgenus Selfia

brookmani 295 Oe55 0.93 3-10 L9 0.76

denningi ay (ey) EO? 3,7-10 Pal 0.88

hieroglyphicus 1.14 0.56 1.03 3,5-10 Zeal 0.83

jacksoni. 1.16 0.54 G93 3-10 Jip 0.80

jamesi 1.21 0.56 1.00 3-10 25 0.87

multipunctatus 0.85 0.55 0.96 3,5-10 Zk 0.86

tenuistylus 1.26 0.58 1.00 3-10 Pad O99

Subgenus Wirthomyia

bottimeri 0.90 0.35 el2 3-10 2.1 0.64

stilobezzioides 1.56 D9 1.87 3-10 Zar 0.69

Subgenus Unplaced, biguttatus Species Group

biguttatus 1.30 0.64 3, 11-15 Zeit 0.85

loisae 1.00 0.60 1.21 3, 11-15 Ziel 0.49

mulrennani 0.95 0.65 0 3,11-15 2.3 0.63

spinosus 1.20 0.64 1.10 3, 11-15 2.6 0.72

sublettei Ihe 0.55 1.07 3, 11-15 2.4 O35

usingeri ] 0.57 Os 3,(7,8),9-15 222 O79

Subgenus Unplaced, stonei Species Group

melleus 1.05 0.60 1.10 3,10-14 Bal 0.63

mortivallis 1.02 0.58 1.12 3,/7-14 22 0.79

niger 1.24 0.58 1.04 3,7-14 2.5 0.90

owyheensis 1.10 0.60 1.00 3,7-14 22 0.83

stonei 1.15 0.57 lene 3-14 25 0.86

tissoti L.02 0.58 1.04 3,7-14 25 0.90

werneri 1.08 0.57 1.02 3,9-14 Zl G79

Subgenus Unplaced, chaetophthalmus Species Group

atchleyi 195 0.66 1.14 3, 11-15 2.2 0.65

saundersi 2550 0.66 1.06 3, 11-15 229 0.93

Miscellaneous Unplaced Species

arubae ete 0.52 0.87 3-10 2.5 0.90

floridensis 0.77 0.65 Ass none 2.0 0.7]

hoquei 1.22 0.57 0.89 3,(7),8-10 2.5 0.94

kibunensis 1.50 0,52 1.40 3-15 2.3 0.89

mohave 1.09 0.54 1.11 3,(6),7-10 2.1 0.67

monoensis Li 0.53 0.97 3,14-15 3.0 0.69

multidentatus 1.00 0.52 0.99 3,(4),5,(6), 7-10 Zel 0.78

nanellus 1.10 0.63 1.02 3-7,9, 11-15 2,0 0.68

nanus 1.00 0.60 1.40 3-15 ey 0.57

posoensis 0.99 O57 1.10 3, 11-15 2.6 0.78

salihi 0.82 0.53 1.12 3-10 2.0 0.69

travisi 1.40 0.62 1.50 3-15 2 0.86

10 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

ATLAS OF SPECIES

Subgenus Culicoides Latreille, cockerellii Species Group

REFERENCE: Wirth and Blanton, 1969a (revision, key)

l. cockerellii (Coquillett), 1901: 603 (Ceratopogon); Colorado. British

Columbia south to California, east to New Mexico and Quebec. Photo:

Woods Landing, Laramie, Wyoming (Denning).

2. freeborni Wirth and Blanton, 1969a: 217; California. California,

Oregon, Washington. Photo: Wawawai Canyon, Whitman Co., Washington

(Jorgensen), paratype.

3. frohnei Wirth and Blanton, 1969a: 219; Alaska. Alaska, south to British

Columbia, east to Labrador. Photo: Whitecourt, Alberta (Kessel), paratype.

4. gregsoni Wirth and Blanton, 1969a: 222; British Columbia. British

Columbia, Oregon. Photo: Exchamsiks, British Columbia (Kessel),

paratype.

5. lahontan Wirth and Blanton, 1969a: 223; California. California, Montana,

Oregon. Photo: Resting Springs, Inyo Co., California (Belkin and

McDonald), paratype.

6. neofagineus Wirth and Blanton, 1969a: 227; Arizona. Arizona,

California, Oregon. Photo: Pinery Canyon, Cochise Co., Arizona

(O'Brien), paratype.

7. neomontanus Wirth, 1976: 15; Utah (Synonym: montanus Wirth and Blanton,

1969a: 225, preoccupied). British Columbia south to California, east to

Manitoba and New Mexico. Photo: Logan, Cache Co., Utah (Knowlton),

paratype. sa

8. saltonensis Wirth, 1952: 173 (as cockerellii ssp.); California. Photo:

North shore Salton Sea, Riverside Co., California (Brenner).

9. sierrensis Wirth and Blanton, 1969a: 232; California. California,

Nevada. Photo: Halleujah Junction, Lassen Co., California

(Schlinger), paratype.

10. sommermanae Wirth and Blanton, 1969a: 234; Alaska. Photo: Anchorage,

Alaska (Sommerman), paratype.

Wirth, Dyce and Peterson: Nearctic Culicoides ll

12 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

ll. sordidellus (Zetterstedt), 1838: 820 (Ceratopogon); Greenland.

Photo: Greenland (no locality) (Harmston).

12. tristriatulus Hoffman, 1925: 294 (cockerellii var.); California.

Coastal; Alaska south to northern California. Photo: Eel Creek Forest

Camp, Coos Co., Oregon (Kessel).

pulicaris Species Group

13. canadensis Wirth and Blanton, 1969a: 211; Minnesota. Alaska south to

British Columbia, east to Quebec and Nova Scotia, Maine, Minnesota,

Wisconsin. Photo: Itasca State Park, Clearwater Co., Minnesota

(Barr).

14. neopulicaris Wirth, 1955: 355; Texas. Louisiana and Texas south to Costa

Rica. Photo: Kerrville, Kerr Co., Texas (Bottimer), paratype.

15. yukonensis Hoffman, 1925: 294 (cockerellii var.); Yukon Territory.

Alberta, Alaska, British Columbia, Northwest Territory, Yukon

Territory. Photo: Palmer, Alaska (Sommerman).

Subgenus Hoffmania Fox

16. insignis Lutz, 1913: 51; Brazil. (Synonym: inamollae Fox and

Hoffman, 1944: 110; Puerto Rico). Alabama, Georgia, and Florida, south to

Argentina. Photo: Orlando, Orange Co., Florida (Stokes).

17. venustus Hoffman, 1925: 290; Maryland. Nebraska and Wisconsin east to

Nova Scotia, south to Louisiana and Florida. Photo: Allegany State Park,

Cattaraugus Co., New York (Wirth).

Subgenus Avaritia Fox

REFERENCE: Jamnback and Wirth, 1963 (revision, key)

18. alachua Jamnback and Wirth, 1963: 187; Florida. Florida, South

Carolina. Photo: Gainesville, Alachua Co., Florida (Wirth).

19. chiopterus (Meigen), 1830: 263 (Ceratopogon); Europe. Holarctic;

entire Canada, U.S.A. and U.S.S.R. Photo: U.S.S.R. (Gutsevich).

20. hirtulus (Coquillett), 1900: 396; Alaska. Coastal; Alaska south

to northern California. Photo: Prince Rupert, British Columbia

(Kessel).

Wirth, Dyce and Peterson: Nearctic Culicoides 13

14 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

21. juddi Cochrane, 1973: 316; New York. Alabama, Florida, Louisiana,

Maryland. Photo: Ringwood Game Preserve, Tompkins Co., New York

(Cochrane), holotype.

22. obsoletus (Meigen), 1818: 76 (Ceratopogon); Europe. Holarctic;

in North America south from Alaska to California and east to

Georgia. Photo: East Concord Bog, Erie Co., New York (Wirth).

23. pechumani Cochrane, 1974: 133; New York. Florida, New York.

Photo: Juniper Springs, Marion Co., Florida (Wirth), paratype.

24. pusillus Lutz, 1913: 52; Brazil. Neotropical; southern Florida to

Argentina. Photo: Ross and Costello Hammock, Dade Co., Florida

(Baranowski).

25. sanquisuga (Coquillett), 1901: 604 (Ceratopogon); Maryland. Alaska east to

Nova Scotia, south to California and Georgia. Photo: Fairland,

Montgomery Co., Maryland (Hubert).

Subgenus Drymodesmyia Vargas

REFERENCE: Wirth and Hubert, 1960 (revision, key; as copiosus group)

26. arizonensis Wirth and Hubert, 1960: 655; Arizona. Arizona,

California, Mexico (Baja California). Photo: Morristown, Maricopa

Co., Arizona (Ryckman), paratype.

27. butleri Wirth and Hubert, 1960: 650; Arizona. Photo: Portal, Cochise Co.,

Arizona (Wirth).

28. byersi Atchley, 1967: 983; New Mexico. California, New Mexico.

Photo: Glenwood, Catron Co., New Mexico (Wirth), holotype.

29. cacticola Wirth and Hubert, 1960: 653; California. Arizona,

California, Mexico, Texas. Photo: San Dimas Canyon, Los Angeles Co.,

California (Christiansen and Fonseca), paratype.

30. copiosus Root and Hoffman, 1937: 171; Mexico. Arizona,

California, south to Texas and northern Mexico. Photo: Brown Canyon,

Pima Co., Arizona (Butler).

Wirth, Dyce and Peterson: Nearctic Culicoides [5

16 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

31. hinmani Khalaf, 1952: 353; Oklahoma. Wyoming south to Texas, east to New

York and Florida. Photo: Gainesville, Alachua Co., Florida (Blanton).

32. insolatus Wirth and Hubert, 1960: 654; Baja California. California,

Mexico (Baja California). Photo: San Felipe, Baja California,

Mexico (Ryckman, Lee and Ames), paratype.

33. jamaicensis Edwards, 1922: 165 (loughnani var.); Jamaica. Neotropical;

Florida, West Indies, Texas south to Colombia. Photo: Negril, Westmoreland

Parish, Jamaica (Woodruff).

34. jonesi Wirth and Hubert, 1960: 650; Texas. Photo: Fredericksburg,

Gillespie Co., Texas (Blanton and Borchers).

35. loughnani Edwards, 1922: 165; Jamaica. Alabama, Bahamas, Cuba, Florida,

Georgia, Jamaica, Texas, and Australia. Photo: Fakahatchee Island,

Collier Co., Florida (Blanton and Meyer).

36. ryckmani Wirth and Hubert, 1960: 656; California. Arizona,

California, Mexico (Baja California), Texas. Photo: San Dimas

Canyon, Los Angeles Co., California (Christiansen and Fonseca),

paratype.

37. sitiens Wirth and Hubert, 1960: 652; California. Arizona,

California, Mexico (Baja California). Photo: San Dimas Canyon,

Los Angeles Co., California (Christiansen and Fonseca), paratype.

38. torridus Wirth and Hubert, 1960: 654; Baja California. California, Mexico

(Baja California). Photo: San Felipe, Baja California, Mexico

(Christiansen), paratype.

Subgenus Diphaomyia Vargas

REFERENCE: Atchley and Wirth, 1979 (revision, key; in part, haematopotus group)

39. baueri Hoffman, 1925: 297; Maryland. Alabama, Colorado, Florida,

Louisiana, Maryland, Tennessee. Photo: Greenbrier, Limestone Co., Alabama

(Snow).

40. bergi Cochrane, 1973: 311; New York. Arizona, California, Colorado, New

Mexico, New York, Texas, Utah, Wisconsin. Photo: Springerville,

Apache Co., Arizona (Wirth).

Wirth, Dyce and Peterson: Nearctic Culicoides 17

Harn ut

i jee Liste iandetsettis A

Al,

42.

43.

44,

45.

46.

47.

49.

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

blantoni Vargas and Wirth, 1955: 33; Mexico. Texas, northern Mexico.

Photo: Cameron Co., Texas (Eads).

defoliarti Atchley and Wirth, 1979: 527; Arizona. Arizona, California

Oregon, New Mexico, Utah, Wyoming. Photo: Manzanita Forest Camp,

Coconino Co., Arizona (Sanderson).

edeni Wirth and Blanton, 1974a: 23; Florida. Bahamas, Florida,

south Carolina. Photo: Rock Springs, Orange Co., Florida (Wirth),

paratype.

erikae Atchley and Wirth, 1979: 532; New Mexico. Arizona, New Mexico.

Photo: Cedar Creek Canyon, Lincoln Co., New Mexico (Atchley), paratype.

footei Wirth and Jones, 1956: 162; Virginia. Kentucky and Mississippi,

east to Quebec and Florida. Photo: Forest Glen, Montgomery Co., Maryland

(Wirth).

haematopotus Malloch, 1915: 302; Illinois. Nova Scotia, Ontario,

entire U.S.A., northern Mexico. Photo: Fairland, Montgomery Co., Maryland

(Hubert).

inyoensis Wirth and Blanton, 1969b: 565; California. Photo: Resting

Springs, Inyo Co., California (Belkin and McDonald, holotype.

Subgenus Unplaced, daedalus Species Group

. daedalus Macfie, 1948: 83; Mexico. Arizona, New Mexico, Mexico, south to

Colombia. Photo: Brown Canyon, Pima Co., Arizona (Butler).

luglani Jones and Wirth, 1958: 89; Texas. Arizona, California,

Florida, Mexico, New Mexico, Texas. Photo: Portal, Cochise Co.,

Arizona (Davis).

pampoikilus Macfie, 1948: 79; Mexico. Arizona, Mexico, south to

Panama and Venezuela. Photo: Portal, Cochise Co., Arizona (Sabrosky).

Wirth, Dyce and Peterson: Nearctic Culicoides 19

ern Sr rad sec ica sc

base

hy a

me it

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Subgenus Unplaced, guttipennis Species Group

REFERENCE: Wirth and Blanton, 1967 (revision, key)

52.

5D:

D4.

DD.

56.

DY.

58.

59:

60.

arboricola Root and Hoffman, 1937: 166; Maryland. Wyoming to Texas, east

to Connecticut and Florida. Photo: Falls Church, Fairfax Co., Virginia

(Wirth).

beckae Wirth and Blanton, 1967: 213; Florida. Alabama, Florida,

Maryland, New York, South Carolina, Virginia. Photo: Atmore,

Escambia Co., Alabama (Blanton).

californiensis Wirth and Blanton, 1967: 215; California. Arizona,

California. Photo: Bakersfield, Kern Co., California (McDonald),

paratype.

cochisensis Wirth and Blanton, 1967: 216; Arizona. Arizona,

California, Mexico, Utah. Photo: Sycamore Canyon, Santa Cruz Co.,

Arizona (Butler), paratype.

flukei Jones, 1956: 30; Wisconsin. New York, Nova Scotia, Rhode Island,

Virginia, Wisconsin. Photo: Dane Co., Wisconsin (Jones), paratype.

quttipennis (Coquillett), 1901: 603 (Ceratopogon); Ohio. Minnesota south

to Oklahoma and Louisiana, east to Nova Scotia and Florida. Photo:

Colesville, Montgomery Co., Maryland (Wirth).

oklahomensis Khalaf, 1952: 355 (villosipennis var.); Oklahoma.

California south to Mexico, east to Oklahoma and Mississippi. Photo:

Fredericksburg, Gillespie Co., Texas (Blanton and Borchers).

ousairani Khalaf, 1952: 354; Oklahoma. Alabama, Florida, Maryland,

New Mexico, Oklahoma, Pennsylvania, Texas, Virginia. Photo:

Kerrville, Kerr Co., Texas (Bottimer). ,

pecosensis Wirth, 1955: 358; Texas. New Mexico, Texas. Photo:

Sanderson, Terrell Co., Texas (Brundrett), paratype.

villosipennis Root and Hoffman, 1937: 165; Maryland. Wisconsin, Nebraska

and Oklahoma, east to Nova Scotia and Florida. Photo: Falls Church,

Fairfax Co., Virginia (Wirth).

Wirth, Dyce and Peterson: Nearctic Culicoides on

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Subgenus Unplaced, piliferus Species Group

REFERENCE: Wirth and Hubert, 1962 (revision, key, eastern North America)

6l. alexanderi Wirth and Hubert, 1962: 190; Massachusetts. (Synonym:

62.

63.

64.

65.

66.

67.

68.

69.

pseudopiliferus Wirth and Hubert, 1962: 189; New York). Michigan east to

Nova Scotia, south to Tennessee and Maryland. Photo: Bittinger 4H

Camp, Garrett Co., Maryland (Wirth).

bickleyi Wirth and Hubert, 1962: 188; Maryland. Wisconsin east to

Quebec, south to Louisiana and Florida. Photo: Snow Hill, Worcester Co.,

Maryland (Anderson).

cavaticus Wirth and Jones, 1956: 166; California. Arizona,

California, Oregon. Photo: Kings River, Fresno Co.,

California (Clark).

chewaclae Glick and Mullen, 1983: 378; Alabama. Alabama, California,

Maryland, Oregon. Photo: Chewacla State Park, Lee Co., Alabama

(Glick), holotype.

denticulatus Wirth and Hubert, 1962: 193; Wisconsin. Minnesota and

Nebraska east to Nova Scotia, south to Maryland. Photo: Washburn Co.,

Wisconsin (Jones), paratype.

doeringae Atchley, 1967: 1014; New Mexico. Arizona, California, Colorado,

Montana, New Mexico, Oregon, Utah. Photo: Mormon Lake Village, Coconino

Co., Arizona (Sanderson).

downesi Wirth and Hubert, 1962: 186; Ontario. Maine, Michigan,

New York, Nova Scotia, Ontario, Quebec. Photo: Rideau River,

Ottawa, Ontario (Wirth), paratype.

franclemonti Cochrane, 1974: 128; New York. Newfoundland, New York,

Nova Scotia, Ontario. Photo: Mer Bleue, Ottawa, Ontario (Wirth),

paratype.

husseyi Wirth and Blanton, 197lb: 76; Florida. Arkansas, Florida,

Maryland. Photo: Torreya State Park, liberty Co., Florida

(Wirth), paratype.

jamnbacki Wirth and Hubert, 1962: 192; New York. Louisiana,

Michigan, New York, North Carolina, Nova Scotia, Ontario.

Photo: Newcomb, Hamilton Co., New York (Jamnback), paratype.

Wirth, Dyce and Peterson: Nearctic Culicoides 25

ibe

72.

ITS.

14,

Wey

76.

IEE

VE.

80.

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

kirbyi Glick and Mullen, 1983: 380; Maryland. Alabama, Maryland.

Photo: Patuxent Wildlife Refuge, Prince Georges Co., Maryland

(Wirth), holotype.

lophortygis Atchley and Wirth, 1975: 1422; California. Photo:

Hopland, Mendocino Co., California (Murphy), paratype.

parapiliferus Wirth and Blanton, 1974b: 71; New York. Wisconsin and

Iowa east to Ontario and Massachusetts, south to Florida. Photo: Montauk,

Suffolk Co., New York (Wirth), holotype.

piliferus Root and Hoffman, 1937: 163; Maryland. Wisconsin east to Nova

Scotia, south to Louisiana and Florida. Photo: Roland Park, Baltimore,

Maryland (Root), paratype.

riggsi Khalaf, 1957: 198; (piliferus ssp.); Oklahoma. Photo: Pushmataha

Co., Oklahoma (Khalaf), paratype.

scanloni Wirth and Hubert, 1962: 187; Virginia. Michigan south to

louisiana, east to Massachusetts and Florida. Photo: Torreya

State Park, Liberty Co., Florida (Blanton).

snowi Wirth and Jones, 1956: 163; Virginia. Illinois south to Mississippi,

east to Quebec and Florida. Photo: Alachua Co., Florida (Smith).

testudinalis Wirth and Hubert, 1962: 191; Pennsylvania. Wisconsin south

to Louisiana, east to Nova Scotia and Florida. Photo: Falls Church,

Fairfax Co., Virginia (Wirth), paratype.

unicolor (Coquillett), 1905: 65 (Ceratopogon); California. British

Columbia, California, Colorado, Oregon, Washington. Photo: Eureka,

Humboldt Co., California (Barber), lectotype.

utowana Jamnback, 1965: 109; New York. New York, Ontario. Photo:

Mer Bleue, Ottawa, Ontario (Wirth).

Wirth, Dyce and Peterson: Nearctic Culicoides 25

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Subgenus Unplaced, palmerae Species Group

REFERENCE: Wirth and Rowley, 1971 (revision, key)

81. calexicanus Wirth and Rowley, 1971: 156; California. California,

GZ.

BS.

84.

85.

86.

87.

88.

89.

90.

Nevada. Photo: North Shore Salton Sea, Riverside Co., California

(Brenner).

davisi Wirth and Rowley, 1971: 157; Washington. Alberta, Washington.

Photo: Othello, Adams Co., Washington (Rowley), allotype.

hawsi Wirth and Rowley, 1971: 159; Washington. California, Oregon,

Utah, Washington. Photo: Othello, Adams Co., Washington (Rowley),

paratype. |

leechi Wirth, 1977: 53; California. Photo: Howell Mountain, Napa Co.,

California (Leech), paratype.

novamexicanus Atchley, 1967: 1012; New Mexico. Photo: Roswell,

Chaves Co., New Mexico (Atchley), allotype.

oregonensis Wirth and Rowley, 1971: 162; Oregon. California, Oregon.

Photo: Corvallis, Benton Co., Oregon (Nelson), allotype.

palmerae James, 1943: 151; Colorado. Washington south to California,

east to Wyoming and New Mexico. Photo: Fort Collins, Larimer Co.,

Colorado (Palmer), paratype.

utahensis Fox, 1946: 246; Utah. Calfornia, Idaho, Nevada, New Mexico,

Oregon, Utah, Washington. Photo: Vale, Malheur Co.. Oregon (Goeden).

wirthi Foote and Pratt, 1954: 36; Montana. British Columbia, Montana,

Oregon, Northwest Territory. Photo: Kamloops, British Columbia

(Curtis).

Subgenus Haematomyidium Goeldi

debilipalpis Lutz, 1913: 60; Brazil. (Synonym: khalafi Beck, 1957:

104; Florida). Neotropical; in U.S.A. from Kentucky and Louisiana east

to Maryland and Florida. Photo: San Felasco, Alachua Co., Florida (Smith).

Wirth, Dyce and Peterson: Nearctic Culicoides

28 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

91. eadsi Wirth and Blanton, 197la: 37; Texas. Cuba, Florida, Mexico,

Texas. Photo: Cameron Co., Texas (Eads), holotype.

92. paraensis (Goeldi), 1905: 137 (Haematomyidium); Brazil. South and

Central America; Colorado and Wisconsin south to Louisiana, east to

Pennsylvania and Florida. Photo: Torreya State Park, Liberty Co., Florida

(Blanton).

93. reevesi Wirth, 1952: 193; California. Arizona, California, New

Mexico. Photo: Needles, San Bernardino Co., California (Mulla).

94. torreyae Wirth and Blanton, 197lb: 74; Florida. Photo: Juniper

Springs, Marion Co., Florida (Wirth), paratype.

Subgenus Oecacta Poey

95. barbosai Wirth and Blanton, 1956: 161; Panama. Coastal; southern Florida

south to Ecuador. Photo: Key Largo, Monroe Co., Florida (Kemp).

96. furens (Poey), 1851: 236 (Oecacta); Cuba. (Synonym: dovei Hall,

1932: 88; Georgia). Coastal; Massachusetts south to Texas, West Indies

and Mexico, south to Ecuador and Brazil. Photo: Sapelo Island, McIntosh

Co., Georgia (Smalley).

97. furensoides Williams, 1955: 271; Michigan. (Synonym: dickei Jones,

1956: 28: Wisconsin). Alaska, Michigan, New York, Ontario, West

Virginia, Wisconsin. Photo: Douglas Lake, Cheboygan Co., Michigan

(Williams).

98. stellifer (Coquillett), 1901: 604 (Ceratopogon); District of Columbia.

Montana south to California, east to Nova Scotia and Florida. Photo:

Falls Church, Fairfax Co., Virginia (Wirth).

Subgenus Monoculicoides Khalaf

REFERENCE: Wirth and Jones, 1957 (revision, subspecies of C. variipennis)

99. gigas Root and Hoffman, 1937: 172; Saskatchewan. Saline and alkaline

habitats: Alaska, British Columbia east to Manitoba, south to Montana

and Nebraska. Photo: Thompson Lake, Burke Co., North Dakota (Wirth).

100. variipennis variipennis (Coquillett), 1901: 602 (Ceratopogon); Virginia.

Forested areas: British Columbia east to Quebec, south to Colorado,

Louisiana and Florida. Photo: Fort Bragg, Cumberland Co., North Carolina.

Wirth, Dyce and Peterson: Nearctic Culicoides 29

aioe

30 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

[variipennis albertensis Wirth and Jones, 1957: 17; Alberta. Saline

habitats; Alberta south to Oklahoma. Photo material not available. ]

101. variipennis australis Wirth and Jones, 1957: 15; Louisiana. Saline

habitats; Arkansas, California, Florida, Kansas, Louisiana, New York,

Texas, Virginia. Photo: Grapeland, Houston Co., Texas (Woodruff).

[variipennis occidentalis Wirth and Jones, 1957: 21; California.

Saline habitats; Arizona, British Columbia, California, Colorado, Mexico

(Baja California), Montana, New Mexico. Photo material not available. }

102. variipennis sonorensis Wirth and Jones, 1957: 18; Arizona. Arizona,

British Columbia, California, Florida, Montana, Nevada, New Mexico,

Tennessee, Texas. Photo: Glenwood, Catron Co., New Mexico (Wirth).

Subgenus Beltranmyia Vargas

103. alaskensis Wirth, 1951: 84; Alaska. Photo: Palmer, Alaska

(Sommerman).

104. bermudensis Williams, 1956: 298; Bermuda. Coastal; Bermuda, Florida,

Louisiana, New Jersey, New York, Texas. Photo: St. Joseph State

Park, Gulf Co., Florida (Wirth).

105. crepuscularis Malloch, 1915: 303; Illinois. Entire continental U.S.A.,

southern Canada, Mexico. Photo: Gainesville, Alachua Co., Florida (Wirth).

106. crepuscularis Malloch. Photo: Forest Glen, Montgomery Co., Maryland

(Wirth).

107. crepuscularis Malloch. Photo: Taos, Taos Co., New Mexico (Wirth).

108. crepuscularis Malloch. Photo: Mecca, Riverside Co., California

(Foulk).

109. crepuscularis Malloch. Photo: Fairbanks, Alaska (Sommerman).

110. hollensis (Melander and Brues), 1903: 13 (Ceratopogon); Massachusetts.

(Synonym: canithorax Hoffman, 1925: 284; Georgia). Atlantic coast; Nova

Scotia south to northern Florida. Photo: Hyannisport, Barnstable Co.,

Massachusetts (Wall).

Wirth, Dyce and Peterson: Nearctic Culicoides 31

eae

PMS Ae a Ay Ak a

32 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

lll. knowltoni Beck, 1956: 136; Florida. Bahamas, Florida. Photo: Vero

Beach, Indian River Co., Florida (Ent. Res. Ctr.).

112. mississippiensis Hoffman, 1926: 158; Mississippi. Gulf coast; Alabama,

Florida, Louisiana, Mississippi, Texas. Photo: Gulf Coast, Baldwin Co.,

Alabama (Smith).

113. sphaqnumensis Williams, 1955: 269; Michigan. Alaska, Minnesota east to

Newfoundland, south to West Virginia. Photo: Douglas Lake,

Cheboygan Co., Michigan (Williams).

114. wisconsinensis Jones, 1956: 32; Wisconsin. Washington and Oregon, east to

Alberta, Michigan and Iowa. Photo: Twin lake, Cherry Co., Nebraska

(Wirth).

Subgenus Selfia Khalaf

REFERENCE: Atchley, 1970 (revision, key, biosystematics)

115. brookmani Wirth, 1952: 179; California. Arizona, California, New

Mexico. Photo: Arroyo Seco, Monterey Co., California (Wirth).

116. denningi Foote and Pratt, 1954: 20; Saskatchewan. Washington and Oregon,

east to Saskatchewan and Nebraska. Photo: Little White River, Mellette

Co., South Dakota (Wirth).

117. hieroglyphicus Malloch, 1915: 297; Arizona. California east to South

Dakota, south to northern Mexico. Photo: Buttonwillow, Kern Co.,

California (Nelson).

118. jacksoni Atchley, 1970: 258; New Mexico. Arizona, New Mexico, Utah.

Photo: Cedar Creek Canyon, Lincoln Co., New Mexico (Atchley),

allotype.

119. jamesi Fox, 1946: 244; Montana. Washington south to California, east to

South Dakota and New Mexico. Photo: Beaver Creek, Lawrence Co., South

Dakota (Wirth).

120. multipunctatus Malloch, 1915: 296; Illinois. Alabama, Illinois,

Kansas, Mexico, Missouri, Oklahoma, Texas. Photo: Fredericksburg,

Gillespie Co., Texas (Blanton and Borchers).

Wirth, Dyce and Peterson: Nearctic Culicoides 33

E22.

2357

124.

125.

126.

127.

128.

129.

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

tenuistylus Wirth, 1952: 178; California. Photo: Hopland, Mendocino

Co., California (Anderson).

Subgenus Wirthomyia Vargas

bottimeri Wirth, 1955: 356; Texas. Arizona, Texas. Photo:

Fredericksburg, Gillespie Co., Texas (Blanton and Borchers).

stilobezzioides Foote and Pratt, 1954: 33; New York. Alaska south to

Washington, east to Minnesota, Nebraska and Newfoundland. Photo: Pelican

Lake, Cherry Co., Nebraska (Wirth).

Subgenus Unplaced, biguttatus Species Group

biguttatus (Coquillett), 1901: 604 (Ceratopogon); District of

Columbia. Wisconsin south to Louisiana, east to Nova Scotia and Florida.

Photo: Douglas Lake, Cheboygan Co., Michigan (Williams).

loisae Jamnback, 1965: 77; New York. Florida, Maryland, New York,

Virginia, West Virginia. Photo: Allegany State Park, Cattaraugus Co.,

New York (Wirth), paratype.

mulrennani Beck, 1957: 103; Florida. Florida, Maryland

Massachusetts, Mississippi, West Virginia. Photo: Wewahitchka,

Gulf Co., Florida (Blanton and Boike).

spinosus Root and Hoffman, 1937: 172; Maryland. Wisconsin south to

Louisiana, east to Nova Scotia and Florida. Photo: Alachua Co., Florida

(Smith).

sublettei Atchley, 1967: 997; New Mexico. California, New Mexico,

Texas, Utah. Photo: Fort Sumner, DeBaca Co., New Mexico (Atchley),

allotype.

usingeri Wirth, 1952: 192; California. California, Oregon, Nevada.

Photo: Hat Creek Ranger Station, Shasta Co., California (Usinger),

allotype.

Subgenus Unplaced, stonei Species Group

REFERENCE: Wirth and Blanton, 197lc (revision, key)

130.

melleus (Coquillett), 1901: 604; Florida. Coastal; Maine south to

Louisiana and the Bahamas. Photo: Brunswick, Glynn Co., Georgia (Hagen).

Wirth, Dyce and Peterson: Nearctic Culicoides 35

Hi nr ac

itn, a

Ime.

loa

32.

153%

134.

135.

156.

isi,

138.

159.

140.

Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

mortivallis Wirth and Blanton, 197lc: 465; California. Photo:

Saratoga Springs, Inyo Co., California (Belkin and McDonald),

holotype.

niger Root and Hoffman, 1937: 168; Maryland. Coastal plain; Massachusetts

south to Florida and Louisiana. Photo: Torreya State Park, liberty Co.,

Florida (Wirth).

owyheensis Jones and Wirth, 1978: 56; Idaho. Photo: Bruneau, Owyhee

Co., Idaho (Jones and Potter), paratype.

stonei James, 1943: 149; Colorado. (Synonym: weesei Khalaf, 1952:

65; Oklahoma). Colorado, Nebraska, New Mexico, Oklahoma, South Dakota,

Texas, Utah. Photo: Fort Collins, Larimer Co., Colorado (Palmer),

paratype.

tissoti Wirth and Blanton, 1966: 279; Florida. Florida, Maryland,

South Carolina. Photo: Snow Hill, Worcester Co., Maryland (Anderson).

werneri Wirth and Blanton, 197lc: 463; Arizona. Arizona, California,

Mexico, Texas. Photo: Quitobaquito, Pima Co., Arizona (Adachi),

holotype.

Subgenus Unplaced, chaetophthalmus Species Group

atchleyi Wirth and Blanton, 1969b: 559; Alaska. Alaska, New Mexico,

Nova Scotia. Photo: Johnson Lake, Kenai Peninsula, Alaska (Sommermann).

saundersi Wirth and Blanton, 1969b: 557; Washington. Alaska, south to

British Columbia, California, Montana, Oregon, Washington. Photo:

Sullivan Lake, Pend Oreille Co., Washington (Jorgensen).

Miscellaneous Unplaced Species

arubae Fox and Hoffman, 1944: 109; Aruba Island, Dutch West Indies.

Coastal; Neotropical; Texas. Photo: Cameron Co., Texas (Eads).

floridensis Beck, 1951: 135; Florida. Bahamas, Bermuda, Florida.

Photo: Gulf Hammock, Levy Co., Florida (Blanton).

Wirth, Dyce and Peterson: Nearctic Culicoides a7

38 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

141. hoguei Wirth and Moraes, 1979: 293; California. California, Mexico (Baja

California). Photo: Seal Beach, Orange Co., California (Moraes),

paratype.

142. kibunensis Tokunaga, 1937: 298; Japan. Holarctic; Arizona, Oregon,

Washington. Photo: Wawawai Canyon, Whitman Co., Washington (Jorgensen).

143. mohave Wirth, 1952: 187; California. California, Mexico (Baja

California). Photo: Vidal, San Bernardino Co., California (Coleman),

paratype.

144. monoensis Wirth, 1952: 193; California. Photo: Mono Lake, Mono Co.,

California (Wirth), paratype.

145. multidentatus Atchley and Wirth, 1975: 1421; California. Arizona,

California, New Mexico. Photo: Hopland, Mendocino Co., California

(Knudsen and Murphy).

146. nanellus Wirth and Blanton, 1969b: 564; California. Photo: Hopland,

Mendocino Co., California (Murphy), holotype.

147. nanus Root and Hoffman, 1937: 165; Maryland. Wisconsin to Colorado and

Texas, east to Ontario and Florida. Photo: Lake Charles, Calcasieu Parish,

Louisiana (Chapman).

148. posoensis Wirth and Blanton, 1969b: 562: California. Photo: Poso

Creek, Kern Co., California (Nelson), paratype.

149. salihi Khalaf, 1952: 351; Oklahoma. Arizona, California, Oklahoma,

Texas. Photo: Fredericksburg, Gillespie Co., Texas (Blanton and

Borchers).

150. travisi Vargas, 1949: 233; Maryland. (Synonym: simulans Root and

Hoffman, 1937: 167; Maryland, preoccupied; horneae Foote and Pratt, 1954:

25; New York). Wyoming south to Arizona, east to Nova Scotia and Florida.

Photo: Wyers Point, Tompkins Co., New York (Blanton).

Wirth, Dyce and Peterson: Nearctic Culicoides 39

i Aigjs PES:

Ardbasestrgars ais

Mai

40 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

LITERATURE CITED

Arnaud, P.H.

1956. The heleid genus Culicoides in Japan, Korea and Ryukyu Islands

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Atchley, W.R. and W.W. Wirth

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Beck, E.C.

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Downes, J.A. and W.W. Wirth

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Goeldi, E.

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Johannsen, O.A.

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42 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

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Jones, R.H. and W.W. Wirth

1958. New records, synonymy, and species of Texas Culicoides (Diptera,

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1978. A new species of western Culicoides of the stonei group (Diptera:

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Jorgensen, N.M.

1969. The systematics, occurrence, and host preference of Culicoides in

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1965. Biting ceratopogonids as vectors of human and animal diseases.

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Khalaf, K.T.

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Wirth, Dyce and Peterson: Nearctic Culicoides 43

Tokunaga, M.

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Williams, R.W.

1955. Two new species of Culicoides from Cheboygan County, Michigan

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1956. A new species of salt marsh sand fly from Florida, the Bahamas,

Panama and Ecuador: its distribution and taxonomic differentiation

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157-162.

1966. A new day-biting sand fly from the southeastern states (Diptera,

Ceratopogonidae). Fla. Ent. 49: 279-28].

1967. The North American Culicoides of the guttipennis group (Diptera:

Ceratopogonidae). Fla. Ent. 50: 207-232.

1969a. North American Culicoides of the pulicaris group (Diptera:

Ceratopogonidae). Fla. Ent. 52: 207-243.

1969b. New species and records of Culicoides from western North America

(Diptera: Ceratopogonidae). Proc. Ent. Soc. Wash. 71: 556-567.

197la. New Neotropical sandflies of the Culicoides debilipalpis group

(Diptera: Ceratopogonidae). Proc. Ent. Soc. Wash. 73: 34-43.

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1971c. New western Culicoides of the stonei group. J. Kans. Ent. Soc.

44: 459-467.

1974a. A new Florida sand fly closely related to Culicoides haematopotus

Malloch (Diptera: Ceratopogonidae). Fla. Ent. 57: 23-26.

1974b. New synonymy and a correction in the Culicoides piliferus group

(Diptera: Ceratopogonidae). Fla. Ent. 57: 71-75.

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1956. A population study of the Culicoides midges of the Edwards Plateau

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44 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

Wirth, W.W. and A.A. Hubert

1960. Ceratopogonidae (Diptera) reared from cacti, with a review

of the copiosus group of Culicoides. Ann. Ent. Soc. Am. 53:

639-658.

1962. The species of Culicoides related to piliferus Root and Hoffman in

eastern North America (Diptera, Ceratopogonidae). Ann. Ent. Soc.

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1956. Three new North American species of tree-hole Culicoides (Diptera,

Heleidae). Proc. Ent. Soc. Wash. 58: 161-168.

1957. The North American subspecies of Culicoides variipennis (Diptera,

Heleidae). U.S. Dep. Agric. Tech Bull. 1170: 1-35.

Wirth, W.W. and A.P.S. Moraes

1979. New records and new species of biting midges from salt marshes in

California and Mexico (Diptera: Ceratopogonidae). Pan-Pacific

Ent. 55: 287-298.

Wirth W.W., Ratanaworabhan, N.C. and F.S. Blanton

1974. Synopsis of the genera of Ceratopogonidae (Diptera). Ann.

Parasit. Hum. Comp. 49: 595-613.

Wirth, W.W., Ratanaworabhan, N.C. and D.H. Messersmith

1977. Natural history of Plummers Island, Maryland. XXII. Biting midges

(Diptera: Ceratopogonidae). 1. Introduction and key to genera.

Proc. Biol. Soc. Wash. 90: 615-647.

Wirth, W.W. and W.A. Rowley

1971. A revision of the palmerae group of the genus Culicoides. J. Kans.

Ent. Soc. 44: 153-171.

Zetterstedt, J.W.

1838. Dipterologia Scandinaviae. Sect. 3: Diptera, pp. 477-868. In

J.W. Zetterstedt, Insecta Lapponica, Lipsiae, 1140 pp.

Wirth, Dyce and Peterson: Nearctic Culicoides 45

INDEX OF SUBGENERIC NAMES (page no.)

Avaritia Fox 6, 12

Beltranmyia Vargas 8, 30

Culicoides Latreille 6, 10

Diphaomyia Vargas 7, 16

Drymodesmyia Vargas 6, 14

Haematomyidium Goeldi 8, 26

Hoffmania Fox 6, 12

Monoculicoides Khalaf 8, 28

Oecacta Poey 8, 28

Selfia Khalaf 9, 32

Wirthomyia Vargas 9, 34

INDEX OF SPECIFIC NAMES (species no.)

alachua Jamnback and Wirth 18

alaskensis Wirth 103

albertensis Wirth and Jones [page 30]

alexanderi Wirth and Hubert 6l

arboricola Root and Hoffman 51

arizonensis Wirth and Hubert 26

arubae Fox and Hoffman 139

atchleyi Wirth and Blanton 137

australis Wirth and Jones 101

barbosai Wirth and Blanton 95

baueri Root and Hoffman 39

beckae Wirth and Blanton 52

bergi Cochrane 40

bermudensis Williams 104

bickleyi Wirth and Hubert 62

biguttatus (Coquillett) 124

blantoni Vargas and Wirth 4]

borinqueni Fox [page 4]

bottimeri Wirth 122

brookmani Wirth 115

butleri Wirth and Hubert 27

byersi Atchley 28

cacticola Wirth and Hubert 29

calexicanus Wirth 81

californiensis Wirth and Blanton 53

canadensis Wirth and Blanton 13

canithorax Hoffman 110

cavaticus Wirth and Jones 63

chaetophthalmus Amosova [page 36]

chewaclae Glick and Mullen 64

chiopterus (Meigen) 19

cochisensis Wirth and Blanton 54

cockerellii (Coquillett) |

copiosus Root and Hoffman 30

crepuscularis Malloch 105-109

daedalus Macfie 48

davisi Wirth and Rowley 82

debilipalpis Lutz 90

defoliarti Atchley and Wirth 42

denningi Foote and Pratt 116

denticulatus Wirth and Hubert 65

dickei Jones 97

doeringae Atchley 66

dovei Hall 96

downesi Wirth and Hubert 67

eadsi Wirth and Blanton 91

edeni Wirth and Blanton 43

erikae Atchley and Wirth 44

floridensis Beck 140

flukei Jones 55

footei Wirth and Jones 45

franclemonti Cochrane 68

freeborni Wirth and Blanton 2

frohnei Wirth and Blanton 3

furens (Poey) 96

furensoides Williams 97

gigas Root and Hoffman 99

gregsoni Wirth and Blanton 4

quttipennis (Coquillett) 56

haematopotus Malloch 46

hawsi Wirth and Rowley 83

hieroglyphicus Malloch 117

hinmani Khalaf 31

hirtulus (Coquillett) 20

_hoguei Wirth and Moraes 14}

hollensis (Melander and Brues) lLIO

horneae Foote and Pratt 150

husseyi Wirth and Blanton 69

inamollae Fox and Hoffman 16

insignis Lutz 16

insolatus Wirth and Hubert 32

inyoensis Wirth and Blanton 47

jacksoni Atchley 118

jamaicensis Edwards 33

jamesi Fox ll9

jamnbacki Wirth and Hubert 70

jonesi Wirth and Hubert 34

juddi Cochrane 21

khalafi Beck 90

kibunensis Tokunaga 142

kirbyi Glick and Mullen 71

knowltoni Beck Ill

lahontan Wirth and Blanton 5

leechi Wirth 84

loisae Jamnback 125

lophortygis Atchley and Wirth 72

loughnani Edwards 35

luglani Jones and Wirth 49

melleus (Coquillett) 130

minutissimus (Zetterstedt) [page 4]

mississippiensis Hoffman Il2

mohave Wirth 143

monoensis Wirth 144

montanus Wirth and Blanton 7

mortivallis Wirth and Blanton 131

mulrennani Beck 126

46 Contrib. Amer. Ent. Inst., vol. 22, no. 4, 1985

multidentatus Atchley and Wirth 145

multipunctatus Malloch 120

nanellus Wirth and Blanton 146

nanus Root and Hoffman 147

neofagineus Wirth and Blanton 6

neomontanus Wirth 7

neopulicaris Wirth 14

niger Root and Hoffman 132

novamexicanus Atchley 85

obsoletus (Meigen) 22

occidentalis Wirth and Jones [page 30]

oklahomensis Khalaf 57

oregonensis Wirth and Rowley 86

ousairani Khalaf 58

owyheensis Jones and Wirth 133

palmerae James 87

pampoikilus Macfie 50

paraensis (Goeldi) 92

parapiliferus Wirth and Blanton 73

pechumani Cochrane 23

pecosensis Wirth 59

pifanoi Ortiz [page 4]

piliferus Root and Hoffman 74

posoensis Wirth and Blanton 148

pseudopiliferus Wirth and Hubert 61

pusillus Lutz 24

reevesi Wirth 93

riggsi Khalaf 75

ryckmani Wirth and Hubert 36

salihi Khalaf 149

saltonensis Wirth 8

sanquisuga (Coquillett) 25

saundersi Wirth and Blanton 138

scanloni Wirth and Hubert 76

sierrensis Wirth and Blanton 9

simulans Root and Hoffman 150

sitiens Wirth and Hubert 37

snowi Wirth and Jones 77

sommermanae Wirth and Blanton IO

sonorensis Wirth and Jones 102

sordidellus (Zetterstedt) 11

sphagnumensis Williams 113

spinosus Root and Hoffman 127

stellifer (Coquillett) 98

stilobezzioides Foote and Pratt 123

stonei James 134

sublettei Atchley 128

tenuistylus Wirth 121

testudinalis Wirth and Hubert 78

tissoti Wirth and Blanton 135

torreyae Wirth and Blanton 94

torridus Wirth and Hubert 38

travisi Vargas 150

tristriatulus Hoffman |2

unicolor (Coquillett) 79

usingeri Wirth 129

utahensis Fox 88

utowana Jamnback 80

variipennis (Coquillett) 100

venustus Hoffman 17

villosipennis Root and Hoffman 60

weesei Khalaf 134

wirthi Foot and Pratt 89

werneri Wirth and Blanton 136

wisconsinensis Jones 114

yukonensis Hoffman 15

A REVIEW OF THE NEOTROPICAL GENUS

STYPOMMISA (DIPTERA: TABANIDAE)!’®

GOR. Fairchild’

Florida State Collection of Arthropods

P.O. Box 1269

Gainesville, FL 32602

and

Richard C. Wilkerson”

Walter Reed Biosystematies Unit

Department of Entomology

National Museum of Natural History

Washington, D.C. 20560

Abstract. The tabanid (Diptera) genus Stypommisa Enderlein is redefined and

28 species are included. Of these 26 are keyed, the remaining 2 omitted for lack

of fresh material. Twenty names associated with the genus at various times are

considered synonyms, nomina nuda or belonging in other genera. Seven new

species are described and figured: S. anoriensis, NW Colombia; S. apicalis, E Peru,

E Bolivia, W Brazil; S. aripuana, Mato Grosso, Brazil; S. changena, mountains of

W Panama and E Costa Rica; S. kroeberi, E Peru, E Bolivia; S. spilota, E peru;

and S. xanthiecornis, Santa Catarina, Brazil. Available material of the deseribed

species is discussed and most species are figured.

INTRODUCTION

This taxonomically difficult group of small species was first recognized by

Lutz and Neiva in 1914, but neither their generic nor specific names were validly

published (Fairchild 1961). Enderlein (1923) proposed the name used here in 1923

with a brief 2 line diagnosis based on the then undescribed Stypommisa punc-

tipennis Enderlein. In 1925 both genus and species were fully described, the latter

from both sexes, but no other species were included. Kroeber (1929) gives a key

to the smaller genera of South American Tabanini, in which Stypommisa is char-

acterized as either lacking ocelli or these single or indistinct, not metallic, males

with thickly pilose eyes, both sexes with thread-like palpi and the wings with an

appendix. He includes 3 species in the genus, one of which he had described on a

previous page in Stypommia. Kroeber (1934) had difficulty with the group, placing

various species in a number of different Enderlein genera, we believe in a desire

1 Florida Agricultural Station Journal Series No./308.

Research Associate, Florida State Collection of Arthropods, Fla. Dept. of Agri-

culture and Consumer Services.

This work supported by National Science Foundation Grant BSR 8211808.

2 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

to follow Enderlein wherever he could. Fairchild (1969, pp. 219, 239) attempted

to define and key the genus, and in 1971 he gave a generic synonymy and listed

23 species of Stypommisa with 8 apparent synonyms. The present list includes 17

of these names as valid, 3, abdominalis Philip, ferruginosus Walker and puncti-

pennis Enderlein are considered synonyms, and 3, boliviensis Kroeber, serena

Kroeber and tantulus Hine, we now believe do not belong in Stypommisa. Since

the publication of the Neotropical Catalogue (Fairchild 1971), Philip has proposed

3 species and 1 subspecies, of which apaches Philip (1969, 1977) is a Stenotabanus,

Stypommisa (Styphocera) antennina Philip is either a good genus, or a subgenus of

Dasybasis while u-nigrum Philip and hypographa neofurva Philip seem valid.

Fairchild (in Wilkerson 1979) described bipuneta as new and Wilkerson (1979) added

Tabanus simplex Walker, previously placed by Fairchild (1971) in Dieladocera,

though we now feel it is better returned to Tabanus. With these deletions and

additions, the number of species in Stypommisa stood at 21, to which we here add

¢ new species, giving a total of 28 species which we now recognize. One other

name, Tabanus maculinevris Macquart 1855 was placed in Stypommisa by Kroeber

(1934), while Fairchild, who studied the poorly preserved type in the BM(NH)

(1956) and felt it was most similar to fulviventris Macquart, nevertheless listed it

as Tabanus in the Catalogue (1971). The possible relationships of this species to

both Stypommisa and to Anerythrops Barretto must await further fresh material

and restudy of Macquart's and Barretto's types.

The genus as treated here consists of small species rarely if ever exceeding a

wing or body length of 13 mm though frequently considerably less than 10 mm in

both dimensions. The included species possess a well marked tubercle at vertex,

often with vestiges of one or more ocelli, few to numerous, rarely without, setae

on the basicosta, well marked frontal calli, only rarely sparsely pilose eyes in the

female, though male eyes often densely pilose. The eyes may be unicolorous

green or bronze, bicolored, or with a median dark stripe, rarely otherwise. Many

of the species have dark clouds on the cross veins and fork of third vein (R A+ )

and often a short to long appendix at the fork. The frons is generally narrow, the

index over 4, and usually nearly parallel sided, though there are exceptions to

both conditions. The generie synonymy is given in Fairchild (1971).

Nothing is Known of the life histories or early stages of any species, and very

little has been recorded as to the habits of the adults. Fairchild (1953) reeords 2

of the Panamanian species as being over 90% arboreal, these at least being at-

tracted to men stationed in the forest canopy. The few known males have usu-

ally been taken at light, though unbaited flight traps have taken a few also. This

last method has yielded the great bulk of our material, in one case, 8 males, 1418

females, of 6 species were taken from a single locality on several occasions, not

exceeding a total of 3 months trapping. Most species seem to be confined to a

forest habitat. We cannot recall having been attacked by Stypommisa on more

than a very few occasions, though men stationed at both ground level and in the

forest canopy to collect mosquitoes have frequently taken Stypommisa. We have

found no records of their biting other animals.

The group as here treated is possibly not monophyletic, since several more or

less distinet groups within the genus seem to exist, as follows: pequeniensis

group--mainly brown species without abdominal markings or these in the form of

median pilose triangles, broad to narrow pilose bands or indistinct median pilose

Stripes, not underlain by pale pollinosity. The wings are tinted, often strongly, but

spots on cross veins and fork are generally small or absent. This group includes

pequeniensis Fairchild, flavescens Kroeber, captiroptera Kroeber, and glandicolor

Lutz; modiea group-—-this includes mainly black species with the first few abdomi-

nal tergites bluish pruinose, with or without white pilose triangles, the wings

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 3

more or less tinted and always with at least a spot on fork of 3rd vein. The

group consists of modica Hine, hypographa Kroeber, hypographa neofurva Philip,

maruccii Fehld., prunicolor Lutz and jaculator Fehld. They also share a black

elavate frontal callus, and most, if not all of the species are arboreal; seythropa

group--these species all have prominent middorsal pale pollinose abdominal trian-

gles, often heavily spotted wings, generally pale body coloration, usually without

pruinosity, and frontal calli usually round and joined above to a slender ridge. In-

cluded are seythropa Schiner., xanthicornis n. sp., fulviventris Macquart.,

paraguayensis Kroeber and rubrithorax Macquart. All are from southern Brazil,

Paraguay and northern Argentina.

ACKNOWLEDGEMENTS

We are most grateful to John E. Chainey, C.B. Philip, Paul Arnaud, and L.L.

Pechuman for the loan of critical specimens under their care, and to H. J. Teskey

and L. L. Pechuman for furnishing much of the long series of specimens which

enabled us to better understand the complexities of the group. We greatly appre-

ciate the careful reviews of this manuscript by L.L. Pechuman, John Burger and

H.J. Teskey, most of whose suggestions have been adopted.

SYNONYMICAL CHECKLIST OF NAMES DISCUSSED

Species believed valid Stypommisa are in boldface, as are all generic names.

abdominalis Philip 1960 (Stypommia) = flavescens (Kroeber) 1930 (Stypommia) (p.

20)

affinis Kroeber 1929 (p. 10)

ney ne Lutz and Neiva 1909 (Tabanus) nomen nudum = rubrithorax (Macquart)

p.287)

anoriensis n. sp. (p. 11)

antennina Philip (as (Styphocera), probably a Dasybasis (p. 2)

apaches Philip 1977 = Stenotabanus (p. 2)

apicalis n. sp. (p. 13)

aripuana n. sp. (p. 14)

beieri Philip 1958 (Hybomitra) = seythropa (Schiner) (p. 39)

bipuneta Fairchild 1979 (in Wilkerson 1979) (p. 16)

boliviensis Kroeber 1930 (Stypommia) = Tabanus s.1. (p. 2)

callicera (Bigot) 1892 (Tabanus) (p. 17)

captiroptera (Kroeber) 1930 (Tabanus) (p. 18)

changena n. sp. (p. 19)

ferruginosus Walker 1850 (Tabanus) = fulviventris (Macquart) (p. 21)

flavescens (Kroeber) 1930 (p. 20)

flavescens: Barretto 1957 Stypommisa = captiroptera (Kroeber) (p. 18)

frondicolus Fairchild 1951 (Stenotabanus) = maruccii (Fairchild) 1947 (p. 30)

fulviventris (Macquart) 1846 (Tabanus) (p. 21)

furva (Hine) 1920 (Tabanus) (p. 22)

glandicolor (Lutz) 1912 (Tabanus) (p. 24)

hypographa (Kroeber) 1930 (p. 25)

jaculator (Fairchild) 1942 (Stenotabanus) (p. 27)

kroeberi n. sp. (p. 28)

lerida (Fairchild) 1942 (Stenotabanus) (p. 30)

4 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

maculinevris Macquart 1855 (in Fairchild (1971) as Tabanus, in Kroeber (1934) as

Stypommisa) Not recognized by us (p. 2)

maculipennis Kroeber 1929 (Stypommia) = furva (Hine) (p. 22)

maruccii (Fairchild) 1947 (Stenotabanus) (p. 30)

modica (Hine) 1920 (Tabanus) (p. 32)

neofurva Philip 1969 (ssp. of hypographa) (p. 25)

paraguayensis (Kroeber) 1930 (p. 34)

pequeniensis (Fairchild) 1942 (Stenotabanus) (p. 35)

pequiensis: Philip 1960 (lapsus) (p. 35)

prunicolor (Lutz) 1912 (Tabanus) (p. 37)

punctipennis Enderlein 1925 = seythropa (Schiner) (p. 39)

punetulipennis Enderlein 1925 (Therioplectes) = rubrithorax (Macquart) (p. 37, 39)

rubrithorax (Macquart) 1838 (Tabanus) (p. 37)

ruficornis Kroeber 1930 = modica (Hine) (p. 32)

seythropa (Schiner) 1868 (Tabanus) (p. 39)

serena Kroeber 1931 (Tabanus). Not Stypommisa (p. 2)

simplex Walker 1850 (Tabanus). Is a Tabanus (p. 2)

spilota n. sp. (p. 40)

tantulus Hine 1920 (Tabanus). Not Stypommisa (p. 2)

u-nigrum Philip 1977 (p. 41)

venosa (Bigot) 1892 (Tabanus) (p. 43)

xanthicornis n. sp. (p. 44)

xenium Fairchild 1947 (Stenotabanus) = captiroptera (Kroeber) (p. 18)

KEYS TO SPECIES

The keys to both sexes presented below are intended only to aid in identifi-

cation and not to suggest phylogenetic relationships. Males of only about half

the known species have been recognized or described, and considerable doubt ex-

ists as to the proper association of a number of these. Generally the male key

seems to work, except in the ease of 3 closely similar species, pequeniensis, cap-

tiroptera and anoriensis, where we have been unable to find unequivocal dif-

ferences among the 8 available specimens, though we feel reasonably certain that

at least 2 and possibly all 3 species are represented. Only study of extensive se-

ries from several localities is likely to resolve the problem.

KEY TO STYPOMMISA FEMALES

1 Wing with weak clouds on fork and end of discal cell, none at ends of

basal cells, but well defined dark apical patch. Legs unicolorous

brown. Abdomen dorsally dark pilose, rarely with small indistinet pale

POLERS EP PTA Sica RSG eRe b AN hes aide adh) ohn. as aaalathider wAREiN ees 2

If strong apical patch present, then very strong spots present on all

LOGY GE EETS HATAC PTOI ihe d we ch darcintin Agana NL ok ch Akis o desde baw hue Ve dae Rees 3

2 Basal plate elongated, dorsal angle approximately 1/3 to 1/2 distance

from base to distal end of plate (Figs. 1-3). Pleura and mesonotum

eoneolorous, at most lower pleura somewhat contrasting gray

pollinose. Vein segment between appendix and Re generally as long

or longer than appendix. Frontal index 7.5-9.9 — .......06. apicalis (p. 13)

3(1)

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) )

Basal plate not elongated, dorsal angle approximately 1/4 distance from

base. Pleura gray pollinose, contrasting with brown mesonotum. Vein

segment at fork nearly always shorter than appendix. Frons wider,

iiMetGt- 164 seaside pequeniensis (p. 35)

Wing including costal cell glass clear. Abdomen above with dark median

integumental streak or spot on tergite 2 and row of small middorsal

pale pollinose triangles. Viewed from behind, abdomen shows promi-

nent dorsolateral pale pruinose spots on all tergites as well as pale

pruinose middorsal triangles. Frontal index 4.6-5.0 .......sccccccssccccescces

Medohs sek pide REa Hats Qua iy es ikon GROEN oO Lanois aripuana (p. 14)

Wing either spotted or tinted, or abdomen without pale pollinose triangles

GP OEY OS hace Tv aieiai sine bail g's Fey g wad UG s CERCA BS TUR Ed cana sacbcces 4

Wing with very large distinct spots at fork and on all ecrossveins, dis-

tinetly demarcated apical wing patch, and posterior margin of wing

with well demarcated smoky brown border. Pale areas of wing yel-

lowish. Abdomen above banded, tergites with broad golden yellow

pilose posterior Marging Ac... Kaaliava. Mestad ds u-nigrum (p. 41)

Wing never with discrete smoky brown posterior border nor, if strongly

spotted, with yellowish membrane. Abdomen various, but not as

ADOVE - dcapucsiieninnedpaihcenho Vaaidewbe din cx hlvupans then enamel de ianalle ses bidiainduindh onscenese +)

Frontal callus clearly higher than wide, occupying at least lower 1/3 of

frons, prominent, shiny black, clavate or drop shaped, tapering above

into narrow ridge nearly reaching ocellar tubercle. Brown to black

species, abdomen, when viewed from behind with extensive blue-gray

pruinosity, especially on first 2 tergites. Frons relatively broad,

frontal index usually less than 5.5. Wings never with appendix at fork

of thie: Vein i: csssujeeindecte cake ian Kaanseeeh visi er hoaxerenannt 6

Not as above. If abdomen bluish pruinose, callus is rectangular or square

with a slender dorsal extension. Often appendix at fork .....ccccccee 11

All tibiae entirely black and black haired. Mesonotum prominently

striped. Abdomen when undenuded with silvery pilose median tri-

angles and posterolateral angles on tergites 2-6 ........ maruccii (p. 30)

At least fore tibiae paler at base, generally with some white hairs.

Mesonotum not, or very weakly, striped. Abdomen with or without

Pale DiLOSe: MAT INAS «(x adeceunesascesidesveees HERR wobath Heed bas ana Canes thn vetesnensads 7

Abdomen gray or brown, in unrubbed specimens with median triangles of

pale hairs on tergites 2 or 3-5. Wing spots small, only prominent on

TOP HL TET VID Milde aals PAE DELI iin oh Sas RR Grav oct acbenntns en 8

Abdomen usually black, without marked middorsal pale triangles, or these

represented by a few pale hairs. Wing spots generally large and

prominent at ‘beth ‘fork and:end Of Ciscal ell. .csivdivcevaeiincacvdsenctaneen’ 9

Area of frons lateral to callus denuded, the lower 1/3-1/2 of frons ap-

pearing as a shiny rectangle with a thin shiny ridge extending through

it. Antennae wholly yellowish orange. Legs dark brown to black,

black pilose, except for bases of fore tibiae. Abdomen dark gray in

SEOTIN GL OLEH inns Ga potieed bat cain sedlbltad winebeiednieniaseliaie ee seaal prunicolor (p. 37)

9(7)

11(5)

Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Callus not as above, gradually tapered and clavate. Antennae with con-

trasting blaek annuli or terminal annulus dusky. Legs yellow to pale

brown and dark pilose. Abdomen light brown, rarely nearly black .....

Pau eUG bis ica CARRE IVT s Viss OUcvauascesevareesBelohue <axs jaculator (p. 27)

Antennae with 3rd segment (flagellum) wholly blackish. Tips of all lon-

gitudinal veins with small dark ClOUdS © ........sceccceeee hypographa (p. 25)

Third antennal segment bicolored or wholly yellow-orange. Longitudinal

VEINISINE £ SO SO LEOG ats ioee. waceaeahi rises dab tains beep RA eRvegse speemEg Heed odscceseres 10

Third antennal segment bicolored, annuli contrastingly black. Legs with

brownish or reddish integuMent .....c.ccccccee hypographa neofurva (p. 25)

Third antennal segment yellow-orange, tip often dusky. Legs black, fore

BIG WEEE SSAA SAL Ys ea Sal baccinccececcasesevocdenscseses modica (p. 32)

Abdomen above with distinct pale pollinose and pilose triangles. Frontal

callus nearly always distinet, rounded, not fused into a clavate ridge

Abdomen above without distinet pale pollinose triangles; at most pale

haired triangles or slight middorsal widening of pale pollinose poste-

rior borders evident. Callus often ridge-like .......c..ccccccccccccvccecccees 16

Antennae wholly yellow; basal plate and style short, annuli about as long

as width of basal plate, the latter with only a slight dorsal angle.

Light brown species with all crossveins and fork with weak infuscated

clouds. Whole wing faintly yellowish Drown tinted ......cccccccccccccceccceee

south gl doe sca acdsee ck s beta ges cieavicans ctiprcneeieass xanthicornis (p. 44)

Antennae not as above, the black annuli usually longer than plate is wide,

basal plate yellow to brown, dorsal angle CiStiNCt ...ccccccccccsccceseees 13

Mesonotum with prominent pale pollinose stripes and spots, consisting of

an anterior dorsomedian slender pair of stripes, a dorsolateral pair of

stripes, a pair of spots in the middle of the mesonotum and a pair of

rounded spots before the secutellum. Abdomen dark brown to nearly

black, the middorsal pollinose triangles nearly equilateral, small.

Crossveins and fork heavily infuscated .......ccccee. paraguayensis (p. 34)

Mesonotum weakly or not striped; if striped, these consist of 3 narrow

bare dark stripes or alternating pale and dark stripes, not as above.

WAGs BG BOCOMeN: VATIONS Fie cance a sccvacweccncnevodianewcasensscntorcccccsecsecece 14

Second abdominal tergite with a small middorsal black integumental mark

(not evident in dark specimens). Wing nearly hyaline, costal cell

hyaline or only slightly tinted yellow. Wing spotting faint to virtually

gnideteciailer widlsicn hadinmeSiumsisidenskeelgeroddeniess rubrithorax (p. 37)

Second abdominal tergite rarely with a black integumental mark. Costal

cell yellow. Wing spotting strong to virtually ADSENt ...ccccccescceccceees 15

Wing without spots on fork and crossveins, but all veins broadly though

faintly brown margined. Larger species (wing length 11-12 mm) with

blaek pilose palpi, and small middorsal abdominal triangles .....ccccccece

[Meets se ds LUT onde boeing iuoalaex fulviventris (p. 21)

16(11)

19(16)

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 7

Wing with small but distinet dark clouds at fork and crossveins. Smaller

species (wing length less than 11 mm). Palpi largely or wholly white

pilose, and abdomen with large prominent white pollinose middorsal

CaS AA ARR A eae seythropa (p. 39)

Wing crossveins and fork of 3rd Vein UNMALKEG ....ccccccccsscccsccsscssccssccsscces 17

Wing variously marked, with at least a distinct, though sometimes small,

Crud at fork of Sr Veit wasteesariedesedocecavssedaradeacstete dav beressecasacncuses 19

Basal callus nearly as wide as frons, shiny and protuberant. Upper exten-

sion forming a widened median callus nearly 1/3 width of frons. An-

tenna short and stout, without a strong dorsal angle, bicolored. Frons

broad, frontal index less than 5.0. Wings faintly smoky, costal cell

yellow ;-appendix very. Short)’ ii eallicera (p. 17)

Basal callus small, drop shaped to ridgclike, not widened above Frons

narrower, index over 5.0. Wings often strongly fumose anteriorly.

Antennae yellow or bicolored, the plate with a strong dorsal angle

Abdomen above black haired with broad golden yellow haired posterior

borders, often widened into middorsal golden yellow haired triangles.

Frons broader, index 5.0-6.5. Fore tibiae dark and dark haired .........

giblint sade saieth on deve bey ead Oueueu sla kwadclunssdl ee eeneeeasaare captiroptera (p. 18)

Abdomen above dark pilose with a more or less distinet median row of

narrow middorsal triangles, often forming a narrow stripe, but with-

out broad golden yellow haired posterior borders. Frons narrow, index

6.5-9.0. Fore tibiae bicolored, basal 1/3-2/3 pale with yellow hairs

é RURAL acce senso eel bc PueWts cue didcone lannwoene gree eeena ale aad glandicolor (p. 24)

Brown species, second abdominal tergite with a small dark integumental

spot, and all tergites with narrow yellowish pilose hind margins. Wing

spots on fork and all crossveins large and distinct, though pale .........

TN Pee AU KP USMC URED FeSIIMER Or Oe Ueber: 0 cae ae oeey | OREN lerida (p. 30)

Without such a dark integumental spot. Abdomen and wings various..... 20

Ground color of thorax black, and of abdomen deep reddish brown to

blackish, steel gray pollinose below. Crossveins distinetly spotted and

often tips of longitudinal veins with small clouds. Frons wide, index

3.5-4.3. Posterior borders of abdominal tergites usually narrowly

white haired. Antenna bDicolored .......cccccccececcccecscccccccccces furva (p. 22)

Without the above combination of characters; never with black abdomen..

Ri nahe eh eR peds 44 GAGEEL Gen eee a aE REAL sta ROOESEAS OTe TTR aaTETRERER (AAW OEE RTA OETR CEES on Reb OCR 21

Mesonotum and scutellum black in ground color, contrasting with dull

yellowish brown abdomen. Wings heavily spotted, with yellow costal

cells and extensive anterior and apical brown tinting. Antennae bi-

COTOP SG Cio steac teva ncekses 240 uo cinacede cists chases vuecsib stacey sank panstaratacegertelieranes 22

Mesonotum and scutellum brown to yellowish, not strongly contrasting

with abdomen. Wings and antennae Various .....scccccccscscscscccscscecseees 23

Appendix at fork short, subequal or less than length of subtending vein

segment. Wings heavily spotted, spots at fork and end of discal cell

much larger than those at ends of basal cells. Veins Ro49) Ry and R.

g Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

often broadly brown margined, the brown along R 4 often extending as

a spur treny fonketo wine NEGRO wesc viskelivasieeans isderes spilota (p. 40)

Appendix at fork of 3rd vein long, twice or more length of the subtending

vein segment. All crossveins with narrow dark clouds and membrane

brownish tinted, especially apically. Mesonotum and scutellum with

black integument; abdomen dark yellowish brown with traces of nar-

row pale pilose hind margins on all tergites. Antennae sharply

WI COIOP OE ttl cds Gein ela ep RG Se RG gue changena (p. 19)

23(21) At least the spot at fork of 3rd vein distinct, round; crossveins usually

also with more or less distinet spots. Abdomen at most with narrow

pale pilose hind margins of tergites or small indistinct pale median

hair tufts. Wing membrane hyaline or brownish tinged. Appendix

shorter than long subtending vein SeGment.’sscssewessssevsssapecdccscccsecese. 24

Wing spots weak and indistinct, often barely visible, irregular. Abdomen

extensively yellow pilose. Wing membrane usually at least partly

yellow. Appendix twice or more length of short subtending vein seg-

WT ETE ee eer Ere L rE reer TTT ere TT TT ePEr TTT TTT PTCTT ETT TTT TTT eT TT OCLC TLE ETT ST eeee 20

24 Costal cell yellow. Clouds on fork and end of discal cell large, conspicu-

ous, that on ends of basal cells hardly visible. Wing apex with a

vague fumose patch. Antenna wholly yellow or the terminal annulus

darkened. Abdomen dull yellowish to brown, with vestiges of pale

HAI SEH IGGOPSAl TPIGNEIES Aeiivcsssrsdecedblscapasbie cnsewdebiiccnes kroeberi (p. 28)

Costal cell hyaline. Clouds on fork and end of discal cell small and faint;

none at ends of basal cells. No apical fumose patch, though tips of

first 2 or 3 longitudinal veins lightly fumose. Antennal style black, in

contrast to yellow basal plate. Abdomen as in kroeber ...........sccceeee

Ore PANES RS RIL EES ERATE ATER. 1 Magee Us AEE TUL OSS Se oR LSE Ee ane neg bipuncta (p. 16)

25(23) Antennae bicolored, the black style contrasting with yellow plate. Wings

with clouds on fork and end of diseal cell small and faint, the wing

yellowish tinted, costal and marginal cells yellow, wing apex smoky.

Pleural pilosity gray, abdomen with brownish pilosity but narrow pos-

terior segmental bands of yellowish hairs, slightly widened in the

middle into broad triangles. West of Andes ........00. anoriensis (p. 11)

Antennae all yellow or terminal annulus blackish | ......ccccccccccsscccsccscscsvees 26

26 Clouds on fork and crossveins pale but mostly large and distinct. Mem-

brane of costal, basal, discal and bases of marginal (1st R) and sub-

marginal (3rd R) cells generally strongly yellow, the apex of wing be-

yond end of discal cell largely grayish hyaline, the line between the

colors irregular. In some the apices of Ro and Re are weakly

clouded. Abdomen with broad yellow pilosé bands. 3 pi and legs

OU OUE OE Ria a Vet Set hah PUY, sc lbbad palepbude dvapeeaeianenwa flavescens (p. 20)

Clouds on fork small and weak, generally absent. When present, only evi-

dent at fork of third vein. Wings lightly to heavily brown fumose,

costal cell orange, often yellow streaks in marginal cell. Abdomen

with broad bands and median triangles of yellow hairs, often entirely

VEUIOW PBS Paves Dies cet ahN clue pt yes vadacheeeednaieeds captiroptera (p. 18)

7(5)

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 9

KEY TO STYPOMMISA MALES

Eyes densely pilose, the upper facets but slightly enlarged and not de-

mareated from the lower small facets. Abdomen with median pale

POLNHOSS: LRIANBIOS> mvsivin QR NS ate ei dda ed UA vee ebesanesees 2

Eyes bare, the upper facets greatly enlarged and clearly demarcated from

the: stall facets pADGOMEN VariOUs Aas. ils I. Sb esi ence seasons 3

Pale brown species with wings weakly or not spotted and no appendix at

fork of third vein. A dark integumental spot on dorsum of second

abdominal segment which is otherwise pale yellowish ........scscccscecceees

Rl povnciob ten Maahedeuavadie pels tnedielbiendeagtlsces Lipa Nt he rubrithorax (p. 37)

Dark brown to blackish species with strongly spotted wings and a short

appendix at fork of third vein not as long as subtending vein segment.

Second abdominal tergite largely black, at most with reddish patches

leteratly (alten RUS RR al Ee aS seythropa (p. 39)

Appendix at fork of third vein long, clearly longer than the short sub-

LOTTI: MOTH BOLIC Gi. volte ven ed db devec as cae Sis Soke eae tunes ees ch leudieesccdivblceds 4

Appendix short or absent, if present, equal or shorter than long sub-

POC VET BOM IMCIT clikscsmncgtnakahcesivaeBeied iisdea canis veddeindddssisWeaw cde 5

Wings strongly spotted at fork and all ecrossveins and all veins more or

less brown margined. Coxae with blackish integument. Mesonotum

and abdomen dark brown, mainly black pilose with but traces of pale

pilose segmental bands on AbdOMeN .......sccsccscescceccees changena (p. 19)

Wings with at most a trace of a spot at fork, the fore border more or

less strongly brown tinged but veins not as above. Coxae with pale

yellowish integument. Mesonotum and abdomen orange brown, mainly

yellowish pilose, the former pale gray POL]INOSE .....cccccccccccccccscccccceces

E want aia vba nib BMH Raa ta hd Pee EON nde ey nd SURE E MIEN OH once UE bine captiroptera (p. 18)

MOSONOLUM PPOITITIONTEY BENTO Ciinencoeiiiicvesnddcnendsnaceseqevecvesdsonceeeaieresibebarans 6

DECSONOUUAT LHIG UFO C1 cwssaianinacipeins ad ydicn uaneabankvees $heeebcete en sneasannadiasse sayswemeuaness 7

Black species, the legs wholly black, the wings faintly tinted, with very

small clouds at fork and apex of discal cell and no appendix at fork.

Abdomen black, bluish pruinose on first 2 segments and with white

pilose median triangles and postero- lateral borders of all tergites

IS "Sy lpn hg MERLE OSES ta va nade ee LOMA AMIS sassicsh sno beieceas maruccii (p. 30)

Brown species, the legs yellowish brown, the wings with large conspicuous

brown spots at fork and all ecrossveins, and a short appendix at fork.

Abdomen brown, not pruinose, with a conspicuous middorsal row of

pale pollinose triangles and narrow posterior pale margins on all seg-

THOS AGA AMO EA AR Ra paraguayensis (p. 34)

Wings clear to faintly brown tinted antero-apically, but without appendix

or spots at fork of third vein or crossveins. A light yellowish brown

species with a broad diffuse median yellow pilose abdominal stripe ...

ONE UEREY ST BTEFES tARS Ak ARADO TEE TOW OEE ESIUETER DS, <Oepiini Cc glandicolor (p. 24)

Wings with at least a spot at fork of third vein. Abdomen without a

middorsahistrige Velie Bietdin ce aR ha 8

10 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

8 Wings without an appendix at fork of third vein. Abdomen with at least

traces of pale pruinosity on first 2 tergites and some pale pilosity on

hind margins of at least the posterior tergites .......cccccsscscscsssscecscees 2)

Wings with a short to long appendix at fork. Abdomen without anterior

prumosity and wholly, dark DiLOSC: sasanssakcudvewweincs spuaweedl dipda vusderanus awe 10

9 Blaek species with black legs, black abdomens, wholly yellow antennae

and strong blackish spots at fork and crossveins at distal end of discal

eell. Costal.cellidark yellows eave. cdlvc aolaata saws modica (p. 32)

Brown species with brown legs, brown abdomens, bicolored antennae and

weak spots at fork and only a trace of spotting at end of diseal cell.

Gpetalieell pale VelOW.« wiccsciseesecsiecansencsannesbcectvaCbadalons jaculator (p. 27)

10 Wings with a strong dark apical patch which includes fork of 3rd vein and

is sharply demarcated from entirely hyaline basal half of 1st sub-

marginal cell (3rd R). Appendix short, shorter than long subtending

vein segment Pleural pilosity DrOWN ...scccccscoeccscescnsceees apicalis (p. 13)

Wings with a weak apical patch. Appendix usually longer than very short

subtending vein.se@ment,.. Pleural pilosityeG@ray vas «csicdsaceveecscoscescesecseacs

Lb caches tenella csuand Seauadko<oluvstawwian si pequeniensis or anoriensis (pp. 35, 11)

DESCRIPTION OF SPECIES

Stypommisa affinis Kroeber

Stypommisa affinis Kroeber 1929. Zool. Anz. 83(1-4): 116-117; holotype

female, Espirito Santo, Brazil (destroyed). Fairchild 1967. Stud. Ent.

9(1-4): 360, fig. 26 (designated lectotype) (Vienna); 1971, Cat. Dipt.

Amer. S. of U.S. Fase. 28: 81; 1975. Proc. Ent. Soe. Washington 77(2):

264.

A small reddish brown species with the wings spotted at the crossveins and

fork and darkened at the fore border. Callus as wide as frons below. Abdominal

tergites with pale hind borders and small pale median triangles.

Female. We have no specimens of this species before us but Fairchild (1967)

discussed and figured it. The following is a translation of the original de-

scription. Length 11-12.8 mm, antennae 1.2-1.3 mm, wing length 12.4-12.9 mm,

wing breadth 4.2-4.3 mm. Extraordinarily like S. punctipennis, but the palpi are

short black-haired. Frons and frontal triangle (subeallus) shiny brownish yellow,

face whitish gray. Eyes unbanded. Frons narrower below than above. Callus tall

rectangular, above with reddish brown pointed furrow, hardly separated from red

brown median callus. Ocellar tuberele clearly marked, black. Vertexal triangle

shining red brown. Thorax entirely red brown with a fine blackish mid-line and 2

whitish lateral stripes. Seutellum paler, white tomentose, pale haired. Pleura

pale red brown. Legs pale red brown, including femora and fore coxae, which are

thickly white tomentose and white-haired. Abdomen entirely brown red, hind

marginal bands moderately broad, enlarged into weak median triangles. All veins

somewhat brown bordered at their ends. Espirito Santo.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 11

Male. Not known.

Material examined. Brazil: 2 females, lectotype and paratype (Vienna); Minas

Gerais State, 1 female, Sete Lagoas (Copenhagen).

Discussion. The above description is not accompanied by a figure and omits

mention of antennae, proportions of frons, or spotting of wings. These supposedly

may be inferred by comparison to S. punctipennis. Specimens determined as S.

affinis by Kroeber in Munich and Warsaw belong to 2 different species, neither

agreeing with paratypes in Vienna. The material agreeing with the Warsaw mate-

rial is described herein as S. xanthicornis n. sp. The holotype was in Kroeber's

collection and destroyed during th war. Since Kroeber labelled at least 2 and

possibly 3 distinet forms as affinis, a question arises whether the lost type was

the same as the paratypes in Vienna. One of these paratypes was labelled lecto-

type by Fairchild (1967).

Based on Fairchild's notes, the lectotype has a frontal index of 6.0, and the

fork of the 3rd vein has a short appendix, all crossveins and tip of anal cell with

small but dark clouds, and the antennae rather slender, yellow with a contrasting

style. The fore border of the wing is also darkened. Fairchild did not note a

shiny subeallus - "stirndreieck...schimmernd"- in any of the Kroeber material

studied. The statement in the original description may only indicate a much de-

nuded specimen.

Fairchild also determined 2 females from Poco Alegre, M. G., Brazil, E. Ra-

bello coll. as S. affinis but as these are not now before us he may very likely

have misdetermined them. The figure in Fairchild (1967) was drawn from the

specimen from Sete Lagoas, now in Copenhagen; it agreed closely with the lec-

totype in Vienna.

We are unable to distinguish this species with certainty from S. seythropa or

S. paraguayensis, and have omitted it from the key.

Stypommisa anoriensis n. sp.

(Figs. 4-6)

Stypommisa n. sp. Wilkerson 1979, Cespedesia 8(31-32): 345-356. fig. 34G,

A slender orange brown species with usually bicolored antennae, yellow legs,

yellow pilose abdominal bands and orange brown fumose wings with small clouds

on fork of 3rd vein and end of diseal cell.

Female. Holotype. Length 11 mm; of wing 10.5 mm. Eyes bare, in life with

lower 1/2 olive drab, upper 1/2 brownish. Head structures as figured. Frons and

subeallus light brown pollinose, the former with short dark proclinate pilosity.

Frontoclypeus and genae light gray, the sparse beard of long whitish hairs mixed

with shorter black hairs. There are also sparse gray hairs seattered over the

frontoclypeus. Frontal callus dark brown, nearly black, shiny. Tubercle at vertex

small, discrete, shiny, with vestiges of at least the anterior ocellus, set at the

lower end of a diffuse shiny stripe which extends over the vertex onto the oc-

ciput. A postocular fringe of several rows of erect black setae is present, dens-

est in back of frons. Antennae with first 2 segments orange, grayish pollinose,

fairly thickly beset with short black setae dorsally, more slender pale setae ven-

trally. Basal plate of 3rd segment bright orange yellow contrasting with the

12 Contrib. Amer. Ent. Inst., vol. 22, no. 5,.1986

densely velvety black terminal annulate portion. Palpi yellowish gray pollinose,

beset with black hairs, except at base below and on lst segment, where they are

yellowish brown and longer. Proboscis dark brown, slightly longer than palpi,

without prominent sclerotized plates in labella.

Mesonotum and scutellum concolorous, dark orange brown, thinly grayish

brown pollinose dorsally becoming denser and grayer towards front and sides.

Pronotal and notopleural lobes slightly paler, gray pollinose. Pilosity of dorsum

of sparse erect dark hairs and recumbent yellow hairs, the latter forming ill-de-

fined stripes. Notopleural pilosity dark, erect. Pleura with integument concol-

orous, but thickly pale gray pollinose and sparsely whitish pilose. Coxae concol-

orous with pleura, white pilose. All tarsi slightly darker. Wings with basicosta

pointed, the black setae slightly less dense than on adjoining costa. Fork of 3rd

vein with an appendix about length of subtending vein segment in left wing, but a

mere stub in right. Costal cell yellow, remainder of wing yellowish brown, the

color most intense along fore border beyond stigma, slightly accentuated along

veins, and with small diffuse slightly darker clouds around fork and crossveins at

end of discal cell. All veins yellowish orange, the setae, where present, black.

Halters with yellow stem and oval orange brown knob.

Abdomen shiny yellowish or orange brown, subtranslucent basally, darker api-

eally. Pollinosity is thick, concolorous, the pilosity dorsally consists of sparse

erect black hairs except for sparse narrow yellow pilose posterior bands on seg-

ments 2 to 5, slightly widened in the middle on some, and represented by sparse

tufts or patches of yellow hairs on segments 1 and 6. These bands are widened at

the extreme sides to cover most of the length of the segments. The abdomen

beneath is pale yellowish pollinose with sparse pale yellow pilosity throughout its

length.

Male. Not known; on further study those listed by Wilkerson (1979) proved to

be S. pequeniensis.

Type data. Holotype female, Colombia, Antioquia Dept., Rio Anori Valley,

La Tirana, 31-V-1971, elev. 570 m leg. C.H. Porter. The specimen also bears a

note as to its eye color, copied above, and a reference to Porter's field notes

reading CHP-71-220. JO4. To be deposited in FSCA. Paratypes, 144 females,

all from western Colombia, as follows: Choco Dept., 16 females, Rio San Juan-

Baudo, II-1976 (R. Wilkerson). Antioquia Dept., 8 females, Rio Anori Valley, La

Tirana, IV-VI-1971 (C.H. Porter). Valle Dept., 1 female, Rio Raposo, Buenaven-

tura, XII-1963 (V.H. Lee); 1 female Buenaventura, 7-X-1964, human bait (V.H.

Lee); 61 females, Lower Anchicaya Dam, I, III, IV, VII, VIII, IX, and XI-1975 (R.

Wilkerson); 2 females, Anchicaya Dam, 10-VIII-73, 500 m elev. (Wilkerson and

Young); 6 females, 41 km E Buenaventura, XI and XII-1975, III-1976 (R. Wilker-

son); 7 females, Rio Bravo, Playa Rica, below dam at Lago Calima, XII-1975, I

and II-1976 (R. Wilkerson); 2 females, Ladrilleros, 30 km N Buenaventura, I-1975

(R. Wilkerson); 12 females, Rio Sabaletas, 4km E Sabaletas, VI, VII, VIII, and X-

1975 (R. Wilkerson); 29 females, Lago Calima, 8km below dan, I, III, IV, VII, IX

and XI-1975 (R. Wilkerson). Paratypes are in FSCA and the authors’ collections.

Some of these will be distributed to CAS, USNM, BM(NH) and CNC.

Discussion. The paratypes are quite variable in intensity of color, size (wing

length 8.5-12 mm) length of appendix at fork of 3rd vein, and frontal index.

Among 25 specimens from western Colombia measured, the frontal index varied

from 4.8 to 5.9 (x = 5.3), the index of divergence from 1.2 to 1.6 (x = 1.4). Other

frontal indices are as follow: Antioquia Dept. (n = 9), 4.9-5.7 (x = 5.2); coastal

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 13

Valle Dept. (n = 10), 4.7-6.8 (x = 5.2); and Lago Calima in Valle Dept. 5.3-5.9 (x =

ona).

This species differs from S. pequeniensis in extensively yellow pilose hind

marginal dorsal abdominal bands, in more yellowish wings with small but distinct

cloud on crossveins at end of discal cell and fork of 3rd vein, and in lacking a

demarcated apical infusecated patch on wings. The antennae are usually sharply

bicolored in S. anoriensis less clearly so or unicolorous in S. pequeniensis. From

S. flavesecens the species differs in bicolored antennae, less well marked clouds

around end of diseal cell, less extensively yellow pilose abdomen, and whitish gray

rather than yellow pleural pilosity. See also discussion under S. flavescens.

Stypommisa apicalis n. sp.

(Figs.. 1-3, 7-9, 72)

A relatively large brown species with narrow convergent frons and wings with

very conspicuous apical patch, similar to members of genus Diachlorus.

Female Holotype. Length 12 mm, of wing 12 mm. Eyes bare, green with 2

narrow blackish stripes and upper and lower margins broadly purple (relaxed).

Head structures as figured. Frontal index 7.5. Frons dull yellowish pollinose, be-

set with short black hairs, longest and densest at vertex. Occipital or post ocular

hairs black, longest in middle behind vertex. Ocellar tubercle with vestiges of a

median yellow ocellus. Frontal callus orange brown. Subcallus concolorous with

frons, appearing paler because of lack of black hairs. Frontoclypeus and genae

slightly grayer than subeallus, with a sparse brownish beard. Antennae orange

brown. The first 2 segments beset with short black hairs, the 3rd segment dark-

ening gradually apically beyond dorsal angle, but style not black and not con-

trasting with basal plate. Palpi dull orange, almost wholly black haired, slightly

shorter than the brown membranous labella.

Mesonotum, scutellum and pleura dull reddish brown, thinly brown pollinose,

mostly sparsely dark haired, but with tufts of whitish hairs on pronotal lobes and

beneath wings, and seattered short yellow hairs dorsally. Basicosta acutely

pointed with at most 2 or 3 strong setae. Wings as figured, the costal cell and

stigma strongly yellow, the crossveins at ends of basal cells with small but dis-

tinet brown clouds, a large and distinet cloud at end of discal cell and the cloud

at fork of 3rd vein lost in the strong infuscation that fills the wing from fork to

apex. Fork with an appendix as long as stem. Legs dark reddish brown, almost

wholly dark brown to black haired, fore tibiae not bicolored.

Abdomen concolorous with thorax on first 2 or 3 tergites, thereafter be-

coming darker, almost blackish, dorsally wholly black haired, ventrally clothed

entirely with short yellowish white hairs, more conspicuous on hind margins of

sternites and posterior lateral corners of tergites.

Male Allotype. Length 11 mm, of wing 10 mm. Eyes holoptic, bare, the area

of enlarged facets reddish, occupying about 2/3 of total eye area, the large

facets well differentiated and demarcated from the small. A pollinose tubercle

sunk between eyes at vertex. Postocular setae longer than in female, curved

forward. Antennae more slender than in female, palpi brown, subfaleate, beset

with long brown hairs. Color of body, legs and wings as in female, the apical

dark patch paler than in holotype, but not paler than in some female paratypes.

14 Contrib. Amer. Ent. Inst., vol. 22, no. 5,1986

Type data. Holotype female, Peru, Cuzco Dept, Quincemil, 2450 ft elev. 16-

31-X-1962, (L.E. Pena). To be deposited in FSCA. Allotype male, same data. In

FSCA. Paratypes. Peru: 14 females, same locality as holotype, IX-XI-1962;

Madre de Dios Dept., 5 females, Avispas, 400 m elev. X-IX-1962, (L.E.

Pena\(there are long series from both the above localities in CNC and CUC); 3

females, Tambopata Reserved Zone, 30 km SW Pto. Maldonado, 290 m elev., 6-

10-XI-1979, (J.B. Heppner), 5 additional females from this collection are in

USNM; 207 females, 5 males, same locality, X-XI-1982 flight trap, (R. Wilkerson).

Brazil: Rondonia Terr., 1 female, highway BR319, km 28 24-X-1980 (J. Arias).

Mato Grosso State, 1 female, X-1976 (Alvarenga leg). Para State, 1 female,

Itinga, Paragominas B.R. 14-I- 1965 (L. Gomes). Bolivia: 4 females, Palos Blan-

cos, Alto Beni(river?), 11-15-1976, 600 m elev. (L.E. Pena). Cochabamba Dept., 1

female, Villa Tunari, 240 m elev., 3-6-IX-1983 (R. Wilkerson). Paratypes will be

deposited in the above mentioned collections and in CAS, INPA, Manaus; Mus.

Goeldi, Belem; BM(NH) and collections of the authors.

Discussion. Paratypes range in wing length from 10.0 to 13.5 mm and in body

color from cinnamon brown with light brown legs to nearly blaek with slightly

bluish black abdomens and black legs. There is considerable variation in the ex-

tent and intensity of the wing pattern, specimens from Cuzco Dept. Peru being

the palest while some from Madre de Dios Dept. are usually darker. The few

Brazilian specimens seen are quite dark, while the 4 from E. Bolivia are like

those from Cuzco. Extremes are quite different appearing, though all in-

termediates occur in our long series from Rio Tambopata. The males from Tam-

bopata are quite uniform, dark brown with nearly black legs and largely black pi-

losity. The allotype from from Cuzco Dept. is lighter, like females from the

same locality. Antennae of all have the same distinetive shape, with the dorsal

angle of basal plate near the middle of the plate. The antennae may be dark

brown to light orange with the style contrasting or not, or in nearly any combina-

tion.

The species is abundantly distinct from all other described species in its com-

bination of ridgelike callus, unique antennae,banded eyes and strong apical wing

pateh. Only S. pequeniensis has a weak apical wing patch, but is smaller and

with different frons and antennal shape, as well as having bicolored fore tibiae.

S. spilota n. sp. has rather heavily marked wings, but the clouds on fork and

ecrossveins are much larger and more prominent, the apical pateh faint and dif-

fuse, the fore tibiae clearly bicolored, and the antennae and frons quite different.

All specimens seen have been taken in the months of September to November

and January, either netted or in flight traps, apparently always in areas of high

rainfall.

Stypommisa aripuana n. sp.

(Figs.. 10-12)

A small brown species with a narrow frons, bicolored antennae, white haired

palpi and yellowish brown legs. Wings hyaline, unspotted. Abdominal integument

dark brown, the Ist two segments pale, semi-translucent, with a middorsal row of

small pale triangles, and when viewed at an angle from behind, with rows of large

sublateral pale pruinose spots and a row of middorsal pruinose triangles on seg-

ments 1 to 5.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 15

Female Holotype. Length 10.5 mm, of wing 9.5 mm. Eyes bare, pattern not

recorded, probably greenish black in life. Head characters as figured. Frontal

index 4.7, index of divergence 0.91, antennal index (basal plate/style) 1.25.

Frontal callus yellowish brown, its surface rugose. Ocellar tubercle flat, bare,

not protuberant, without clear ocelli, surrounded by a diffuse area of dark polli-

nosity. Remainder of frons dark yellowish pollinose, without pilosity. Postocular

setae short and pale, in several irregular rows. Subcallus thinly pollinose, slightly

paler than frons, its median groove widened in the middle and with a small bare

area. Frontoclypeus and genae with reddish integument, gray pollinose, very

sparsely gray pilose. Pedicel and scape orange, grayish pollinose, dark pilose

above, paler below. Basal plate orange, slightly dusky at distal end, the style

deep black. Palpi yellowish orange, gray pollinose, both segments wholly pale pi-

lose, the hairs rather short. Proboscis with stylets 1/2 longer than palpi, the

theea blackish, subshiny, the labella brown, membranous.

Mesonotum with brown integument, subshiny dorsally, yellowish gray polli-

nose, with vague indications of a broad median stripe and dorsolateral darker

Stripes and a darker median streak on posterior third. Pilosity of short sparse

yellowish and dark hairs, forming vague stripes. Scutellum concolorous, its apex

paler, reddish, thinly gray pollinose, sparsely yellowish pilose. Pleura and coxa,

including notopleural and pronotal lobes, slightly paler, gray pollinose, sparsely

whitish pilose. Legs with entirely yellow integument, the coxae and femora

wholly pale pilose, the tibiae pale pilose on mid pair, dark pilose on fore and hind

pair, except basally on fore pair. All tarsi dark pilose. Wings hyaline, the costal

cell weakly yellowish tinted, stigma yellow, veins yellowish brown, no appendix at

fork of 3rd vein, and only the faintest indication of dark clouds at fork or any

crossveins, not visible without high magnification. Basicosta pointed, with 1 or 2

black setae. Halters brown, the stem slightly paler.

Abdomen with integument of first 2 tergites semitranslucent yellow at sides,

the first dusky beneath scutellum, the second with a broad median inverted V-

Shaped dark brown mark, and posterior border brown. Remaining tergites brown

with paler hind borders and narrow lateral margins. Pollinosity is thin, dark

basally and pale on margins of tergites, and forming large median pale triangles

on tergites 2 to 6, plus a small spot on 1. Pilosity sparse and short, dark on dark

areas, pale on pale areas. Superimposed on this pattern, only visible from di-

rectly behind, is a pattern of white pruinosity consisting of large dorsolateral

connected patches forming a weakly interrupted pair of wide stripes and a median

row of unconnected triangles. The hind borders of all tergites are also white

pruinose. Beneath the abdomen is entirely whitish pollinose and sparsely pale pi-

lose.

Male. Not known.

Type data. Holotype female, Brazil, Mato Grosso State, Rio Aripuana, Hum-

boldt, 99° 27'W, 10°10'S, 12-16-VIII-74, flight trap (D.G. Young). To be deposited

in Museu Emilio Goeldi, Belem, Para, Brazil.

Paratypes, 60 females, same data as holotype, In FSCA, MCZ, CNC, BM,

USNM, CAS and collections of the authors.

Discussion. We had at first thought this taxon was but a variant of

S.rubrithorax to which it is very similar. It differs, however, in consistently nar-

row frons, index 4.6 to 5.0 (x = 4.8, n = 10), a slightly less divergent frons, index

0.9 to 1.3 (x = 1.05), and relatively longer antennal style in relation to the basal

plate of antenna, index 1.2 to 1.5 (x = 1.3). It also entirely lacks an appendix at

16 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

fork of 3rd vein and seldom shows any trace of a wing spot there or elsewhere.

The abdomen when viewed from behind shows a striking dorsolateral row of large

white pruinose spots, totally lacking in any other species we have seen. The

paratypes vary somewhat in intensity of coloring and size, wing length varying

from 8.0 to 9.5 mm.

Stypommisa bipuncta Fairchild

(Figs.. 13-15, 73)

Stypommisa bipuncta 1979, in Wilkerson 1979, Cespedesia 8(31-32): 334-

335, Colombia, Valle, Rio Frio (MCZ).

A small pale brown species with the hyaline wings bearing two small brown

spots, on fork of 3rd vein and on crossveins at apex of discal cell.

Female. Length 9.5 mm, of wing 10 mm. Eyes bare, color in life not

recorded. Head structures as figured. Frontal index 5.4. Frons yellowish gray

pollinose, darker and sparsely black-haired at vertex. Frontal callus and median

ridge shiny, dark brown, as is ocellar tubercle. Subeallus coneolorous with frons,

frontoclypeus and genae paler, more grayish. The sparse beard whitish. Palpi

yellowish tan, beset with short black hairs. Antennae orange yellow, the scape

and pedicel beset dorsally with reddish brown hairs, the style black, contrasting

with basal plate. Proboseis brown, membranous, the stylets longer than the palpi.

Mesonotum pale cinnamon brown in ground color, the seutellum yellowish at

apex, both beset with mixed semi-erect black hairs and recumbent shiny pale

hairs. Pleura paler, whitish pollinose and white haired. Legs yellowish brown,

the foretibiae apically darkened and black haired, the foretarsi blackish. Mid legs

missing. Wing including costal cell hyaline, slightly brownish tinted in anteroapi-

cal area, and with pale brown clouds around fork of 3rd vein and crossveins at

apex of discal cell, but not on basal crossveins. Appendix at fork slightly shorter

than long subtending vein segment. Basicosta with sparse black macrotrichiae.

Abdomen yellowish brown, subshiny, denuded, beset with short dark hairs

above, pale hairs beneath. Tergites 4-6 bear remnants of pale-haired middorsal

broad triangles, apparently extended laterally on hind borders of segments.

Male. Not known.

Material examined. Holotype female and three female paratypes, Colombia,

Valle del Cauca, Rio Frio, La Carmelita 13-VI-1942 (J. Renjifo).

Discussion. Only the holotype bears a date, and it is the only specimen re-

taining a whole antenna. Some of the paratypes are otherwise better preserved,

showing the abdomen with a dorsal row of broad median sparsely pale-haired tri-

angles. In one the frontal calli are black. One wing of the holotype is slide

mounted No. St-36. We have seen no additional specimens.

The species differs from S. venosa as treated here, in having a discrete basal

frontal callus rather than an evenly tapered clavate Keel, in pale forecoxae, and

in nearly wholly glass clear wings except for the diserete small clouds on fork of

3rd vein and apex of discal cell. In S. venosa there-is considerable diffuse

brownish infuseation of the wings, larger clouds on fork and apex of discal cell,

Small clouds around ends of basal cells and a yellow costal cell.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 17

Stypommisa ecallicera (Bigot)

(Figs. 16-18)

Tabanus callicera Bigot 1892, Mem. Soc. Zool. France 5: 686 (BM(NH)).

Fairchild 1956, Smiths. Miscell. Colls. 131(3): 12; 1967, Pacific Ins.

9(2): 244, fig. 3 (Places in Stypommisa).

Stypommisa callicera: Fairchild 1971 Cat. Dipt. Amer. 8S. of U.S., Fase.

28: 81.

A small unmarked light brown species with wings unspotted but lightly

brownish tinted in costal cell and anteroapically, frons broad for the genus and

with stumpy bicolored antennae.

Female. ‘Translation of original description: Female, length 9.5 mm. An-

tennae tawny, the third segment black at extremity, without coneavity above and

with an obtuse very little projecting tooth. Palpi tawny, brownish above and at

the extremity. Face dull reddish sprinkled with whitish hairs. Beard short,

white. Frons dull gray, callosity black, narrow, linear, abruptly dilated and nearly

quadrangular below. Vertex with very small ocelli. Thorax and scutellum reddish

tawny, pleura with whitish pollinosity. Calypters and halters reddish brown, the

club pale tawny at its extremity. Legs reddish, extremity of tibiae and tarsi

black. Wings very pale grayish, stigma a little reddish, fork of 3rd vein with very

short appendix. Brazil, one specimen.

Male. Not known.

Material examined. Brazil. Rio de Janeiro State, 1 female, Rio de Janeiro,

II-1939; 1 female, Teresopolis, Parque Nacional da Serra dos Orgaos, 1000 m elev.,

17-IX-1948 (P. Wygodzinsky)

Discussion. The type in British Museum is somewhat rubbed and shrunken,

perhaps collected in aleohol. There are sparse macrotrichiae on the basicosta.

The head characters are as figured by Fairchild (1967), the mesonotum light cin-

namon brown, pleura gray pollinose and white-haired. Fore tibiae obscurely bi-

colored, due mainly to vestiture rather than integumental color, the remaining

legs cinnamon brown with darker tarsi. Abdomen yellowish brown, completely

denuded. |

The female from Rio was compared to the type in 1964 and labeled "not in

agreement though close." It measures 7.5 mm in length. The female from Tere-

sopolis measures slightly over 10 mm in length and agrees both with Bigot's de-

scription and with Fairchild's (1967) figure and brief description. We believe now

that the Rio specimen above is probably only an undersized specimen of the

species. The eyes (relaxed) are without pattern, probably greenish in life. Both

specimens above have the yellowish brown abdomen darkened apically, apparently

due to a blood meal, almost entirely but sparsely clothed with short dark hairs,

but with a few pale hairs on the midlines of some tergites. The fore femora and

apical halves of foretibiae are quite dark, thickly black haired, contrasting with

the white haired basal halves of foretibiae in the Teresopolis specimen, though in

the Rio specimen this is not so evident. In both specimens the wings are lightly |

yellowish brown tinted, the stigma darker anteriorly and with the costal cell

darker yellow, the stigma orange brown. There is a short appendix in the wings

of both specimens, but no vestige of wing spotting. The frontal indices are 4.0

and 4.4, the divergence indices 1.1 and 1.2, and the antennal indices 1.4 and 1.6.

18 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

The very wide, square basal callus, nearly as wide as frons, and the somewhat

basally inflated palpi suggest that this species may be related to Stenotabanus,

though the basicostas bear a few macrotrichiae in both specimens.

The species seems most similar to S. fulviventris in general appearance, but

is smaller, frons broader, antennae shorter and strikingly bicolored, and the ab-

domen appears to lack all but faint traces of pale median triangles or pale polli-

nose hind margins of tergites, evident in all specimens of S. fulviventris seen.

Stypommisa captiroptera (Kroeber)

(Figs. 19-21, 74)

Tabanus (Macrocornus) captiropterus Kroeber 1930, Zool. Anz. 87: 10, fig.

8, holotype female, Venezuela (Vienna) [examined].

Stypommisa captiroptera: Fairchild 1966, Stud. Ent. 9(1-4): 364; 1971,

Cat. Dipt. Amer. S. of U.S. Fase. 28: 81. Hogue and Fairchild 1974,

Rev. Biol. Trop. 22(1):22, Costa Rica. Wilkerson 1979, Cespedesia

8(31-32): 355-337, fig. 112, female, male, Colombia. Philip 1977,

Proc. Ent. Soe. Wash. 79(1): 28.

Stipommisa (sic) flavescens Barretto 1957, Rev. Brasil. Malar. (1956),

8(1): 82, male, British Guiana. Not flavescens Kroeber 1930.

Stenotabanus xenium Fairchild 1947, Ann. Ent. Soe. Amer. 39(4): 568, pl.

1, fig. 5,a,b, female, Panama (MCZ).

A light yellowish brown species, the wings rather evenly fumose, darkest

along foreborder and apex, the fork of 3rd vein with an appendix longer than sub-

tending vein segment and with a small, usually discrete, dark cloud, often faint,

but not clouds on apices of discal or basal cells. Pilosity of pleura and abdomen

yellow below, and forming broad segmental bands and large middorsal triangles

above. Legs yellow, mainly yellow pilose, except for usually dark pilose hind

tibiae and all tarsi.

Material examined. Fairchild has studied Kroeber's type in Vienna and the

male from British Guiana reported by Barretto. Both of us have studied the fol-

lowing specimens: Nicaragua: 1 female, Eden, 4-VI-1922 (W. Huber). 1 female,

Jinotega, Rio Potecas, 10-VIII-1953 (J. Boshell). Costa Rica: 1 female, Finca

Margarita, V-1938. Limon Prov., 2 females, Finca La Lola, VII-1963(Hogue); 1

female, Guapiles, 18-VI-1965. Panama: Bocas del Toro Prov., 1 female, Almi-

rante Yellow Fever Sta., 16-V-1951 (H. Trapido); 2 females, Miramar, UV light

trap, 29-V and 11-VII-1979 (H. Wolda). San Blas Prov., 6 females, Rio Mandinga-

Rio Tangandi Yellow Fever camp, 1-27-V-1957 (P. Galindo). Darien Prov., 1 fe-

male, Cana, VII-1965 (O.R. Henriquez); 1 female, Rio Canita, Rio Bayano, 13-

VIII-1949, at light (P. Galindo). Colon Prov., 1 female, Rio Pequeni, 23-VI-1944,

(Fairchild). Colombia: Choco Dept., 1 female, Teresita, 26-VI-1967 (D.G. Young

leg.); 1 female, Curiche River, 4-VIII-1967 (D.G. Young leg.). Antioquia Dept., 2

males 1 female, Rio Anori Valley, La Tirana 9-111-1971 (C.H. Porter); 1 fe-

male, vic. Zaragoza, IX-1970 (D.G. Young). Valle Dept., long series from various

localities as listed in Wilkerson 1979 under S. eaptiroptera. Ecuador: Pichincha

Prov., 1 female Tinalandia, E of Alluriquin, 4-8-V-1976 (T.E. Rogers); 2 females,

Quito, 1930, (R. Benoist). Peru: Cuzeo Prov., 1 female, Quincemil, 16-31-X-1962

(L. Pena). Amazonas Prov. 1 female, Iquitos, III and IV-1931 (R. C. Wilkerson).

Bolivia: Alto Beni. 2 females, S. Inicua river, 15-18-1-1976, 1100 m (L.E. Pena).

Brazil: Para State, 13 females, Municipio Benevides, Fazenda Morelandia, 23-VI-

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 19

1981, horse bait and flight trap (G.B. Fairchild); 1 female, Belem, 7-8-VI-1971;

1 female, APEG Forest, Belem, VII-VIII-1974 (D. G. Young); 1 female, Bujaru, 3-

XII-1979 (R.B. Neto); 1 female, Belem, 1949, (R. Damasceno leg.); 1 female,

Marajo ld., no date. 1 female, VII-1931 (R. C. Shannon). Amazonas State, 9 fe-

males, Manaus, Reserva Ducke, I, II, II, V, VII-1978 (J. Arias and N. Penny).

Rondonia State, 3 females, Vilhena, 6-XI-1979 (J. Arias). In addition Fairchild de-

termined a specimen from Chiapas Mexico, as Stenotabanus xenium for Philip, as

reported by him (1977).

Discussion. As can be seen by the above, the species ranges widely, from

Mexico, Nicaragua and Honduras south to Ecuador west of the Andes and to Bo-

livia and Para, Brazil, east of the Andes. There is some geographical variation,

specimens from Para and the Amazon basin tending to be darker than those from

west of the Andes and Central America. The species usually can be separated

from S. pequeniensis by the yellow abdominal pilosity and lack of a clear demar-

cation of the dark apical wing patch, and from S. flaveseens by lacking any trace

of dark clouds at the apex of discal cell. Stypommisa anoriensis is very similar in

wing and body color, but the antennae of S. anoriensis are smaller, shorter, and

Sharply bicolored, with yellow basal plate and black style.

The male was described by Wilkerson (1979), who also figured the wing of the

female, while Fairchild (1947) figured the head characters of the female. There

is some indication that the species is ecrepuscular or even nocturnal, as notations

on labels often state that the specimens were taken at light or in the evening.

The eyes in life are unpatterned, dull green to golden bronze, rarely with a faint

median dark stripe.

Stypommisa changena n. sp.

(Figs. 22-24, 75)

A moderate sized dull brown species with wings both spotted and the veins

brown-margined, and the abdominal tergites with obscure pale hind margins or

small median pale hair triangles.

Female. This species is best described by comparison with S. lerida. It is a

more slender species and the overall coloration is darker. The scutellum is

darker blackish brown, (with at least the apex usually red in S. lerida). The frons

is slightly narrower in S. changena (frontal index 5.3-6.2 [n = 8, x = 6.0] vs 4.5-

5.3 [n = 7, x = 4.9] for S. lerida). The indices of divergence (frontal width at

vertex divided by frontal width at base) are not significantly different, ranging

from 1.1 to 1.5 in both species. The wings of S. changena have the appendix usu-

ally longer than the subtending vein segment due to shortness of the latter, while

in S. lerida the vein segment is often longer than the appendix. The wings in S.

changena have a noticeable anteroapical brown tinting and all the veins are nar-

rowly brown margined. In S. lerida the wing is lightly and evenly tinted without

noticeable darkening apically and the veins not obviously brown margined.

Stypommisa lerida has a prominent but diffuse median blackish integumental spot

on abdominal tergite 2, wholly lacking in S. changena, where the abdomen is uni-

formly dark brown. Stypommisa changena has but faint traces of pale hair

fringes on the tergites, while these are prominent in S. lerida. The two species

have different times of flight, February, March and April for S. lerida, May to

November for S. changena. The eyes of both species are green with a single pur-

plish transverse stripe.

20 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Male. Color and wing pattern as in the female. Eyes with enlarged upper

facets bare, well differentiated from small facets, but without a sharp line of

demarcation, the larger facets occupying about 2/3 to 1/2 total eye area. Small

facets extend in a narrow band to vertex behind and there is a small tubercle

sunk to eye level at vertex with vestige of at least the anterior ocellus. The oc-

ciput bears a sparse fringe of long forwardly curved dark hairs.

Type data. Holotype female, Panama, Bocas del Toro Prov., Rio Changena

Yellow Fever Camp, 2400 ft. elev., 26-IX-1961, (G.B. Fairchild). Allotype male,

Panama, Chiriqui Prov., La Fortuna, 10-VI-77, blacklight trap (H. Wolda)

[collected in alcohol]. Paratypes: 47 females same locality as holotype, various

dates in August and September 1961, human bait and Shannon trap. Panama:

Bocas del Toro Prov., 3 females, Rio Claro-Rio Changena, 910 m elev., 20-21-V-

1966 (C. Meyers); 3 females, Rio Uri, Changuinola, VIII-IX-1968 (R. Hartmann); 1

female, Robalo trail, 6500 ft, 6-VIII-47. Chiriqui Prov.,l female, Calderas-

Chiriquicito trail, 600-1400 ft elev., in tree tops, 26-X-5-XI- 1955 [genitalia on

Slide RW-300]; 3 males 5 females, same locality as allotype, U.V. light trap, IV-

VIU-1977 and VII-VIII-1978. Costa Rica: Puntarenas Prov.,l male 1 female, Mon-

teverde area, 1400-1700 m elev., 6-14-VI-73 (Erwin and Hevel); 1 female, same

locality, 21-V-76 (Wasbauer). Alajuela Prov., Chomogo area, 1620 m elev, 13-VI-

73 (Erwin and Hevel). Cartago Prov., 1 female, Hacienda Moravia de Chirripo, 7-

10-VII-1964 (R. McDiarmid) [genitalia on slide RW-299]; 1 female, Carillo, no

date (C. F. Underwood) (wing on slide St. 32). We have also seen 19 more speci-

mens from Fortuna, Chiriqui which are in too poor condition to be labelled as

paratypes. There is also a female paratype, Costa Rica, Heredia Prov., 19-20-

VII-1975, Fisher coll. in C.B. Philip's collection in CAS, and 26 females from

Monteverde area in U.C., Berkeley which we believe are this species, though they

are not now before us. :

Stypommisa flavescens (Kroeber)

(Figs. 25-27, 76)

Stypommisa flavescens Kroeber 1930. Zool. Anz. 86: 249, holotype female

"Peru" (destroyed).

Stypommisa flavescens: Fairchild 1967, Stud. Ent. (91-4): 359 (1966),

selected neotype (Munich); 1971, Cat. Dipt. Amer. S. of U.S., Fase.

28: 81; 1975; Proce. Ent. Soe. Washington, 77(2): 264, Ecuador.

Stypommisa abdominalis Philip 1960, Proc. Calif. Acad. Sci., Ser. 4, 31(3):

88, holotype female [examined]. Fairchild 1971, Cat. Dipt. Amer. S.

of U.S. Fase. 28: 81.

A predominantly yellowish species with yellowish pleural and abdominal pi-

losity, the wings extensively yellowish tinted, with fuseus clouds around end of

discal cell and fork of 3rd vein, and more or less infuseated anteroapical area.

Antennae yellow to orange, with at most the terminal annulus infuscated. Legs

wholly yellow, fore tarsi only slightly dusky.

Material examined. Colombia: Putumayo Int., 1 female, Mocoa, 650 m elev.,

Very Wet Tropical Forest, flight trap with COo; 15-IV-1976 (R. Wilkerson).

Ecuador: Pastaza Prov., 8 females, 17.2 km. SE Puyo, 5-XI-1981 (R. Wilkerson).

Zamora Prov., 1 female, 12 km S Zamora, bait horse, 5-7-111-1982 (R. Wilker-

son). Peru: Cuzco Prov., 3 females, Quincemil, 753 m elev. X-1962 (L.E. Pena).

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 21

Discussion. This species has a limited distribution along the eastern base of

the Andes from Colombia to Peru. It is very similar to both S. eaptiroptera and

S. anoriensis, being generally separable from the first by having the 1st marginal

cell (Ist R) largely yellow, and fuscus clouds around both crossveins at end of

discal cell and fork of 3rd vein. From S. anoriensis it differs chiefly in lacking

contrastingly black antennal styles. We suspect that S. anoriensis and S.

flavescens may be only trans- and cis-andean subspecies of a single taxon, but

lack specimens from enough localities to be certain.

Fairchild (1967) studied specimens determined by Kroeber in Munich and se-

lected one from unknown locality (mislabelled Espirito Santo) as Neotype. The

type was from Peru, presumably from Callanga in the Pinipini Valley, but was in

Kroeber's collection and destroyed during World War II. Subsequently (1975),

Fairchild was able to study another Kroeber specimen from Sta. Inez. Ecuador in

Warsaw which appears to be the same. Both these Kroeber determined specimens

agreed with a female from Peru, Cuzco, Quincemil, which we also found to agree

with the type of S. abdominalis Philip on loan from Dr. Philip.

Stypommisa fulviventris (Macquart)

(Figs. 28-30)

Tabanus fulviventris Macquart 1846, Dipt. Exot., Suppl. 1, p. 164. holo-

type female, Brazil [BM(NH)] [examined].

Stypommisa fulviventris: Kroeber 1930, Zool. Anz. 86(9-10): 254-255, fig.

3. Fairchild 1971. Cat. Dipt. Amer. S. of U.S. Fase. 28: 81.

Tabanus ferruginosus Walker 1850, Dipt. Saunders. 1: 40, holotype female,

South America [BM(NH)] [examined]. NEW SYNONYM.

Tabanus (Macrocornus) ferruginosus: Kroeber 1930, Zool. Anz. 87(1-2): 2,

4-5, fig. 2, Espirito Santo.

Stenotabanus ferruginosus: Fairchild 1956, Smiths. Miscell. Colls., 131(3):

16.

Stypommisa ferruginosa: Fairchild 1971, Cat. Dipt. Amer. S. of U.S.

Fase. 282 3).

A small grayish brown species with lightly tinted but unspotted wings, slender

antennae, and a faint row of small pale middorsal abdominal triangles.

Female. Length 11.5 to 13 mm, of wing 12 to 13 mm. Eyes bare, no pattern

evident. Head structures as figured. Frontal index of the type of S. fulviventris

9.8. Frons grayish brown pollinose, with sparse short recumbent brown hairs.

Callus and tubercle orange brown, the latter with vestiges of ocelli. Subeallus

and upper genae concolorous with frons, becoming steel gray on frontoclypeus and

lower genae. Tentorial pits prominent. Beard sparse, gray. Antennae orange

yellow, first 2 segments black haired, style black, contrasting. Palpi pale brown,

black haired, nearly as long as brown pollinose proboscis. Mesonotum brown,

brown pollinose, with notopleural sutures faintly paler, beset with sparse short

erect black hairs and short recumbent pale hairs. Seutellum and notopleural lobes

concolorous, pleura, sternum and coxae paler, more grayish, sparsely pale haired.

Legs yellowish brown, tarsi darker, fore tibiae yellow haired basally, black-haired

apically. No hind tibial fringe. Wings rather long and slender, appendix present,

variable in length. Wings faintly brownish, stronger along veins, but no definite

clouds on crossveins or fork. Costal cell and stigma yellow. Basicosta with 6 to

10 setae. Halters pale brown. Abdomen brown, slightly paler on anterior tergites,

22 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

the hind margins of tergites not noticeably paler. Hairs black, except for small

medial pale-haired triangles and posterolateral corners, the hairs easily lost. The

2 Specimens measured have frontal indices of 5.2 to 5.8, divergence indices of 1.2

to 1.4 and antennal indices (Basal plate/style) of 1.0 to 1.3. They agree among

themselves otherwise. One of these was compared and agreed with Macquart's

type in London, and was found in good agreement with Walker's type on loan in

1983. This is the specimen figured here.

Male. Not known.

Material examined. In addition to the types of both names: Brazil: Rio de

Janeiro State, 1 female, Itatiaya, Estacion Biologica, 1670 m elev., 15-I-1936; 1

female, Itatiaya, 800 m elev., VI-1931 (D. Mendes); 1 female, Lago Azul, Itatiaya,

750 m elev., 17-31-VIII-1958 (R. Barth).

Discussion. Recent reexamination of the respective holotypes of Tabanus

ferruginosa and Tabanus fulviventris, kindly lent to us by J.E. Chainey indicates

that they are not separable. Neither type is in optimum condition, but all diag-

nostie structures are present, and both agree equally well with one of our avail-

able specimens from Itatiaya. Two other specimens from the same area vary

slightly in size and distinctness of abdominal pilose markings, but are believed to

be conspecific. Our specimens are labelled Dec., Jan., and June, all in different

years. Kroeber (1930) reported one female of S. ferruginosus from Espirito Santo,

but whether his description and figure was drawn from the type in London or his

specimen is unclear.

Stypommisa furva (Hine)

(Figs. 37-39, 77)

Tabanus furvus Hine 1920, Ohio J. Sci. 20(6): 189, holotype female,

Songo, Bolivia (Columbus) [examined].

Stenotabanus furvus: Philip 1958, J. Kansas Ent. Soc. 31(2): 180; 1961,

Pan. Pac. Ent. 37(2): 112. Peru, Bolivia,makes S. maculipennis Kroe-

ber a synonym.

Stypommisa furva: Philip 1969, Acta. Zool. Anz. 83(104): 52-53.

Stypommisa maculipennis: Kroeber 1929, Zool. Anz. 83(1-4): 117, Bolivia

holotype female (destroyed); 1930, op. cit. 83(9-10): 253. Fairchild

1966, Stud. Ent. 9(1-4): 359-360, fig. 24.

A dull blackish species with heavily spotted wings, largely brown legs with

fore tibiae bicolored, frontal callus quadrangular, abdomen with narrow pale pi-

lose segmental margins.

Female. Length 11 to 12 mm, of wing 11 to 12 mm. Eyes bare or sparsely

short pilose, blue green in life, unbanded (Kroeber 1929) or with a single dark

band (Philip 1969) (but see under discussion below). Head characters as figured.

Frontal index 3.5 to 4.3 (n = 12, x = 4.0). Frons brownish gray pollinose, callus

black. Tuberele at vertex shiny black with vestiges of 3 yellowish ocelli. Sub-

callus, frontoeclypeus and genae brownish gray, beard gray. First 2 antennal seg-

ments yellow, gray pollinose, densely short black haired. Third segment with

basal plate yellow to orange, the style black, contrasting. Palpi light brown

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 23

pollinose, black haired. Proboscis brown pollinose, considerably exceeding palpi,

the labella large and membranous.

Mesonotum and scutellum brownish black, thinly gray pollinose, sparsely beset

with mixed dark and pale hairs, unstriped. Notopleural lobes and, variably, sides

of mesonotum and scutellum yellowish brown. Pleura blackish, gray pollinose,

largely pale-haired. Legs with coxae, fore femora, basal halves or less of mid

and hind femora blackish, remainder of femora and most of mid and hind tibiae

light to dark reddish brown, more or less darkened apically. Fore tibiae bicol-

ored, basal third yellowish, white haired, the remainder black and black-haired, as

are all tarsi. Wings as figured, sometimes with small clouds near ends of all

veins, or faint brown margins along apical segments of some or all veins, in addi-

tion to the large brown clouds on all crossveins and fork. Costal cell yellowish.

Fork with a short appendix, usually not longer than adjoining short vein segment.

Abdomen dark brown to black, often vaguely lighter and more reddish on sides of

tergites 2 and 3, dark gray pollinose, and with some bluish-white pruinosity on

first 2 tergites when viewed from behind. There are sparse pale-haired fringes

indistinctly widened in middle and at sides on all tergites; otherwise dorsum

wholly black haired. Venter reddish to black, gray pollinose, entirely sparsely

pale-haired.

Male. Not known.

Material examined. All from Bolivia: 6 female paratypes of T. maculipennis

in BM(NH); 4 females labelled "Sammlung Hartmann," no locality, one det. Kroe-

ber 1930, in Munich; 2 females Songo, one the holotype of T. furvus, the other

unlabelled, in Ohio State Univ. coll. Cochabamba Dept., 3 females, Cristal Mayu,

VIII-1949, IX-1956 (L.E. Pena) (1 female compared and agreeing with paratype of

T. maculipennis in BM(NH), another compared and agreeing with holotype of T.

furvus); 8 females, El Palmar, Chapare, XI- 1956 and X-1943 (L.E. Pena); 240 fe-

males, km 95 on road from Cochabamba to Villa Tunari, 1800 m elev., 2-IX-1983

(R. Wilkerson). La Paz Dept., 1 female, Mapiri, S. Carlos, 800 m I-1903; 1 fe-

male, Yungas von Coroico, 1800 m X-1906; 1 female, ?Yungasweg 1200-2500 m

X-1906; 2 females, Yungas La Paz, Yolera a Caranavi, X-1968 (S. Cosearon); 1

female, N Yungas Prov., Caranavi, IX-1971 (J. Velaseo); 5 females, S Yungas

Prov., S. Antonia, XI-1971 (J. Velasco); Dept. not known: 1 female, Amazon

Basin, 1914, in Paris Mus. Others seen but not now before us. 5 females, Cha-

pare, El Palmar IX-1956 (L. Pena) in CNC; 1 female, Osuntom, VII-1921, Mulford

Exped. det. maculipennis by Stone, in USNM; 1 female, Rio Colorado, IX-1921

(Mann), in USNM; 1 female, Cordillera, 3000 m XII-1902

Discussion. Little variation in structure is shown, but intensity of color in

body and wings varies considerably. The eye of a specimen preserved in alcohol

from San Pedro, Coroico is figured (Fig. ). It is bronzy green with a dark me-

dian line which is well defined below but diffuse above, giving the impression that

upper half of eye is darker than lower half. S. furva is very similar to S. hy-

pographa neofurva Philip, a largely sympatric species, but differs in the structure

of frons, in having evident dark clouds at ends of basal cells and white pilose pos-

terior margins to all tergites. The species ranges from Peru, Dept. Huanuco,

south to Cochabamba, Bolivia, along the wet eastern slopes of the Andes up to

3000 m. Our records, and those in the literature, are all for the months of Au-

gust to January, inclusive.

24 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Stypommisa glandicolor (Lutz)

(Figs. 40-45)

Tabanus glandicolor Lutz 1912, Comm. Linh. Telegr. Estrat. Mato Grosso-

Amazonas, Annexo 5, Hist. Nat., Tabanideos, p. 5, fig 3, lectotype

female, Mato Grosso, Brazil (Rio de Janeiro). Fairchild 1961, Mem.

Inst. Oswaldo Cruz 59(2): 231 (designates lectotype).

Stypommisa glandicolor: Fairchild, 1971. Cat. Dipt. Amer. S. of U.S.,

Fase. 28: 81.

A small slender pale yellowish brown species with very narrow frons, thread-

like eallus, hyaline or faintly tinted and unspotted wings, and a faint pale pilose

mid abdominal stripe or series of connected triangles.

Female. Wing lengths of 8-12 mm, and body lengths slightly greater. Eyes

bare, greenish black, without pattern (relaxed). Frontal indices 6.0-9.3 and di-

vergence indices 1.4-2.0. Frontal callus threadlike or oval below and threadlike

above. Ocellar tuberele present, usually inconspicuous, seldom showing vestiges of

ocelli. Basicosta pointed, with few to numerous dark setae, rarely bare, always

with setae sparser than on adjoining costa. Integument of thorax and scutellum

yellow to cinnamon brown, the scutellum often paler. Vestiture gray, yellowish

or brownish. Legs yellow to pale brown, the fore femora sometimes dusky, tarsi

often darker and fore tibiae obscurely to sharply bicolored. Antennae yellow to

orange, the style concolorous to contrastingly black. Wings yellowish to smoky,

rarely nearly entirely clear and rarely with a faint cloud at fork. Appendix at

fork inconsistent, usually short, often absent. Abdomen yellowish to brown, dark

pilose dorsally, pale ventrally, and generally with at least vestiges of a middorsal

pale pilose stripe or series of contiguous triangles which are easily lost and not

underlain by pale pollinosity.

Male. Available males, 1 from Costa Rica, 2 from Amazonas, 1 from Peru,

are structurally indistinguishable, though the Costa Rican specimen is slightly

darker. The eyes are bare, holoptic, the enlarged upper facets well differentiated

but not sharply demarcated from the small facets and without an occipital border

of small facets. There is a small tubercle sunk between the eyes at vertex. The

palpi are porrect, pointed, and clothed with long pale hairs. Basicostas are

pointed, bare, or with a few black setae.

Material examined. In addition to Lutz’ types the following specimens have

been studied. Costa Rica: Puntarenas Prov., 2 females, Palmar, no date, (P.and D.

Allen); 1 female 1 male, Osa Peninsula, 1.8 mi W of Rineon, 27-11-1971 (Donahue

and Hogue); 2 females, same locality, III-1965 (McDiarmid). Colombia: Meta

Dept., 8 females, Restrepo, III-1935 (P.C.A. Antunes); 2 females, Villavicencio,

carretera al puente Guayuriba, 12-III-1939. Venezuela: Bolivar State, 2 females,

San Juan Manipiar, 30-IJI-1958 (Ortiz). Peru: Cuzco Dept., 2 females, Quincemil,

2450 ft elev., IX-X-1962 (L.E. Pena). Madre de Dios Dept., 15 females, Avispas,

IX-X-1962 (L. E. Pena). 30 females, Tambopata Reserved Zone, Pto. Maldonado,

XI-1979 (J. Heppner); 1 male, 1075 females, VII,VIII,X,XI-1982, (R. Wilkerson).

Huanuco Dept., Tingo Maria, 1 female, Rio Huallaga, 700 m elev. IX-1946

(Weyrauch). Surinam: 7 females, Phedra, X-XI-1964 (D.C. Geijskes). Brazil:

Amazonas State, 2 females, Manaus, Reserva Ducke, X-1968, (R.L. Dressler); 2

males 5 females, same locality, IX, X, XI-81 (J.A. Rafael); 1 female, same local-

ity 6-IX-78 (Arias and Penny); 1 female, Manaus, University Campus, 17-X-78,

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 23

(J.A. Rafael); 1 female, CEPLAC AM 010, km 30, 3-X-75 (J. Arias); 2 females,

Manacapura, IX-1936 (Worontzow). Para State, 1 female, km 165 W of Altamira,

6-XI-74 (J.F. Reinert); 5 females, 70 km W Monte Dourado, 27-29- -XII-1980, (R.

Knell). Mato Grosso aa 26 females, Rio Aripuana, Humboldt, 59° 27'w,

10° 19'S, 12-16-VIII-74 (D. Young); 2 females, Sinop, X-1976 (M. Alvarenga

leg.)s y females, Rio tie: Abuna, no date (Mann and Baker). Bolivia:

Cochabamba Dept., 2 females, Villa Tunari, 240 m elev. 2-6-IX-1963 (R. Wilker-

son).

Discussion. Under this name we include a variable and widespread taxon

which we at one time considered as belonging to 5 distinet taxa, but with addi-

tional material, showing overlapping and intermediate character states, has con-

vineed us all belongs to a single locally variable species. Future collecting may

enable definition of local races or subspecies, but at present we cannot consis-

tently separate the forms.

Specimens from south of the Amazon river, the Mato Grosso and Rio Madeira

localities, have the antennae all yellow or with the terminal annulus dusky, those

from north of the Amazon usually have the antennal style black, contrasting with

the plate. The single Par&é specimen, however, has a dark style. Specimens from

eastern Peru may have the antennae all yellow or the style variably darkened to

all black. Material from Surinam, Colombia and Costa Rica have the antennae

Strongly bicolored. Specimens from the Amazon and Surinam are generally yel-

low with more infuseated wings than those from Venezuela, Colombia and Costa

Rica, which have nearly clear wings.

The discontinuous distribution which skips Panama is probably not an artifact,

as Panama has been quite extensively collected, and there are a number of other

instances of such discontinuity both in Tabanidae and other groups.

The species is likely to be confused only with S. flaveseens from which it dif-

fers in generally smaller size, narrower frons with threadlike callus, lack of a

long appendix at fork and usually no cloud surrounding fork of 3rd vein, or apex

of discal cell. The abdomens when undenuded bear a row of middorsal, often

contiguous, pale pilose triangles forming a more or less conspicuous slender

stripe.

Stypommisa hypographa (Kroeber)

and

S. hypographa neofurva Philip

(Figs. 31-33, 78)

Stypommia hypographus Kroeber 1930, Zool. Anz. 86: 252-253. holotype

female, Venezuela (Vienna) [examined]; 1934, Rev. Ent. 4(2): 274,

Peru. Fairchild 1967, Stud. Ent. (1966) 9(1-4): 359, fig. 32.

Stypommisa hypographa: Fairchild, 1972, Cat. Dipt. Amer. S. of US.,

Fase.28: 81.

Stypommisa hypographa neofurva Philip 1969, Acta Zool. Lilloana, Tu-

cuman (1967) 22: 123-124, holotype female, fig. 6, Bolivia (IML).

A small black or nearly black species without conspicuous pale pilose mark-

ings, the abdomen bluish pruinose, the legs nearly wholly blackish, and wings with

small clouds on fork and crossveins and moderately strong anterior and apical

tinting.

26 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Female. Translation of original description of S. hypographa (Kroeber).

Length 12 mm, antenna ?, wing length 12.8 mm, wing breadth 4 mm. Frons about

2 times as high as wide, parallel sided, silky brownish gray, pale haired. Vertical

triangle slender, long, truneated below. Ocellar spots without clear ocelli, with

scattered black hairs. Frontal callus long oblong, slender, separated from eyes,

shining blaek, moderately strongly swollen. On this rests a broad flat wedge

which merges into a slender ridge which reaches nearly to the ocellar spot.

Frontal triangle (subeallus) brownish gray. Face whitish gray, white haired. Palpi

slender, light brown, finely white pollinose, thickly black bristled, especially to-

wards the tip. Antennae reddish brown, first and second segments thickly black

bristled, third with broad black plate without real tooth, only a very small shortly

black-bristled angle. Style deep black, with short broad segments. Back of head

whitish gray, short black haired. The whole body black, moderately dull, with

bluish white bloom (pruinosity). Sides of thorax and scutellum brownish. Pleura

and coxae whitish gray haired; legs brown, bases of fore tibia whitish yellow, sil-

ver white haired; hind tibia outwardly silver white haired; base of mid tibia yel-

lowish, above black haired, below white haired; hind tibia outwardly silver white

haired, otherwise black haired. Femora silver white haired, calypters and halter

brown, stem reddish yellow. Wing somewhat brownish tinted. All veins in apical

part moderately clearly brown margined, so that to the naked eye a large brown

apical patch appears. Crossveins somewhat brown spotted, especially the fork

with nearly circular brown spot, and the hind margin (apex) of discal cell. The

ends of all longitudinal veins and apex of anal cell are dark brown becoming thus

spotted. Abdomen above and below bluish pollinose without pale segmental bands

or pattern. Hairs below and above at the sides of the hind margins white, other-

wise black. Tergites 1, or 1 and 2, notably light whitish blue pruinose. Bolivia,

Mapiri, 26.1 ; Peru, Pichis-Weg, Puerto Yessup 23.XII.

Male. Not known.

Material examined. All ssp. neofurva. Bolivia: La Paz Dept., 2 females,

Mapiri, S. Carlos, 800 m, 9-1-03, in Dresden Mus.; 5 females, S. Inicua Riv., Alto

Beni, 1100 m 15-18-1-1976 (L.E. Pena); Peru: Junin Dept., 1 female, Chan-

chamayo, 14-II-49 (J. Schunke). Ecuador: 1 female, no further data (L. Leon);

Pastaza Prov., 1 female, 17.2 km SE Puyo, 5-XI-81 (R. Wilkerson). Colombia: 5

females, Int. Putumayo, Mocoa, 650 m, Very Wet Subtropical Forest, flight trap

with CO,, IV-15-76 (R. Wilkerson); 2 females, Pto. Asis, 240 m, Very Wet Tropi-

cal Forest, flight trap with CO,, IV-15-76 (R. Wilkerson). Panama: Panama

Prov., 2 females, Cerro Jefe, 28-14 and 8-V-1968 (R. Dressler).

Discussion. The type in Vienna from Venezuela now lacks antennae. Whether

Kroeber's description was drawn wholly from the Venezuelan type or whether he

included differing characteristics from his Bolivian and Peruvian specimens is un-

known. His description of the antennae as black suggests that the Vienna type

possessed these structures when described, although Kroeber placed a query after

the antenna in the original description. Considerable material from Bolivia and

Peru seen by us all have bicolored yellow or orange and black antennae. Philip

(1969) differentiated his subspecies neofurva from Bolivia on the bicolored anten-

nae and less evident pale pilosity on the bases of fore tibiae and sides of ab-

domen. Our specimens, listed above, show considerable variation in the amount

of pale pilosity, and the intensity of the dark wing markings, though all but three

have sharply bicolored antennae. These three have the basal plates of antennae

more or less apically infuseated, so that they appear less sharply contrasting with

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 27

Style. One of these is from Colombia, Putumayo, Mocoa, the other from Panama,

Cerro Jefe, and all also have the wing spotting reduced in extent. Fairchild's

(1967) figure of the type from Venezuela shows a more heavily spotted wing than

any we have seen. Several other specimens from Putumayo have sharply bicol-

ored antennae and larger wing spots, like those from Bolivia and Peru. One of our

Ecuadorian specimens is in poor condition and lacks antennae; the other closely

matches Bolivian specimens. Frontal indices of our S. neofurva specimens range

from 4.2-5.5 (x = 4.9). We believe that S. neofurva had best be maintained as a

subspecies, at least until Venezuelan specimens with antennae can be studied.

Philip (1969) records the eyes as being bicolored, purple above, green below, which

agrees with our specimens.

This species is very similar to S. modiea, but has less extensively silver pilose

fore tibiae, bicolored or dusky antennae, and lacks the silver pilose terminal ab-

dominal tergites of S. modiea. The wing spotting of S. modica is usually less ex-

tensive, though some neofurva have indistinguishable wings. The eye pattern of

S. modica is complicated and quite different from the bicolored eyes of ssp. neo-

furva. Stypommisa hypographa neofurva can be separated from the sympatric and

similar S. furva as detailed under the latter.

Stypommisa jaculator (Fairchild)

(Fig. 79)

Stenotabanus jaculator Fairchild 1942, Ann. Ent. Soe. Amer., 35(3): 309,

figs. 9, 9a, 9b, holotype female, Panama (MCZ); 1951, Ann. Ent. Soe.

Amer., 44(3): 452-453, female, male; 1953, Proe. Ent. Soc. Washing-

ton, 55(5): 241.

Stypommisa jaculator (Fairchild). Barretto 1956, Rev. Brazil. Malar.

Doeneas Trop. 8(1): 82, Br. Guiana. Fairchild, 1971, Cat. Dipt.

Amer.S. of U.S., fase. 28: 81. Wilkerson 1979, Cespedesia 8(31-32):

339-340, fig. 108.

Female. A small cocoa brown or grayish brown species. Length 7-11 mm, of

wing 7-10 mm. Wings tinted along fore border and apex, with small clouds at

fork of 3rd vein, and more faintly at apex of discal cell. No appendix at fork.

Abdomen cocoa brown, all tergites with broad silvery-haired triangles not under-

lain with pale pollinosity. Legs brown, the fore tibiae weakly bicolored. Anten-

nae orange, the style sometimes darker, rarely black and contrasting. Eye in life

greenish bronze with a single narrow and often faint median dark transverse

stripe.

Male. The male is similar in color, and easily associated with the female.

Its eyes are holoptic, the brown upper enlarged facets well differentiated and

demarcated from the small bronzy green lower facets, occupying about 1/2 eye

area and entirely bare. There are no long occipital hairs.

Material examined. Panama: Canal Zone, 2 females, Bohio, and 2 females

Las Caseadas (both A.H. Jennings), (holotype and paratypes). 1 female, Gamboa,

20-V-1953 at light; . Panama Prov., 2 males, 7 females, La Victoria, Cerro Jefe,

2200 ft elev., 20-IV-1949, in light trap; 14 females same locality, 24-ITI-1946; 50

females same locality, II-IV-1949, mostly taken attacking man at platforms from

23 to 71 feet above ground level in the forest canopy; 1 female, La Victoria,

Cerro Jefe, 20-IV-1958; 1 female, Cerro Azul, 24-V-1962; 1 female same locality,

28 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

20-V-1964. Cocle Prov., 2 females, E] Valle, 16- VI-1946 and 10-V-1951. Darien

Prov., 2 females, Upper Rio Pucro, 16-V--1958; 1 female, Rio Tuira Yellow Fever

Sta., 5-III-1958, tree tops; 1 female, Rio Paya Yellow Fever Sta., 24-IV-1958; 1

female, La Laguna, Tacarcuna, 4100 ft 18-III- 1959; 3 females, Cerra Quia, Rio

Mono, 2500 ft elev., 8-III-1970 human bait in tree canopy 60 ft above ground

level; 2 females, Morti River, 7-IV-1967; 1 male, Cerro Horeona, 17-III-80. 1 fe-

male, Candelaria Hydro Sta., Rio Pequeni, V-1966. Colombia: Choco Dept., 2

females, Teresita, 21 and 29 -III-1967; 2 females, Curiche River, 12 and 25-IV-

1967. British Guiana (Guyana): 1 female, New River, 12-V-1938, in BM(NH), the

specimen reported by Barretto (1956).

Discussion. The species is highly arboreal and appears to have a short flight

season, so that it may have a greater range than at present known, since arboreal

collections may not have been made at the right time of year to reveal its pres-

ence. Most collections have come from areas of high rainfall and heavy forest,

from near sea level to over 4000 ft. This species is very variable in size, wing

lengths of available specimens ranging from 7 to 11 mm.

Stypommisa jaculator is similar to S. prunicolor but can be separated on: 1)

eolor of antenna, bicolored or with at least last annuli dusky in S. jaculator,

wholly yellow in S. prunicolor; 2) brown middle and hind femora of S. jaculator

which are black in S. prunicolor; 3) yellow and black haired legs of S. jaculator,

all black haired legs in S. prunicolor; 4) the callus of S. jaculator is evenly ta-

pered above, while larger and rectangular in S. prunicolor; 5) frontal index of S.

jaculator is 4.5-5.6 (x = 5.05), and that of S. prunicolor 4.2-4.9 (x = 4.6). The

Species can be separated from S. hypographa and S. modiea by having middorsal

abdominal triangles of white hair and less heavily marked wings.

A single specimen before us from Colombia, Antioquia Dept., Rio Anori Val-

ley, La Tirana, 13-IV-1971, C.H. Porter leg., CHP-71-172 DO1, does not agree

with the concept of either S. jaculator or S. prunicolor. In overall coloration it is

quite like a dark S. jaculator but has a callus similar to S. prunicolor. In his ex-

tensive collecting in Antioquia Porter found this species to be commonly taken in

the forest canopy. Since we have only one of his specimens and no ready access

to his series, we defer making a decision on the status of this form.

Stypommisa kroeberi n. sp.

(Figs. 34-36, 80)

?Stypommia venosa: Kroeber, in part, 1930 Zool. Anz. 86(-10): 250, fig.

1. Not Bigot 1892. j

A pale yellowish brown species with large clouds around fork of 3rd vein and

end of diseal cell, small clouds or none at ends of basal cells and apical infusca-

tion beyond stigma.

Female Holotype. Length 11.5 mm, of wing 10.5 mm. Head structures as

figured. Eyes bare, no pattern (relaxed), probably greenish in life. Frons and

subeallus grayish brown pollinose, concolorous. Frontoclypeus and genae paler,

pinkish gray. Frontal callus reddish brown. Antennae orange yellow, only the tip

of the terminal annulus blackish. Palpi yellowish brown pollinose, wholly black

haired except for sparse brownish hairs at base of terminal segment beneath.

Beard sparse, white, proboseis brown, the labella membranous, pollinose.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 29

Integument of thorax brown, thinly gray pollinose dorsally, more thickly white

pollinose laterally and ventrally. Scutellum paler, apically yellowish. Dorsally

the vestiture consists of sparse recumbent mixed black and brassy short hairs,

longer and black on notopleural lobes, long and white or pale gray on pleura and

coxae. Legs largely orange yellow, the coxae, femora and at least ventral as-

pects of mid and hind tibiae yellow haired, the apical half of fore tibiae, apices

and dorsal surfaces of mid and hind tibiae, and all tarsi more or less dark pilose.

Fore tibial integument apically dusky, the vestiture white basally, black apically,

appearing bicolored. Wings as figured, the basicosta pointed, black setose, the

setae nearly as dense as on adjoining costa. Venation normal, a short appendix on

fork of 3rd vein, shorter than the long subtending vein segment. Veins brown,

costal cell yellow, clouds brownish, apical infuscation paler brown, rest of wing

nearly hyaline. Halters yellowish orange, knob and stem concolorous.

Abdominal integument orange yellow, dorsally subshiny, beset with short

black hairs, the posterior margins of all terga narrowly and sparsely pale yel-

lowish pilose, more evident in middle and at sides. Beneath the hairs are longer

and entirely whitish except for longer outstanding black hairs on last visible ster-

nite.

Male. Not known.

Type data. Holotype female, Peru: Huanuco Dept., Tingo Maria, 670 m, IX-

47 (Weyrauch leg.), deposited in MCZ. Paratypes, 2 females, same data as holo-

type, 1 det. venosus Bigot by Fairchild 1956. 1 female, Tingo Maria, Rio Hual-

laga, 700 m Peru (W. Weyrauch), labelled as same as homotype of venosus in coll.

C.B. Philip, except that basal plate longer, style shorter, both yellow, dusky

throughout in homotype. Right wing mounted on slide St. 6. Cuzco Dept., 1 fe-

male, Quincemil, 740 m elev., VIII-1962 (L.E. Pena); 2 females, Quincemil, 2450

ft., VIII-1962 (L.E. Pena). Madre de Dios Dept., Avispas, 1000-1300 ft., 10-30-IX-

1962 (L.E. Pena). Bolivia: Sara Prov. 1 female, (Steinbach). Cochabamba Dept.,

1 female, Cristal Mayu. 23-XI-1949 (Luis E. Pena); 21 females, Villa Tunari, 2400

m elev. (R. Wilkerson). Paratypes in CNC, Cornell, FSCA and collections of

authors.

Discussion. There is not much variation in the paratypes, except that frontal

callosities vary from yellow to nearly black, and the intensity of wing markings

may be greater or less than shown. One specimen nearly lacks a dark cloud at

end of discal cell. Some have the abdomen darker, with evident middorsal pale

haired triangles. One has frontal callus evenly tapered, clavate. Frontal indices

of 9 specimens: 5.1-5.9 (x = 5.4), divergence index 1.2-1.6 (x = 1.4).

This appears to be, at least in part, the species treated as Tabanus venosus

Bigot by Kroeber (1930). It differs from T. furva Hine in lacking strong spots at

ends of basal cells, in narrower frons, in pale brownish rather than black in-

tegument, and in nearly wholly yellow antennae. S. spilota n. sp. differs in more

extensive dark spotting of the wings, black integument of thorax and scutellum,

and bicolored antennae. S. changena n. sp. also has bicolored antennae and much

darker thoracic integument, as well as smaller and less defined wing spots.

From. S. pequeniensis and S. captiroptera this species differs in much more heav-

ily spotted wings, although some populations of S. pequeniensis and all S. ecap-

tiroptera have similarly colored antennae. It is quite possible that more exten-

sive material may show this, S. venosa, S. spilota and possibly S. changena to be

but geographic forms of a single species.

30 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Stypommisa lerida (Fairchild)

(Fig. 81)

Stenotabanus lerida Fairchild 1942, Ann. Ent. Soc. Amer. 35(3): 307, fig.

14, 14a, 14b., Panama Chiriqui (MCZ) [examined].

Stypommisa lerida: Fairchild 1971. Cat. Dipt. Amer. S. of U.S. Fase. 28:

A small stout brown species with prominently spotted wings, banded abdomen

with an integumental middorsal black spot on tergite 2, bicolored antennae and

fore tibiae.

Female. Length 10-11.5 mm, of wing 10.5-11 mm. Eyes bare, apparently

green with a single purple band in life. Frons a little over five times as high as

basal width, slightly narrowed below, yellowish brown pollinose. Frontal callus

dark brown, higher than wide, narrower than frons, and prolonged in a slender line

2/3 the distance to the vertex. Tubercle at vertex with traces of three ocelli.

Subeallus yellowish pollinose, with sparse long whitish hairs. Frontoeclypeus and

genae gray, sparsely grayish haired. First two antennal segments yellowish, black

and pale haired. Basal part of 3rd segment orange, longer than wide, the dorsal

tooth small and near base. Annulate portion black, shorter than basal portion.

Palpi pale brownish, slender, mostly black haired.

Mesonotum and seutellum brown, the latter with reddish apex, indistinctly

grayish striped, and with sparse golden hairs. Pleura, sternum and coxae gray

pollinose, rather abundantly gray haired. Legs light brown, the tarsi and apices

of fore tibiae darker. Femora pale haired, tibiae and tarsi mostly dark haired.

Wings with basicosta bare. Wings faintly smoky, darker along costa and hind mar-

gin, and with prominent clouds on all ecrossveins. Upper branch of 3rd vein long

appendiculate, costal cell yellow. Abdomen orange brown, darker towards apex,

the hind margins of all tergites paler and yellow haired, rest of tergites dark

haired. The second tergite, and rarely the third, bears a median black in-

tegumental spot. Venter pale reddish brown, mostly pale haired.

Material examined. Holotype female and 7 female paratypes, Panama:,

Chiriqui Prov., Boquete, III-IV-1941 (Mrs. T.B. Monniche); 9 female paratypes,

Chiriqui Prov., Cerro Punta, II-III-1940, 6,000 ft (T.T. Howard); Chiriqui Prov., 1

female, Finca Lerida, Boquete, III-1956 (G.B. Fairchild); 4 females, Palo Santo, II-

III-1951 (Galindo leg.).

This species is structurally like S. furva, but differs in brown abdomen and

legs. From S. changena n. sp. it differs in having a black integumental spot on

2nd abdominal tergite, a generally shorter appendix at fork, in being confined to

the Pacifie slope of the Chiriqui mountains, and in being on the wing during the

dry season from Feb. to April.

Stypommisa maruccii (Fairchild)

(Fig. 82)

Stenotabanus marucceii Fairchild 1947, Ann. Ent. Soc. Amer. 39(4): 568,

fig. 8a, b, (1946), holotype female, Panama (MCZ); 1958, op. cit.

51(6): 529, synonymy. Lee, Fairchild and Barreto, 1969, Caldasia,

Bogota, 10(49): 450-451, Colombia.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 31

Stenotabanus frondicolus Fairchild 1951, Ann. Ent. Soe. Amer. 44(3): 453-

454, figs 5b., Panama; 1953, Proc. Ent. Soe. Washington 55(5): 241.

Stypommisa maruccii (Fairchild) 1971, Cat. Dipt. Amer. S. of U.S., fase.

28: 82. Wilkerson 1979, Cespedesia, 8(31-32): 339-340, figs. 109.

A small blackish species with prominently striped thorax, usually bicolored

antennae, wholly black legs, bluish pruinose abdomen with silvery pilose median

triangles and faintly tinted wings with small clouds at fork and end of diseal cell.

Female. We give below the original description of Stenotabanus frondicolus,

as it is more complete that that of S. maruccii. The figure accompanying this

description shows the frons slightly narrower than usual. The figure of S.

maruccii is better.

Length, 9.5-11 mm, wing, 10-11 mm. _ Frons dark golden yellow pollinose.

Frontal callus black, nearly as wide as frons, gradually narrowed above into a

Slender ridge nearly reaching vertex. Vertical tubercle prominent, with 3 well

marked ocelli, the vertex with unusually dense stout black setae. Eyes bare, in

life bright brick red, with a single narrow green transverse band, fading on death

to dull bronzy. Subeallus, frontoclypeus and genae yellow pollinose, the last with

sparse whitish beard. Antennae moderately slender, brownish, black haired. Pro-

boseis short, much less than head height, but little longer than palpi, the labella

fleshy. |

Mesonotum blackish with four somewhat gray pollinose stripes, whose inten-

sity varies with the light incidence. Hairs black on dark parts, brassy on the pale

Stripes. Scutellum pale pollinose, with dark hairs in the middle and brassy hairs

around the margin. Pleura steel gray pollinose, mostly pale haired, except a tuft

of black hairs in the middle. Legs wholly black and black haired. Wings with

subepaulet pointed and with fairly abundant macrotrichiae. Wings grayish hyaline

with small but distinct clouds on fork of third vein and end of discal cell.

Abdomen above black in ground color, rather thickly overlaid with somewhat iri-

descent bluish pruinosity. This is very pronounced on the first tergite, less so on

succeeding segments, where it is more or less confined to the posterior two-thirds

of the tergites. The tergites are mainly black haired, except the first, which is

wholly silvery haired at the sides. There is also a series of median silvery haired

triangies on the second to sixth tergites and lateral patches of silvery hairs on

the sides of the same tergites. Beneath the abdomen is black, mostly bluish pru-

inose and with all sternites with complete transverse bands of silver hairs. The

insect in life bears a remarkable resemblance to a sarcophagid fly.

Holotype female, Progreso, Chiriqui Prov., Panama 20 July 1950; 9 female

paratypes, same locality, 16 June (1). 28 July (1), 3 August (3), 17 August (1). 31

August (1), 9 Nov. (1), 16 Nov. (1); 7 paratypes, Sta. Fe.Veraguas Prov., Panama,

18-23 May (1), 29 May (1), 31 May (3), July (1), 10 Aug. (1).

With a few exceptions, all the Panama specimens were taken at platforms

built in the tree tops in heavy forest, from 36 to 83 feet above ground level.

Colombian examples were mos‘'y taken in flight traps.

In addition to the above description, measurement of the fronts of 19 speci-

mens from Nicaragua (1), Panama (12), and Colombia (6) show the frontal index

varying from 3.8 to 5.5 (x = 4.95).

Male. The male, based on a specimen from Rio Palenque, Ecuador, is easily

associated with the female on color characters. The eyes are bare, the upper

facets much enlarged and well demarcated from the small, occupying fully 2/3

32 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

eye area. There is a small tubercle between the eyes at vertex and the antennae

are bicolored.

Material examined. Only a sample, including 9 paratypes of frondicalus, of

the material discussed in the cited references is now before us. Additional

records are Colombia: Choco Dept., Curiche River, 18-IX, 26-X-1967; Valle

Dept., 25 and 41 km E of Buenaventura, 12-VIII-1973 and 19-III-1976 (Wilkerson

and Young); Anchicaya Dam, 10-VIII-1973 (Wilkerson and Young); Nicaragua:

Villa Somoza, 7-VII-1953. See Wilkerson (1979) for additional Colombian records.

The species appears to range from Nicaragua to Ecuador west of the Andes.

Discussion. The species is predominantly arboreal (Fairchild 1953) and a good

mimic of a Sareophagid fly (flesh fly), even to the red eyes. Attempts to revive

the eye color give only a green eye without the median band seen in fresh speci-

mens. The species appears to range from Nicaragua south to western Ecuador

and flies throughout the year. We have records from every month save April. It

ean be distinguished from similar musecoid mimics such as S. hypographa neofurva

and S. modiea by the strongly striped thorax, unicolorous dark legs, median sil-

very pilose abdominal triangles and weakly spotted wings. Several other tabanids,

belonging to other genera also seem to be mimics of similar sarcophagid models.

Philipotabanus (Mimotabanus) plenus Hine is larger with white beard and with a

dark wing patch below the stigma, but is very like S. maruccii in other respects.

Eutabanus pictus Kroeber has the same striped mesonotum, white median abdomi-

nal triangles and unicolorous black legs, but head structures are quite different.

The wings are hyaline with a narrow transverse fuscus stripe below stigma and a

Slender apical fuscus stripe along the margin from stigma to apex. Myiotabanus

sarcophagoides Lutz has the same general appearance, but is much smaller, less

than 10 mm, and the mid abdominal pale triangles are contiguous, forming a

Stripe. The head structures are again distinctive, with partly bare subcallus, pro-

boseis equalling head height and short inflated palpi. M. muscoideus Hine is also

very similar, but specimens are not at hand for detailed comparison.

Stypommisa modiea (Hine)

(Fig. 83)

Tabanus modicus Hine 1920, Ohio H. Sci. 20(6): 188, British Guiana, holo-

type female (Columbus) [examined].

Stypommisa modica: Fairchild 1971, Cat. Dipt. Amer. S. of U.S., fase.

28: 82, synonymy.

Stypommia hypographa var. ruficornis Kroeber 1930 Zool. Anz. 86: 253,

Iquitos, Peru, holotype female (Halle) [examined]. Fairchild 1967,

Studia Ent. 9(1-4): 350, fig. 33 (1966), gives notes on types, syn-

onymized with modiea Hine. Philip, 1969 Acta Zool. Lilloana, Tu-

ecuman, Argentina, 22: 124 (1967), confirms syn. with modica Hine.

A small black species with bluish pruinose thorax and anterior abdomen, the

wings with small clouds on crossveins and fork and apically brown tinted. Fore

tibiae basally with some white, antennae wholly orange yellow.

Female. Hine's description is as follows: "Total length of the body 11 mm.

Whole body black with a whitish bloom especially on the thorax and base of the

abdomen. Palpi rather slender, pale, with sparse dark colored pile, somewhat

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 33

Shorter than proboscis. Antenna yellow, Ist segment slightly produced forward

above and furnished with black pile, 3rd segment angulate above near the base

and with a tip of black pile but not drawn out into a process. Front moderately

narrow, frontal callosity below practically as wide as the front, shining black,

widest portion distinctly longer than wide, then gradually narrowed to a line

which connects with a shining area at the vertex. Thorax without apparent

stripes, wing fuliginous, darker along costal border and on the margins of the

crossveins at the apices of the basal and discal cells and at the fureation of the

3rd vein. First posterior cell wide open, anterior branch of 3rd vein without an

appendix. Legs black, front tibiae narrowly white at the base. Holotype female

from British Guiana in my collection."

The eyes in life are red or reddish with a broad median green band, tapered

outwardly, through which a narrow bright blue stripe runs, the green being con-

siderably wider below the stripe than above it. There is only minor variation in

color.

Male. The male, hitherto undescribed, is similar to the female in wing and

body colors, but almost lacks white pilosity at base of fore tibiae. The eyes are

bare, the upper facets greatly enlarged and well demarcated from the lower small

facets. In both dried specimens available the large facets are bright reddish

brown with a double band of paler facets enclosing a darker central area. The

lower pale band is almost white, lies close to the lower border of the large

faceted area, but is separated from the blackish small facets by a narrow strip of

reddish facets. There is a small pedunculate black and sparsely pilose tubercle

between the eyes at vertex. The palpi are dull yellowish, densely long dark pi-

lose. Both specimens were taken in a flight trap set in the forest canopy at

Tambopata Reserved Zone, Madre de Dios, Peru.

Material examined. In addition to Hine's and Kroeber's types studied by

Fairchild, we have seen the following specimens. Peru: Pasco Dept., 1 female,

Miriatiriani, Camino de Pichis, 9-VII-1920, Cornell Univ. coll., comp. with types

of hypographa var. ruficornis Kroeb. Madre de Dios Dept., 2 males, 20 females,

Tambopata Reserved Zone, 30 km SW Puerto Maldonado, X-XI-1982 (R. Wilker-

son). Ecuador: Napo Prov., 1 female, Limoncocha, 18-25-V-76 (D.G. Young).

Brazil: Para State, 2 females, Belem, APEG forest, 13-VII and 2-XII-1970, in for-

est canopy (Aitken and Toda). Four females, same locality, 29-VII to 6-VIII-1974,

flight trap, and VIJ-1981, arboreal flight trap (G.B. Fairchild leg.). There is also

1 female, not now before us, from Chapare, Bolivia in coll. Sixto Cosearon. The

frontal indices of 6 specimens ranged from 4.6 to 5.25.

Discussion. Kroeber's description of his "varietat ruficornis" is brief, trans-

lated as follows: Type Halle, from Amazonas. Equal in all respects to the stem-

form (typical hypographa). The antennal style is light reddish yellow, only the

terminal annulus is somewhat browned. Wing with uniformly brownish apical

part. The vein endings not darker bordered, but fork and hind border (distal end)

of discal cell brown spotted. Abdomen from 8rd tergite on with little white hairs

in the middle of the hind margins, on tergites 6 and 7 the whole margins are

white haired. Iquitos, Fonteboa, Rio Negro.

From the limited data available, it appears that the species flies throughout

the year and may be primarily arboreal. Stypommisa modica can be separated

from the similar S. hypographa and ssp. neofurva by nearly wholly yellow anten-

nae, slightly narrower frons and smaller wing spots. It differs from S. prunicolor

and S. jaculator in lacking any trace of dorsal pale triangles on abdomen. The

34 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

distribution indicated by the material we have seen is no doubt less than the ac-

tual range, as little arboreal collecting has been done except in a few localities.

Stypommisa paraguayensis (Kroeber)

(Figs. 49-51, 84)

Stypommia paraguayensis Kroeber 1930, Zool. Anz. 86(9-10): 250-251,

holotype female, Paraguay, Hohenau (Dresden) [examined].

Fairchild 1967, Stud. Ent. 9(104): 360, fig. 25.

Stypommisa paraguayensis: Fairchild 1971, Cat. Dipt. Amer. S. of U.S.

Fase. 28: 82.

A small dark brown species resembling S. seythropa but with contrasting

whitish pattern on mesonotum, more heavily spotted wings, different frons, and

bare-eyed males. Our material has frontal indices of 3.8-4.8 and divergence in-

dices of 1.2-1.4. |

Female. Translation of original description: "Type: Dresden, von Paraguay,

Hohenau, 25. X. Length 10.6 mm; antenna 1.4 mm; wing length 10.2 mm; wing

breadth 3.7 mm. Frons about 5 times as high as broad, parallel sided, cinnamon

brown, yellow haired, at vertex brown, black haired. Ocellar spot small, ocelli

indistinct. Frontal callus triangular with upper prolongation double its length,

pale brown. Subeallus and face reddish cinnamon brown next to the eyes, oth-

erwise yellow brown and so haired. Antenna bright reddish yellow; lst and 2nd

segments yellow brown, with fine short black hairs. Style deep black. Palpi long

as in Esenbeckia, outwardly short and fine black and yellow haired, slightly bent.

First segment long yellow haired. Back of head gray, at eye border yellow, yel-

low haired. Thorax dark reddish brown with the beginnings of 2 pale longitudinal

stripes and a finer dark midline. Hairs yellowish. Scutellum brown, blackish

basally, the hind border white tomentose. Pleura paler yellow brown, in part

gray, yellow haired. Halters and squamae yellowish brown, the knob dark brown.

First and 3rd femora blackish brown, fore predominantly yellow haired, knees

yellowish. Tibiae yellowish, yellowish brown haired. Tibia black at apex. Tarsi

brown, the fore pair black. Wings nearly hyaline, lightly brown tinged. Veins

finely brown margined, ecrossveins and fork intensely brown spotted. Appendix (at

fork of 3rd vein) shorter than basal segment of upper branch of fork. First to 3rd

tergites yellow-brown, 4 to 7 black brown, all hind margins clearly yellowish

haired with middorsal triangles. Otherwise the hairs are black. Sternites 1 to 3

pale yellow brown, the rest blackish, yellowish tomentose and haired."

Male. Length 11 mm, of wing 10 mm. Eyes bare, holoptic, the upper en-

larged facets sharply demarcated and differentiated from the lower small facets,

occupying about 2/3 of eye area, the small facets margining the larger outwardly

but not reaching vertex. Large facets in the dried specimen reddish brown, with

a broad yellowish band on lower margin. Small facets black. A small scale-like

tubercle between eyes at vertex. Frontal triangle golden brown pollinose. An-

tennae yellow orange, style contrasting black, more slender than female. Palpi

porrect, long oval, slightly faleate at tip, pale yellow pollinose and beset with

long pale hairs. Thorax, wings and abdomen as in female, the abdomen paler

brown, more pointed, and with longer denser hairs, especially laterally. Legs red-

dish yellow, the femora only slightly darker.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 30

Material examined. Brasil: Minas Gerais State, plesiotype male, Sapu-

caimirim, Cidade Azul, 1400 m, 6-IX-1953, (Travassos F., Periera and Madeiros),

right wing slide mounted. Sao Paulo State, 5 females, Cantareira, VIII-1958 (J.

Lane); 1 female, Horto Florestal, Chapada, VII-46 (F. Lane); 1 female, Cotia, VII-

61 (Rabello); 1 female, Est. Biol. Boraceia, Salesopolis, 850 m, 24-VII-61, isca

humana, 17-19 horas, (Rabello). 1 female, Serra da Bocaina, Faz. do Bonito, 25-

30-IX-1913, (A. Lutz).

Discussion. In spite of not having both sexes from the same locality, we as-

sociate our single male on the basis of agreement on wing and body color, espe-

cially the prominently striped thorax. The females can usually be separated from

dark specimens of the punctipennis form of S. seythropa by the more heavily

spotted wings, yellow triangular frontal callus, and frons with median patch of

dark pollinosity as well as such an area surrounding the ocellar tubercle. In addi-

tion, S. seythropa may have a median denuded groove in the subeallus appearing

to be a slender prolongation of the usually round basal callus. We associate our

specimens with S. paraguayensis on the basis of Kroeber's description and

Fairchild's figure (1967) of the holotype. We have seen no Paraguayan specimens

like these.

Stypommisa pequeniensis (Fairchild)

(Figs. 46-48, 85)

Stenotabanus (Stenotabanus) pequeniensis Fairchild 1942, Ann. Ent. Soc.

Amer. 35(3): 308-309, fig. 13, a, b., holotype female, Panama (MCZ);

1961, Rev. Biol. Trop. 9(1): 29, Costa Rica.

Stenotabanus pequiensis Philip (lapsus) 1960, Proce. Calif. Acad. Sci., Ser.

4, 31(3): 77, Peru.

Stypommisa pequeniensis Fairchild 1971, Cat. Dipt. Amer. S. of U.S.

Fase. 28: 82. Wilkerson, 1979, Cespedesia 8(31-32): 340-42, fig. 110,

111. Hogue and Fairchild 1974, Rev. Biol. Trop. 22(1): 22, Costa

Rica.

Female. A small brown species, wing length 8.5 mm to 11.0 mm., the dorsum

of abdomen entirely black pilose, the venter, pleura and beard white to gray

pollinose and pilose. Legs brown, the tarsi, especially fore pair, nearly black.

Wings apically fumose, the dark area clearly demarcated from hyaline discal

area. Costal cell brown. Fork of 3rd vein appendiculate and with a distinct

small dark cloud; ecrossveins at end of discal cell also usually with faint to strong

clouds. Antennae variable, yellow to brown or bicolored.

Male. The male is like the female in color and vestiture, though the wings

are usually paler and clouds at end of discal cell fainter. Upper eye facets bare,

greatly enlarged and well demarcated from the lower facets, occupying fully 2/3

eye area,making the head distinctly wider than thorax. Antennae more slender

than in female. A small tubercle is sunk between the eyes at vertex and there is

a marked post occipital fringe of forward pointing hairs.

Material examined. In the following enumeration of specimens we have not

included material identified or reported by Fairchild in previous years unless the

Specimens are currently before us, since the complexity of the group was not

then appreciated, and some determinations may have been erroneous. We believe

36 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

that reports of the species from Costa Rica in Hogue and Fairchild (1.c.) are

probably correct, though no Costa Rican specimens are now before us.

Panama: Bocas del Toro Prov., 1 female, Almirante, Yellow Fever Camp,

Shannon trap, 20-V-1952 (Galindo leg.); 2 females, same locality, VII-1951; 2 fe-

males, Almirante, 16,17-V-51 (Trapido); 1 female, Rio Terribe, Timishik, V-1962.

Panama Prov., 1 female paratype, Rio Pequeni, 21-VIII-1940 (G.B. Fairchild); 1

female, Candelaria Hydrographic Station, Rio Pequeni, V-1966 (Galindo); 1 fe-

male, Rio Mamoni, Salto de Madrono, 20-V-1957 (Johnson); 1 female, Cerro Cam-

pana, on hostel window, 6-IX-56, (G.B. Fairchild); 1 female, Cerro Azul, La

Eneida, 23-V-61 (Galindo leg.) Colon Prov., 1 female, Mandinga, V-1964 (Galindo

leg.). San Blas Prov., 1 female, Sasardi, Interoceanice Canal, Study No. 7078, 1-

X-1967 (D.G. Young). Darien Prov., 1 female, Morti River, 21-IX-67, I.C.S. No.

1997; 2 males, 3 females, Rio Tacarcuna Yellow Fever Camp, 1900 ft elev. 21-VI,

11,12-VII-1963 (G.B. Fairchild). Colombia: . Antioquia Dept., 1 female, Rio

Anori Valley, Providencia, 30-V-70 (C.H. Porter); 1 female, SW Zaragoza, IX-1970

(D.G. Young); 1 male, La Tirana, Providencia, 25 km W of Zaragoza, 30-V-71, in

house at night (D.G. Young); 2 males, Rio Anori Valley, La Tirana, 17-IX and 20-

V-1971, at light (C.H. Porter). Valle Dept., 3 females, Anchicaya Dam, 500 m

elev. 10-VII and 18-VIII-75 (R. Wilkerson); 3 females, Anchicaya Dam, 500 m elev.

10-VIII-73 (Wilkerson and Young); 1 male, 50 females, Lower Anchicaya, Very Wet

Forest, 400 m elev., 28-1, 5-II, 2-IV, 10, 11, 14-VI, 16-VIII-1975 (R. Wilkerson); 2

females, 25 km E of Buenaventura, 50 m elev., 12-VIII-1973 (Wilkerson and

Young); 6 females, 41 km E of Buenaventura, 140 m elev., 5-XII-75 and 19-III-76

(R. Wilkerson). Cauca Dept., 1 female, Guapi, 10 m elev., 10-V-76 (R. Wilker-

son); 69 females from various localities in Valle and Cauca, III, VII, VI, XI-1977

(M.A. Tidwell). Intendencia Putumayo, 4 females, Mocoa, 650 m elev., Very Wet

Subtropical Forest, flight trap with CO2, 15-IV-76 (R. Wilkerson); 2 females,

Puerto Asis, 1240 m elev., 15-IV-76 (R. Wilkerson). Ecuador: Pastaza Prov., 6

females, vic. Puyo, 5-XI-81 (R. Wilkerson). Guayas Prov., 1 female, Balao Chico,

23-29-IV-1963 (L. Pena); 1 female, Bucay, 13-11-73, 13-VI-1977 (D.L. Vincent); 1

female, Rio Cononaco, Anglo Air Strip, 70 mi SE Limoncocha, 29-V-76 (J. Cohen);

1 female, Puerto Montufar, 28-IV-76 (J. Cohen). Peru: Cuzco Dept., 1 female,

Quineemil, 2450 ft elev. 16-31-X-1962 (L.E. Pena). Huanuco Dept., 1 female,

Tingo Maria, 670 m elev., I-1942 (Weyrauch). Madre de Dios Dept., 7 females,

Tambopata Reserved Zone, 30 km SW Puerto Maldonado, 290 m elev., X-XI-1982

(R. Wilkerson). Loreto Dept., 1 female, Iquitos, III-IV-1930 (R.C. Shannon).

Bolivia: Beni Dept., 7 females, S. Inicua River, 1100 m elev., 15-18-1-1976 (L.E.

Pena).

Diseussion. St. pequeniensis forms one of a group of four similar species, S.

flavesecens Kroeber., S. ecaptiroptera Kroeber and S. anoriensis n. sp. being the

other three. From all of these it is distinguished by lack of yellow pilosity, ei-

ther on the abdomen or elsewhere, all pale areas being whitish or gray pollinose

and pilose. The apical dusky area on the wing is usually all dark and distinctly

demareated from the more hyaline discal area. Our series of pequeniensis,

though limited, shows that most specimens from east of the Andes have largely

yellow antennae and most of those from west of the main chain have bicolored

antennae, with the whole style black. Specimens from Panama may have anten-

nae all yellow, all brownish or bicolored.

This species is apparently not arboreal nor crepuscular. It has been taken at-

tacking man and in flight traps, while males have been taken at light. Most col-

leetions have been made in heavy forest in areas of high rainfall, mostly at low

to moderate elevations.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 37

Stypommisa prunicolor (Lutz)

(Figs. 52-54, 86)

Tabanus prunicolor Lutz 1912, Tabanideos, in Comm. Linhas Telegr. Es-

trat. Mato Grosso ao Amazonas, Annexo No. 5, Historia Natural, 6,

pl. 1, fig. 5, lectotype female Brazil, Mato Grosso (Rio). Fairchild:

1961, Mem. Inst. Oswaldo Cruz, 59(2): 232, pl. 2, fig. 1 designates

lectotype; 1971, Cat. Dipt. Amer. S. of U.S., fase. 28: 100.

A small grayish black species with wholly yellow antennae, large, rectangular,

frontal callus filling width of frons, basally blackish femora, and wings with a

single small brown spot at fork of 3rd vein and fore border apically brownish.

Male. Not known.

Material examined. The types were studied by Fairchild in the Instituto Os-

waldo Cruz in 1959 and figured in 1961 (1.¢c.). Aside from these specimens, we

have seen the following material. Brazil: Maranhao State, 1 female, Marmelo, at

human bait, 8-XI-62 (Werner). Mato Grosso State, 6 females, Sinop, X-1976 (M.

Alvarenga leg.); 8 females, Humboldt, Rio Aripuana, 59° 27'w, 10°10'S, 12-16-VIII-

74 (D.G. Young). 1 female, Rio Madeira, Abuna (Mann and Baker) Peru: Madre

de Dios Dept., 6 females, Tambopata Reserved Zone, 30 km SW Puerto Maldon-

ado, 290 m elev., 15-IX-82, 15-30-VIII-82, 20-31-X-82 (R. Wilkerson).

Discussion. The species is very similar to S. jaculator, but is darker, with all

femora at least blackish at base, broader frons (index 4.2-4.9, x = 4.6) with the

tapered callus set in a broad oblong bare area as wide as frons and about 1.3 to

1.2 height of frons. Antennae are wholly yellow, usually bicolored in S. jaculator.

Otherwise color and wing pattern are as in S. jaculator.

There is also considerable resemblance to S. modica, but the latter has wholly

black legs except bicolored fore tibiae, a strong cloud on crossveins at apex of

discal cell, frontal callus separated from sides of frons, and the abdomen without

large pale pilose triangles or broad sutural bands, being generally entirely black

pilose except at extreme sides and sometimes with tiny silvery hair tufts on last

few segments dorsally.

Stypommisa rubrithorax (Macquart)

(Figs. 55-58)

Tabanus rubrithorax Macquart 1838, Dipt. Exot. 1(1): 142, Minas Gerais,

Brazil, holotype female (Paris) [examined]. Kroeber, 1934, Rev.

Ent. 4(3): 314.

Stenotabanus rubrithorax: Fairchild 1956, Smiths, Miscell. Coll. 131(3):

yee

Therioplectes punctulipennis Enderlein 1925, Mitt. Zool. Mus. Berlin 11(2):

262, Mato Grosso, Brazil, Lectotype female (Berlin) [examined].

Enderleiniella punctulipennis: Kroeber 1932, Stettin Ent. Zeitg. 93: 153.

Dudaella punctulipennis: Strand 1932, Fol. Zool. Hydrobiol., Riga 4: 142.

Stypommisa rubrithorax: Fairchild 1966, J. Med. Ent. 3(1): 16, fig. 28 (as

punctulipennis Enderlein); 1971, Cat. Dipt. Amer. S. of U.S., Fase. 28:

82.

38 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

Stypommisa n. sp. aff. seythropa Schiner: Strickman 1982, J. Med. Ent.

19(4): 402, Paraguay.

A small slender pale brownish species usually with bicolored antennae, narrow

frons, unicolorous pale legs, very faintly spotted or unspotted wings, and yellow-

ish brown abdomen with a black integumental mark on 2nd tergite and pale me-

dian dorsal pollinose triangles on tergites 2 to 6.

Female. Length 9-10 mm, wing length 8-9 mm. Eyes bare, pattern not

recorded. Head structures as figured. Frontal index, 4.1-4.4 (x = 4.3, n = 10); di-

vergence index, 1.0-1.2 (x = 1.1). Mesonotum grayish brown, obscurely striped.

Legs yellowish, pale haired, except apex of fore tibiae and all tarsi darker haired.

Wings including costal cell yellowish hyaline, with barely perceptible clouds on

fork and apex of diseal cell, basicosta rarely bare, usually with 1-12 setae. Ap-

pendix on fork of 3rd vein shorter than base of vein, rarely absent. First poste-

rior cell widely open. Abdomen light brown in ground color, with a usually

prominent dark integumental triangle or streak on 2nd tergite and 3rd and suc-

ceeding tergites progressively darker. Tergites 2 to 6 with prominent pollinose

middorsal triangles and narrow pale-pollinose hind margins, slightly widened at

Sides.

Male. Length 9.5 mm, of wing 8.5 mm. Eyes with facets in center slightly

enlarged, but not well differentiated or demarcated from small facets. Eye dor-

sally short pilose, the pilosity denser and longer on upper and central part of eye,

nearly absent on lower 1/4. Palpi porrect, long and slender. Coloration and

wings as in female and easily associated with it. Matches specimen from Brazil,

Coqueiros Cajuru, Sao Paulo, 27-IX-1954 (Barretto and Duret).

Material examined. In addition to the types of both names: Brazil: Sao Paulo

State, 1 male, 2 females, Coqueiros, Cajuru, 27-IX- to 1-X-1954 (Barretto);

Goyaz State, 2 females, Anapolis, 23-IX-1936 (homotype of rubrithorax Macq.)

and 17-X-1936 (G.B. Fairchild); 4 females, 1907 (Baer). Mato Grosso State, 23

females, Rio Aripuana, Humboldt, VIII-1974 (D.G. Young); 4 females, Aracaju,

XII-1937 and IX-1937, (1 female homotype of punctulipennis End.) (Fairchild and

Shannon); 1 female, Chapada (Lane). Paraguay: Amambay Dept., 3 females, Par-

que Nacional Cerro Cora, 16-XIII-1980 (D. Strickman). Argentina: Jujuy Prov., 1

female, Ing. Ledesma, near General San Martin, 10-X-1968 (L. Pena). Bolivia:

Santa Cruz Dept., 2 females, 10 km E Warnes, 19-22-VIII-83 (R. Wilkerson). 1

female without data except "2945". One of us has also seen considerable addi-

tional material from Goyaz and southern Mato Grosso, not now before us.

Discussion. This little species ean generally be separated from others in the

group by having the wing spots very faint or rarely absent, the appendix at fork

of 3rd vein short or nearly absent, the legs pale, except occasionally the femora

dusky, the second abdominal tergite yellow to brown, usually with a small median

dark integumental streak, and all tergites with conspicuous median pale pollinose

triangles. The specimens from Sao Paulo, Goyaz and Mato Grosso mentioned

above all have entirely yellowish legs and largely yellow to light brown abdomens;

the three from Paraguay are much darker, with femora dark brown to nearly

black. We detect no significant differences in structure or frontal indices, how-

ever, and believe these are but local variants. The specimen from Argentina is

not now before us, but we believe it was pale like those from Mato Grosso.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 39

Stypommisa scythropa (Schiner)

(Figs. 62-64, 87)

Tabanus scythropus Schiner, 1868, Reise Novara, Zool. 2(1,B) Diptera pp.

92-93, holotype female, Rio de Janeiro, Brazil(Vienna) [examined].

Fairchild, 1967, Pacific Ins. 9(2): 251.

Sciladynus seythropus: Kroeber 1930, Zool. Anz. 86(9-10): 256.

Dudaella seythropus: Kroeber 1934, Rev. Ent. 4(2): 276.

Hybomitra beieri Philip 1958, J. Kansas Ent. Soc. 31(2): 180, holotype

female, Cantareira, S. Paulo, Brazil (Vienna) [examined]. Fairchild

1967, Pacific Ins. 9(2): 244, synonymized with seythropa.

Stypommisa seythropa: Fairchild 1971, Cat. Dipt. Amer. S. of U.S., Fase.

28: 82.

Stypommisa punctipennis Enderlein, 1925, Mitt. Zool. Mus. Berlin 11(2):

369, lectotype female, Rio Grande do Sul, Brazil (Berlin). Kroeber

1929, Zool. Anz. 83: 115, fig. 4 (wing). Fairchild 1966, J. Med. Ent.

3(1): 16, fig. 28, designates lectotype; 1971, Cat. Dipt. Amer. S. of

U.S., Fase. 28: 82 (NEW SYNONYM).

Therioplectes punctulipennis Enderlein 1925, Mitt. Zool. Mus. Berlin 11(2):

262, male only. Fairchild 1966, op. cit. 3(1): 16.

Sziladya punctulipennis: Kroeber 1930, Zool. Anz. 86(9-10): 555.

A small light to dark brown species with heavily spotted wings and a promi-

nent middorsal row of pale pollinose triangles on abdomen. Antennae bicolored

and at least the fore tibiae bicolored.

Female. Length 9-12.5 mm, of wing 8.5-10 mm. Eyes bare, no pattern

recorded. Frontal index 3.7 to 4.5 Head structures as figured. Antennal plate

yellow to dark orange, sometimes apically blackish or with a black streak, the

Style black. Palpi yellow, black pilose. Mesonotum blackish brown, clearly to ob-

securely striped. Pleura grayish pollinose and pilose. Legs with femora and coxae

black to dark brown, contrasting with yellow tibiae, the fore pair bicolored. Wing

as figured, the appendix shorter than stem, often absent, spots on crossveins and

fork prominent and usually some tinting of wing apically. Basicosta with few to

many setae, but these always smaller and sparser than on costa. Abdomen brown

to nearly black, darkest on midline, especially first 2 segments, with sharp dis-

crete pale triangles on tergites 1 to 6 and narrow pale hind margins widening

slightly at sides.

Male. Length 10.0-11.5 mm of wing 9-10 mm. Eyes holoptic, densely long

pilose, the facets but slightly differentiated in size and not demarcated, the pi-

losity longest on upper part of eye. A small tubercle between eyes at vertex, not

extending above eye level. Frontal triangle silvery white pollinose, contrasting

with brown black-haired face. Palpi brown, long dark haired. Antennae more

Slender than female, plate yellow to dark red, style black. Mesonotum subshiny,

blackish, rarely with traces of anterior gray stripes. Legs as in female, the tib-

lae nearly black. Wings as in female, but spots on fork and crossveins smaller

and paler. Abdomen conical, colored as in female, but median pale triangles

broader and smaller and sides of first tergites often contrasting more with cen-

tral blackish area.

Material examined. Brazil: Santa Catarina State. 3 females, 5 males, Nova

Teutonia, X-XI, (F. Plaumann). A female and male were compared and found to

40 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

agree with the types of punctipennis on loan from Berlin Museum. Rio de_

Janeiro State, 5 females, Tingua, no date or collector, one compared and agreeing

with types of T.seythropa and H. beieri in Vienna; 1 female, Mangaratiba, VIII-

1938. Argentina: 1 female, 1 male, Iguassu. Paraguay: Amambay Dept., 1 fe-

male, 16-VIII-1980 (D. Strieckman). 1 male, Villa Rica, (F. Schade). In addition

Fairchild has determined as this species the following material: 1 male, Pelotas,

Rio Grande do Sul, Brazil in Coll. M. Leclercq; 71 males, 3 females, Nova Teuto-

nia, Sta. Catarina, Brazil, in CNC.

Discussion. Specimens from Santa Catarina and Sao Paulo States are gen-

erally larger and darker than those from Rio de Janeiro, and our homotypes of

punctipennis and scythropa are like the larger and smaller specimens respectively.

There are, however, specimens of intermediate size and coloration, and we are

unable to point out any consistent differences among our series. Our dated

Specimens indicate that the species flies from Aug. to Nov., late winter to early

Spring in the latitude of Sao Paulo.

The species can be separated from S. paraguayensis by the less boldly marked

mesonotum,, dark femora, and usually less heavily spotted wings, though the most

Striking differences are shown in the male eye, densely pilose and without en-

larged upper facets in S. seythropa, bare and with greatly enlarged upper facets

in S. paraguayensis. In addition, we have 3 females from Paraguay, Villarica, Col.

Independencia and 1 female from Maracaju, Mato Grosso, which are probably this

species, but have broader frons, indices 3.3-3.9, are paler, reddish, with weaker

wing spots. All are either old and faded or otherwise poorly preserved. One

agreed structurally with type of S. paraguayensis but was much paler. A further

3 females from Rio de Janeiro, Campos Murundu, Aug. 1978, Alvarenga leg. have

much narrower frons, indices 5.1-5.8, no appendix, small wing spots but dark yel-

low costal cell, pale legs and body. They are not well preserved, but may repre-

Sent an additional species. We are probably dealing here with a complex of

species, but available material is neither abundant enough nor sufficiently well

preserved to enable us to sort them out.

Stypommisa spilota n. sp.

(Figs. 65-67, 88)

A moderate sized dark brown species with a blackish mesonotum, lighter

brown abdomen, slender bicolored or all yellow antennae, and heavily spotted

wings which have the spot on fork often extended as a spur along R. to wing

margin.

Female. Length 12 mm, of wing 11 mm. Eyes bare, greenish black without

pattern (revived). Head structures as figured. Frons yellowish gray pollinose,

Slightly darker at vertex, where it is beset with numerous short black hairs.

Ocellar tubercle shiny, black, with vestiges of two lateral ocelli. Callus black,

shiny, merging above into a broad ridge. Subeallus thinly pollinose, econeolorous

with frons; frontoclypeus slightly more yellowish, genae gray, sparsely gray

bearded. Antennae orange, the scape and pedicel beset with short black setae,

the dorsal angle of basal plate with a few short black setae, the style slightly

darker, the terminal 2 annuli black or nearly so. Palpi pale tan, whitish pollinose,

black haired. Proboscis blackish, membranous, slightly exceeding palpi.

Mesonotum and scutellum black in ground color, the notopleural lobes brown-

ish, thinly gray pollinose, unstriped, sparsely beset with erect black and recum-

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 41

bent brassy hairs. Pleura and coxae black to dark brown, gray pollinose, white

haired. Last 2 two pairs of legs brown, the femora mainly pale haired, the tibiae

mainly dark haired, the tarsi dusky. Forelegs darker, the femora nearly black,

the fore tibiae bicolored, the basal half yellowish, sparsely dark haired, the apical

half black, densely black haired, as are tarsi. Wings as figured. Costal cell yel-

low, wing membrane otherwise hyaline except for prominent brown clouds around

erossveins and fork of 3rd vein, much larger around end of diseal cell and fork,

where the cloud is often extended along R, nearly or quite to wing margin.

There is also a brown diffuse pateh in wing apex beyond the brown stigma. Ap-

pendix at fork shorter than the long subtending vein segment. Basicosta pointed,

beset with short black setae, though less densely setose than adjoining costa.

Abdomen pale brown in ground color, semitranslucent on tergites 1 to 3,

darker opaque brown on remaining segments, beset throughout with short black

hairs except for small, faint, sparse patches of pale hairs on middle of posterior

borders of tergites 2-5, and complete hind border of 6. Lateral corners of ter-

gites 4-6 also with some pale hairs. Pollinosity sparse, brown, the whole dorsum

subshiny. Venter paler, more evidently pale pollinose and entirely sparsely pale

haired.

Male. Not known.

Material examined. Holotype female, Peru, Cuzco Dept., Quincemil, 13-3-

VIII-1962, 780 m elev. (L. Pena). Paratypes, 85 females, same locality and col-

lector with dates in August and September, 1962. Holotype in coll. of L.L.

Pechuman, paratypes in coll. Peehuman, CNC, FSCA and the authors.

Discussion. This species resembles S. changena, S. bipuncta and S. kroeberi in

having the dark clouds around fork and apex of discal cell much more prominent

than those at ends of basal cells. From S. changena it differs in having the ap-

pendix at fork of 3rd vein (R iH ) short, shorter than the long subtending vein

segment, in generally paler wings, especially the costal cell, and in narrower

frons. From S. bipuncta and S. kroeberi it differs in blackish mesonotum and

scutellum,, brown and yellow in 2 latter species, and in more heavily infuscated

wings. Stypommisa bipuncta also differs in much smaller size and quite faint and

small wing spots, while S. kroeberi has wholly yellow antennae. Frontal indices

of 7 specimens of S. spilota range from 5.7 to 7.2 with a mean of 6.3. The name

is from the Greek spilota,meaning stained or spotted, in reference to the heavily

marked wings.

Stypommisa U-nigrum Philip

(Fig. 89)

Stypommisa u-nigrum Philip 1977, Proc. Ent. Soe. Washington, 79(1): 28-

30, figure 1.

A large species for the genus, with prominently yellow banded abdomen and

yellow wings with very large and prominent brown spots on all crossveins and

fork, and apex and hind margin grayish infuseated.

The original description is as follows: "Holotype female, 12.5 mm. Eyes

bare, unbanded (relaxed). Frons brown pollinose, narrow, index 1/5.8, nearly

parallel-sided, patch of coarse black bristles at vertex above small but distinct

42 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

tubercle with 3 compact vestigial ocelli; callosity dark brown, taller than broad,

tapered abruptly above into slender keel, separated narrowly from eye margins

and from yellow pollinose subecallus below, the usual suture across top of latter

indistinct. Face and cheeks gray pollinose with mixed gray and some black hairs.

Two basal antennal segments brown with black hairs, scape a little longer than

tall, plate red with low dorso-basal angle, subequal in length to sharply black

style. Palpi dusky gray, black-haired, long and slender but blunt, nearly as long

as unsheathed stylets.

Thorax pale pollinose and mostly pale pilose, accentuated in the prescutellar-

seutellar area behind a peculiar brown, black-haired, wide U-shaped marking and

a narrow midline forward from it. Fore legs dark brown, black-haired, tibiae

reddish with pale hairs in basal thirds. Two hind pairs of legs reddish with mostly

pale hairs, more black on dorsums of femora and in hind tibial fringes. Wing with

unusual pattern of whitish membrane around prominent clouds, the apical and

hind margins smoky, accentuated in certain lights. Spur veins as long as stems.

Halters pale brown. Basicostas with a few setulae.

Abdomen reddish brown, darker caudally, black-haired across bases of terga,

the incisures broadly reddish yellow (as much as 1/2 of 3 and 4) with yellow hairs;

venter reddish, entirely yellow-haired.

Holotype and 2 paratype females, Mexico: Chiapas, Montebello National Park,

17-V-69. H.J. Teskey. In CNC (Canadian National Collection) No. 12925; one in

CAS (California Academy of Sciences). Wear in the latter has reduced the

prominence of the ineisural bands and the scutal marking. This U-shaped marking

in front of the pallid scutellum and the peculiar whitish wing membranes sur-

rounding unusually heavy clouds with smoky apical and hind margins distinguish

this from related species. The banded abdomen is unusual in the genus."

Material examined. One female same data as holotype and paratypes. (The

specimen bears notes as to frontal index ete. apparently in Philip's hand, though

not labelled as paratype. It was one of ten specimens sent for identification

from Dr. Teskey, the other nine being topotypes of Stenotabanus apaches Philip

with identical data to those recorded for the holotype of that species). Mexico:

Chiapas State, six females, Lagos Montebello, 19-20-V-1970 (R.L. Dressler). All

lack most of abdomens due to a visitation of ants shortly after they were col-

lected. Frontal and divergence indices of this material are as follows: 4.8-5.9

(x = 5.32, n = 7) and 1.3-1.6 (x = 1.4).

Discussion. The original description stresses a dark U-shaped mark on

mesonotum, but this is visible only in some of the above specimens and prominent

in none. The costal cell is strongly yellow, as is the wing membrane between the

large brown spots, not white in any of our specimens. The frontal calli of the

Lago Montebello specimens are black, only the topotype having it brown, as fig-

ured for the holotype. All known specimens of this species are from this locality,

all taken in May. This is one of only 2 Stypommisa recorded from Mexico, the

other being S. eaptiroptera.

The specimens of Stenotabanus apaches mentioned above were the first we

had seen of this species. Unlike Philip, we believe these are better placed in

Stenotabanus, in spite of the strongly spotted wings. The frontal characters with

eallus as wide as frons, vestiges of black hair pateh surrounding median callus,

bare basicosta, and well developed ocellar tubercle are very similar to those of

Stenotabanus flavidus Hine, which is even larger. Both frons and wing spotting

are like the smaller Stenotabanus brunettii and St. batesi from the Antilles. The

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 43

two green eye stripes in St. apaches also would be most unusual in Stypommisa,

but expected in Stenotabanus.

Stypommisa venosa (Bigot)

(Figs. 68-71)

Tabanus venosus Bigot 1892, Mem. Soc. Zool. France, 5: 685, Brazil,

Para.

Stypommisa venosus: Kroeber 1930, Zool. Anz. 86(9-10): 251-252, fig. 1;

1934, Rev. Ent. 4(2): 275.

Stenotabanus venosus: Fairchild 1956, Smiths. Miscell. Colls. 131(3): 31.

Wrongly synonymized Stypommia maculipennis Kroeb. 1929 with this

Species.

Stypommisa venosa: Fairchild 1971, Cat. Dipt. Amer. S. of U.S. Fase.

28¢:82%

A pale yellowish brown species with paler scutellum, mainly dark haired ab-

domen and wings with strong round brown clouds at fork of 3rd vein and apex of

diseal cell but not at ends of basal cells.

Female. In 1980 we borrowed Bigot's type, courtesy of J.E. Chainey, from

the British Museum and append here a description of it.

Length 10.5 mm, of wing 11.5 mm. Head characters as figured. Eyes bare,

pattern not discernible. Frons and subeallus reddish brown pollinose. Callus

nearly black. Antennae reddish brown, dorsally dark haired, the style blackish,

contrasting with basal plate. Palpi light brown, pale pollinose, entirely long black

pilose. Face and cheeks gray pollinose, sparsely pale haired. Mesonotum and

scutellum reddish brown in ground color, the vestiture lost so that no pattern is

evident. Pleura apparently pale pollinose, dark haired except for tufts of pale

hairs on propleura and beneath wing insertions, but much denuded and encrusted

with dirt. Legs reddish, largely pale haired, the fore coxae infuseated and tarsi

and apices of at least fore tibiae dusky and dark haired. Halters with orange

knobs. Wings as figured, costal cell orange, stigma yellow, membrane quite dirty,

apparently clear except for brown clouds around crossveins.

Abdomen much denuded and crusted, at least the first 2 segments clear yel-

lowish brown, without integumental dark markings or visible pollinose markings.

Discussion. We cannot exactly match this specimen with any in our posses-

sion. It comes closest in structure to S. fulviventris, but that species lacks

strong wing spots and has a narrower frons. S. changena is very similar in most

respects, but has shorter and stubbier 3rd antennal segment and integument of

thorax and scutellum are blackish and knob of halters dark brown, differences

seemingly too great to be explained by fading of the type of S. venosus. No

Brazilian species we have seen is like S. venosus, and in view of Bigot's known

disregard of localities, the type may have come from elsewhere.

Specimens determined by Kroeber in Dresden Museum from eastern Peru are

not the same, nor does his description and figure published under S. venosus

(1930) agree at all with the type. A specimen from Nanegalito, Ecuador, was

thought to agree with Bigot's type by Philip (in litt.) and we have an old specimen

from Gualea, Pichincha, Ecuador, which also agrees fairly well with the type, but

it lacks antennae. Both these specimens are closest to S. furva in structure and

wing pattern, but are light brown rather than bluish black. We give a figure of

44 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

the type, but lack of fresh material prevents us from placing the species in the

key. It will probably key out with S. kroeberi or S. bipuncta.

The light brown species from Bolivia and eastern Peru which seems to repre-

sent the form identified as S. venosa by Kroeber is described earlier in this publi-

cation as S. kroeberi. It differs from the type of S. venosa Bigot by shorter an-

tennal style relative to the basal plate, narrower frons with smaller and more

slender callus, usually with wing apex beyond fork more or less infuseated, and

has pale ecoxae and yellowish scutellum.

The original description is of little use. In 1953, Fairchild studied the type in

British Museum and in addition to the above comments his notes are appended

here, made partly at the time and partly later.

Type: 1 female in BM(NH) Red circled Type label, an indecipherable Bigot

locality (?) label, a Bigot hand label with "Tabanus venosus female n. sp. inedit

Alger Fevr. 1890 J. Bigot Bresil" and a Bigot coll.-- BM accession label. The

type is a female as indicated by the description and Bigot's label, not a male as

published. The specimen is fairly well preserved, lacking one antenna and being

somewhat dirty. I ean detect no structural differences between this and mac-

ulipennis Kroeb. (= furvus Hine) In color the type is lighter, being a uniform light

cinnamon brown. The specimens labelled "Stypommia venosa Big." from Peru,

Urubamba fl. and Pichis, Pto. Yessup, Schnuse coll. ex Dresden Museum, are not

the same in my opinion, though they apparently formed part of the material

studied by Kroeber (1930). His figure seems to have been taken from another

Species entirely as the antennae, palpi and frons are not at all like those of the

Type."

Stypommisa xanthicornis n. sp.

(Figs. 59-61)

A small brown species with faintly spotted wings and wholly yellow antennae.

Female. Length 10 mm, of wing 10 mm. Eyes bare, no pattern evident.

Head characters as figured. Frontal index 5.8; divergence index 1.5. Frons

grayish yellow, without darker areas or evident hairs. Callus and tubercle at ver-

tex reddish brown. Subeallus and upper genae concolorous with frons. Fron-

toclypeus and lower genae gray. Beard sparse and short, pale brown. Antennae

wholly yellow, the first 2 segments sparsely brown haired dorsally. Palpi rather

inflated, white pollinose, beset with short black hairs. Proboscis black, membra-

nous, unusually short, hardly longer than palpi. Occiput silver gray pollinose,

without marked postocular fringe of setae.

Mesonotum reddish brown, subshiny, without marked stripes. Scutellum con-

eolorous. Notopleural lobes concolorous, slightly gray pollinose, as are pleura and

sternum. Legs yellow, tips of hind tibiae, distal half of fore tibiae and all tarsi

darkened. Wings faintly yellowish hyaline, costal cell and stigma yellow,

erossveins and fork of 3rd vein with faint diffuse brownish clouds. Appendix at

fork as long as basal segment in one wing, much shorter in the other. Basicosta

pointed, with 2 small black setae. Halters light yellowish brown.

Abdomen light orange brown, subshiny, the hind margins of tergites vaguely

paler, and faint indications of pale median pollinose triangles, only clearly evi-

dent on Ist and 2nd tergites. Hairs sparse, long reddish, pale yellowish on median

triangles and margins. Beneath abdomen gray pollinose, wholly pale haired.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 45

Type data. Holotype female, Brazil, Santa Catarina State, (Luderwaldt), in

FSCA. The specimen bears a Kroeber det. label 1930 as Stypommisa affinis and

was received from the Mus. Zool. Polonicum, Warsaw, through the kindness of Dr.

Trojan. It probably formed part of the collections of the Stettin Museum. it

agrees neither with a paratype of S. affinis in Vienna, with Kroeber's description

of S. affinis, nor with a specimen in Munich determined as S. affinis by Kroeber.

Discussion. The species differs from other regional species, particularly S.

fulviventris, by the shorter wholly yellow antennae, smaller size, more evident

wing spots, and stouter palpi. There were supposed to be other specimens of this

Species in Stettin, but we do not know if they were the same, as only this one

was sent. |

LITERATURE CITED

ENDERLEIN, G. 1923. Vorlaufig. Diagnosen neuer Tabanidengenera. Deutsch.

Ent. Zeitsehr. 4-545. 1923(5): 54.

ENDERLEIN, G. 1925. Studien an blutsaugenden Insekten I. Grundlagen eines

neuer Systems der Tabaniden. Mitt. Zool. Mus. Berlin 11(2): 255-409.

FAIRCHILD, G. B. 1947. Additional notes on The Tabanidae of Panama. Ann.

Ent. Soe. Amer. (1946) 39(4): 564-575, 8 figs. ;

FAIRCHILD, G. B. 1953. Arboreal Tabanidae in Panama. Proc. Ent. Soe. Wash-

ington 55(5): 239-243.

FAIRCHILD, G. B. 1961. The Adolpho Lutz collection of Tabanidae (Dipt.) I.

The described genera and_ species, condition of the collection and selection

of lectotypes Mem. Inst. Oswaldo Cruz 59(2): 185-249, 2 pls.

FAIRCHILD, G. B., 1967. Notes on neotropical Tabanidae IX. The species de-

scribed by Otto Kroeber. Studia Entom., Sao Paulo. 9(1-4): 329-379, 33 figs.

(1966).

FAIRCHILD, G. B. 1967a. Notes on neotropical Tabanidae X. The species de-

scribed by J.R. Schiner and others. Pacific Ins. 9(2): 243-256.

FAIRCHILD, G. B. 1969. Notes on Neotropical Tabanidae XII. Classification and

distribution with keys to genera and subgenera. Arq. Zool. Sao Paulo.17(4):

199-255.

FAIRCHILD, G. B. 1971. A catalogue of the Diptera of the Americas south of

the United States. Fasc. 28, family Tabanidae. Mus. Zool. Univ. S. Paulo, 1-

163.

FAIRCHILD, G. B. 1975. Notes on neotropical Tabanidae: Genus Philipotabanus

Fehld., Subgenus Mimotabanus Fchld. Ann. Ent. Soc. Amer. 68(4): 689-694, 2

pls.

HOGUE, C. L.,and G. B. FAIRCHILD 1974. A revised check list of the Tabanidae

of Costa Rica. Rev. Biol. Trop. 22(1): 11-27.

KROEBER, O. 1929. Ueber einige kleinere Gattungen der sudamerikanischen Ta-

banini. Zool. Anz. 83: 47-63, 115-137, 13 figs.

KROEBER, O. 1930. Nachtrage zu den Kleinen Gattungenden der su-

damerikanischen Tabanini. Zool. Anz. 86: 248-265, 8 figs.

KROEBER, O. 1934. Catalogo dos Tabanidae do America do Sul e Central, in-

cluindo o Mexico e as Antilhas. Rev. Ent., Rio de Janeiro 4(2-3); 222-276,

291-333.

LUTZ, A. and A. NEIVA, 1914. As Tabanidae do Estado do Rio de Janeiro. Mem.

Inst.Oswaldo Cruz 6(2): 69-80.

46 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

PHILIP, C. B. 1969. Deseriptions of new neotropical Tabanidae and new records

for Argentina. Acta. Zoologica Lilloana, XXII (1967): 105-132.

PHILIP, C. B. 1969a. New or little known neotropical Tabanidae. Pan-Pacific Ent.

45(2): 147-152.

PHILIP, C. B. 1977. New North American Tabanidae (Diptera) XXII. Additional

new Diachlorini from Mexico. Proc. Ent. Soe. Washington 79(1): 28-32, 3 figs.

WILKERSON, R. C. 1979. Horseflies (Diptera: Tabanidae) of the Colombian de-

partments of Choco, Valle and Cauca. Cespedesia (Cali) 8(31-32): 87-435, 119

figs.

EXPLANATION OF FIGURES 1-18

Figs. 1-3, S. apiealis, holotype: 1, antenna; 2, palp; 3, frons. 4-6, S. anorien-

sis, holotype: 4, antenna; 5, palp; 6, frons. 7-9, S. apicalis, paratype: 7, an-

tenna; 8, palp; 9, frons. 10-12, S. aripuana, paratype: 10, antenna; 11, palp; 12,

frons. 13-15, S. bipuneta, holotype: 13, antenna; 14, palp; 15, frons. 16-18, S.

eallicera, Brazil, Parque Nacional da Serra dos Orgaos, Teresopolis: 16, antenna;

17, palp; 18, frons.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) 47

48 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 19-36

Figs. 19-21, S. eaptiroptera, Panama, Canal Zone, 23-VI-1944: 19, antenna;

20, palp; 21, frons. 22-24, S. changena, holotype: 22, antenna; 23, palp; 24,

frons. 25-27, S. flavescens, Peru, Quincemil, Cuzeo, 16-31-X-1962: 25, antenna;

26, palp; 27, frons. 28-30, S. fulviventris, Brazil, Sao Paulo, Itatiaya, Est. Biolo-

gia, 15-I-1936: 28, antenna; 29, palp; 30, frons. 31-33, S. hypographa neofurva,

Bolivia, Alto Beni, S. Inicua, 15-18-I-1976: 31, antenna; 32, palp; 33, frons. 34-

36, S. kroeberi, paratype, Bolivia, Sara: 34, antenna; 35, palp; 36, frons.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae)

ca tele ye mae

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20 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 37-54

Figs. 37-39, S. furva, Bolivia, Dept. Cochabamba, Prov. Chapare, El Palmar,

8-X-1943: 37, antenna; 38, palp; 39, frons. 40-42, S. glandicolor, Brazil, Abuna,

Rio Madeira: 40, antenna; 41, palp; 42, frons. 43-45, S. glandicolor, Colombia,

Meta, Villavicencio, Guayuriba, Carretera al Puente: 43, antenna; 44, palp; 45,

frons. 46-48, S. pequeniensis, Peru, Quincemil, Cuzco, 16-31-X-1962: 46, an-

tenna; 47, palp; 48, frons. 49-51, S. paraguayensis, Brazil, Sao Paulo, E.S.P.,

Cantareira, VIII-1958: 49, antenna; 50, palp; 51, frons. 52-54, S. prunicolor,

Brazil, Mato Grosso, Rio Aripuana, 12-16-VIII-1974: 52, antenna; 53, palp; 54,

frons.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae)

au Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 55-71

Figs. 55-58, S. rubrithorax, holotype of S. punctulipennis: 55, antenna; 56,

palp; 57, frons; 58, wing. 59-61, S. xanthicornis, holotype: 59, antenna; 60, palp;

61, frons. 62-64, S. seythropa, Brazil, Rio de Janeiro, Tingua: 62, antenna; 63,

palp; 64, frons. 65-67, S. spilota, holotype: 65, antenna; 66, palp; 67, frons. 68-

71, S. venosus, holotype: 68, antenna; 69, palp; 70, frons; 71, wing.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae) ‘(ae

o4 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 72-76

Fig. 72, S. apiealis, Peru, Cusco, Quincemil, X-1962. 73, S. bipuneta, holo-

type. 74, S. eaptiroptera, Colombia, Valle, Rio Zapaletas. 75, S. changena,

paratype, Panama, Bocas del Toro, 16-IX-1961. 76, S. flaveseens, Peru, Cusco,

Quinecemil, X-1962.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae)

290

06 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 77-81

Fig. 77, S. furva, Bolivia, Yungasweg, 12-X-1906. 78, S. hypograhpa neo-

furva, Bolivia, Mapiri, S. Carlos, 9-I-1903. 79, S. jaculator, Colombia, Choco,

Curiche River, 25-IV-1967. 80, S. kroeberi, Peru, Tingo Maria, Rio Huallaga. 81,

S. lerida, Panama, Chiriqui, Boquete, Finca Lerida, II-1956.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae)

58 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 82-86

Fig. 82, S. marucii, Colombia, Valle, Buenaventura, 12-VIII-1973. 83, S.

modiea, Peru, Miriatiriani, Cord. de Pichis, 9-VII-1920. 84, S. paraguayensis,

Brazil, Mato Grosso, Sapucai-Mirim, Cidade Azul, 8-IX-1953. 85, S. pequeniensis,

Colombia, Valle, 41 km E Buenaventura, 19-III-1976. 86, S. prunicolor, Brazil,

Marmelo, 8-XI-1962.

Fairchild and Wilkerson: Neotropical Stypommisa (Tabanidae)

60 Contrib. Amer. Ent. Inst., vol. 22, no. 5, 1986

EXPLANATION OF FIGURES 87-89

Fig. 87, S. seythropa, Brazil, Rio de Janeiro, Tingua. 88, S. spilota, Peru,

Cuzeo, Quincemil, [X-1962. 89, S. u-nigrum, Mexico, Chiapas, Lagos Montebello,

20-V-1970.

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ON THE CORRECT APPLICATION OF RHIMPHOCTONA FOERSTER

(HYMENOPTERA: ICHNEUMONIDAE)

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, K1S 5B6

ABSTRACT

An examination of the type-species of Rhimphoctona Foerster, R.

grandis (Fonscolombe), has shown it to belong in Pyracmon Holmgren.

The next available namein place of Rhimphoctonais Xylophylax Kriechbaumer.

Pyracmon shares a common ancestor with the sister-pair of Xylophylax and

Nemeritis.

INTRODUCTION

The Foerster genus Rhimphoctona was fixed by the inclusion of R.

rufipes Tschek by Tschek in 1871. This species has subsequently become

regarded as a junior synonym of R. grandis (Fonscolombe), 1852.

The characters of grandis were carefully examined in preparation for a

revision of the nearctic species of Rhimphoctona which showed it to be differ-

ent from all known species of this genus. Comparisons with species belonging

to Pyracmon Holmgren showed that grandis has many features in common with

this genus.

This paper presents an analysis of 12 characters and their character

states and their distribution among grandis, Rhimphoctona, and Pyracmon

and provides conclusions as to the correct application of the name Rhimphoc-

tona.

RESULTS

Table 1 lists twelve characters and their character states as they are

expressed among the various species of Rhimphoctona, Pyracmon, and

grandis, and demonstrates their presence or absence in Pyracmon.

Table 2 shows the distributions of the character states (from table 1)

among Pyracmon, grandis, and Rhimphoctona..

The species of Rhimphoctona (excluding grandis) can be clearly divided

into two subgenera (subgenus 1 and subgenus 2 in table 2) which are named

and described in the following paper.

Table 2 shows that grandis agrees entirely with Pyracmon but that it

differs in a number of character states from species of Rhimphoctona

These similarities and differences are discussed below with respect to their

phylogenetic significance and their value in assessing the systematic place-

ment of grandis.

4 Contrib. Amer. Ent. Inst. , vol. 22, no. 6, 1986

Table 1. Twelve selected characters and their character states as they are

expressed among species of Rhimphoctona and Pyracmon with

presence (+) or absence (-) in Pyracmon indicated.

Character Character State Presence or Absence

in Pyracmon

1. Apical margin of | Moderately oblique +

antennal scape Strongly oblique -

2. Vertex Short +

Long ee

3. Clypeus Broad (antero-lateral corners oa

extend beyond tentorial pits)

Narrow (antero-lateral corners -

not extending beyond tentorial pits)

4. Clypeus Anterior margin produced medially +

as a small tooth

Anterior margin not produced 2

medially as a small tooth

5- Mandible Long, weakly tapering +

Short, strongly tapering -

6. Mandible Lower tooth as long as or a

shorter than upper tooth

Lower tooth longer than upper “=

7. Temple Longest ventrally (near level of +

mandible)

Longest medially (at mid-height -

of eye)

8. Petiolar seg. of Short, stout +

abdomen Long, slender ~

9. Ovipositor Depth at midlength at least 0.5 x the +

width of hind basitarsus, dorsal notch

distinct

Depth at midlength less than0.5x the -

width of hind basitarsus, dorsal notch

indistinct

10. Ovipositor Tip upturned xy

Tip not upturned -

1. Fore tibia Apex produced as a small +

tooth externally

Apex not produced as a small tooth =

Sanborne: Application of the name Rhimphoctona (Ichneumonidae) 3

Table 2. continued

Character Character State Presence or Absence

in racmon

12. Face of Male Entirely black +

Black with yellow markings -

Table 2. Distribution of character states (from Table 1) among Pyracmon,

Rhimphoctona, and R. grandis.

Character Character States

Pyracmon grandis Rhimphoctona

subgenus 1 subgenus 2

1 + + — + ~

2 + + = +

3 + + ~ +

4 - + + ~

5 + + ~ +

6 + + - -

7 + + - -

8 4. + +,- ~

9 + + ~

10 + 4. og ES

i + + + +

12 - + ~ -

4 Contrib. Amer. Ent. Inst. , vol. 22, no. 6, 1986

DISCUSSION

The following is an analysis of each of the twelve characters listed

above, in the order that they appear in the tables. The analysis is based on

the assumption that Pyracmon shares a common ancestor with the sister-

pair of Rhimphoctona and Nemeritis and that Pyracmon is the more general-

ized group.

Character 1 demonstrates a fundamental difference between Pyracmon

and Rhimphoctona.. The apical margin of the antennal scape is strongly

oblique in the latter and this is considered to be a derived state based on

out-group comparisons with other Porizontinae, including the closely relate-

ed genus Nemeritis. A character state similar to the postulated ancestral

state found in Pyracmon and grandis is found in a small species-group (2

species) within subgenus one of gushes 8 This is interpreted as being

a character reversal.

Character 2 sheds no light on possible relationships as a short vertex

is widely distributed among the Porizontinae and is therefore considered

ancestral. The long vertex possessed by species of subgenus one of Rhim-

phoctona is a uniquely derived state.

The broad clypeus (character 3) possessed by all species of Pyracmon ,

grandis, and all species of subgenus two of Rhimphoctonais also observed

in Nemeritis and is considered to be a shared derived feature indicative of

a close relationship. The narrow clypeus of species of subgenus one of

Rhimphoctonais a uniquely derived feature.

In character 4, the presence of a median tooth on the anterior margin

of the clypeus in grandis appears to be a derived feature which links over to

subgenus one of Rhimphoctona and has subsequently been lost in subgenus

two and Nemeritis.

Weakly tapering mandibles (character 5 ) are widely distributed within

the Porizontinae and are therefore considered ancestral and cannot be used

to establish relationships. The short, strongly tapering mandibles of subgenus

one of Rhimphoctona are unique and are probably an adaptation for chewing

wood and or bark during emergence from twigs and logs (their hosts are sub-

cortical coleopterous larvae).

Character 6 demonstrates another fundamental difference between

Pyracmon and grandis, and Rhimphoctona. The lower tooth is longer than the

upper tooth in all species of the latter genus, regardless of the length and

taper of the mandible. The lower tooth of Pyracmon and grandis is as long

or shorter than the upper tooth. This type of mandible is considered ancest-

ral as it is widely distributed within the Porizontinae.

Character 7 illustrates another fundamental difference between Pyracmon

and grandis, and Rhimphoctona . The temple of Rhimphoctona is unusually

long in comparison with other Porizontinae, with its greatest length occuring

opposite the mid height of the compound eye. The temple of grandis and all

Pyracmon has its greatest length occuring at the level of the mandible.

The ancestral state of character 8 appears to be with the petiolar seg-

ment of the abdomen short and stout. Long petiolar segments have arisen

twice in Rhimphoctona, once within a small species group within subgenus

one (the same species-group discussed under character 1 ) and it is found in

all species of subgenus two. This character cannot be used to establish

relationships as the ancestral state is distributed among all three taxa being

compared.

Sanborne: Application of the name Rhimphoctona (Ichneumonidae) +)

The ovipositor (character 9 ) exhibits two fundamental differences

between Pyracmon and grandis, and Rhimphoctona. The ovipositor of

Rhimphoctona is slender, with its depth at mid length less than half the width

of the hind basitarsus, and with the dorsal notch indistinct. This type of ovi-

positor functions in the manner of a drill to reach a host larva. The ovipos-

itor of Pyracmon and grandis has a depth at mid length which is greater than

half the width of the hind basitarsus and the dorsal notch is distinct. This

type of ovipositor is operated as a probe, which explains its relative stout-

ness. In Pyracmon, it is used to search out soil-dwelling elaterids (Barron, 1983).

Characters 10 and 11 show that the same character states are possessed

by all the species of the groups being compared. These states ( ovipositor

tip upturned, apex of fore tibia produced as a small tooth ) are considered to

be derived with respect to most other porizontine genera (their occurence

elsewhere is considered to be convergence and is rare ) and are indicative

of a close relationship.

Character 12 appears to demonstrate another fundamental difference

between Pyracmon and grandis, and Rhimphoctona. The faces of the males

of all species of Rhimphoctona are marked with white or yellow except for a

few highly melanic individuals of lucida (a palearctic species). The faces of

all males of Pyracmon and grandis are entirely black. Black faces with white

or yellow markings are considered to be derived states and have arisen

several times within the porizontines.

The hosts of Pyracmon and Rhimphoctona were briefly discussed earlier.

An account given byQzols, 1928 , indicates that the host of grandis is Saperda

carcharias (Cerambycidae). This could be interpreted as evidence for

placing grandis in Rhimphoctonaif morphological evidence were ignored. No

mention was made as to how grandis larvae were observed and a host-para-

site association cannot be presumed in all certainty. It is possible that an-

other sub-cortical species was acting as host, such as an elaterid. If the

host association is correct, it could be interpreted as a transitional stage

from one way of life to another ( see below),

SUMMARY AND CONCLUSION

Table 2 shows that grandis shares all twelve character states with

Pyracmon but shares similar character states with some Rhimphoctona in

only seven characters. It differs from all Rhimphoctona in characters 1 (a

similar charcter state in two species within subgenus one of Rhimphoctona

is interpreted as being a character reversal), 6, 7, 9, and12. The ancestral

states of characters 2, 5, and 8 are distributed among all three taxa being

compared ( not all species ) and cannot be used to infer relationships. Their

derived states can be used to separate subgenera within Rhimphoctona and in

one instance, a species-group within subgenus one. The ancestral state of

character 3 (derived with respect to other Porizontinae) and the states ex-

pressed in characters 10 and 11 indicate the closeness of the relationship

between Pyracmon, grandis, Rhimphoctona and Nemeritis.

The ancestral hosts of all four taxa are coleopterous larvae but some

Nemeritis have switched to the sub-cortical larvae of Rhaphidiidae (Neurop-

tera).

The morphological evidence supports the view that Pyracmon is the

most generalized of the three genera discussed in that it possesses many

ancestral character states and few derived ones.

6 Contrib. Amer. Ent. Inst. , vol. 22, no. 6, 1986

Morphological and biological evidence supports the view that Pyracmon

shares a common ancestor with Rhimphoctona and Nemeritis .

Morphological evidence strongly supports the view that grandis belongs

in Pyracmon and if biological observations are proven to be correct, then

this species can be interpreted as being a link between Pyracmon and the

Rhimphoctona-Nemeritis sister pair .

I herewith transfer grandis to Pyracmon based upon careful consideration

of the evidence presented in this paper. This transfer results in the removal

of Rhimphoctona as the generic name of the species previously included

under it, and this means that Rhimphoctona, 1868 becomes a junior synonym

of Pyracmon, 1859.

The next available name inplaceof Rhimphoctona Foerster is

Xylophylax Kriechbaumer, 1878 (Type: Pyracmon (Parapyracmon) rufocoxalis

Clément, designated by Townes, 1970). I have examined the type material of

this species and found it to be conspecific with species previously included

under Rhimphoctona.

ACKOWLEDGEMENTS

The following colleagues and institutions kindly lent specimens needed

for the completion of this paper: H. K. Townes (H.& M. Townes collection,

world-wide Xylophylax, male grandis ),Gainesville, Fla.; J. R. Barron

(Biosystematics Research Institute, world-wide Xylophylax and Pyracmon),

Ottawa, Ont.; K. Horstmann (Horstmann collection, female grandis),

Wirzburg, W. Ger.Dr. S. B. Peck provided support in the form of an NSERC

( Natural sciences and engineering research council) operating grant

(Dept. of Biology, Carleton Univ., Ottawa). The remainder of the support

came from a two year NSERC postgraduate fellowship.

REFERENCES

Barron, J. R. , G. S. Walley. 1983. Revision of the holarctic genus Pyracmon

(Hymenoptera: Ichneumonidae). Can. Ent. 115: 227-241.

Foerster, A. 1868. Synopsis der Familien und Gattungen der Ichneumonen.

Verh. naturh. Ver. Rheinl. 25: 135-221.

Fonscolombe, E. L. J. H. , Boyer de. 1852. Ichneumonologie provencale ou

catalogue des ichneumonides qui se trouvent aux environs d'Aix et

description des especes inedites. Ann. Soc. ent. France.(3) 2:497-520.

Holmgren, A. E. 1858. Conspectus generum Ophionidum Sueciae. Ofvers.

Svenska Vetensk. Akad. Forh. 15: 321-330.

Kriechbaumer, J. 1878. Bitte an die Ichneumonologen. Ent. Nachr. 4: 210-

211.

Ozols, E. 1928. Ichneumoniden aus bekannten Wirtstieren. Konowia. 7: 135-

146.

Townes, H. K. 1970. Genera of Ichneumonidae, Part 3. Mem. Amer. Ent.

Inst., no. 13. p. 162.

Tschek, C. 1871. Ichneumonologische Fragmente. I. Verh. zool.-bot. Ges

Wien. 21: 37-68.

THE SUBGENERA OF XYLOPHYLAX KRIECHBAUMER

(HYMENOPTERA: ICHNEUMONIDAE)

WITH DESCRIPTIONS OF TWO NEW EASTERN PALEARCTIC SPECIES

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, KiS 5B6

ABSTRACT

The subgenera of Xylophylax Kriechbaumer (Hymenoptera: Ichneumonidae)

are described and include Euraspidion, new subgenus, and Xylophylax

Kriechbaumer. Two new eastern palearctic species are described: nigrifemur

(Xylophylax), bimaculata (Euraspidion).

I have had the opportunity to examine all the known species of Xylophylax

Kriechbaumer, including two previously undescribed eastern palearctic

species. It became readily apparent that the genus could be classified into

two clearly defined subgenera and these are described below.

Subgenus Xylophylax Kriechbaumer

Xylophylax Kriechbaumer, 1878. Ent. Nachr. 4: 210

Type-species: Pyracmon (Parapyracmon) rufocoxalis Clément.

Designated by Townes, 1970.

Antennal scape with inner margin 1.8-2-4 x as long as outer margin

(except Longicauda group and Pectoralis group which have inner margin 1.3-

1.4 x as long as outer margin). Vertex unusually long, with portion produced

behind lateral ocelli approx. 0.8-1.9 x as long as length of ocellar triangle.

Lower temple very long. Clypeus narrow, with antero-lateral corners not

extending beyond tentorial pits, with anterior margin usually produced med-

ially as a small tooth (weakly or not produced in Longicauda group, not pro-

duced in Pectoralis group). Mandible short, strongly tapering, with lower

tooth slightly to distinctly longer than upper tooth. Abdomen of both sexes

entirely black.

This subgenus comprises one holarctic, eight palearctic, and seven

nearctic species in four species-groups (see following paper for descriptions

and keys to world species-groups).

Euraspidion, new subgenus

Type~species: Xylophylax (Euraspidion) vancouverensis

(Harrington). By present designation.

2 Contrib. Amer. Ent. Inst., vol. 22, no. 7, 1986

Antennal scape with inner margin approx. 1.5-1.9 x as long as outer

margin. Vertex moderately long, with portion produced behind lateral ocelli

approx. 0.5-1.0 x as long as length of ocellar triangle. Lower temple mod-

erately long. Clypeus broad, with anter0-lateral corners extending beyond

tentorial pits, its anterior margin not produced medially as a small tooth.

Mandible long, weakly tapering, with lower tooth distinctly longer than upper

tooth. Abdomen partially to entirely rufous except one new species from

palearctic and macdunnoughi (nearctic) which have entirely black abdomens.

This subgenus comprises one palearctic, and five nearctic species in

three species groups. Its name is derived from the greek words 'eury' mean-

ing wide, and 'aspidus' meaning lip, in reference to the broad clypeus.

The following are descriptions of two new eastern palearctic species

and includes the first species of the subgenus Euraspidion recorded from this

region

Illustrations depicting these subgenera can be seen in the paper immed-

iately following this (Sanborne, 1986c).

Xylophylax (Xylophylax) nigrifemur, new sp.

Clypeus with antero-lateral corners not extending beyond tentorial pits.

Propodeum with petiolar area about as long as wide, with median longitudinal

carinae distinctly angulate medially. Ovipositor approx. 2.2 x as long as hind

femur. Hind femur mostly black.

HOLOTYPE. Female. Antennal flagellum with 34 articles. Antennal scape

with inner margin approx. 1.6 x as long as outer margin. Cmpd. eye with

inner margin weakly diverging ventrad. Face strongly shagreened, matte,

with small punctures which are separated by distance less than dia. of single

puncture. Clypeus with antero-lateral corners not extending beyond tentorial

pits, its anterior margin weakly, broadly produced medially. Gena approx.

0.9 x as wide as basal width of mandible. Temples, in dorsal view, inflated.

Mandible with lower tooth much longer than upper tooth. Mesopleuron with

speculum smooth, polished, impunctate. Lower 0.5 of mesopleuron with small

punctures which are separated by distance approx. equal to dia. of single

puncture. Propodeum with carinae complete, strong. Areola about as long as

wide, completely confluent with petiolar area. Petiolar area about as long as

wide, with median longitudinal carinae distinctly angulate medially. Hind femur

approx. 4.1 x as long as deep. Hind tarsal claw with moderately strong teeth

which extend to base of tip. First abdominal segment with petiole approx.

1.1 x as long as postpetiole. Ovipositor approx. 2.2 x as long as hind femur.

Fore wing approx. 7.0 mm. long.

Head, thorax, abdomen, black. Median area of mandible yellowish.

Palpi, fore and middle femora, fore and middle tibiae, ferrugineous. Tegula,

coxae, basal segment of hind trochanter, hind femur, hind tibia, hind tarsi,

darkly fuscous. Wings hyaline.

Sanborne: Subgenera of Xylophylax (Ichneumonidae) 3

Range and Variation. Known only from holotype.

Male. Fore wing approx. 5.5 mm. long. Hind femur approx. 4.7 x as

long as deep. Vertex, upper temple, thorax, petiolar segment of abdomen,

black. Hind coxa, abdominal tergites 2 to apex, dark fuscous. Front of scape

and pedicel, face, clypeus, lower temple, median area of mandible, fore and

middle trochanters, ventral surfaces of fore and middle coxae, yellowish.

Fore and middle femora, fore and middle tibiae, ferrugineous. Dorsal sur-

face of fore and middle coxae, basal segment of hind trochanter, hind tibia,

hind tarsi, fuscous. Wings weakly infuscated.

Holotype. Female. Jozankei, Hokkaido, Japan, VII-10-1954, Townes family.

Deposited in H. & M. Townes collection, Gainesville, Fla.

Paratype. Male. Yokohama, Japan, V-10-1934, Kaku Sato. Deposited in H. &

M. Townes collection, Gainsville, Fla.

Etymology. The species is named for the colour of the hind femur.

Host and Biology. Unknown.

Remarks. This species belongs in the subgenus Xylophylax, and is most

similar to the palearctic species obscuripes . The petiolar area of obscuripes

differs in being longer than wide and its ovipositor is relatively much shorter

than that of nigrifemur.

Xylophylax (Euraspidion) bimaculata, new sp.

Clypeus with antero-lateral corners extending beyond tentorial pits, its

anterior margin not produced medially. Mandible with lower tooth distinctly

longer than upper tooth. Temples, in dorsal view, strongly sloping inwardly.

Abdomen black. Face with white spot below each antennal socket (spots some-

times absent.

HOLOTYPE. Female. Antennal flagellum with 28 articles. Antennal scape

with inner margin approx. 1.7 x as long as outer margin. Cmpd. eyes with

inner margins weakly diverging ventrad. Face strongly strongly shagreened,

matte, with medium sized punctures which are separated by distance less

than dia. of single puncture. Clypeus with antero-lateral corners extending

beyond tentorial pits, its anterior margin not produced medially. Gena approx.

0.9 x as wide as basal width of mandible. Temples, in dorsal view, strongly

Sloping inwardly. Mandible with lower tooth distinctly longer than upper

tooth. Mesopleuron with speculum smooth, polished, impunctate, its lower

0.5 shagreened, weakly polished, with small punctures which are separated

by distance 1.0-1.5 x the dia. of single puncture. Propodeum completely

areolated, the carinae very strong, with areola confluent with petiolar area.

Hind femur approx. 4.2 x as long as deep. Hind tarsal claw with moderately

strong teeth which extend to base of tip. First abdominal segment with pet-

iole approx. 1.4 x as long as postpetiole. Fore wing approx. 7.3 mm. long.

Hind wing with nervellus intercepted near basal 0.8.

Head (except spots below antennal sockets), thorax, abdomen, coxae,

4 Contrib... Amer... Ents, Inst, vols :22, nos.:7%;:1986

hind femur, black. White spot under each antennal socket. Median area of

mandible yellowish. Palpi, tegula, legs (except hind femur), fuscous. Wings

weakly infuscated.

Range and Variation. Antennal flagellum with 26-28 articles. Hind femur 3.8-

4.2x as long as deep. Fore wing 5.6-7.3 mm. Face with or without white

spot under each antennal socket. The preceeding is based upon one other

specimen.

Male. Unknown.

Holotype. Female. Sapporo, Japan, July 6, 1954, Townes family. Deposited

in H. & M. Townes collection, Gainesville, Fla.

Paratype. Female. Same data as above.

Etymology. The species is named for the white spots occuring beneath the

antennal sockets of the holotype specimen.

Host and Biology. Unknown.

Remarks. This species is the only known representative of the subgenus

Euraspidion in the palearctic region. It is readily distinguished from all

known palearctic species by the broad clypeus (antero-lateral corners extend

beyond tentorial pits) and from all known species of the genus by the presence

of white spots beneath each antennal socket of some females. Its is most

closely related to the nearctic species macdunnoughi.

ACKNOWLEDGEMENTS

I thank Dr. H. K. Townes for lending me the material upon which these

descriptions are based. Support for this project was in the form of an oper-

ating grant administered by N.S.E.R.C. (Natural Sciences and Engineering

Research Council) and held by Dr. S. B. Peck, Carleton Univ., Ottawa, Ont.

and additional support came from the N.S.E.R.C. post-doctoral fellowship

awards program.

REFERENCES

Harrington, W. H. 1894. Canadian Hymenoptera No. 6. Can. Ent. 26:246.

Kriechbaumer, J. 1878. Bitte an die Ichneumonologen. Ent. Nachr. 4:210-211.

Townes, H. K. 1970. Genera of Ichneumonidae, Part 3. Mem. Amer. Ent.

INSC... ida OO DD.

REVISION OF THE NEARCTIC SPECIES OF XYLOPHYLAX KRIECHBAUMER

(HYMENOPTERA: ICHNEUMONIDAE)

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, K1S 5B6

ABSTRACT

The nearctic species of Xylophylax Kriechbaumer (Hymenoptera: Ich-

neumonidae) are revised, including seven new combinations - alaskensis

(Ashmead) n. comb., aldrichi (Davis) n. comb., longicauda (Horstmann) n.

comb., macdunnoughi (Viereck) n. comb., macrocephala (Provancher) n.

comb., separata (Viereck) n. comb., vancouverensis (Harrington) n. comb.,

six new species - albicoxa, breelandi, brevicauda, genalis, montana, rufic-

incta, and two new subspecies - vancouverensis maculosa, vancouverensis

tibialis . Three species names are placed in synonymy - atrocoxalis (Ash-

mead) and xanthognatha (Rohwer) = aldrichi (Davis), lipoparia (Viereck) =

alaskensis (Ashmead). In addition, the following paiearctic species become

new combinations ~ lucida (Clement) n. comb., megacephala (Gravenhorst)

n. comb., melanura (Holmgren) n. comb., obscuripes (Holmgren) n. comb.

pectoralis (Kriechbaumer) n. comb., rufocoxalis (Clément) n. comb.,

xoridiformis (Holmgren) n. comb., xoridoidea (Strobl). The genus Xylo-

phylax is redefined and a key to the world sub-genera, species-groups, and

nearctic species is provided. Descriptions, illustrations, and distribution

maps are given for each nearctic species and there is a summary of the

biology of the genus.

INTRODUCTION

Thirteen nearctic species of Xylophylax (Kriechbaumer) (formerly

Rhimphoctona) (Ichneumonidae) are revised in this paper, including seven new

combinations - aldrichi (Davis), longicauda (Horstmann), macdunnoughi

(Viereck), macrocephala (Provancher), separata (Viereck), vancouverensis

(Harrington) , six new species - albicoxa, breelandi, brevicauda, genalis,

montana, ruficincta, and two new subspecies - vancouverensis maculosa,

vancouverensis tibialis .

Replacement of the name Rhimphoctona Forster with Xylophylax Kriech-

baumer results in all the palearctic species becoming new combinations -—

lucida (Clément), megacephala (Gravenhorst), melanura (Holmgren),

obscuripes (Holmgren), pectoralis (Kriechbaumer), rufocoxalis (Clément),

xoridiformis (Holmgren), and xoridoidea (Strobl).

A key to the world sub-genera, species groups and nearctic species is

presented. All known species world-wide are arranged in species groups in

what is believed to be a natural order. Each nearctic species is described,

illustrated (scanning electron photomicrographs) and its distribution plotted

on a map.

A summary of what is known of the hosts of Xylophylax species is

2 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

presented in table form. Biological information is sparse and little is known

of the eastern palearctic fauna so that a detailed phylogenetic and zoogeogra-

phic analysis is premature at this time. Phylogenetic relationships at the

generic level are briefly discussed and a cladogram (Fig. A.) is presented

depicting the distribution of derived character states (Ssynapomorphs, autap-

omorphs) and nearness of relationship among the species groups of Xylophylax.

The following institutions and private collectors (with abbreviations as

they appear in the text with regards to deposition of type material) kindly

lent specimens used in this study.

American Museum of Natural History (AMNH). New York, New York

(Marjorie Favreau)

California Academy of Sciences (CAS). San Francisco (W. J. Pulawski)

Canadian National Insect Collection (CNC). Ottawa, Ont. (J. Barron)

Clement Dasch Collection (CDC). New Concord, Ohio (C. Dasch)

Cornell University Collection (CUC). Ithaca, New York (Q. D. Wheeler)

David Wahl Collection (DWC). Lawrence, Kansas (D. Wahl)

Henry and Marjorie Townes Collection (HMT). Gainesville, Fla.

(H. K. Townes)

Oregon State University (OSU). Corvallis, Oregon (John D. Oswald)

P. M. Sanborne (PMS). Ottawa, Ontario (P. M. Sanborne)

University of Alberta Collection (UAC). Edmonton, Alberta (D. Shpeley)

University of California, Berkley (UCB). Berkley, California

(L. E. Caltagirone)

University of California, Davis (UCD). Davis, California (R. O. Schuster)

University of California, Riverside (UCR). Riverside , California

(Jack Hall)

University of Guelph Collection (UGC). Guelph, Ontario (S. Marshall)

University of Kansas, Snow Museum (KSM). Lawrence, Kansas

(G. Byers)

United States National Museum (USNM). Washington, D. C. (S. R. Shaw)

Nlustrations. Each species is illustrated with the aid of scanning electron

photomicrographs using uncoated specimens.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 3

Measurements. The way in which measurements are made are given in my

revision of Sinophorus (Sanborne, 1984) and are not repeated here. Additional

measurements were made in the following manner. To obtain the ratio of

length of the inner margin of the antennal scape vs the length of the outer

margin, the inner margin must be measured first, then the eye-piece micro-

meter must be lined up exactly parallel to that measurement along the outer

margin (i.e., the curvature of the ventral portion of the outer margin is not

taken into account as it is variable from species to species). The length of

the teeth of the mandible is taken from a line passing perpendicularly through

the bases of the teeth at the point where the inner margins of the teeth meet in

Biology

What little that is known of the biology of Xylophylax indicates that the

species are primary internal parasites of wood-boring coleopterous larvae

of the families Cerambycidae, Curculionidae, and possibly Scolytidae. This

information is summarized in table form below. Some of the plant associations

are inferred from knowledge of the host larva's identity (i.e., it is assumed

that species of Xylophylax will probably attack preferred hosts throughout

their geographic range in whatever tree species the hosts chose).

Table 1. Summary of hosts and host-plant associations for the world species

of Xylophylax.

Species Plant Associations Hosts

Obscuripes Group

macrocephala Picea glauca Tetropium cinnamopterum

Pinus resinosa

obscuripes unknown unknown

rufocoxalis unknown unknown

Longicauda Group

longicauda Pinus contorta unknown

Picea, Abies

montana unknown unknown

Megacephala Group

alaskensis Picea glauca Tetropium parvulum

Tetropium velutinum

aldrichi Pseudotsuga menziesii Tetropium velutinum

Pseudotsuga taxifolia Orthotomicus lasiocarpa

Larix occidentalis

Abies concolour

Abies shastensis continued....

a Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Table 1. continued.

Species Plant Associations

lucida Picea

megacephala Fagus, Ostrya, Quercus

Salix

nigrifemur unknown

separata Picea glauca

xoridiformis unknown

xoridoidea probably conifer

Pectoralis Group

albicoxa Carya, Juglans nigra

Cercis canadensis

genalis unknown

pectoralis Picea, Abies

Vancouverensis Group

breelandi unknown

ruficincta Pseudotsuga menziesii

vancouverensis unknown

Macdunnoughi Group

bimaculata unknown

macdunnoughi Picea mariana

Brevicauda Group

brevicauda unknown

Hosts

Tetropium fuscum

Tetropium gabrieli

Clytus arictis

Plagionotus arcuatus

Pyrrhidium sanguineum

unknown

Tetropium fuscum

Magdalis (Curculionidae)

Tillomorpha geminatus

unknown

Caenoptera minor

unknown

* specimens of melanura were not available for study and

the species-group to which it belongs cannot be ascertained

at this time, therefore, it has been left out of the above

table.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) +)

Phylogeny

Table 2 below lists seventeen characters of Xylophylax with their postulated

ancestral and derived states (based upon out-group comparisons with its

sister group Pyracmon, and the closely related genus Nemeritis). The infor-

mation presented in the table is superimposed on a cladogram below, depict-

ing nearness of relationship. No attempt has been made to include a zoo-

geographic analysis or a time scale on this cladogram because of a lack of

biological information and an incomplete knowledge of the palearctic fauna,

particularly the eastern palearctic component.

The generic relationships of Xylophylax:alluded toabove are now gener-

ally accepted and are not dealt with here. Detailed discussions of this matter

can be found in Sanborne, 1986a, and Barron and Walley, 1983.

Table 2. Seventeen characters of Xylophylax and their postulated ancestral

(a) and derived (d) states (superscripts ', '', ''' indicate 1st, 2nd, and 3rd

derived states).

1 a. Vertex short.

d. Vertex moderately long ', '' (arose twice); long '''.

2 a. Antennal scape with apical margin weakly oblique.

d. Antennal scape with apical margin moderately oblique '; strongly

oblique '', ''' (arose twice).

3 a. Temples flaring.

d. Temples not flaring (sloping inwardly in dorsal view) ', '' (arose twice).

4 a. Gena with width less than basal width of mandible.

d. Gena as wide or wider than basal width of mandible '; secondarily nar-

rowed to less than basal width of mandible ''.

5 a. Clypeus broad, flat, with small, median projection on anterior margin.

d. Clypeus broad, flat, without a median projection '; narrow, with small

median projection on anterior margin ''; narrow, without median proj-

ection '"'.

6 a. Mandible with lower tooth as long as or shorter than upper tooth.

d. Mandible with lower tooth 1.8-2.3x as long as upper tooth '; lower

tooth 1.5-1.6 x as long as upper tooth ', '' @rose twice).

. Mandible long, weakly tapering.

. Mandible short, strongly tapering '.

a. Head about as long as wide.

d. Head wider than long in frontal view '.

. Thorax not dorso-ventrally flattened.

. Thorax dorso-ventrally flattened '.

. Petiolar segment short, stout. continued....

6 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Table 2. continued.

10 d. Petiolar segment elongate ', '' (arose twice); very short '''.

It a. Ovipositor long, approx. 2.5-3.5 x as long as hind femur.

d. Ovipositor of medium length, approx. 1.5-2.3 x as long as hind femur '.

Ovipositor short, approx. 1.5 x as long as hind femur ''.

12 a. Ovipositor with depth at mid length approx. 0.5 x as wide as hind basi-

tarsus or more (measured at mid length of basitarsus).

d. Ovipositor with depth at mid length less than 0.5 x the width of hind

basitarsus '.

13 a. Ovipositor with dorsal notch distinct.

d. Ovipositor with dorsal notch indistinct '.

14 a. Ovipositor with tip not upturned.

d. Ovipositor with tip upturned! .

- Face of males entirely black.

. Face of males yellow with black markings to entirely yellow '.

a. Abdomen of both sexes entirely black.

d. Abdomen of both sexes partially to entirely rufous '.

. Clypeus of females entirely black.

. Clypeus of females partially to extensively reddish‘

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae)

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8 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

TAXONOMY

The Genus Xylophylax Kriechbaumer

Xylophylax Kriechbaumer, 1878. Ent. Nachr. 4: 210

Type: Pyracmon (Parapyracmon) rufocoxalis Clément. Designated by

Townes, 1970. Type material examined in 1984.

Fore wing 3.7 to 10.0 mm. long. Body moderately slender. Inner marg-

ins of cmpd. eyes not distinctly indented opposite antennal sockets, weakly

diverging ventrad. Gena 0.7 to 1.4 x as wide as basal width of mandible.

Clypeus narrow to broad, its antero-lateral corners not extending to, extend-

ing to, or extending beyond tentorial pits, its anterior margin blunt, evenly

convex or more or less angulate, sometimes with a small tooth medially.

Mandible long, weakly tapering to short, strongly tapering, with lower tooth

slightly to distinctly longer than upper tooth. Temple of moderate length to

very long, usually swollen and sometimes flaring so that head is wider across

temples than across eyes. Genal carina joining oral carina. Mesopleuron

matte, its punctures fine to moderately coarse, usually separated by distance

less than or equal to dia. of single puncture. Posterior mesosternal carina

complete. Propodeum long, with carinae moderately strong to strong, with

costulae and/or lateral longitudinal carinae rarely absent, with areola small

to moderately large, as long as or longer than wide, sometimes partially to

entirely separated from petiolar area by a transverse carina. Propodeal

spiracle circular or elliptic. Fore tibia with apex produced as a small tooth

outside. Hind basitarsus without a mid-ventral row of closely spaced hairs.

Tarsal claws moderately long with teeth extending from base to midlength or

sometimes beyond. Fore wing with areolet moderately large, rarely petiolate

above, receiving second recurrent vein distad of the middle. Nervulus oppo-

site basal vein. Nervellus intercepted by discoidella at or below middle,

usually pigmented. Petiolar segment of abdomen short, stout to moderately

long and weakly and evenly decurved, with petiole 1.0 to1.5 x as long as

postpetiole. Glymma large, deeply impressed. Thyridium small, a little

elongate. Ovipositor approx. 1.5 to 3.0 x as long as hind femur, unusually

slender, cylindric, its tip upcurving, the dorsal notch indistinct or absent.

This is a moderate sized genus with a holarctic distribution. The genus

comprises two subgenera with twenty two described species in seven species

groups. Their hosts, as far as is known, are wood-boring coleopterous

larvae in the families Cerambycidae, Curculionidae, and possibly Scolytidae.

Most species are rare in collections and this is probably due to the fact that

their distribution and numbers within a given habitat are dependent upon a

rather patchy resource, i.e., timber in the proper stage of decay bearing

suitable hosts. See table 1 for a summary of the biology of the genus.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 9

Key to the Subgenera, Species-Groups, and Nearctic Species of Xylophylax

This key will correctly place all known species of Xylophylax into sub-

genera and species-groups but will identify only nearctic species.

1. Vertex short to moderately long, with length behind lateral ocelli 0.5 -

1.1 x the length of ocellar triangle. Clypeus broad, with antero-

lateral corners extending beyond tentorial pits, its anterior margin

without a small median tooth though sometimes broadly angulate.

Mandible long, weakly tapering. Abdomen usually partially to entire-

ly rufous or, rarely, entirely black.. Euraspidion Sanborne (p. 28). 2.

Vertex moderately long to long, with length behind lateral ocelli 0.8 -

1.9 x the length of ocellar triangle (almost always greater than 1.0).

Clypeus narrow, with antero-lateral corners not extending beyond

tentorial pits, its anterior margin usually with a small median tooth

(sometimes absent or indistinct in males). Mandible short, strongly

tapering. Abdomen uniformly black. . Xylophylax Kriechbaumer (p.11).

2. Vertex moderately long, with length behind lateral ocelli approx. 0.8 -

1.1 x the length of ocellar triangle. Temples, in dorsal view, dist-

inctly inflated. Mandible with lower tooth approx. 1.8 - 2.0 x as long

as upper tooth. Abdomen extensively to entirely rufous. Nearctic...

se Sea ole Oe ea eek Cbs een eas Vancouverensis Group (p.30).3.

Vertex short, with length behind lateral ocelli approx. 0.5 -0.8 x the

the length of ocellar triangle. Temples, in dorsal view, not inflated,

moderately sloping inwardly. Mandible with lower tooth approx. 1.0

- 1.5 x longer than upper tooth (nearctic species), 1.8 x longer

(palearctic species). Abdomen extensively rufous to entirely black.

3. Coxae extensively to entirely black. Abdomen usually fuscous to black

POI OAL, Pree dt aecck Baie ailes Baw Kalla eh ee «Wes Bcc Cs

Coxae rufous. Abdomen entirely rufous. Cascade-Sierra Nevada Ranges.

CRE WAD 0 ww Beeia oe Cie PEA s Cibola ede Odea sc eee Cine a a Sp. (p-35).

4. Propodeum with areola about as long as wide, the propodeal carinae very

strong. Mesopleuron usually somewhat polished, with medium to

large sized punctures. Pacific Northwest s. to Colorado......ee...

Cee eee rere ecco ce cccveseoess Vancouverensis (Harrington) (p. 31).

Propodeum with areola distinctly longer than wide, the propodeal carinae

not as strong as above. Mesopleuron matte, with fine punctures....

Kentiol, £0.00, 2e.% Shit: < ate be PR a ate dy breelandi n. sp. (p. 36).

9. Ovipositor approx. 1.3 x as long as hind femur. Abdomen rufous........

Nearctic. Brevicauda Group....... ew <iaeaie brevicauda n. sp. (p. 28).

Ovipositor approx. 2.8 - 3.0 x as long as hind femur. Abdomen black.

10.

Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Holarctieur olla 964% sats ete ani Macdimneughi-Group (p87).

Mandible with lower tooth approx. 1.0-1.1 x as long as upper

tooth. Boreal Forest, Rocky Mts. to Mexico.....+..sseeee-

Siw. Wolo Ge we. low Imaedunnough? (Atereck): (ps8) -

Antennal scape with apical margin strongly oblique, with inner margin

1.6 - 2.2.x as long as outer margin. Clypeus with antero-median

tooth usually present. Medium to large sized species...........7.

Antennal scape with inner margin 1.3 -1.4 x as long as outer margin.

Clypeus without an antero-median tooth, the margin evenly convex.

SpA IBD OCI CS oe ile. da R Es 6 SEMEN SUV W OWS 6 Ma es See eee S10.

Mandible with lower tooth approx. 1.3 - 1.5 x as long as upper tooth.

Ovipositor approx. 1.3 - 2.6 x as long as hind femur. Holarctic...

he eWAW Mer wees tine ke coke ees bbls ONSCULIPCS Group “(p.11).

Ovipositor approx. 2.1 - 2.6 x as long as hind femur. Coxae

of female rufous. Inner orbits of male with yellow extending

posteriorly to middle of antennal socket. Face of male yell-

ow with postero-median fuscous spot. Alaska e. to Nova

Scotia, s. to N. Carolina..macrocephala (Provancher) (p.11).

Mandible with lower tooth approx. 1.8 - 2.3 x as long as upper tooth.

Ovipositor approx. 2.5 - 3.5 x as long as hind femur (2.0 - 2.1x

in separata). Holarctic.......... Megacephala Group (P-14).8.

Ovipositor approx. 2.0 - 2.1 x as long as hind femur. Head of female

with reddish spot (often faint) opposite each antennal socket. Face

of male yellow, with large postero-median fuscous patch. Hind tib-

ia (both sexes) ferruginous medially, with weak basal and apical

infuscation. Boreal Forest...........-..separata (Viereck) (p.15)

Ovipositor 2.5 - 3.5 x as long as hind femur. Head of females without

reddish spots opposite antennal sockets. Face of male entirely

yellow or yellow with large postero-median fuscous patch. Hind

HUG PROSE FOIA ras iia + eels vs eee bree eid wales aS reir. Baek Ny Ba

Ovipositor approx. 2.5 - 2.8 x as long as hind femur. Coxae of female

extensively to entirely black. Face of male yellow with large

postero-median fuscous patch. Pacific Northwest...... gy te ee Z

‘reveveee e+ CTs VER Ce eRe ba ee ee aldrichi (Davis)(p.14)

Ovipositor approx. 3.3 - 3.5 x as long as hind femur. Coxae of female

rufous. Face of male yellow, with at most a faint, minute postero-

median fuscous spot. Alaska s. to shi e. to Gaspe, s. to Penn-

BVIVETIAN eas EWS Yo b His aes -alaskensis (Ashmead) (p.18 )

Gena approx. 0.7 - 0.8 x as wide as basal width of mandible. Lower

tooth of mandible approx. 1.8 - 2.0 x as long as upper tooth. Thor-

ax somewhat dorso-ventrally flattened. Abdominal petiole short,

Stout FOAL wr sysre ) bi 5is dee » eee Pectoralis Group (p. 24) .1l.

Gena approx. 1.0 -1.1 x as wide as basal width of mandible. Lower tooth

of mandible approx. 1.3 - 1.4 x as long as upper tooth. Thorax not

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) am

noticeably dorso-ventrally flattened. Abdominal petiole elongate,

slender. Holarctic ..... ot IC US & oh oeeeeee LONGICAUdA Group.12.

11. Fore coxa pale to dark ferruginous. Propodeal carinae not unusually

strong. Propodeal trough without coarse transverse wrinkling.

Face of male yellow with a large postero-median fuscous patch.

Thorax of male entirely black. w. N. Amer.. .genalis n. sp.(p.- 25)

Fore coxa ivory. Propodeal carinae unusually strong. Propodeal trough

with several coarse, transverse wrinkles. Face of male ivory.

Thorax of male with ivory markings. e. N. AMELr....ceeesseveces

12. Propodeal trough usually with distinct transverse wrinkling. Tegula

often infuscated. Male with thorax entirely black. w. N. Amer....

eeeeoeeoe@eeeeseVeoeoeeeeeoeeeeseeeeeeeee @ ans eid eee lain eR OTe TP. sp. (p. 21)

Propodeal trough with granulose surface and weak transverse wrinkling

usually present. Tegula white. Male with yellow markings on

thorax. Holarctic (transcontinental in conifer forests in N. Amer.)

ccc seetiensdieececéwccecoesencs ese se LONDIGAUda (Horstmann) (Pp. 20)

Subgenus Xylophylax Kriechbaumer

The description of this subgenus is found in the paper immediately pre-

ceeding this one and is not repeated here. Salient features of the group are

presented in the key above.

Obscuripes Group

Antennal scape with inner margin approx. 1.8 - 2.2 x as long as outer

margin. Vertex long, with length behind lateral ocelli approx. 1.3 -1.6x as

long as ocellar triangle. Temples, in dorsal view, strongly inflated. Clypeus

with antero-lateral corners not extending beyond tentorial pits, its anterior

margin usually with a small, blunt, median tooth. Mandible short, strongly

tapering, with lower tooth approx. 1.3 - 1.5 x as long as upper tooth. Ovi-

positor approx. 1.5 - 2.3 x as long as hind femur. Clypeus sometimes with

anterior margin narrowly reddish, otherwise, entirely black. Abdomen

black.

This is a holarctic group comprising two palearctic species, rufocoxalis

(Clément), n. comb., obscuripes (Holmgren) n. comb., and one nearctic

species, macrocephala (Provancher) n. comb. What is known of the biology

of this group is that they attack larvae of Cerambycidae in various conifers.

Xylophylax macrocephala (Provancher), n. comb.

Figs. 1, 15

Limneria macrocephala Provancher, 1874. Nat. Canadien, 6: 149. Female.

Quebec (McGill Univ.). Examined in 1984.

12 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Cubocephalus nigricornis Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 203.

Female. Preoccupied in Cubocephalus by Provancher, 1875. Alaska

(USNM). Examined in 1984.

Pyracmonoides succineum Viereck, 1925. Can. Ent. 57(7):178 (key); 57(12):

~ 800. Male. Smith's Cove, Nova Scotia (CNC). Examined in 1984.

Pyracmonoides trochantericum Viereck, 1925. Can. Ent. 57(7): 178 (key); 57

(12): 299. Male. Teulon, Manitoba (CNC). Examined in 1984.

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin usually with a small, blunt, median tooth. Mandible with

lower tooth much larger than upper tooth. Ovipositor approx. 2.0 - 2.2xas

long as hind femur. Coxae rufous. Abdomen black. Face of male yellow, with

median fuscous spot and inner orbits with yellow extending a short distance

behind antennal sockets.

Female. Antennal flagellum with 40-48 articles. Antennal scape with inner

margin approx. 2.0-2.4 x as long as outer margin. Cmpd. eyes with inner

margins weakly diverging ventrad. Face shagreened, matte, with fine punc-

tures which are separated by distance approx. equal to dia. of single punc-

ture. Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin usually with a small, blunt, median tooth. Gena approx.

1.0-1.2 x as wide as basal width of mandible. Temples in dorsal view, inflated.

Mandible short, strongly tapering, with lower tooth approx. 1.4.x as long as

upper tooth. Mesopleuron with speculum weakly to moderately shagreened,

weakly polished, sparsely punctate. Lower 0.5 of mesopleuron shagreened,

matte, with small punctures which are separated by distance 1.0-1.5 x the

dia. of single puncture. Propodeum completely areolated but costula often

weak. Areola longer than wide, almost always completely confluent with

petiolar area. Median longitudinal carina almost always strongly angulate

medially. Hind femur approx. 3.7-4.0 x as long as deep. Hind tarsal claw

with moderately strong teeth which do not extend to base of tip. First Abd.

segment with petiole approx. 1.1-1.2 x as long as postpetiole. Ovipositor

approx. 2.1-2.6 x as long as hind femur. Fore wing approx. 5.8-8.5 mm.

long. |

Head, thorax,abdomen, black. Median area of mandible, tegula, whitish.

Palpi fulvous. Coxae, trochanters, fore and middle femora, fore tibia, ferr-

ugineous. Middle tibia ferrugineous, with apex sometimes infuscated. Hind

femur ferrugineous, with apical dark band or, entirely infuscated. Hind tibia

extensively to entirely infuscated. Hind tarsus fuscous.

Male. Hind femur approx. 4.2-4.7 x as long as deep. Front of scape and

pedicel, inner orbits, face, clypeus, lower temple, lateral lobes of pro-

sternum, front and middle coxae, front and middle trochanters, yellow.

Palpi, tegula, white. Remainder of legs coloured as in female except that

front and middle legs generally paler.

Distribution. CANADA. New Brunswick: Bathhurst. Nova Scotia: Digby.

Ontario: Constance Bay, Frater, Mississaugi, One-Sided Lake, Sault Ste.

Marie. Quebec; Gaspé Co., Laniel. Saskatchewan: Carrot R. Yukon Territory:

La Force Lake, Mile 87 (Dempster Hwy.). UNITED STATES. Alaska:

Bonanza Cr., Fairbanks. Maine: Lincoln Co. New York: Green Co., Uphill

Brook, Mt. Marly. North Carolina: Pisgah Ridge. ADDENDA. Alberta: Seebe

Host and Biology. The only recorded host is Tetropium cinnamopterum

(Cerambycidae) in red pine. It has also been reared from an unknown host

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 13

in white spruce. Adults are active from June through August, with peak ab-

undance in July.

Material Examined. 26 males, 50 females

Remarks. This species is very closely related to the palearctic species

rufocoxalis but its closest relative in North America is alaskensis. Females

of macrocephala are easily distinguished from alaskensis by the longer ovi-

positor. Males of macrocephala have yellow faces with a dark median spot

and the faces of alaskensis are entirely yellow.

TOT Se eae = he: 4

REM fe

macrocephala

500

14 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Megacephala Group

Antennal scape with inner margin approx. 1.8-2.2 x as long as outer

margin (1.6-1.7 x in megacephala). Vertex long, with length behind lateral

ocelli 1.3-1.9 x as long as ocellar triangle (0.9-1.0 in nigrifemur). Temples,

in dorsal view, strongly inflated. Clypeus with antero-lateral corners not

extending beyond tentorial pits, its anterior margin usually with a small,

median tooth. Mandible short, strongly tapering, with lower tooth approx.

1.7-2.3 x as long as upper tooth. Ovipositor approx. 2.5-3.5 x as long as

hind femur. Clypeus often with anterior 0.5 yellowish-red, otherwise, entire-

ly black. Abdomen (both sexes) black.

This is a holarctic group with 5 palearctic species ( lucida, megacephala,

nigrifemur, xoridiformis, xoridoides) and 3 nearctic species ( alaskensis,

aldrichi, separata). The hosts of this group are larvae of Cerambycidae

occuring in Pinaceae, except megacephala which is associated with Fagaceae

and Juglandaceae.

Xylophylax aldrichi (Davis), n. comb.

Figs. 2, 15

Pyrachmon (!) aldrichi Davis, 1898 (1897). Amer. Ent. Soc. Trans. 24: 363.

Male, Female. Moscow, Idaho (Academy of Nat. Sci. Phila.). Exam-

ined in 1984. |

Cubocephalus atrocoxalis Ashmead, 1902. Wash. Acad. Sci. Proc. 4: 203.

Female. Alaska (U.S.N.M.). New synonymy. Examined in 1984.

Helcostizidea xanthognatha Rohwer, 1914. Ent. Soc. Wash. Proc. 15 : 186.

Female. Columbia Falls, Montana (U.S.N.M.). New synonymy. Exam-

ined in 1984.

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin usually with small, median tooth. Face with punctures

separated by distance less than dia. of single puncture. Mandible with lower

tooth approx. 2.0 x as long as upper tooth. Hind trochanter rufous. Hind

coxae extensively to entirely black. Abdomen black. Male face yellow with

black markings in form of median spot or stripe which covers less than 1/3

surface area of face.

Female. Antennal flagellum with 48-52 articles. Antennal scape with inner

margin approx. 1.8-2.1 x as long as outer margin. Cmpd. eyes with inner

margins weakly diverging ventrad. Face shagreened, matte, with small punc-

tures which are separated by distance less than dia. of single puncture. Cly-

peus with antero-lateral corners not extending beyond tentorial pits, its ant-

terior margin usually with a small, median tooth. Gena approx. 1.3-1.4 x as

wide as basal width of mandible. Temple, in dorsal view, distinctly inflated.

Mandible short, strongly tapering, with lower tooth approx. 2.0 x as long as

upper tooth. Mesopleuron with speculum smooth, polished, impunctate. Low-

er 0.5 of mesopleuron with fine to small punctures which are separated by

distance 1.0-2.0 x dia. of single puncture. Propodeum with costula present

as a trace, the lateral longitudonal carina absent, the median longitudinal

carina strong. Areola approx. 1.5-2.0 x as long as wide, rarely separated

from petiolar area by transverse carina. Hind femur approx. 3.8-4.2 x as

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 15

long as deep. Hind tarsal claw with moderately strong teeth. First abdominal

segment with petiole approx. 1.1-1.2 x as long as postpetiole. Ovipositor

approx. 2.6-2.8 x as long as hind femur. Fore wing approx. 6.5-9.5 mm.

long.

Head, thorax, abdomen, coxae (except apices), black. Median area of

mandible yellow. Palpi fulvous. Tegula white. Apices of coxae, trochanters,

fore and middle femora, fore and middle tibiae, ferrugineous. Hind femur

ferrugineous, with apex sometimes infuscated. Hind tibia with ground colour

ferrugineous but often extensively to entirely infuscated. Hind tarsus ferrug-

ineous to fuscous. wings weakly infuscated.

Male. Hind femur approx. 4.0-4.5 x as long as deep. Front of scape, face

(except median fuscous spot), clypeus, median area of mandible, apical mar-

gins of lateral lobes of prosternum, most of fore and middle coxae, fore and

middle trochanters, pale yellow (prosternum rarely entirely black). Fore and

middle femora, fore and middle tibiae, yellow-ferrugineous. Hind coxa fusc-

ous. Hind femur ferrugineous to fuscous. Hind tibia and tarsus ferrugineous

with variable degree of infuscation.

Material Examined. 81 males, 39 females

Distribution. CANADA. Alberta: Banff; Edmonton. British Columbia: Canoe;

Duncan Lake; Howser Ridge; Lower Post; Radium; Robson; Squamish; Trinity

Valley; Vernon. UNITED STATES. California: Buck's Lake, Plumas Co.;

Camp Connell; Carrville; Dardanelle; Echo Lake, Eldorado Co.; Giant Forest,

Tulare Co.; Lake Tahoe; Pinecrest; Vade; Yosemite Pk. Colorado: Burke.

Idaho: Krassel, Valley Co.; McCalf, Brundage Mt. Montana: Columbia Falls;

Lake Roman. Oregon: Caves; Clover Cr., Grant Co.; Mt. Hood; Mt. Jeffer-

son; Ochoco Cr. Washington: Easton.

This species is confined to the pacific northwest in the Cascade-Sierra

Nevada ranges and the Rocky Mts. from Alaska to Colorado.

Host and Biology. The species attacks cerambycid larvae in firs and larch.

Host records include Tetropium velutinum and Orthotomicus lasiocarpa . It

has been found in association with Larix occidentalis, Pseudotsuga menziesii,

_P. taxifolia, Abies concolor, and A. shastensis. Adults are active from June

to September with a peak abundance in July.

Remarks. This species appears to be closely related to separata but is typ-

ically larger, has a longer ovipositor and the face of the female is entirely

black. See further remarks following the description of separata below.

Xylophylax separata (Viereck), n. comb.

Figs.3,16

Pyracmonoides separatum Viereck, 1925. Can. Ent. 57(7):178 (key); 57(12):

298. Male and Female. Nordegg, Alberta (CNC). Examined in 1984.

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its margin with a small, median tooth. Temples, in dorsal view, weakly in-

flated. Mandible with lower tooth approx. 2.3 x as long as upper tooth. Thorax

weakly flattened dorso-ventrally. Petiolar segment of abdomen approx. 1.1-

16 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

1.2 x as long as postpetiole. Female with faint reddish spot beside each ant-

ennal socket (outside). Face of male yellow, with a median, elongate-triang-

ular fuscous patch, the yellow extending along the orbits a short distance be-

hind the antennal sockets.

Female. Antennal flagellum with 36-39 articles. Antennal scape with inner

margin approx. 1.8-2.0 x as long as outer margin. Inner margins of cmpd.

eyes weakly divergent ventrad. Face shagreened, matte, with fine punctures

which are separated by distance 1.5-2.0 x the dia. of single puncture. Clypeus

with antero-lateral corners not extending beyond tentorial pits, its anterior

margin with a small, medain tooth. Gena approx. as wide as basal width of

mandible. Temples, in dorsal view, weakly inflated. Mandible short, strongly

tapering, with lower tooth approx. 2.3 x as long as upper tooth. Thorax, in

lateral view, weakly dorso-ventrally flattened. Mesopleuron with speculum

smooth, polished, impunctate. Lower 0.5 of mesopleuron shagreened, weak-

ly polished, with fine punctures which are separated by distance 1.0-2.5 x

the dia. of single puncture. Propodeum with areola distinctly longer than

wide, weakly to strongly separated from petiolar area by a transverse carina.

Costula and lateral longitudinal carinae absent. Hind femur approx. 3.9-4.2

x as long as deep. Hind tarsal claw with strong teeth which do not extend to

base of tip. First abdominal segment with petiole approx. 1.1-1.2 x as long

as postpetiole. Ovipositor approx. 1.7-1.8 x as long as hind femur. Fore

wing approx. 4.8-7.3 mm. long. Hind wing with nervellus weakly intercepted

at basal 0.2-0.4

Head, thorax, abdomen, coxae, black ( most specimens have a faint

reddish spot between eye and antennal socket). Tegula, palpi, whitish. Tro-

chanters, femora, ferrugineous, the hind femur sometimes infuscated. Fore

tibia, ferrugineous; middle tibia ferrugineous to weakly fuscous; hind tibia

and hind tarsus weakly to strongly infuscated.

Male. Hind femur approx. 4.4-4.6 x as long as deep. Propodeum with areola

usually approx. 2.0 x longer than wide, bounded by weak carinae. Front of

scape and pedicel yellow. Face with yellow on inner orbits extending a short

distance behind antennal sockets, with median elongate-triangular fuscous

patch, the remainder yellow. Tentorial pits fuscous. Clypeus yellow. Tegula

white. Fore and middle coxae yellow apically, the remainder black. Hind

coxa entirely black. Fore and middle trochanters entirely yellow. Hind tro-

chanter with basal segment ferrugineous to fuscous, the apical segment

ferrugineous. Fore and middle femora ferrugineous. Hind femur ferrugineous,

with an apico-dorsal fuscous patch.

Material Examined. 21 males, 7 females

Distribution. CANADA, Alberta: Nordegg. British Columbia: Stone Mt.

Northwest Territories: Ft. McPherson; Reindeer Depot, Mackenzie Delta;

Yellowknife. Ontario: Frater. Quebec: Indian House Lk.: Knob Lk. Saskatch-

ewan: Carrot R. Yukon: Fort Cudahy. UNITED STATES, Alaska: Fort Yukon.

This species is confined to the Boreal Forest region of Alaska and Can-

ada.

Host and Biology. There are no rearing records for this species but it has

been associated with white spruce in Saskatchewan. Adults are active from

June through August, with a peak abundance in July.

Remarks. This species appears to be closely related to aldrichi. Females

of separata are easily distinguished from those of aldrichi by the much shorter

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 7

ovipositor and by the faint reddish spots present beside each antennal socket

(sometimes beside each mandible as well). Males of separata are difficult to

separate from aldrichi but typical specimens of the former are smaller, the

areola is relatively longer and bordered by weaker carinae, the hind tibia is

usually much paler. Some males can be separated bydistribution as those of

aldrichi occur west of the rockies while those of separata continue on through

the boreal forest.

aldrichi @

Sseparata ©

: 500

18 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Xylophylax alaskensis (Ashmead), n. comb.

Figs. 4, 18

Cubocephalus alaskensis Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 203.

Female. Alaska (U.S.N.M.). Examined in 1984.

Pyracmonoides lipoparium Viereck, 1925. Can. Ent. 57(7): 128 (key); 57(12):

299. Female. Sudbury, Ont. (CNC). New synonymy. Examined in 1984.

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin usually with a small, median tooth. Mandible with lower

tooth approx. 2.0 x as long as upper tooth. Coxae rufous. Ovipositor approx.

2.8 xas long as hind femur. Face of male entirely yellow, the inner orbits

with yellow not extending behind antennal sockets. Abdomen (both sexes)

black. :

Female. Antennal flagellum with 38-45 articles. Antennal scape with inner

margin approx. 1.7-1.9 x as long as outer margin. Inner margins of eyes

weakly divergent ventrad. Face strongly shagreened, matte, with medium

sized punctures which are separated by distance less than dia. of single punc-

ture. Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin usually with a small, median tooth. Gena approx. as wide

as basal width of mandible. Temples, in dorsal view, distinctly inflated.

Mandible short, strongly tapering, with lower tooth approx. 2.0 x as long as

upper tooth. Mesopleuron with speculum weakly to strongly shagreened, im-

punctate. Lower 0.5 of mesopleuron strongly shagreened, matte, with small

to medium sized punctures which are separated by distance 1.0-2.0 x dia. of

single puncture. Propodeum with areola usually about 2.0 x as long as wide,

sometimes a little longer, often partially separated from petiolar area by

traces of transverse carina laterally. Petiolar area usually broadly horse-

shoe shaped. Hind femur approx. 4.3-4.7 x as long as deep. Hind tarsal claw

with moderately strong teeth which do not extend to base of tip. First abdom-

inal segment with petiole approx. 1.0-1.1 x as long as postpetiole. Ovipositor

approx. 3.3-3.5 x as long as hind femur. Fore wing approx. 5.0-8.5 mm.

long.

Females indistinquishable from those of macrocephala except as follows:

apical 0.5 of clypeus ferrugineous; genae, adjacent to mandibles, ferrugineous;

hind femur with apex usually more extensively infuscated.

Male. Front of scape and pedicel yellow. Head with yellow on inner orbits

extending back to anterior ocellus. Face entirely yellow or with a very small

fuscous dot medially. Lower temple and clypeus entirely yellow. Prosternum

yellow in front of each coxa, the remainder of thorax black. Fore and middle

coxae, fore and middle trochanters, extreme bases of femora, yellow, re-

mainder of femora pale ferrugineous. Fore and middle tibiae pale ferrugineous.

Hind femur ferrugineous beyond base, with apical infuscation. Hind tibia dark

fuscous basally, externally, and apically, dark ferrugineous interno-medially.

Hind tarsus fuscous.

Material Examined. 59 males, 37 females

Distribution. CANADA. British Columbia: Hixon; Lower Post; Paint Potts,

oo mi. N. Radium; Robson. Labrador: Hopedale. Northwest Territories:

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 19

Ft. McPherson; Reindeer Depot, Mackenzie Delta; Yellowknife. Ontario:

Frater; Mississagi; Stittsville; Westree. Quebec: Gaspé; Joliette; Laniel.

Yukon: Hootalingua. UNITED STATES. Alaska: Kowak R., Kotzebue Sound.

Idaho: Leesburg. New York: Syracuse. Pennsylvania: Charter Oak.

This species occurs in conifer forests from Alaska south to Idaho, east

to New Brunswick (probably to Maritimes), south to Pennsylvania.

Host and Biology. The known hosts are larvae of Tetropium (Cerambycidae)

occuring in Pinaceae ( T. parvulum , T. velutinum ). Adults are active from

June through August but are most abundant in July.

Remarks. The closest relative of this species appears to xoridiformis from

the palearctic region. In North America it most closely resembles macro-

cephala (see remarks following macrocephala). This species is moderately

common in collections but is often misidentified as macrocephala .

alaskensis

r $00

20 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Longicauda Group

Antennal scape with inner margin approx. 1.3-1.4 x as long as outer

margin. Vertex long, with length behind lateral ocelli 0.8-1.4 x as long as

ocellar triangle. Temple long, weakly inflated. Clypeus with antero-lateral

corners not extending beyond tentorial pits, its anterior margin evenly con-

vex and lacking a median tooth. Mandible short, strongly tapering, with low-

er tooth approx. 1.3-1.4 x as long as upper tooth. Abdomen with petiolar seg.

moderately elongate. Ovipositor approx. 2.8-3.1x as long as hind femur.

Abdomen black.

This is a holarctic group with one holarctic species , longicauda(Horst-

m and one nearctic species, montana n. sp. The biology of this group is

unknown. but is associated with conifers.

Xylophylax longicauda (Horstmann), n. comb.

Figs. 5, 19

Rhimphoctona longicauda Horstmann, 1980. Nach bayer Ent. 29(2): 22. Fem-

ale. Oberbayern Ellmau (Munich).

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior margin evenly convex, without a median tooth. Mandible with

lower tooth slightly longer than upper tooth. Propodeal trough granulose,

with weak transverse wrinkling or sometimes wrinkling absent. Tegula white.

Abdomen black. Male with face entirely yellow and mesopleuron with yellow

markings.

Female. Antennal flagellum with 27-29 articles. Antennal scape with inner

margin approx. 1.3-1.4 x as long as outer margin. Cmpd. eyes with inner

margins weakly diverging ventrad. Face shagreened, matte, with small punc-

tures which are separated by distance less than or equal to dia. of single

puncture. Clypeus with antero-lateral corners not extending beyond tentorial

pits, its anterior margin evenly convex and lacking a median tooth. Gena

approx. 1.1-1.2 x as wide as basal width of mandible. Temples, in dorsal

view, moderately inflated. Mandible short, strongly tapering, with lower

tooth slightly longer than upper tooth. Mesopleuron with speculum polished,

smooth, impunctate. Lower 0.5 of mesopleuron shagreened, matte, with

fine punctures which are separated by distance 2.0-3.0 x the dia. of single

puncture. Propodeum completely areolated, with areola usually confluent

with petiolar area. Propodeal trough strongly matte, usually distinctly

granulose, usually with weak transverse wrinkling. Hind femur approx. 4.3-

4.7x as long as deep. Hind tarsal claw with moderately strong teeth which

often extend to base of tip. First abdominal segment with petiole approx.

1.2-1.3 x as long as postpetiole. Ovipositor approx. 2.8-3.2 x as long as

hind femur. Fore wing approx. 4.4-4.7 mm. long.

Head, thorax, abdomen, black. Median area of mandible yellow. Palpi,

tegula, fuscous. Coxae fuscous to black. Trochanters with basal segments

fuscous, apical segments ferrugineous. Fore and middle femora, fore and

middle tibiae, ferrugineous. Hind femur ferrugineous, with apex often in-

fuscated. Hind tibia, hind tarsus, fuscous. Wings weakly infuscated.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) Al

Male. Hind femur approx. 4.5-5.0 x as long as deep. Front of scape, pedicel,

inner orbits, face, clypeus, lower 0.7 of temple, median area of mandible,

palpi, fore and middle trochanters, pale yellow. Baso-lateral corners of

pronotum, tegula, sub-tegular ridge, lower 0.5 of mesopleuron, meso-

sternum, fore and middle coxae, ivory to pale yellow. Hind coxa yellow, with

dorsal surface partially to entirely fuscous. Fore and middle femora, fore

and middle tibiae, pale yellow-ferrugineous. Hind femur ferrugineous with

apex sometimes infuscated. Hind tibia with basal 0.6-0.7 ferrugineous, api-

cal 0.3-0.4 fuscous. Hind tarsus ferrugineous to fuscous.

In some males the temple is entirely yellow, the pronotum entirely

yellow laterally and in melanic specimens the mesosternum can be extensive-

ly marked with fuscous.

Material Examined. 22 males, 40 females.

Distribution. CANADA. Alberta: Banff; Calgary; Edmonton; Villenuve.

British Columbia: Hixon; Lac La Hache; Stanley; Stone Mt. Pk.; Squamish.

Labrador: locality ?. New Brunswick: Fredricton. Newfoundland: Raleigh;

South Branch. Ontario: Aylmer West; Ottawa; Westree. Quebec: Cascapedia

R.; Gaspé Co.; Old Chelsea; Laniel; Kam. co., Park Reserve; Stoneham.

Yukon: Dawson. UNITED STATES. Colorado: Steamboat Spr. Idaho: Moscow.

Maine: Lincoln Co. Michigan: Iron River. New Hampshire: Lancaster; Mast-

yard; Pinkham Notch. New York: Bemus Pt.; Cranberry Lake. Oregon: Grant

Co.3; Mt. Jefferson.

Host and Biology. This species has been observed ovipositing in timber in-

fested with Scolytus ventralis in Idaho. This could be misleading as small

species of Cerambycidae could also have been present. The species has been

associated with several conifer species, including spruce, fir and lodgepole

pine. Adults are active from May through August with peak activity in July.

Remarks. This is a holarctic species, as noted above. Specimens from North

America are typically smaller than pal earctic specimens and usually have the

hind tibia more infuscated. The closest relative in North America is montana

which has more extensive wrinkling on the propodeal trough and generally

much stronger propodeal carinae. Males of montana have faces which are

yellow with dark markings while the faces of male longicauda are entirely

yellow. The species montana is restricted to western North America while

longicauda has a widespread trans-continental distribution .

Xylophylax montana, new sp.

Figs. 6, 20

Similar to longicauda except that propodeal carinae stronger, propodeal

trough with distinct transverse wrinkling, tegula white to brown, faces of

males yellow with black markings and male mesopleuron entirely black,

confined to western North America.

HOLOTYPE. Female. Antennal flagellum with 30 articles. Antennal scape

with inner margin approx. 1.4 x as long as outer margin. Cmpd. eyes with

inner margins weakly diverging ventrad. Face strongly shagreened, matte,

with small punctures which are separated by distance less than dia. of single

puncture. Clypeus with antero-lateral corners not extending beyond tentorial

pits, its anterior margin evenly convex and lacking a median tooth. Gena

approx. 1.1 x as wide as basal width of mandible. Temples, in dorsal view,

inflated. Mandible short, strongly tapering, with lower tooth slightly longer

than upper tooth. Mesopleuron with speculum weakly shagreened, polished,

22 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

impunctate. Lower 0.5 of mesopleuron with small punctures which are sep-

arated by distance 1.0-2.0 x dia. of single puncture. Propodeum completely

areolated, the carinae strong. Areola slightly longer than wide, with coarse

wrinkling laterally, completely confluent with petiolar area. Petiolar area

with coarse transverse wrinkling. Median longitudinal carina strongly angul-

ate medially. Hind femur approx. 4.2 x as long as deep. Hind tarsal claw

with moderately strong teeth which extend to base of tip. First Abdominal

segment with petiole approx. 1.2 x as long as postpetiole. Ovipositor approx.

2.6 x as long as hind femur. Fore wing approx. 6.0 mm. long.

Head,thorax, abdomen, coxae, black. Palpi, tegula, fulvous. Basal

segments of trochanters fuscous, apical segments ferrugineous. Femora,

fore tibia, ferrugineous, the middle tibia ferrugineous with apex weakly

infuscated. Hind tibia dark fuscous basally, apically and internally, the ex-

terno-median surface dark ferrugineous. Hind tarsus fuscous. Wings very

weakly infuscated. .

Range and Variation. Antennal flagellum with 29-30 articles. Antennal scape

with inner margin 1.3-1.4 x as long as outer margin. Clypeus with anterior

margin sometimes very weakly produced medially. Areola sometimes weakly

separated from petiolar area by weak, incomplete transverse carina. Hind

femur approx. 4.1-4.4 x as long as deep. Abdominal petiole approx. 1.2-1.3

x as long as postpetiole. Ovipositor approx. 2.6-2.8 x as long as hind femur.

Fore wing 4.5-6.5 mm. long. Tegula white to btown. Palpi fulvous to fuscous.

Apical segments of trochanters ferrugineous to fuscous. Hind femur sometimes

infuscated apically.

Male. Hind femur approx. 4.3-4.8 x as long as deep. Front of scape yellow.

Face yellow with 3 vertical fuscous bars medially. Clypeus yellow medially,

fuscous along anterior and posterior margins. Palpi pale fulvous. Tegula

ivory. Thorax entirely black. Fore coxa, most of middle coxa, yellow. Hind

coxa black. Fore and middle trochanters, yellow, hind trochanter with basal

segment fuscous, apical segment yellow with some infuscation. Fore and

middle femora yellow ferrugineous, hind femur ferrugineous, with infuscated

areas apically and ventrally. Hind tibia paler than in female, usually with a

weak banding pattern.

Holotype. Female. nr. Glacier Pt., Yosemite Pk., California, July 19, 1948,

H. and M. Townes. Deposited in Townes collection, Gainesville, Fla.

Paratypes. CANADA. British Columbia: Hixon, 20-VI-1965 (1 female), E.D.A.

Dyer (CNC); Stone Mt. Pk., 3500-3800', VII-19-26-1973 (2 males, 2 females)

H. & M. Townes (HMT); Summit Lake, 4600', 16-VII- 1959 (1 female), R. E,.

Leech (CNC). Alberta: Banff, 6-VIIT-1924 (ifemale), E. Hearle (CNC) and

4700', Aug. 16, 1962 (i female), K. C. Hermann (CNC). UNITED STATES.

Alaska: Anchorage, VI-30-VII-12-1976 (1 male, 3 females) Peter A. Rush

(HMT); McKinley Hotel, VI - 25-1954 (1 female), W.C.F. (HMT). California:

Cisco, July 31, 1948, (1 female), H.M.G.D.&J Townes (HMt); Echo Lk.,

Eldorado Co., VII-1-1954 (ifemale), W.W. Middlekauff (UCB); nr. Glacier

Pt., Yosemite Pk., July 16-18-1948 (63 males, 14 females), H.M.G.D. & J.

Townes (HMT); May LK., 10,500', Yosemite Pk., July 27, 1948 (4 males, 1

female), H.M.G.D. & J. Townes (HMT); Sagehen Cr., VII-10-12-1978 (1 male,

2 females), R. M. Bohart (HMT); Snow Flat, Yosemite Pk., 8700', July 26-

27, 1948 (4 males, 1 female), H.M.G.D. & J. Townes (HMT). Idaho: Lowman,

Aug. 9, 1978 (1 female), H. & M. Townes (HMT); nr. Stanley, Aug. 3, 1978

(1 male), H. & M. Townes (HMT). Washington: Mt.Ranier, 2900', July 12,

1940 (1 female), H. & M. Townes (HMT).

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 23

Distribution. Rocky Mts., Aka. to Idaho and Sierra-Nevada range, Wash. to

Calif.

Etymology. The species epithet is in reference to the montane habitats this

species seems to prefer.

Host and Biology. There are no rearing records for this species. Adults are

active from June through August with a peak of activity in July.

Remarks. This species is very closely related to longicauda. See remarks

under longicauda .

longicauda @

montana e

24 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Pectoralis Group

Antennal scape with inner margin approx. 1.3-1.4 x as long as outer

margin. Vertex long, with length behind lateral ocelli approx. 0.9-1.4 x as

long as ocellar triangle. Temple not or weakly inflated. Clypeus with antero-

lateral corners not extending beyond tentorial pits, its anterior margin even-

ly convex and lacking a median tooth. Gena narrow, approx. 0.7 as wide as

basal width of mandible. Mandible short, strongly tapering, with lower tooth

1.9-2.1 x as long as upper tooth. Thorax, in lateral view, somewhat dorso-

ventrally flattened. Abdomen with petiolar segment short, stout. Ovipositor

approx. 3.0 x as long as hind femur. Abdomen (both sexes) black.

This is a holarctic group with one palearctic species (pectoralis) and

2nearctic species (albicoxa, genalis) . Hosts and host-plant associations

are known only for albicoxa (Curculionidae and Cerambycidae in hardwoods)

and pectoralis (Cerambycidae in Pinaceae).

Xylophylax albicoxa, new sp.

Fries: 7, 22

Face strongly convex. Clypeus with antero-lateral corners not extend-

ing beyond tentorial pits, its anterior margin not produced medially. Prop-

odeal carinae unusually strong and propodeal trough with very coarse trans-

verse wrinkling. Coxae white to yellow. Abdomen black. Face of male white,

mesopleuron with white areas, and lower division of metapleuron white.

HOLOTYPE. Female. Antennal flagellum with 29 articles. Antennal scape

with inner margin approx. 1.4 x as long as outer margin. Inner margins of

cmpd. eyes very weakly diverging ventrad. Face strongly convex, strongly

shagreened, matte, with minute punctures which are separated by distance

approx. equal to dia. of single puncture. Clypeus with antero-lateral corners

not extending beyond tentorial pits, its anterior margin not produced medially.

Gena approx. 0.7 x as wide as basal width of mandible. Temples, in dorsal

view, not inflated, moderately sloping inwardly. Mandible with lower tooth

approx. 2.0 x as long as upper tooth. Thorax, in profile, moderately dorso-

ventrally flattened. Mesopleuron with speculum finely wrinkled, polished,

impunctate. Lower 0.5 of mesopleuron strongly shagreened, matte, with

small punctures which are separated by distance less than or equal to dia. of

single puncture. Propodeum with unusually strong carinae. Areola distinctly

longer than wide and completely confluent with petiolar area. Propodeal

trough with approx. 11 very coarse transverse wrinkles. Hind femur approx.

4.4x as long as deep. Hind tarsal claw with strong teeth which extend to

base of tip. First abdominal segment with petiole approx. as long as post-

petiole. Ovipositor approx. 3.0 x as long as hind femur. Fore wing approx.

4.0 mm. long. Hind wing with nervellusweakly intercepted at basal 0.3.

Head, thorax, abdomen, black. Mandible, tegula, white. Fore coxa

white tinged with ferrugineous. Middle and hind coxae, femora, tibiae (ex-

cept extreme apex of hind), ferrugineous. Extreme apex of hind tibia, hind

basitarsus, fuscous. Wings hyaline.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 25

Range and Variation. Antennal flagellum with 29-32 articles. Antennal scape

with inner margin approx. 1.3-1.4 x as long as outer margin. Areola some-

times separated from petiolar area by transverse carina. Hind femur approx.

4.1-4.4 x as long as deep. Fore wing approx. 4.0-5.0 mm. long. Hind wing

with nervellus intercepted near basal 0.3-0.4. Fore coxa pale ferrugineous

to ivory. Fore trochanter pale ferrugineous to ivory.

Male. Generally much paler than female. Black. Scape, pedicel, inner orbits,

temple, face, clypeus, mandible, tegula, pronotum, most of mesopleuron,

metasternum, lower division of metapleuron, coxae, trochanters, fore and

middle femora, tibiae (except apex of hind which is weakly infuscated), hind

tarsal segments (except apices which are weakly infuscated), ivory. Hind

femur ivory to pale ferrugineous.

Holotype. Female. Gulfshores, Alabama, April 25, 1968, H. & M. Townes.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. Alabama: one male with same data as holotype (HMT). Connecticut:

South Woodstock, ex Magdalis olyra on hickory, date? (1 female), coll. ?

(USNM). Georgia: St. Catherines Is., Liberty Co., April 24-28-1972, (1 male),

Thompson and Picchi (AMNH). Kansas: Lawrence, Douglas Co., V-3-1981

(1 female), D. B. Wahl (DWC). Louisiana: Mansura, IV-13-1909 (1 female),

W. D. Pierce (USNM). Michigan: Ann Arbor, VI-16-1980 (1 female), H. & M.

Townes (HMT). North Carolina: Cedar Mt., 29-IV-1913 (3 females), ex

Tillamorpha geminata, F. C. Craighead (USNM); Durham, III-15-1943 (1 male)

on Juglans nigra, Wm. Haliburton, J. A. Beal (USNM). Tennessee: Bolivar,

Hardeman Co., Apr. 2-6-1974, ex Cercis canadensis (6 males, 2 females),

R. D. Ward (HMT). Unknown locality: 1 male, 1 female, no further data

(USNM).

Host and Biology. This species has been reared from Magdalis olyra (Curc-

ulionidae) in hickory and Tillomorpha (= Tilloclytus) geminatus (Cerambycid-

ae). Other plant associations include Juglans nigra (black walnut) and Cercis

canadensis (redbud). It appears that this species attacks, in general, small

sub-cortical coleopterous larvae in a variety of hardwoods. Adults are active

in late spring in the northern part of its range, and early spring in the south-

ern portions.

Etymology. The species is named for the ivory colouration of the coxae of

the males.

Remarks. The small size, convex face,sculpturing of the propodeum, extens-

ive ivory colouration of males, and association with hardwoods will easily

distinguish this species from all other nearctic Xylophylax . Its closest

nearctic relative appears to be genalis ( see remarks following genalis) .

Xylophylax genalis, new sp.

Pige, 3, 21

Clypeus with antero-lateral corners not extending beyond tentorial pits,

its anterior not produced medially. Temples, in dorsal view, moderately

sloping inwardly. Mandible with lower tooth approx. 2.0 x as long as upper

tooth. Coxae of female ferrugineous. Abdomen black. Abdominal petiole

approx. as long as postpetiole. Face of male white to yellow, with a median

26 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

infuscated spot. Male thorax entirely black.

HOLOTYPE. Female. Antennal flagellum with 29 articles. Antennal scape

with inner margin approx. 1.3 x as long as outer margin. Inner margins of

cmpd. eyes parallel ventrad. Face strongly shagreened, matte, with fine

punctures which are separated by distance approx. equal to dia. of single

puncture. Face, in lateral profile, moderately convex. Clypeus with antero-

lateral corners not extending beyond tentorial pits, its anterior margin not

produced medially. Gena approx. 0.7 x as wide as basal width of mandible.

Temples, in dorsal view, moderately sloping inwardly. Mandible with lower

tooth approx. 2.0 x as long as upper tooth. Thorax, in profile, weakly dorso-

ventrally flattened. Mesopleuron with speculum smooth, polished, impunctate.

Lower 0.5 of mesopleuron moderately shagreened, polished, with small

punctures which are separated by distance less than or equal to dia. of single

puncture. Propodeum with areola distinctly longer than wide, weakly separated

by weak transverse carina from petiolar area. Petiolar area with numerous

moderately coarse irregular wrinkles. Hind femur approx. 4.1.x as long as

deep. Hind tarsal claw with weak teeth which do not extend to base of tip.

First abdominal segment with petiole approx. as long as postpetiole. Ovipos-

itor approx. 3.0 x as long as hind femur. Fore wing approx. 5.1 mm. long.

Hind wing with nervellus weakly intercepted near basal 0.3.

Head, thorax, abdomen, black. Median area of mandible, palpi, tegula,

white tinged with ferrugineous. Fore and middle coxae, apex of hind coxa,

trochanters, femora (except apico-dorsal patch of hind), fore and middle

tibiae, ferrugineous. Basal 0.7 of hind coxa, apico-dorsal patch of hind fem-

ur, hind tibia, fuscous. Wings hyaline, with darkly pigmented veins.

Range and Variation. Antennal flagellum with 29-31 articles. Antennal scape

with inner margin approx. 1.3-1.4 x as long as outer margin. Lower 0.5 of

mesopleuron polished to matte. Hind femur approx. 3.8-4.3 x as long as deep.

Fore wing approx. 4.0-5.5 mm. long. Middle and hind coxae ferrugineous to

fuscous. Hind tibia ferrugineous with apex fuscous to entirely fuscous.

Male. Vertex, median spot on face, thorax, abdomen, black. Fore coxa

ivory to pale ferrugineous. Middle coxa pale to dark ferrugineous. Hind coxa

ferrugineous to fuscous. Legs coloured as in female.

Holotype. Female. Lake Wohlford, California, 1V-27-1974, H. & M. Townes.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. CANADA. British Columbia: Stone Mt. Pk., 3800'. VII-22-1973

(1 female), H. &M. Townes (HMT). UNITED STATES. California: Alpine Cr.,

Tahoe, VII-5-1915 (1 female), E. P. Van Duzee (UCB); Lake Wohlford, IV-

22-30-1974 (31 males, 19 females), H. & M. Townes (HMT). Idaho: Stanley,

Aug. 7, 1978 (1 female), H. & M. Townes (HMT). Oregon: Corvallis, June 4,

1978 (1 male), H. & M. Townes (HMT); Pinehurst, VI-21-1978 (1 female), H.

& M. Townes (HMT).

This species is distributed throughout the Cascade and Rocky Mt. ranges.

Host and Biology. There are no rearing records or plant associations for this

species. Adults are active from early spring to mid-summer.

Remarks .The somewhat dorso-ventrally flattened thorax, moderately oblique

apical margin of scape, narrow gena, and short, stout petiolar segment with

petiole as long as postpetiole demonstrate a close relationship to albicoxa .

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 27

The weaker propodeal carinae, darker colouration of the females and entire-

ly black thorax of the males will distinguish this species from albicoxa.

albicoxa @

genalis ©

\ 500

28 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

Subgenus Euraspidion Sanborne

The description of this subgenus can be found in Sanborne, 1986b, imm-

ediately preceeding this paper. Salient features of the group are presented

earlier in this paper in the identification key.

Brevicauda Group

Antennal scape with inner margin approx. 1.5-1.6 x as long as outer

margin. Vertex short, with length behind lateral ocelli approx. 0.5-0.7 x as

long as ocellar triangle. Temple moderately long, not inflated, strongly

sloping inwardly in dorsal view. Clypeus with antero-lateral corners extend-

ing beyond tentorial pits, its anterior margin not produced medially. Mandible

long, weakly tapering, with lower tooth approx. 1.4-1.5 x as long as upper

tooth. First abdominal (petiolar) segment moderately elongate. Ovipositor

short, approx. 1.3 x as long as hind femur. Abdomen (both sexes) partially to

entriely rufous.

This is a nearctic group with a single species, brevicauda .

Xylophylax brevicauda, new sp.

Figs. 12, 26

Clypeus with antero-lateral corners extending beyond tentorial pits, its

anterior na rgin not produced medially. Mandible with lower tooth approx.

1.4-1.5 x as long as upper tooth. Temples, in dorsal view, strongly sloping

inwardly. Ovipositor approx. 1.3 x as long as hind femur. Abdomen (both

sexes) partially to entirely rufous. Face of male yellow with black markings.

HOLOTYPE, Female. Antennal flagellum with 30 articles. Antennal scape

with inner margin approx. 1.6 x as long as outer margin. Inner margins of

cmpd. eyes weakly diverging ventrad. Face shagreened, matte, with medium

sized punctures which are separated by distance less than dia. of single punc-

ture. Clypeus polished, with numerous small punctures, with antero-lateral

corners extending beyond tentorial pits, its anterior margin not produced

medially. Gena approx. as wide as basal width of mandible. Temples, in

dorsal view, strongly sloping inwardly. Mandible long, weakly tapering, with

lower tooth approx. 1.5 x as long as upper tooth. Mesopleuron with speculum

smooth, polished, impunctate. Lower 0.5 of mesopleuron weakly shagreened,

more or less polished, with medium sized punctures which are separated by

distance less than or equal to dia. of single puncture. Propodeum with areola

weakly separated from petiolar area by a transverse carina, the propodeal

trough rather broad and with strong carinae. Hind femur approx. 3.8 x as

long as deep. Hind tarsal claw with strong teeth which extend to base of tip.

First abdominal segment with petiole approx. 1.3 x as long as postpetiole.

Ovipositor approx. 1.3 x as long as hind femur. Fore wing approx. 7.7 mm.

long. Hind wing with nervellus distinctly intercepted near middle.

Head, thorax, coxae, abdominal petiole, basal 0.6 of postpetiole, black.

Tegula, basal segments of all trochanters, apical 0.4 of middle and hind

tibiae, fuscous. Femora, basal 0.6 of hind tibia, apical 0.4 of postpetiole,

abdominal tergites 2 to apex, rufous. Fore tibia yellow-ferrugineous. Hind

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 29

tarsal segments 1-4 yellow, 5th segment fuscous. Fore wings with yellow

tinge, particularly toward bases.

Range and Variation. Antennal flagellum with 27-30 articles. Antennal

scape with inner margin 1.5-1.6 x as long as outer margin. Propodeum with

areola often strongly separated from petiolar area by transverse carina. Hind

femur approx. 3.5-4.5 x as long as deep. Ovipositor approx. 1.3-1.5 x as long

as hind femur. Fore wing 6.1-7.3 mm. long. Postpetiole sometimes entirely

rufous. Coxae varies from fuscous to black. Basal segments of trochanters

vary from ferrugineous to fuscous.

Male. Hind femur approx. 4.0-4.5 x as long as deep. Colour as in female

except as follows: front of scape and pedicel, inner orbits, lower temple,

ventral surfaces of fore and middle coxae, fore trochanter, yellow; face

entirely yellow or sometimes with median fuscous spot; clypeus yellow marked

with fuscous basally and medially or entirely yellow; hind femur often with

infuscated areas.

Holotype. Female. King Mt., Old Chelsea, Quebec, VI-22-26-1977, P.M.

Sanborne. Deposited in Canadian National Insect Collection, Ottawa, Ont.

Paratypes. CANADA. Alberta: Bilby, 25-VI-1924 (1 female), G. Salt (USNM).

Ontario: Havelock, May 27, 1977 (2 males), J. W. McCreadie (PMS). Quebec:

Old Chelsea, King Mt., 1150', 21-VI-1965 (1 male), J. R. Vockeroth (CNC).

UNITED STATES. Illinois: Glen Ellyn, 30-V-1949 (1 male), A. L. Melander

(UCR). Iowa: Sioux City, date? (1 female), C. N. Ainslie (USNM). New Hamp-

shire: Pinkham Notch, June 25, 1938 (1 male), H. & M. Townes (HMT). Ohio:

Ashtabula Co., V-26-?, (1 male) C. H. Kennedy (HMT); Hocking Co., May,

192? (2 females) (PMS &USNM) and June 192? (1 male) (HMT), (1 female)

(USNM); Jefferson St. Pk., June 8, 1952 (1 female), C. Dasch (CDC); Oak

Pk., 5 mi. W. New Concord, May 27, 1968 (2 females), C. Dasch (CDC).

West Virginia: Bowden, VI-6-7-1980 (1 female), C. Dasch (CDC); Cranberry

Gls, 2-VI-1955 (1 female), H. V. Weems (HMT).

This species is distributed from Alberta south to Iowa, east to New

Hampshire, and south to West Virginia.

Host and Biology. There are no rearing records for this species. Adults are

active during May and June.

Etymology. The species epithet is in reference to the short ovipositor.

Remarks .The broad clypeus and red abdomen indicates a relationship with

the Vancouverensis Group. The strongly sloping temples and unusually short

ovipositor will easily distinguish this species from all other nearctic Xylo-

phylax and are my reasons for placing it in its own species group. Though

widely distributed, it is very rare in collections.

30 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

brevicauda

\ 500

Vancouverensis Group

Antennal scape with inner margin approx. 1.5-1.6 x as long as outer

margin. Vertex moderately long, with length behind lateral ocelli approx.

0.8-1.1 x as long as ocellar triangle. Temples, in dorsal view, distinctly

inflated. Clypeus with antero-lateral corners extending beyond tentorial pits,

its anterior margin not produced medially. Mandible long, weakly tapering,

with lower tooth approx. 1.8-2.0 x as long as upper tooth. Ovipositor approx.

2.8 x as long as hind femur. Face of females entirely black. Abdomen (both

sexes) partially to entirely rufous.

This is a nearctic group comprising 3 species (breelandi, ruficincta,

vancouverensis). The group is associated with conifers and although there

are no rearing records, the hosts are presumed to be wood-boring larvae of

Cerambycidae.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 31

Xylophylax vancouverensis (Harrington), n. comb.

Figs. 10, 24

Pyracmon vancouverensis Harrington, 1894. Can. Ent. 26: 246. Female.

Vancouver, British Columbia (CNC). Examined in 1984.

Clypeus with antero-lat eral corners extending beyond tentorial pits, its

anterior margin not produced medially. Temples distinctly inflated in dorsal

view. Mandible with lower tooth approx. 1.8-2.0 x as long as upper tooth.

Mesoplueron polished, with medium to large sized punctures. Propodeum

with areola approx. as long as wide. Coxae extensively to entirely black.

Abdomen extensively rufous. Face of male entirely yellow (Rocky Mts.), with

black markings (Cascade-Sierra Nevada Ranges)

Female. Antennal flagellum with 29-33 articles. Antennal scape with inner

margin approx. 1.5-1.6 x as long as outer margin. Cmpd. eyes with inner

margins weakly diverging ventrad. Face shagreened, matte, rather flat, with

medium sized punctures which are separated by distance less than dia. of

single puncture. Clypeus with antero-lateral corners extending beyond tent-

orial pits, its anterior margin not produced medially. Gena approx. 0.8 x as

wide as basal width of mandible. Temples distinctly inflated in dorsal view.

Mandible long, weakly tapering, with lower tooth approx. 1.8-2.0 x as long

as upper tooth. Mesopleuron with speculum smooth to weakly wrinkled, pol-

ished, impunctate. Lower 0.5 of mesopleuron with medium sized punctures

which are separated by distance less than or equal to dia. of single puncture.

Propodeum completely areolated, the carinae moderately strong to strong.

Areola approx. as long as wide, sometimes separated from petiolar area by

transverse carina. Hind femur approx. 4.0-4.6 x as long as deep. Hind tar-

sal claw with weak teeth which do not extend to base of tip. First abdominal

segment with petiole approx. 1.2-1.3 x as long as postpetiole. Ovipositor

approx. 2.8 x as long as hind femur. Fore wing approx. 5.5-8.0 mm. long.

Head, thorax, abdominal petiole, black. Postpetiole partially rufous to

entirely black. Abdominal tergites 2 to apex partially to entirely rufous.

Tegula, basal segments of trochanters, apical 0.3-1.0 of hind tibia, fuscous.

Femora, fore and middle tibiae, ferrugineous. Wings very lightly infuscated.

Male. Hind femur approx. 4.4-5.5 x as long as deep. Face entirely yellow

to yellow with black markings. Thorax entirely black to black with yellow

markings. Remaining colouration similar to female.

There are three subspecies which can be identified by the following key.

1. Petiolar segment of abdomen bicoloured. Second abdominal tergite entire-

ly rufous. Hind femur approx. 4.0-4.5 x as long as deep in females,

4.5-5.0 x in males. Face of males entirely yellow. Thorax of males

with yellow markings. Rocky Mts. e. to Saskatchewan, s. to Idaho.

Petiolar segment of abdomen entirely black or with extreme apical mar-

gin of petiole reddish. Second abdominal tergite fuscous to black

basally, rufous apically. Face of male yellow with fuscous or black

markings. Thorax of males entirely black.....ccccccccscccceccede

32 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

2. Occuring in southern Rocky Mts. Hind femur approx. 4.0-4.5 x as long

as deep in females, 4.5-5.0 x in males (estimated as male unknown).

Hind tibia extensively to entirely darkly infuscated..... Se are sea

bop Cele 006 Ocoee e eee eens ececcoeece se -v. tibialis, new subspecies.

Occuring in Cascade-Sierra Nevada ranges. Hind femur approx. 4.5-

4.8 x as long as deep in females, 5.0-5.5 x in males. Hind tibia

ferrugineous, with apical 0.2-0.3 weakly infuscated......ce.cccces

ccc cree ccs coevecccceccccscveeccess Ve maculosa, new subspecies.

Xylophylax vancouverensis vancouverensis (Harrington)

Pyracmon vancouverensis Harrington, 1894. Can. Ent. 26: 246. Female.

Vancouver, B. C. (CNC).

Female. Hind femur approx. 4.0-4.5 x as long as deep. Abdomen with petiole

and basal 0.5-0.7 of postpetiole black. Second abdominal tergite entirely ruf-

ous. Hind tibia with apical 0.3 weakly infuscated.

Male. Hind femur approx. 4.4-5.0 x as long as deep. Face and clypeus entir-

ely yellow. Pronotum with latero—median yellow spot. Mesopleuron with sub-

tegular ridge and antero—ventral areas yellow. Mesosternum, fore and middle

coxae, yellow. Hind tibia coloured as female.

Distribution. CANADA. Alberta: Banff; Drumheller. British Columbia: Adam's

Lk.; Clinton; Kamloops; Keremeos; Osoyoos; Richter Pass Rd., 7 mi. W.

Osoyoos. Saskatchewan; Elbow. UNITED STATES. California:Carrville, Trin-

ity Co.3; Sequoia Nt. Pk. Colorado: Boulder. Idaho: Bear Valley, Valley Co.;

Bear Pass Cr., Butte Co.; Idaho City. Washington: Blue Mts.

This subspecies is distributed in the Rocky Mts. from British Columbia

to Colorado, east to Saskatchewan, with a few specimens taken in the Cascade-

Sierra Nevada ranges.

Host and Biology. Unknown. Adults are active from late May to late July.

Remarks. The bicoloured petiolar segment of both sexes , entirely yellow

face of male, and yellow markings of male thorax will easily distinguish this

subspecies. It could be argued that this is a good species as its range extends

into the ranges of the other two subspecies. It is my belief that this represents

a recent expansion of the range of v. vancouverensis. Tests for hybridization

will have to be performed with specimens taken from contact zones.

Xylophylax vancouverensis tibialis, new subspecies

Female. Similar to v. maculosa except that hind femur distinctly more swol-

len and hind tibia extensively to entirely dark fuscous.

Male. Unknown but presumed to be similar to v. maculosa except that hind

femur distinctly more swollen and hind tibia extensively to entirely dark

fuscous.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 33

Holotype. Female. Boulder, Colorado, June 18, 1933, M. T. James. Depos-

ited in Townes collection, Gainsville, Florida. |

Paratypes. Colorado: Boulder, 5800', Flagstaff Canyon, June 10, 1961 (2 fem-

ales), coll. ? (CNC); Cottonwood Pass, Eagle Co., 8200', VIU-2-1961 (1 female)

R.J. Joe (HMT); Unknown locality, date? (1 female) C. F. Baker (USNM).

New Mexico: Ute Park, Cimarron Canyon, on pastinaca flowers, July 1-4,

1960 (1 female), Burks and Kinzer (HMT).

Distribution. Southern Rocky Mts. (Colorado, New Mexico).

Host and Biology. There are no rearing records for this subspecies. Adults

are active in June and July.

Etymology. The subspecies name draws attention to the hind tibia which is

extensively to entirely fuscous.

Remarks. The extensively dark fuscous coloured hind tibia distinguishes this

subspecies from the other two. _

Xylophylax vancouverensis maculosa, new subspecies

Female. Hind femur approx. 4.6-5.0 x as long as deep. Abdomen with pet-

iolar segment entirely black or, at most, with extreme apical margin red-

dish. Second abdominal tergite with basal 0.4-0.5 fuscous to black. Hind

tibia with apical 0.3 weakly fuscous.

Male.Hind femur approx. 5.0-5.5 x as long as deep. Face with large median

triangular fuscous patch. Clypeus yellow with antero-median fuscous spot, the

extreme anterior margin fuscous. Thorax entirely black. Fore and middle

coxae with apico-ventral surfaces yellow. Hind tibia and abdomen coloured as

in female.

Holotype. Female. Corvallis, Oregon, May 18, 1978, H. & M. Townes.

Deposited in Townes collection, Gainsville, Florida.

Paratypes. Oregon: 4 mi. s. Canyonville, Douglas Co., May 4, 1936 (1 female)

G. Ferguson (USNM); Cornelius, May 6, 1936 (1 male), Gray and Schuh (OSU);

Corvallis, 5-8-98 (1 male), coll. ? (USNM) and (1 male, 1 female), coll? (OSU)

and May 8, 1925 (1 female), H. A. Scullen (USNM) and May 10, 1925 (1 male)

H. A. Scullen (USNM) and May 8, 1932 (1 male), coll. ? (USNM); Dixie Cr.,

May 30, 1937 (1 female), G. Thompson (OSU) and May 14, 1938 (1 male), Gray,

Schuh (OSU); Hyatt Reservoir, VI-19-1978 (1 female), H- & M. Townes (HMT);

Seneca, July 11, 1978 (1 male), H. & M. Townes (HMT); Sulphur Springs, 10

mi. n. Corvallis, May 16, 1952 (1 male), Vincent Roth (OSU) and 5 mi. n.

Corvallis May 14, 1963 (1 male), David R. Smith (OSU).

Distribution. Known only from Oregon

Host and Biology. There arenorearing records for this species but it has been

associated with douglas fir. Adults are active from May through July.

Etymology.The subspecies name refers to the large triangular fuscous patch

on the face of the male.

Remarks. This subspecies is easily distinguishable from the nominal subspecies

34 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

by the black petiolar segment and partially black 2nd abdominal tergite, and

by the colouration of the male. It is distinguished from v. tibialis by its

distribution and much paler hind tibia.

v. vancouverensi

v. tibialis

v. maculosa

t 500

Sanborne: Revisionf of Nearctic Xylophylax (Ichneumonidae) 35

Xylophylax ruficincta, new sp.

Pigs. 9, 23

Clypeus with antero-lateral corners extending beyond tentorial pits, its

anterior margin not produced medially. Mandible with lower tooth approx.

1.8-2.0 x as long as upper tooth. Temples ditinctly inflated in dorsal view.

Coxae rufous. Abdomen entirely rufous. Face of male entirely yellow.

HOLOTYPE. Female. Antennal flagellum with 35 articles . Antennal scape

with inner margin approx. 1.6 x as long as outer margin. Inner margins of

cmpd. eyes weakly diverging ventrad. Face shagreened, flat, with medium

sized punctures which are separated by distance less than dia. of single punc-

ture. Clypeus weakly shagreened, polished, with antero-lateral corners ex-

tending beyond tentorial pits, its anterior margin not produced medially.

Gena approx. 0.8 x as wide as basal width of mandible. Temples distinctly

inflated in dorsal view. Mandible long, weakly tapering, with lower tooth

approx. 1.8-2.0 x as long as upper tooth. Mesopleuron with speculum polish-

ed, impunctate. Lower 0.5 of mesopleuron densely, coarsely punctate. Pro-

podeum with carinae very strong, the areola completely confluent with petiol-

ar area. Hind femur approx. 4.7x as long as deep. Hind tarsal claw with

weak teeth which extend to base of tip. First abdominal segment with petiole

approx. 1.5 x as long as postpetiole. Ovipositor approx. 2.8 x as long as hind

femur. Fore wing approx. 9.0 mm. long. |

Head, thorax, apex of mandible, baso-dorsal patch of petiole, black .

Tegula fuscous. Coxae, trochanters, femora, abdomen, rufous. Hind tibia

fuscous basally, grading to black apically. Hind tibial spurs, hind basitarsus,

yellow. Wing veins very darkly pigmented.

Range and Variation. Antennal flagellum with 30-35 articles. Antennal scape

with inner margin approx. 1.5-1.6 x as long as outer margin. Hind femur

approx. 4.6-4.7 x as long as deep. Petiole approx. 1.5-1.6 x as long as post-

petiole. Fore wing approx. 6.5-9.0 mm. long. Hind tibia with basal 0.5

rufous to fuscous. Apical tergites of abdomen sometimes infuscated.

Male. Hind femur approx. 4.9-5.0 x as long as deep. Colour as in female

except as follows: front of scape, inner orbits, face, lower temple, clypeus,

fore coxa, fore trochanter, ventral surface of middle coxa, much of middle

trochanter, yellow.

Holotype. Female. Oregon, Corvallis, June 21, 1978, H. & M. Townes.

Deposited in Townes collection, Gainsville, Florida.

Paratypes. CANADA. British Columbia: Royal Oak, 24-VII-1917 (1 male), W.

Downes (CNC). UNITED STATES. California: Blocksburg, 6-6-1935 (1 male),

E. W. Baker (USNM); Summit Mt. Canyon, Solano Co., V-30-1960 (1 male),

A. S. Menke (UCR). Oregon: Corvallis, 28, June, 1936 (1 female), G. Ferg-

uson (HMT) and July 15, 1978 (1 female), H. & M. Townes (HMT); Mt. Hood,

VII-26-1978, 3500' (1 male), H. &M. Townes (HMT). Washington: Signal Pk.,

July 4, 1950 (1 male), A. R. Rolfs (USNM).

Distribution. This species is confined to the Cascades-Sierra Nevada ranges.

Host and Biology. There are no rearing records for this species but it has

36 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

been associated with douglas fir.

Etymology. The species epithet refers to the red abdomen of both sexes.

Remarks. The entirely rufous coxae distinguishes this species from its

close relative, vancouverensis.

ruficincta

t 500

Xylophylax breelandi, new sp.

Figs. 1, 25

Clypeus with antero-lateral corners extending beyond tentorial pits.

Mandible with lower tooth approx. 2.0 x as long as upper tooth. Temples

inflated in dorsal view. Thorax weakly dorso-ventrally flattened. Meso-

plueron matte, with fine punctures. Propodeum with areola longer than wide.

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 37

Coxae extensively black. Abdomen extensively rufous.

HOLOTYPE. Female. Antennal flagellum with 31 articles. Antennal scape

with inner margin approx. 1.5 x as long as outer margin. Inner margins of

cmpd. eyes very weakly diverging ventrad. Face shagreened, matte, with

fine punctures which are separated by distance approx. equal to dia. of single

puncture. Clypeus shagreened, matte, with antero-lateral corners extending

beyond tentorial pits. Gena approx. 0.8 x as wide as basal width of mandible.

Temples distinctly inflated in dorsal view. Mandible long, weakly tapering,

with lower tooth approx. 2.0 x as long as upper tooth. Mesopleuron with

speculum finely wrinkled, weakly polished, impunctate. Lower 0.5 of meso-

pleuron strongly shagreened, matte, with small punctures which are separated

by distance approx. equal to 1.0-1.5 x the dia. of single puncture. Propodeum

with areola distinctly longer than wide, the areola separated from petiolar

area by a weak transverse carina. Lateral longitudinal carina present only as

trace. Hind femur approx. 4.3 x as long as deep. Hind tarsal claw with strong

teeth which extend to base of tip. First abdominal segment with petiole approx.

1.5 x as long as postpetiole. Ovipositor approx. 2.8 x as long as hind femur.

Fore wing approx. 5.3 mm. Hind wing with nervellus weakly intercepted at

basal 0.3

Head, thorax, coxae, black. Tegula, basal segments of trochanters,

hind tibia, petiole, basal 0.5 of postpetiole, apical bands of tergites 4 to 6,

fuscous. Apical segments of fore and middle trochanters, fore and middle

tibiae, fore and middle femora, testaceous. Hind femur ferrugineous. Hind

tarsus white, with apices of segments infuscated.

Range and Variation. Known only from holotype.

Male. Unknown

Holotype. Female. Golden Pond Kentucky, V-26-VI-10-1964, S. G. Breeland.

Deposited in Townes collection, Gainesville, Florida.

Etymology. This species is named in honour of its collector, S. G. Breeland.

Remarks. This species is closely related to vancouverensis , differing prim-

arily in features of the propodeum ( areola longer than wide, absence of

lateral longitudinal carina) and in the much smaller punctures of the head and

thorax.

Macdunnoughi Group

Antennal scape with inner margin approx. 1.7-1.9 x as long as outer

margin. Vertex short, with length behind lateral ocelli approx. 0.5-0.8 x as

long as ocellar triangle. Temples sloping inwardly in dorsal view. Clypeus

with anter-lateral corners extending beyond tentorial pits, its anterior mar-

gin not produced medially. Mandible long, weakly tapering, with lower tooth

approx. 1.1-1.8 x as long asupper tooth. Ovipositor approx. 2.8-3.0 x as long

as hind femur. Face of females entirely black or with white spot under each

antennal socket. Face and clypeus of males ( males known for macdunnoughi

only) entirely yellow. Thorax of nearctic males black with yellow markings.

Abdomen of both sexes entirely black.

This is a holarctic group with one nearctic and one palearctic species

38 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

(macdunnoughi, bimaculata, respectively)

Xylophylax macdunnoughi (Viereck), n. comb.

Figs. 138, 14

Pyracmon macdunnoughi Viereck, 1925; 1926. Can. Ent. 57(7): 180 (key); 58

(1); 5. Female. Nordegg, Alberta (CNC). Examined in 1984.

Campoplex (Ecphora) alternatus Viereck, 192531926. Can. Ent. 57(12): 202

(key); 58( ): 282. Female. Seven Isles, Quebec (CNC). Examined in

Clypeus with antero-lateral corners extending beyond tentorial pits.

Mandible with lower tooth approx. 1.1 x as long as upper tooth. Temples

moderately sloping inwardly in dorsal view. Abdomen entirely black in both

sexes. Faceof male entirely yellow. Thorax of male black with yellow mark-

ings.

Female. Antennal scape with inner margin approx. 1.6-1.8 x as long as outer

margin. Antenna with 30-32 articles. Cmpd. eyes with inner margins moder-

ately diverging ventrad. Face shagreened, matte, with medium sized punct-

ures which are separated by distance less than dia. of single puncture. Cly-

peus polished, with antero-lateral corners extending beyond tentorial pits,

its anterior margin not produced medially. Gena approx. 1.0-1.2 x as wide as

basal width of mandible. Temples moderately sloping inwardly in dorsal view.

Mandible long weakly tapering, with lower tooth approx. 1.1 x as long as up-

per tooth. Mesopleuron with speculum shagreened, polished, impunctate.

Lower 0.5 of mesopleuron shagreened, matte, with medium sized punctures

which are separated by distance less than dia. of single puncture. Propodeal

carinae very strong. Areola approx. as wide as long, rarely separated from

petiolar area by transverse carina. Hind femur approx. 4.3-4.8 x as long as

deep. Hind tarsal claw with weak teeth which do not extend to base of tip.

First abdominal segment with petiole approx. 1.4-1.5 x as long as postpetiole.

Ovipositor approx. 2.1-2.8 x as long as hind femur (very rarely less than

2.5x). Fore wing approx. 6.0-7.5 mm. long. Hind wing with nervellus inter-

cepted near basal 0.3.

Head, thorax, abdomen, black. Palpi, tegula, fuscous. Coxae, basal

segments of trochanters, fuscous to black. Apical segments of trochanters,

fore and middle femora, fore tibia, ferrugineous. Hind femur ferrugineous,

with apical infuscation. Middle tibia ferrugineous, with apical infuscation.

Hind tibia and tarsus fuscous.

Male. Black. Scape, pedicel, face, lower temple, palpi, tegula, subtegular

ridge, anterior margin of mesopleuron, anterior margin of mesosternum,

fore and middle coxae, ventral surface of hind coxa, fore and middle trochan-

ters, apical segment of hind trochanter, yellow. Legs paler than in female

with hind tibia exhibiting a weak banding pattern, the basal and apical bands

fuscous, median band ferrugineous. In some males the mesosternum is

entirely yellow.

Material Examined. 3 males, 53 females

Sanborne: Revision of Nearctic Xylophylax (Ichneumonidae) 39

Distribution. CANADA. Alberta: Banff; Edmonton. British Columbia: Atlin;

Racing River; Stone Mt. Pk. Labrador: Cartwright; Goose Bay: Northwest

Territories: Norman Wells. Quebec: Knob Lake; Lac Huerd; Laniel. Yukon:

Dawson; La Force Lake; mi. 87, Dempster Hwy. UNITED STATES. Alaska:

Delta Junction; Fairbanks; Nehana; Unalakleet. Colorado: Gould. MEXICO.

El Salto, Durango.

This species is distributed throughout the boreal forest region of Alaska

and Canada, and the Rocky Mts. from Alaska into Mexico.

Host and Biology. There are no rearing records for this species but it has

been associated with black spruce. Adults are active from May to September

with a peak abundance in July.

Remarks. This species is closely related to bimaculata from Japan. It can be

distinguished from all North American species of the subgenus Euraspidion

in having the abdomen of both sexes entirely black. It is also unique in having

the lower tooth of the mandible only slightly longer than the upper tooth.

macdunnoughi

f 500

40 Contrib. Amer. Ent. Inst., vol. 22, no. 8, 1986

ACKNOWLEDGEMENTS

This project was funded in part by an NSERC (Natural Sciences and

Engineering Research Council of Canada) operating grant held by Dr. S. B.

Peck of the Department of Biology, Carleton University, Ottawa. Additional

funding came from the NSERC post-doctoral fellowship program. Their

support is gratefully acknowledged. Luis Ling of Carleton Univ. was res~-

ponsible for producing the S. E. M. photomicrographs.

REFERENCES

Ashmead, W. H. 1902. Papers from the Harriman Alaska expedition. 28.

Hymenoptera. 4: 117-274

Barron, J. R., G. S. Walley. 1983. A revision of the holarctic genus Pyrac-

mon (Hymenoptera: Ichneumonidae). Can. Ent. 115: 227-241.

Davis G. C. 1897. Descriptions of new species of Trigonalidae, Stephanidae,

and Ichneumonidae. Amer. Ent. Soc., Trans. 24: 349-372.

Harrington, W. H. 1894. Canadian Hymenoptera, No. 6. Can. Ent. 26: 245-

250.

Horstmann, K. 1980. Revision der europaischen Arten der Gattung Rhimph-

octona Forster (Hymenoptera: Ichneumonidae). Nach. Bayer. Ent. 29

(2): 17-24. |

Provancher, Abbe L. 1874. Les ichneumonides de Québec. Nat. Can. 6:143-

179.

Rohwer, S. A. 1914. Descriptions of new parasitic hymenoptera. Ent. Soc.

Wash. 15:180-188.

Sanborne, Michael. 1984. A revision of the world species of Sinophorus

(Ichneumonidae). Mem. Amer. Ent. Inst., no. 38. pp. 1-403.

Sanborne, Michael. 1986a. On the correct application of Rhimphoctona

Foerster (Hymenoptera: Ichneumonidae). Contrib. Amer. Ent. Inst.

vol. 22 (6): 1-6

ibid. .1986b. The subgenera of Xylophylax Kriechbaumer

(Hymenoptera: Ichneumonidae), with descriptions of two new eastern

palearctic species. Contrib. Amer. Ent. Inst., vol. 22 (7): 1-4

Townes, H. K. 1970. Genera of Ichneumonidae, Part 3. Mem. Amer. Ent.

Inst., 18. 307 pp.

Viereck, H.L . 1925. A preliminary revision of the Campopleginae in the

Canadian National Collection. Can. Ent. 57(7): 176-181.

ibid. 1925. A preliminary revision of the Campopleginae in the

Canadian National Collection. Can. Ent. 57(12): 296-303.

ibid. 1926. A preliminary revision of the Campopleginae in the

Canadian National Collection. Can. Ent. 58(1): 2-8.

Addenda

Kriechbaumer, J. 1878. Bitte an die Ichneumonologen. Ent. Nachr. 4: 210-

211.

Xylophylax spp.: Figs. 1-5, frontal view of head: 1, macrocephala; 2, aldrichi; 3, separata;

4, alaskensis; 5, longicauda.

Xylophylax spp.: Figs. 6-8, frontal view of head: 1,

montana; 2, albicoxa; 3,

genali Ss.

Xylophylax spp.: Figs. 9-13, frontal view of head: 9, ruficincta; 10, vancouverensis; 11,

breelandi; 12, brevicauda; 18, macdunnoughi. Fig. 14, propodeum: 14, macdunnoughi.

Xylophylax spp.: Figs. 15-20, propodea: 15, macrocephala; 16, separata; 17, aldrichi; 18,

alaskensis; 19, longicauda; 20, montana.

Xylophylax spp.: Figs. 21-26, propodea: 21, genalis; 22, albicoxa; 23, ruficincta; 24,

vancouverensis; 25, breelandi; 26, brevicauda.

A REVISION OF THE NEARCTIC SPECIES OF CYMODUSA HOLMGREN

(HYMENOPTERA: ICHNEUMONIDAE)

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, K1S 5B6

Abstract

The nearctic species of Cymodusa Holmgren (Hymenoptera: Ichneumon-

idae are revised. Five species are re-described: cruentata (Gravenhorst);

distincta (Cresson); melanocera Viereck; nigripes (Viereck); partis (Viereck).

Four species and one subspecies are new to science: columbiensis; montana ;

nicolei; ruficincta; partis ocularis. One new synonym was created: provancheri

Viereck = distincta (Cresson).

Introduction

The genus Cymodusa is widely distributed throughout the holarctic.

oriental, and neotropical regions. The palearctic species are being revised

by Dbar (Leningrad). The paper which follows represents the first revision

of the nearctic species and includes nine species in two species-groups. Four

species and one subspecies are new to science. There is a key to the species-

groups and species from the nearctic region along with illustrations (S.E.M.

photomicrographs) and distribution maps.

The biology of Cymodusa is essentially unknown and many of the world's

species remain undescribed and, for these reasons, no attempt has been

made to produce a phylogeny or provide a zoogeographical analysis.

Cymodusa Holmgren

Cymodusa Holmgren, 1859 (1858). Svenska Vetensk-Akad. Ofvers. af.....

Forhandl. 15: 325.

Type-species: Cymodusa leucocera Holmgren, by original desig.

Thersitia Schmiedknecht, 1907. Hym. Mitteleuropas: 598.

Type-species: Thersitia egregia Schmiedknecht ( = C. leucocera Holm-

gren), by monotypy.

Fore wing 3.0-5.5 mm. long. Body elongate. Eye moderately indented

opposite antennal socket. Cmpd. eyes distinctly hairy, with inner margins

weakly convergent ventrad in males, strongly convergent ventrad in females.

Lower facets of cmpd. eyes of females 1.4-1.8 x as wide as upper facets, of

equal size in males. Base of female flagellum of some species white on inner

side. Tentorial pits (= clypeal foveae) separated from inner margins of eyes

2 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

by distance less than or equal to dia. of single pit in females, or sometimes

touching the inner margin. Clypeus moderately convex, with anterior margin

evenly, moderately convex. Mandible short, its lower edge with a high car-

ina or narrow flange, with teeth approx. equal length. Temple short. Genal

carina joining oral carina. Lower 0.5 of mesopleuron matte, with fine to

medium sized punctures and usually with distinct rugosities. Propodeum

moderately long to long, with apex projecting between bases of hind coxae a

distance equal to 1.2-1.4 x the length of hind coxa. Areola usually confluent

with petiolar area, sometimes partially or completely separated by incom-

plete or complete transverse carina. Propodeal spiracle round. Hind basi-

tarsus without a median ventral row of closely spaced hairs. Tarsal claws

pectinate only at base. Areolet of fore wing closed (except nicolei). Nervulus

opposite basal vein or distad by as much as 0.2 x its length. Discoidella inter-

cepting nervellus, not pigmented. First abdominal segment moderately long,

slender, weakly to moderately decurved distad of spiracle, with petiole weak-

ly, evenly dorso-ventrally tapering from spiracle to base, its sides usually

smooth, polished. Glymma absent. Second abdominal tergite 1.3-3.0 x as

long as wide in females, 1.5-5.0 x in males. Ovipositor stout, compressed,

nearly straight (except nigripes where it is slender, moderately upcurving),

approx. 1.5-2.2 x as long as hind femur.

There are nine species known from the nearctic region. They occur

mostly in grassy meadows but there are no rearing records. The few rearing

records from the palearctic region indicates that the hosts are small lepid-

optera.

Key to the Species-Groups and Species of Nearctic Cymodusa

The males of ruficincta and columbiensis are unknown but are placed in

the key on the basis of characters they are presumed to have.

1. Cmpd. eyes strongly converging ventrad in females, with lower facets

approx. 1.4 x as wide as upper facets. Tentorial pit of females never

touching inner margin of eye. Second abdominal tergite approx. 1.3-

1.8 x as long as wide in females, 1.5-2.2 x in males. Antennal flag-

ellum of some females white on inner surface of basal articles..... F

Cc dN GRAS AAC NRE A RO OR Ge Gee Ces RD a ewloee wees ChbeNntaAta Group. 2.

Cmpd. eyes very strongly converging ventrad in females, with lower

facets approx. 1.8 x as wide as upper facets. Tentorial pit of females

always touching inner margin of eye. Second abdominal tergite approx.

2.0-3.0 x as long as wide in females, 2.5-5.0 x in males. Antennal

flagellum of females uniformly black........Melanocera Group. .15.

Be POMS OG sos eve a i ah ena bic BR chmactl es g SEES & cid acbaratlage Scar stants we bhee wees

PREIS rack gta hig Sl alee ey gta aU eat a ek nal eal iMate lees eee (glial a Reomneaie ye g Og

3. Antennal flagellum with basal article(s) white on inner surface.......4.

antenna) tiagel lum aniformly, DIRK. ecu euce oie dee eieverds kwreeemrers od doe Be

4. Ovipositor approx. 1.5-2.0 x as long as hind femur, stout, more or less

SUVA Lo yiviace se w ae'ale teats @eee72e8ee8ee@ @eeee0eeoe2u12eeee8e eee @ @e@eeeee7s#4e#e7e#e7#2e8ee¢e80 0 Be

10.

Ta.

13.

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 3

Ovipositor approx. 2.1-2.2 x as long as hind femur, slender, moderate-

ly upcurving. eastern N. America....... ---enigripes (Viereck)(p. 4)

Abdominal tergites 2-5 black basally, rufous apically. Hind coxa black,

with at most some ferrugineous at extreme apex. Hind femur exten-

sively infuscated to entirely black. Boreal forest, Rocky Mts.......

Pa ge i eemcontinagigs aie siti aia aK ai eck aleve clue a partis (Viereck) (p. 6).

Abdominal tergites not as above, with tergites 4,5 entirely rufous. Hind

coxa with apical 0.5-0.7 ferrugineous. Hind femur usually entirely

ferrugineous. B.C. 6G. #0 Ore... Cs, 10 Neo.. os tO Ellice ccarceese sss

Eh Sa dt cian mune. Laila nag Ceili fan e mallhe)eea so cob goa. a cue Gee Nee (Cresson) (p. 10)

Abdominal tergites with alternating banding pattern or entirely black..7.

Abdominal tergites 2 to apex entirely rufous. Pacific Northwest.........

Cece c cee c reece cence crecccecoescocsoesvoruticincta, New sp. (p.12)

Abdominal tergites 2-4 (sometimes only 2 and 3) black basally, rufous

218 SOC) en I RE ip THE SD SR NESE gta he y(n al Mme Amn a AR:

Abdominal tergites uniformly fuscous to black. B. C...ccrccccesoveces

<a. 00ie's 0 0 4 sps:s ele e wade 964 20/6 e000 oe < eee COLUMDICNSIS, DEW Sp. (p-14)

Hind femur ferrugineous, with a small baso-ventral infuscated area....

Holarctic (Boreal Forest, n. U.S.)....cruentata (Gravenhorst) (p.16)

Hind femur more extensively infuscated than above to entirely black....

Boreal Forest, Rocky Mts......++-.+-.+++... partis (Viereck) (p. 6)

Abdominal tergites 4,5 entirely PU OME Ree et aoe hie es giae a glia ae Ug catego aie heb

Abdominal tergites 4, 5 black and rufous banded or entirely black... .12.

Abdominal tergites 2 to apex entirely rufous (apical tergites sometimes

intuecated dorsally).. Hind coxa DIAck,.s s< 00s p06 oe aetegecmecett »

Abdominal tergites 2 to apex never entirely rufous. Hind coxa with

apical 0.3-0.5 ferrugineous..............- distincta (Cresson) (p.10)

Hind femur ferrugineous. Pacific Northwest...ruficincta, new sp. (p.12)

Hind femur extensively infuscated to entirely black. eastern N. America

seer eccerorcccccccccccceccccccccccscves lliPripes ( Viereck) (p. 4)

Abdominal tergites 2-4 black basally (sometimes only 2,3), rufous

OE itt, a go Cebit es eee a eee a kL. eee de Ne

Abdominal tergites entirely fuscous to black... .cccccccccsscccccccelt.

Hind femur extensively infuscated to entirely black. Boreal Forest.....

ROCKY MGB. ele tiew Gk Corecesscosscvececcecss partis (Viereck) (p. 6)

Hind femur ferrugineous, with a small baso-ventral infuscated area....

4 Contrib. Amer. Ent. Inst., vol. 22, no, 9, 1986

Holarctic (Boreal Forest, n. U. S.)..cruentata (Gravenhorst) (p.16)

14. Hind femur ferrugineous to weakly infuscated. B.C......ccccccccccces

liek Wises als oa ca a Nicks ah ea or eereseeeese-eeeColumbiensis, new sp. (p.14)

Hind femur extensively to entirely black. Colorado....... hr ae Ge Ne “*

@e0eoee#e Cees er ee rerccroressrerecercccscceee partis (Viereck) (p. 6 )

1s, LOMAS. sa s-a6% 4:0 tie ike aes 6g bik wy Ar oo a i dhs ey how aca ivi ee eae eer iM

TAG ae ln oes «ain @e@eee¢8#8ee«ee¢ @esevees#eesecstcee@eeese0e58seseen#ee#e#*eescsrsteeesvrmeenvesv es @ Pegenssamd, |e

16. Fore wing with areolet closed. Second abdominal tergite 1.6-2.4 x as

OTA Pe SSC aia os de hs a ev ge. bes eg ak as Sas 598 A BLN Seb oehich a: aae eeehis

Fore wing with areolet open. Second abdominal tergite approx. 3.0 x as

long as wide. N. and S. Carolina, Tenn.......nicolei, new sp. (p- 22)

17. Antennal flagellum with basal article approx. 5.0-6.0 x as long as wide.

Ovipositor approx. 1.7-1.9 x as long as hind femur. Widespread in

North, Middle, South America...........melanocera Viereck (p.18 )

Antennal flagellum with basal article approx. 4.7-5.0 x as long as wide.

Ovipositor approx. 1.5-1.7 x as long ashind femur. Texas, Mexico,

POUR spk whee see Oe ed hes Ouse d oe aae se RICANS, new sp. (p.19)

18. Fore wing with areolet closed. Second abdominal tergite approx. 2.4-

Bel MOS [ONT BS WIK0, ss. ewes ee sense Cee Rak ein SU? FLEE oo pale

Fore wing with areolet open. Second abdominal tergite approx. 5.0 xas

long as wide. Tenn., N. and S. Carolina......nicolei, new sp. (p.22)

19. Basal article of antennal flagellum approx. 3.0-3.5 x as long as wide.

Second abdominal tergite approx. 3.0-3.1 x as long as wide. Texas,

Mexico, PVN Vela.a 5k os Vibes ese e 60 8 aon oa ee 42 OA, new Sp. (p -19)

Basal article of flagellum approx. 3.5-4.0 x as long as wide. Second

abdominal tergite approx. 2.4-2.5 x as long as wide. Widespread in

North, Middle, South America...........melanocera Viereck (p.18)

Cruentata Group

The salient features of this group are given in the first couplet of the

key above.

Cymodusa nigripes (Viereck)

Figs. 1, 3

Neonortonia nigripes Viereck, 1925. Can. Ent. 57: 179 (key); 57: 300. Male.

des. Holotype. Grey Mills, New Brunswick (CNC). Examined in 1984.

Neonortonia laevissima Viereck, 1925. Can. Ent. 57: 179 (key); 57: 301. Male.

des. Holotype. Annapolis, Nova Scotia (CNC). Examined in 1984.

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 5

Cmpd. eyes strongly covergent ventrad in females, distinctly hairy.

Fore wing with areolet closed. Ovipositor approx. 2.1-2.2 x as long as hind

femur, slender, moderately upcurving. Antennal flagellum of female with

inner surface of basal articles white. Mandible often entirely black. Hind

coxa entirely black. Abdominal tergites 3 to apex rufous.

Female. Antennal flagellum with 28-33 articles, the basal article approx.

3.6-4.0 x as long as wide. Antennal scape with inner margin approx. 1.4 x

as long as outer margin. Cmpd. eyes strongly covergent ventrad. distinctly

hairy, with lower facets approx. 1.4 x as wide as upper facets. Face strongly

shagreened, matte, with minute, close-set punctures. Tentorial pit separated

from inner margin of eye by distance less than or equal to dia. of pit, never

touching inner margin. Clypeus strongly convex, with anterior margin mod-

erately convex. Lower 0.5 of mesopleuron moderately to coarsely rugoso-

punctate. Propodeum moderately long, with apex projecting between bases of

hind coxae approx. 0.3 x the length of hind coxa. Propodeal carinae very

strong, complete. Areola often separated from petiolar area by transverse

carina. Petiolar area with coarse reticulate pattern of wrinkling. Hind femur

approx. 5.2-5.5 x as long as deep. Hind tarsal claw with tip strongly bent,

the shank straight. Petiolar segment of abdomen moderately decurving. Pet-

iole approx. 1.5-1.6 x as long as postpetiole, with sides smooth, polished.

Second abdominal tergite approx. 1.6-1.7 x as long as wide, with thyridium

positioned at basal 0.2. Ovipositor approx. 2.1-2.2 x as long as hind femur,

slender, moderately upcurving, its depth at mid length distinctly less than

width of hind basitarsus. Fore wing approx. 4.5-6.5 mm. long. Hind wing

with nervellus intercepted.

Antennal flagellum with 3-7 basal articles white on inner surface. Head,

thorax petiolar segment of abdomen, basal 0.6-0.7 of 2nd tergite, coxae,

basal segments of all trochanters, bases of fore and middle femora, entire

hind femur, black. Mandible extensively to entirely black. Palpi fulvous.

Tegula fuscous to black. Apical segments of all trochanters white. Apices of

fore and middle femora ferrugineous to fuscous. Fore tibia ferrugineous;

middle tibia ferrugineous to fuscous; hind tibia black basally and apically,

ferrugineous medially. Hind tarsus fuscous. Apical 0.3-0.4 of 2nd abdomin-

al tergite, abdominal tergites 3 to apex, rufous. In some specimens the hind

femur is paler than described and abdominal tergites 4 to apex are sometimes

weakly infuscated dorsally.

Male. Hind femur approx. 4.4-4.8 x as long as deep. Palpi white. Fore and

middle trochanters mostly white. Fore and middle femora ferrugineous, hind

femur ferrugineous with apical black band and black baso-ventral patch. Fore

and middle tibiae white externally, ferrugineous internally, hind tibia with

narrow basal and apical dark bands, the median band dark ferrugineous. Ab-

domen coloured as in female.

Material Examined. 55 males, 46 females

Distribution. CANADA. British Columbia: Agassiz. New Brunswick: Grey

Mills. Ontario: Belleville; Constance Bay; Gloucester; One Sided Lake; Stitts-

ville. Quebec: Chambly Co.; Lac Mondor; Mt. Albert; Ste. Therese.

Host and Biology. There are no rearing records for this species. Adults are

active from late August through November, with a peak abundance in October.

Its habitat appears to be grassy meadows.

6 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Remarks. This species appears to be very closely related to ruficincta

from the pacific northwest. Females of nigripes are distinguished by the

much longer ovipositor, white inner surface of basal articles of antennal flag-

ellum, and distinctly more melanic legs.

nigripes

500

Cymodusa partis (Viereck)

Figs. 5, 6

Campoplex (Diadegma) partis Viereck, 1925, 1926. Can. Ent. 57(9): 227 (key)3

58(11): 280. Male. des. Holotype: Aylmer, Quebec (CNC). Examined in

1984.

Cmpd. eyes strongly covergent ventrad in females, distinctly hairy.

Fore wing with areolet closed. Ovipositor approx. 1.61.8 x as long as hind

femur. Antennal flagellum entirely black or often with inner surface of basal

articles white (females). Hind coxa black. Hind femur extensively to entirely

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 7

black, rarely ferrugineous. Abdominal tergites 2 to apex with alternating

black and rufous bands (p. partis) to entirely black (many p. ocularis).

Female. Antennal flagellum with 25-28 articles, with basal article approx.

3.4-4.0 x as long as wide. Antennal scape with inner margin approx. 1.4-1.5

x as long as outer margin. Cmpd. eyes strongly convergent ventrad, distinctly

‘hairy, with lower facets approx. 1.4 x as wide as upper facets. Face strongly

shagreened, matte, with minute punctures. Tentorial pit separated from in-

ner margin of eye by distance less than or equal to dia. of pit, never touch-

ing inner margin. Clypeus moderately convex, with anterior margin moderate-

ly convex. Lower 0.5 of mesopleuron moderately to coarsely rugoso-punctate.

Propodeum moderately long, with apex projecting between bases of hind coxae

approx. 0.2-0.3 x the length of hind coxa. Propodeal carinae weak to moder-

ately strong, the costula weak or absent. Areola completely confluent with

petiolar area. Petiolar area with moderately coarse reticulate wrinkling.

Hind femur approx. 5.0-6.0 x as long as deep. Hind tarsal claw straight, with

tip moderately to strongly bent. Petiolar segment of abdomen moderately de-

curving. Petiole approx. 1.2-1.3 x as long as postpetiole. Second abdominal

tergite approx. 1.4-1.8 x as long as wide, with thyridium at basal 0.2. Ovi-

positor approx. 1.5-1.8 x as long as hind femur, stout, with depth at mid

length equal to or greater than width of hind femur. Fore wing 3.2-5.0 mm.

long. Hind wing with nervellus intercepted.

Antennal flagellum uniformly black or with inner margins of basal art-

icles white. Head, thorax, petiolar segment of abdomen, bases of abdominal

tergites 2-4 or sometimes to apex or entire abdomen, hind coxa, black. Pal-

pi fulvous. Tegula white to fuscous. Fore coxa black with apex ferrugineous;

middle coxa coloured as fore coxa or entirely black. Fore trochanter varying

from yellow to ferrugineous to black and ferrugineous to black and fuscous;

middle trochanter coloured as fore; hind trochanter with basal segment fus-

cous to black, apical segment yellow to fuscous. Ground colour of femora

and tibiae ferrugineous, with highly variable degree of infuscation, some-

times hind femur entirely black. Hind tarsus fuscous.

Male. Coloured as in female.

There are two subspecies which can be distinguished in the following key.

Cmpd. eye approx. 1.6-1.7 x as wide as long. Antennal flagellum often

with inner surface of basal articles white (females). Abdominal ter-

gites 2-4 or sometimes to apex with alternating black and rufous

bands. Widely distributed........ .-.. partis partis (Viereck) (p. )

Cmpd. eye approx. 1.8-2.0 x as wide as long. Antennal flagellum of fe-

males entirely black. Abdominal tergites 2-4 with black and rufous

bands, the apex black, or abdomen entirely black. Colorado........

PL ee Pore oo ee ee CN ee -»-partis ocularis, new subsp. (p. )

Cymodusa partis partis (Viereck)

Fig. 5

Campoplex (Diadegma) partis Viereck, 1925, 1926. Can. Ent. 57(9): 227 (key);

08(11): 280. Male. des. Holotype: Aylmer, Quebec (CNC).

8 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

The characters given in the key are sufficient to describe this sub-

species.

Material Examined. 62 males, 96 females

Distribution. CANADA, Alberta: Banff; Elkwater; Kananaskis; Lethbridge;

Norton; onefour; Orion. British Columbia: Oliver; Osoyoos; Sawmill Lake;

Stone Mt. Lake; Summit Lake; Terrace; Vancouver; West Bank. Manitoba:

Carberry; Reynolds. New Brunswick: Jacquet R. Newfoundland: Port au

Basques; St. Vincents; South Branch. Northwest Territories: Pangnirtung.

Ontario: Hamilton; Marmora; One Sided Lake; Ottawa; Stittsville; Thunder

Bay. Quebec: Lac Mondor; La Verendreye Pk.; Val Marie. Yukon Territory:

North Fork Crossing. UNITED STATES, Alaska: Anchorage; Tok. California:

Beatrice; San Bruno. Colorado: Boulder; Steamboat Springs. District of

Columbia: Washington. Idaho: Galena Summit; Lowman; Mt. Pleasant; Stanley.

Massachusetts: Fitchburg; Reading Highlands. Michigan: Iron River; Yellow

Dog Plains. Minnesota: Big Fork. Montana: Bozeman; Glacier Nat. Pk.

Nevada: Tuscarora. New Hampshire: Hanover. New York: Bemus Pt.; Oswego.

Ohio: Columbus. Oregon: Calapooya; Cascadia; Ochoco Cr.; Selma; Sparks

Lk.; Three Cr. Lake; Three Sisters. Pennsylvania: Cookshire. Vermont:

Lake Willoughby. Virginia: Spruce Knob. Washington: Orens Is. West Virgin-

ia: Dolly Sodo Wild. Area.

Host and Biology. There are no rearing records for this subspecies. Adults

are active from May through August with a peak abundance in July. It occurs

in grassy meadows.

Remarks.There are two colour forms with regards to the basal articles of

the antennal flagellum of females of this subspecies. One form has the inner

surface of three to seven articles coloured white. It occurs primarily east

of the Manitoba-Ontario border in Canada, and east of the Mississippi R. in

the United States but is also widely scattered in western North America

where it is rare (see map below). The second form has the antennal flagellum

entirely black and occurs primarily in western North America.

Cymodusa partis ocularis, new subsp.

The characters given in the key above are sufficient to describe this

subspecies.

Holotype. Female. Summit Lake, Mt. Evans, 12, 800', Colorado, July 16,

1961, S. M. Clark. Deposited in Canadian National Insect Collection.

Paratypes. Colorado: nr. Estes Pk., June 15, 1948 (1 female), H.,M.,G.,D.,

J. Townes (HMT); Independence Pass, Lake Co., 12,100', VII-31-1961 (1 female)

B. H. Poole (CNC); Loveland Pass, 12,000', VIII-7-1961 (1 female), B. H.

Poole (CNC); Mt. Evans, VII-27-VIII-4-1961 (7 males, 3 females), 12, 000-

14,000', S. M. Clark, J. G. Chillcott, W. R. M. Mason, B. H. Poole, J. R.

Stainer (CNC); Summit Lake, Mt. Evans, 12,800', Vii-16-17-1961, (2 males,

1 female), S. M. Clark and C. H. Mann (CNC) and July 18, 1964, 12, 700'

(2 males, 2 females), C. Dasch (CDC).

Etymology. The species epithet draws attention to the modified cmpd. eyes.

Host and Biology. There are no rearing records for this subspecies. All

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae)

specimens have been taken on alpine tundra at altitudes of 12, 000-14, 000

feet during July and August.

Remarks. The short, wide eyes and slightly produced mouthparts will

distinguish this subspecies from p. partis.

p. partis “™

(white antenna) ¢°><-

p- partis @

(black antenna

p- oculariso :

10 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Cymodusa distincta (Cresson)

Fig. 4

Mesoleptus subrubidus Cresson, 1864. Proc. Ent. Soc. Phila., 3: 264. Male.

des. Holotype: Illinois (ANSP). Examined in 1983.

Limnerium vigilis Viereck, 1906. Trans. Amer. Ent. Soc., 32: 179. Male.

des. Holotype: Lawrence, Kansas (KSM). Examined in 1983.

Idechthis biconjunctus Viereck, 1906. Trans. Amer. Ent. Soc., 32:180. Male.

des. Holotype: Lawrence, Kansas (KSM). Examined in 1983.

Campoplex (Nepiera) amassecontorum Viereck, 1917. Bull. Conn. Geol. Nat.

Hist. Surv., 22: 266. Male. des. Holotype: New Haven Connecticut.

Examined in 1983.

Cymodusa plesius Viereck, 1925,1926. Can. Ent. 57(7): 180 (key); 58(1): 4.

Female. des. Holotype: Oliver, B. C. (CNC). Examined in 1984.

Cymodusa provancheri Viereck. 1925,1926. Can. Ent. 57(7):180 (key); 58(1):

4. Female. des. Holotype: Kentville, Nova Scotia (CNC). New synonomy.

Examined in 1984.

Cmpd. eyes strongly convergent ventrad in females, distinctly hairy.

Fore wing with areolet closed. Ovipositor approx. 1.6-1.7 x as long as hind

femur, stout, more or less straight. Antennal flagellum of female with inner

surface of basal articles white. Hind coxa black with apical 0.5-0.7 ferrug-

ineous. Abdominal tergites without regular banding pattern, tergites 3-5

usually entirely rufous.

Female. Antennal flagellum with 26-33 articles, the basal article approx.

4.3-4.4x as long as wide. Antennal scape with inner margin approx. 1.4 x

as long as outer margin. Cmpd. eyes strongly covergent ventrad, distinctly

hairy, with lower facets approx. 1.5 x as wide as upper facets. Face strongly

shagreened, matte, with minute, close-set punctures. Tentorial pit separated

from inner margin of eye by distance less than or equal to dia. of pit. Clypeus

strongly convex, its anterior margin moderately convex. Lower 0.5 of meso-

pleuron strongly shagreened, with small to medium sized close-set punctures.

its surface weakly to strongly rugose. Propodeum moderately long, with apex

projecting between bases of hind coxae 0.3-0.4 x the length of hind coxa. Pro-

podeal carinae moderately strong, complete. Areola sometimes partially to

completely separated from petiolar area by transverse carina. Petiolar area

granulose, sometimes with weak wrinkling. Hind femur approx. 5.5-6.0x

as long as deep. Hind tarsal claw straight, with tip strongly bent. Petiolar

segment of abdomen weakly decurving, with sides smooth, polished. Petiole

approx. 1.4-1.5 x as long as postpetiole. Second abdominal segment approx.

1.6-1.8 x as long as wide, with thyridium at basal 0.2. Ovipositor approx.

1.6-1.7 x as long as hind femur, more or less straight, with depth at mid

length equal to or slightly greater than width of hind basitarsus. Fore wing

approx. 3.2-5.5 mm. long. Hind wing with nervellus intercepted.

Antennal flagellum with 3-7 basal articles white on inner surface or,

very rarely, the flagellum entirely black. Head, thorax, abdominal petiole,

basal 0.5-1.0 of postpetiole, bases of abdominal tergites 1,2, basal 0.3-0.5

of hind coxa, black. Palpi white to fulvous. Tegula white to ferrugineous.

Fore and middle coxae extensively to entirely pale ferrugineous; hind coxa

with apical 0.5-0.7 ferrugineous. Fore and middle trochanters pale yellow;

hind trochanter with basal segment ferrugineous to fuscous, apical segment

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 1

pale yellow. Femora ferrugineous. Fore and middle tibiae ferrugineous in-

ternally, whitish externally; hind tibia ferrugineous, with weak sub-basal

and apical fuscous bands. Hind tarsus fuscous. Abdominal tergites 2,3 with

apices rufous; tergites 4, 5 entirely rufous; tergite 6 to apex partially infusc-

ated, sometimes entirely rufous.

Male. Hind femur approx. 4.5-5.0 x as long as deep. Coloured as in female

except that antennal flagellum always black and there is a tendency for the

apical tergites to be darker .

Material Examined. 988 males, 1035 females

Distribution. CANADA. Alberta: Banff; Edmonton. British Columbia: Douglas;

North Vancouver; Okanogan Falls; Oliver; Selkirk Mts.; Shennon Falls;‘ Terr-

ace. New Brunswick: Grey Mills. Newfoundland: Port au Basques; Raleigh;

South Bend. Nova Scotia: Bridgewater; Truro. Ontario: Almonte; Ancaster;

Ariss; Belleville; Bell's Corners; Brighton; Chalk River; Chatterton; Christie

Lake; Constance Bay; Elmira; Finland; Fort Erie; Guelph; Hamilton; Innisville;

Jockvale; Jordan; London; Madoc; Marmora; Merivale; Niagara Falls; Norman-

dale; Ottawa; Rockport; Shannonville; St. Davids; St. Lawrence Nat. Pk;

Southhampton; South March; Stittsville; Strathroy; Thamesville; Tobermory;

Tweed; Waubamie. Prince Edward Island: Alberton; Brackley Beach Nat. Pk.;

Dalvay HouseNat. Pk. Quebec: Aylmer; Breckenridge; Kirk's Ferry; Montreal;

Ste. Flore. Saskatchewan: Elbow. UNITED STATES, California: Bear Valley;

Laguna Beach. Connecticut: Canterbury; Green Falls; Hamden; Ledyard; Lyme;

Sterling; Stonington; Voluntoown. District of Columbia: Washington. Florida:

Gainesville. Georgia: Atlanta; Hiawassee. Illinois: Urbana. Indiana: Tippe -

canoe Co.lowa: Ames; Mt. Pleasant. Kansas: Lawrence. Louisiana: Opel-

ousas; Shreveport. Maine: Casco; Dryden; Mt. Desert Is. Maryland: Bowie;

Colesville; Cumberland; Glen Echo; Laurel; Silver Springs; Takoma Pk;

Wheaton. Massachusetts: Chicopee; South Natick; Wellesely. Michigan: Ann

Arbor; Detroit; Iron Co.; Nahma. Missouri: St. Leuis. New Hampshire:

Crawford Notch; Hampton; Mt. Washington; Pinkham Notch; Randolph; Stin-

son Lake. New Jersey: Cape May; Kancocas; Lakehurst; Moorestown. New

York: Bemus Pt.; Blackbrook; Blasdell; Brainard; Buffalo; Colden; East

Aurora; Fall Cr.; Farmingdale; Hamburg; Hancock; Hunter; Ithaca; McLean

Bog; Medina; Middleton; Napeague; Oswego; Pelkham Bay; Poughkeepsie;

Remsen; Rome; Sea Clift; Shokan; Slaterville; Spencer Lake; Spring Lake;

Yankee Lake . North Carolina: Asheville; Cedar Mt.; Crabtree Meadows;

Elon College; Flat Rock; Franklin; Hendersonville; Highlands; Mt. Mitchell;

West Jefferson. Ohio: Cleveland; Columbus; New Concord; Otsego; Saline-

ville; Seneca Lake; Steubenville; West Lancaster; Willoughby; Wooster.

Oregon: Corvallis. Pennsylvania: Bartonsville; Castle Rock; Ft. Loudon;

Gaines; Spring Brook; Stillwater. Rhode Island; Block Is.; Hopington; Kingston;

Westerly. South Carolina: Cleveland; Greenville. Tennessee: Knoxville;

White House. Vermont: Woodstock. Virginia: Black Pond; Falls Church;

Galax; Glencarlyn; Great Falls; Mountain Lake; Rocky Knob; Shenandoah Nat.

Pk.; Skyline Dr.; Vienna. West Virginia: Bowden; Spruce Knob.

Host and Biology. This is one of the commonest of species in the nearctic

region yet there are no rearing records. It occurs in grassy meadows. Adults

are active from May through September and there appears to be two generat-

ions in many areas.

12 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Remarks. This species is easily distinguished from the similar partis partis

and cruentata by the lack of a regular banding pattern on the abdomen. I have

very carefully examined the type specimen of provancheri and compared it

to specimens of distincta. They are identical except for the fact that the

antennal flagellum of provancheriis entirely black. Since then, a number of

other specimens have shown up with similarly dark antennae and no other

discernable differences. I have concluded that distincta and provancheri_ are

conspecific and that this is the identical situation found in partis where the

flagellum also displays two colour forms. Therefore, provancheri becomes

a junior synonym of distincta .

distincta

\ 500

Cymodusa ruficincta, new sp.

Fig. 8

Cmpd. eyes strongly covergent ventrad in females, distinctly hairy.

Antennal flagellum uniformly black in both sexes. Fore wing with areolet

closed. Ovipositor approx. 1.4-1.6 x as long as hind femur, stout, straight.

Abdominal tergites 2 to apex entirely rufous.

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 13

HOLOTYPE. Female. Antennal flagellum with 29 articles. Antennal scape

with inner margin approx. 1.3 x as long as outer margin. Basal article of

flagellum approx. 4.3 x as long as wide. Cmpd. eyes strongly convergent

ventrad, distinctly hairy, with lower facets approx. 1.3 x as wide as upper

facets. Face strongly shagreened, matte, with small punctures which are

separated by distance less than dia. of single puncture. Tentorial pit separ-

ated from inner margin of eye by distance slightly less than dia. of pit. Cly-

peus moderately convex, its anterior margin moderately convex. Lower 0.5

of mesopleuron strongly shagreened, matte, rugolose, with punctures separ-

ated by distance less than dia. of single puncture. Propodeum with apex only

slightly projecting between bases of hind coxae. Areola granulose, matte,

separated from petiolar area by weak transverse carina. Petiolar area shag-

reened, somewhat polished, with numerous, moderately strong, irregular

transverse wrinkles. Hind femur approx. 5.5 x as long as deep. Hind tarsal

claw weakly curving. Petiolar segment of abdomen moderately decurving,

with petiole approx. 1.5 x as long as postpetiole. Second abdominal tergite

approx. 1.3 x as long as wide, with thyridium at basal 0.2. Ovipositor approx.

1.6 x as long as hind femur, straight, with depth at mid length distinctly greater

than width of hind basitarsus. Fore wing approx. 4.5 mm. long. Hind wing

with nervellus intercepted near basal 0.2.

Antennal flagellum, head, thorax, abdominal petiole, basal 0.5 of post-

petiole, coxae (except apices), black. Palpi, tegula, fulvous. Apices of coxae,

fore and middle trochanters, apical 0.5 of basal segment and entire apical

segment of hind trochanter, femora, tibiae, hind tarsal segments 1-3, ferr-

ugineous. Basal 0.5 of basal segment of hind trochanter, tarsal segments 4,

5, fuscous. Apical 0.5 of postpetiole, tergites 2~apex rufous. Wings weakly

infuscate.

Range and Variation. Antennal scape with inner margin approx. 1.3-1.4 x as

long as outer margin. Lower 0.5 of mesopleuron shagreened, matte to dis-

tinctly rugolose. Propodeum with areola weakly separated from to completely

confluent with petiolar area. Hind femur approx. 5.4-5.7 x as long as deep.

Ovipositor approx. 1.4-1.6 x as long as hind femur. Fore wing approx. 4.5-

0-0 mm. long. No significant colour variation in specimens at hand.

Male. Unknown but probably resembling nigripes .

Holotype. Female. Corvallis, Oregon, Oct. 6, 1931, Collector unknown.

Deposited in United States National Museum (USNM).

Paratypes. California: Del Puerto Canyon, Stanislaus Co., X-4-1975 (1 fem-

ale), N. J. Smith (UCB); Dos Palos, Mercedes Co., VIII-31-1947 (1 female),

V. M. Stern (UCB).Oregon: Corvallis, Aug. 6, 1965 (1 female), C. Dasch

(CDC). Utah: Petersboro, July 4, 1958 (1 female), K. C. Tilley (USNM).

Host and Biology. There are no rearing records for this species. Adults are

active in late summer.

Etymology. The species epithet refers to the extensively rufous abdomen.

Remarks. This species appears to be closely related to nigripes but the

ovipositor of ruficincta is shorter and stouter and the antennal flagellum

of ruficincta females is entirely black.

14 Contrib. Amer. Ent. Inst. , vol. 22, no. 9, 1986

ruficincta

f 500

Cymodusa columbiensis, new sp.

Fig. 7

Cmpd. eyes strongly convergent ventrad in females, moderately hairy.

Antennal flagellum uniformly black in both sexes. Fore wing 3.5-3.7 mm.

long, with areolet closed. Abdominal tergites uniformly fuscous.

HOLOTYPE. Female. Antennal flagellum with 26 articles, the basal article

approx. 3.6 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes strongly convergent ventrad,

moderately hairy, with lower facets approx. 1.4 x as wide as upper facets.

Tentorial pit separated from inner margin of eye by distance less than dia .

of pit. Face strongly shagreened, matte, with minute punctures. Clypeus

strongly convex, with anterior margin moderately convex. Lower 0.5 of

mesopleuron strongly shagreened, matte, with small punctures which are

separated by distance less than or equal to dia. of single puncture. Propod-

eum moderately long, with apex projecting between bases of hind coxae approx.

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 15

0.3x the length of hind coxa . Areola completely confluent with petiolar area.

Hind femur approx. 5.4 x as long as deep. Hind tarsal claw straight, with

tip moderately bent. Petiolar segment of abdomen weakly decurving, with

petiole approx. 1.3 x as long as postpetiole. Second abdominal tergite approx.

1.9 x as long as wide, with thyridium positioned at basal 0.2. Ovipositor

approx. 1.8 x as long as hind femur, weakly upcurving, with depth at mid

length approx. equal to width of hind basitarsus. Fore wing approx. 3.5 mm.

long. Hind wing with nervellus intercepted near basal 0.2.

Antennal flagellum, head, thorax, black. Dorsal surface of fore coxa,

middle and hind coxae, basal segments of all trochanters, abdomen, fuscous.

Apical segments of trochanters, legs, pale ferrugineous. Tegula fulvous.

Range and Variation. Fore wing approx. 3.5-3.7 mm. long. No significant

morphological or chromatic variation apparent in other specimens at hand.

Male. Unknown but abdomen probably uniformly fuscous as in female.

Holotype. Female. Summit Lake, mi. 392, Alaska Hwy., British Columbia.

4500', 28-VII- 1959, E. E. Macdougall. Deposited in Canadian National Insect

Collection, Ottawa (CNC).

Paratypes. One female. Same data as for holotype except 4700', VII-15-1959

(CNC).

Etymology. The species is named for the holotype collecting region.

Host and Biology. There are no rearing records for this species.

columbiensis

16 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Cymodusa cruentata (Gravenhorst)

Figs. g

Campoplex cruentatus Gravenhorst, 1829. Ichneumonologia europaea, 3: 575.

Male, female. des. Holotype: Female. Poland, Wroclaw (Wroclaw).

Cmpd. eyes strongly covergient ventrad, ditinctly hairy. Fore wing with

areolet closed. Antennal flagellum uniformly black in females. Legs mostly

ferrugineous. Abdomen with alternating black and rufous bands on tergites

2 to apex.

Female. Antennal flagellum with 29-32 articles, the basal article approx.

3.8-4.0 x as long as wide. Antennal scape with inner margin approx. 1.4-1.5

x as long as outer margin. Cmpd. eyes strongly convergent ventrad, distinctly

hairy, with lower facets approx. 1.4 x as wide as upper facets. Face strongly

shagreened, matte, with minute, close-set punctures. Tentorial pit separated

from inner margin of eye by distance less than dia. of pit. Clypeus moderately

convex, with anterior margin moderately convex. Lower 0.5 of mesopleuron

very strongly shagreened, matte, weakly to distinctly rugoso-punctate. Pro-

podeum moderately long, with apex projecting between bases of hind coxae

approx. 0.3-0.4 x the length of hind coxa. Propodeal carinae weak to moder-

ately strong, complete. Areola often partially separated from petiolar area

by transverse carina. Surface of petiolar area strongly shagreened, with

weak reticulate wrinkling. Hind femur approx. 4.4-4.8 x as long as deep.

Hind tarsal claw straight, with tip weakly bent. Petiolar segment of abdomen

moderately decurving. Petiole approx. 1.4-1.5 x as long as postpetiole, the

sides of petiole smooth, polished. Second abdominal tergite approx. 1.6-1.8

x as long as wide, with thyridium at basal 0.2. Ovipositor approx. 1.8-1.9 x

as long as hind femur, straight to weakly upcurving, with depth at mid length

approx. equal to width of hind basitarsus. Fore wing approx. 4.0-5.5 mm.

long. Hind wing with nervellus intercepted near base.

Antennal flagellum, head, thorax, petiolar segment of abdomen, bases

of abdominal tergites 2 to apex, fore and middle coxae (except apices), hind

coxa, basal segment of hind trochanter, black. Palpi fulvous. Tegula, apices

of fore and middle coxae, fore and middle trochanters (basal segments some-

times infuscated), apical segment of hind trochanter, femora, tibiae, ferrug-

ineous (hind fmeur sometimes with baso-ventral infuscated patch). Hind tar-

sus fusco-ferrugineous.

Male. Hind femur approx. 4.4-4.9 x as long as deep. Coloured as in female

except that tergites 3 or 4 to apex sometimes entirely black and hind femur

generally with greater degree of infuscation.

Material Examined. 61 males, 32 females

Distribution. CANADA. Alberta: Banff. British Columbia: Clinton; Lac La

Hache; mi 434, Alaska Hwy; Oliver; Stone Mt. Pk.; Summit Lake. Manitoba:

Churchill. Northwest Territories: Baffin Is., Clyde Inlet; Pangnirtung; Sal-

mita Mines; Tuktoyaktuk. Ontario: Smoky Falls, Mattagami R.. UNITED

STATES. Alaska: Anchorage; Granite Cr.; Isabel Pass; mi206, Richardson

Hwy. Idaho: Roberts. South Dakota: Fox Ridge. Also Colorado: Pagosa Spr.

This species is widely distributed throughout the Holarctic region in

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 17

arctic, sub-arctic, and cool-temperate areas.

Host and Biology.There are no rearing records for this species from the

nearctic region. It has been reared from Alsophila aescularia according to

labels from palearctic specimens. Adults are active from June to September,

with a peak abundance in July.

Remarks. The combination of entirely black antennal flagellum, black and

rufous banded abdominal tergites and ferrugineous legs will distinguish this

species from all other in the Cruentata Group.

cruentata

f 500

18 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Melanocera Group

The characters given in the key are sufficient to describe this group.

It is comprised of three species in the nearctic region: melanocera, montana,

nicolei. The biology of this group is unknown but the species are presumed

to attack small lepidoptera in grassy areas and meadows.

Cymodusa melanocera Viereck

Figs. 2, 10

Cymodusa melanocera Viereck, 1925, 1926. Can. Ent., 57(7): 180 (key); 58(1):

3. Male. des Holotype: Saanich District, British Columbia (CNC). Exam-

ined in 1984.

Cymodusa gracilicornis Viereck, 1925, 1926. Can. Ent., 57(7): 180 (key); 58(1):

4. Female. des. Holotype: Georgetown, Ontario (CNC). Examined in 1984.

Cmpd. eyes very strongly covergent ventrad in females, distinctly hairy.

Fore wing approx. 3.0-4.5 mm. long, with areolet closed. Second abdominal

tergite approx. 2.0-2.4 x as long as wide. Ovipositor approx. 1.7-1.9 x as

long as hind femur, straight, stout. Antennal flagellum of both sexes uniformly

black. Abdominal tergites 2-4 with apical reddish bands, sometimes abdomen

uniformly fuscous.

Female. Antennal flagellum with 28-32 articles, the basal article 5.5-6.0 x

as long as wide. Antennal scape with inner margin approx. 1.4 x as long as

outer margin. Cmpd. eyes very strongly convergent ventrad in females, dis-

tinctly hairy, with lower facets approx. 1.8 x as wide as upper facets. Face

strongly shagreened, matte, with minute punctures. Tentorial pit touching

inner margin of eye. Clypeus strongly convex, with anterior margin moder-

ately convex. Lower 0.5 of mesopleuron strongly shagreened, matte, with

minute punctures. Propodeum long, with apex projecting between bases of hind

coxae approx. 0.4 x the length of hind coxa. Propodeal carinae rather weak,

with costula often absent. Areola and petiolar area completely confluent.

Petiolar area granulose, matte, without distinct wrinkling. Hind femur approx.

9-0-6.0 x as long as deep. Hind tarsal claw straight with tip moderately to

strongly bent. Petiolar segment of abdomen straight to weakly decurving, the

petiole approx. 1.1-1.2 x as long as postpetiole. Second abdominal tergite

approx. 2.0-2.2 x as long as wide, with thyridium at basal 0.3. Ovipositor

approx. 1.7-1.9 x as long as hind femur, straight, with depth at mid length

distinctly greater than width of hind basitarsus. Fore Wing approx. 3.0-4.5

mm. long, with areolet closed. Hind wing with nervellus weakly intercepted.

Antennal flagellum, head, thorax, petiolar segment of abdomen, hind

coxa (except extreme apex which is ferrugineous), black. Palpi, tegula, white.

Fore coxa pale ferrugineous dorsally, white ventrally; middle coxa ferrugin-

eous. Fore trochanter with basal segment ferrugineous, apical segment white;

middle trochanter with basal segment ferrugineous to pale fuscous, apical

segment white to ferrugineous; hind trochanter with basal segment fuscous to

black, apical segment white to ferrugineous. Fore femur ferrugineous; middle

femur ferrugineous, with or without infuscated areas; hind femur ferrugineous

to fuscous. Fore tibia ferrugineous internally, white externally; middle tibia

with pale fuscous basal and apical bands, the median band yellow to ferrugin-

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 19

eous; hind tibia coloured as middle tibia. Hind tarsus pale fuscous.

Male. Hind femur approx. 4.5-5.0 x as long as deep. Coloured as female

except that fore and middle coxae often more extensively white.

Material Examined. 756 males, 780 females

Distribution. CANADA. Alberta: Frank; Kananaskis. British Columbia: Kaslo;

Kelowna; Keremeos; Robson; Victoria. Manitoba: Cranberry; Portage. Nova

Scotia Truro. Ontario: Brittania; Latta; Stittsville. Quebec: Aylmer; Hull;

Low; Timagami. UNITED STATES. Alabama: Longdale. Arizona: Cave Cr.

Flagstaff; Negalis; Oak Cr.; Pocket Cr.; Portal; Ruby; Sierra Vista; Huachaca

Mts.; Tucson. California: Almanor; Beatrice; Dario; Davis; Lake Wohlford;

Legget; Leona Heights; Mill Valley; Mono Lake; Oakland; White Cloud; Wood- ~

side. Colorado: Bond; Boulder; Gould; Phantom Valley. District of Columbia:

Washington. Georgia: Atlanta. Idaho: Bliss; Idaho City; Osgoode; St. Martin;

Stanley; Tuttle. Kansas: Hodgeman Co. Maryland: Hall; Laurel; Takoma Pk.

Wheaton. Massachusetts: Sherborne. Michigan: Ann Arbor; Huntington Woods;

Nahma. Montana; Greenough. Nebraska: Valentine Refuge: New Jersey:

Moorestown. New Mexico: Albuquerque; Cherry Cr. Camp; Gila Nat. Forest;

Sandia Mts; Water Canyon Camp. New York: Bemus Pt.; Farmingdale; Ithaca;

Oswego; Wilmington; Yonkers. North Carolina: Crabtree Meadows; Highlands;

Mt. Mitchell. Ohio: New Concord. Oregon: Corvallis; Hyatt Reserve; Lapine;

Mt. Angel; Ochoco Cr.; Pinehurst; Selma; Tumalo; Vale. Rhode Island: West-

erly. South Carolina: Cleveland; Greenville. Tennessee. Knoxville. Virginia:

Bluemont; Middle Mt.; Skyline Dr. Washington: Rochester. West Virginia:

Bolivar; Spruce Knob. Wisconsin: Gurney. MEXICO. Guerraro; Vera Cruz.

ECUADOR. Ona.

Host and Biology. There are no rearing records for this species in spite of

its abundance in collections. Adults are active from May through August and

there appear to be two generations in some areas.

Remarks. This species is distinguished from its close relative, montana ,

by the longer basal article of the antennal flagellum, and the longer second

abdominal tergite. It is distinguished from nicolei in having the areolet of

the forewing closed and the second abdominal tergite shorter.

Cymodusa montana, new sp.

Fig. li

Cmpd. eyes very strongly covergent ventrad in females, distinctly

hairy. Propodeum long, distinctly projecting between bases of hind coxae.

Antennal flagellum uniformly dark coloured in both sexes. Fore wing approx.

4.5-5.0 mm. long, with areolet closed. Second abdominal tergite approx.

2.9 Xas long as wide. Abdomen with alternating black and ferrugineous

bands or with black extending dorso-medially to apical margins of tergites,

the apical lateral corners ferrugineous.

HOLOTYPE. Female. Antennal flagellum with basal article approx. 4. 7x

as long as wide. Antennal scape with inner margin approx. 1.5 x as long as

outer margin. Cmpd. eyes very strongly convergent ventrad, distinctly

20 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

hairy, with lower facets approx. 1.4 x as wide as upper facets. Tentorial pit

touching inner margin of eye. Face strongly shagreened, matte, with minute,

close-set punctures. Clypeus moderately convex, with anterior margin mod-

erately convex. Lower 0.5 of mesopleuron strongly shagreened, matte, with

small punctures which are separated by distance 1.0-1.5 x dia. of single

puncture. Propodeum long, with apex projecting between bases of hind coxae

approx. 0.4 x the length of hind coxa. Areola completely confluent with pet-

iolar area, the surface of both areas granulose, matte. Hind femur approx.

6.8 x as long as deep. Hind tarsal claw with tip moderately bent. Petiolar

segment of abdomen weakly decurving, with petiole approx. 1.6 x as long as

postpetiole. Second abdominal tergite approx. 1.9 x as long as wide, with

thyridium at basal 0.2. Ovipositor approx. 1.7 x as long as hind femur,

straight, with depth at mid length distinctly greater than width of hind basi-

tarsus. Fore wing approx. 5.0 mm. long, with areolet closed. Hind wing

with nervellus intercepted near basal 0.2.

Antennal flagellum, head, thorax, petiolar segment of abdomen, bases

of tergites, coxae (except apex of fore coxa), black. Palpi fulvous. Tegula

ivory. Fore and middle trochanters extensively yellowish, hind trochanter

with basal segment fuscous, apical segment yellowish. Femora (except apex

of hind which is infuscated), tibiae, apical bands of tergites 2 to apex, ferr-

ugineous. Hind tarsus fuscous.

Range and Variation. Hind femur approx. 5.9-6.8 x as long as deep. Abd-

ominal petiole approx. 1.4-1.5 x as long as postpetiole. Second abdominal

tergite 1.9-2.2 x as long as wide. Ovipositor approx. 1.5-1.7 x as long as

hind femur. Fore coxa yellow to extensively black; middle coxa ferrugineous

to black. Hind femur with variable degree of infuscation. Abdominal tergites

black and ferrugineous banded or with black extending dorso-medially to

apical margins of tergites, the apico-lateral corners ferrugineous.

Male. Face with dense silvery pubescence. Hind femur approx. 5.2-5.4 x

as long as deep. Petiolar segment of abdomen straight. Fore and middle

coxae, fore and middle trochanters, apical segment of hind trochanter, pale

yellow. Hind coxa, abdomen, black. Hind femur extensively infuscated to

black. Hind tibia with weak fuscous sub-basal and apical bands, the median

band fulvous to ferrugineous.

Holotype. Female. Mexico, Mexico, 24.5 mi. nw. Toluca, July 30, 1962,

Naumann and Marston. Deposited in Snow Museum, University of Kansas.

Paratypes. UNITED STATES. Texas: Guadalupe Mts, 7750! VII-21-1975 (4

males), S. & J. Peck (HMT). MEXICO. 15.5 mi. e. Toluca, 9500', July 6,

1961, U. Kans. Mex. Exped. (1 female), (KSM); Hidalgo, 5 mi. n. Pachuca,

8750', Aug. 25, 1962 (1 male), Ordway, Marston (KSM); Linares, X-5-1962

(1 male), H. & M. Townes (HMT); Mexico no. 5, 7.5 mi. se. Amecameca,

9,600', July 1, 1961 (2 males), G. W. Byers (KSM). PANAMA, Cerro Punta,

VI-1-1977 (1 male), H. & A. Howden (HMT).

Ktymology. The species epithet refers to the high altitudes at which this

species occurs.

Host and Biology. There are no rearing records for this species. There —

appear to be two generations as specimens have been taken in July and October.

Remarks. See remarks under melanocera .

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae)

melanocera

mo eae ee

reg

montana

f 500

22 Contrib. Amer. Ent. Inst., vol. 22, no. 9, 1986

Cymodusa nicolei, new sp.

Fig. 12

Cmpd. eyes very strongly covergent ventrad in females, distinctly hairy.

Petiolar segment of abdomen straight. Propodeum distinctly projecting between

bases of hind coxae. Fore wing with areolet open. Second abdominal tergite

approx. 5.0 x as long as wide. Abdominal tergites uniformly fuscous.

HOLOTYPE. Female. Antennal flagellum with 34 articles, the basal article

approx. 5.5 x as long as wide. Antennal scape with inner margin approx.

1.6 x as long as outer margin. Cmpd. eyes very strongly covergent ventrad,

distinctly hairy, with lower facets approx. 1.8 x as wide as upper facets.

Face strongly shagreened, matte, with minute punctures. Tentorial pit touch-

ing inner margin of eye. Clypeus strongly convex, its anterior margin mod-

erately convex. Lower 0.5 of mesopleuron strongly shagreened, matte, with

minute punctures. Propodeum long, with apex projecting between bases of

hind coxae approx. 0.4 x the length of hind coxa. Areola approx. 0.9 x as

long as petiolar area, the areola and petiolar area completely confluent.

Surface of petiolar area with reticulate pattern of wrinkling laterally. Hind

femur approx. 5.6 x as long as deep. Hind tarsal claw with tip strongly bent.

Petiolar segment of abdomen straight, with petiole approx. 1.3 x as long as

postpetiole. Second abdominal tergite approx. 3.0 x as long as wide, with

thyridium at basal 0.4. Ovipositor approx. 1.6 x as long as hind femur,

straight, with depth at mid length approx. equal to width of hind basitarsus.

Fore wing approx. 4.2 mm. long, with areolet open. Hind wing with nervellus

intercepted very near base.

Antennal flagellum, head, thorax, hind coxa (except apex), petiolar seg-

ment of abdomen, black. Abdominal tergites 2 to apex, fuscous. Palpi, tegula,

fore and middle coxae, fore and middle trochanters, apical segment of hind

trochanter, outer surface of fore tibia, ivory. Fore and middle femora, inner

surface of fore tibia, entire middle tibia, pale ferrugineous. Hind femur,

hind tibia, ferrugineous. Hind tarsus with ground colour ferrugineous but

uniformly infuscated.

Range and Variation. Antennal flagellum with 33-36 articles. Antennal scape

with inner margin 1.5-1.6 x as long as outer margin. Basal flagellar article

approx. 5.4-5.6 x as long as wide. Hind femur 5.3-5.7 x as long as wide.

Fore wing 4.5-6.0 mm. long. No significant colour variation in specimens

at hand.

Male. Hind femur 4.8-5.3 x as long as deep. Fore and middle legs paler

than in female. Hind femur more or less completely, evenly infuscated.

Hind tibia ferrugineous medially, infuscated basally and apically. Hind tar-

sus fuscous.

Holotype. Female. Cleveland, South Carolina, VI-4-1961, G. F. Townes.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. North Carolina: Mt. Pisgah, 3,000', Sept. 5, 1939 (1 male, 1 fem-

ale), H. & M. Townes (HMT). South Carolina: Cleveland, VI-19-VII-17, 1961

(33 males, 5 females), G. F. Townes (HMT) and July 3, 1971, (3 males),

G. Townes family (HMT); Columbia, VIII-14-1951, G. & L. Townes (HMT).

Tennessee: Lexington, Natchez St. Pk., 15-19, June, (1 male), and 23, June,

1972 (6 males), G. Heinrich (CNC).

Sanborne: Revision of Nearctic Cymodusa (Ichneumonidae) 23

Etymology. This species is named in honour of Nicole Hughes in recognition

of the many hours she spent mounting and labelling specimens for my res-

earch.

Host and Biology. There are no rearing records for this species. Adults

are active from June to September and there may be two generations.

Remarks. This species is most closely related to melanocera, its sister

species is unknown. It is easily distinguished from all north american

Cymodusa by the unusually elongate, uniformly coloured abdomen, and

the open areolet in the fore wing.

nicolei

t 500

Acknowledgements

This research was funded, in part, by an NSERC (Natural Sciences and

Engineering Research Council of Canada) operating grant held by Dr. S. B.

Peck, Dept. of Biology, Carleton University, Ottawa. Additional funds came

from an NSERC postdoctoral fellowship held by the author. Luis Ling of

Carleton produced the S.E.M. photomicrographs.

24. Contrib. Amer. Ent. Inst. , vol. 22, no. 9, 1986

The sources for specimens used in this research and the abbreviations

used in the text to indicate deposition of type material are identical to those

found in my Xylophylax revision immediately preceeding this paper in this

contribution.

References

Cresson, E. T. 1864. Descriptions of North American Hymenoptera in the

collection of the Entomological Society of Philadelphia. Ent. Soc. Phil. ,

Proc., 3: 257-321.

Gravenhorst, J. L. C. Ichneumonologia europaea. Vratislaviae, 3. 1097 pp.

Holmgren, A. E. 1858. Conspectus generum Ophionidum Sueciae. Ofv. K.

Vet. Akad. Forh., 15: 321-330.

Viereck, H. L. 1906. Notes and descriptions of hymenoptera from the west-

ern United States. Amer. Ent. Soc., Trans. 32: 173-248.

.1916. In The hymenoptera, or wasp-like insects of Connecti-

cut. Guide to the insects of Connecticut, Part III. Conn. Nat. Hist.

Surv., Hartford Bull., 22: 824 pp.

1925. A preliminary revision of the Campopleginae in the

Canadian National Insect Collection. Can. Ent. 57(7): 176-181; 57(12):

296-3038.

. 1926. A preliminary revision of the Campopleginae in the

Canadian National Insect Collection. Can. Ent. 58(1): 2-83 98(11): 276-

283.

Cymodusa spp.: Figs. 1-2, frontal view of head: 1, nigripes; 2, melanocera. Figs. 3-6,

propodea: 3, nigripes; 4, distincta; 5, partis partis; 6, partis ocularis.

itty

Cymodusa spp.: Figs.7-12, propodea: 7, columbiensis; 8, ruficincta; 9, cruentata; 10,

melanocera; 11, montana; 12, nicolei.

A REVISION OF THE NEARCTIC SPECIES OF CYMODUSOPSIS VIERECK

(HYMENOPTERA: ICHNEUMONIDAE)

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, KiS 5B6

ABSTRACT

The nearctic species of Cymodusopsis Viereck (Hymenoptera: Ichneu-

monidae are revised and includes aristoteliae Viereck and eight species new

to science: albiscapa; asperata; brevicauda; latifacies; macdunnoughi; melan-

osoma; nigricincta; variabilis.

INTRODUCTION

This revision describes eight new species in the nearctic fauna and

provides a key to the identification of adult males and females as well as

species groups. Each species is illustrated by means of scanning electron

photomicrographs and their distributions plotted on maps.

The genotype, aristoteliae Viereck, was the only previously described

species of Cymodusopsis prior to this publication

MATERIALS AND METHODS

The sources of specimens used in this study are the same as those

listed in my revision of Xylophylax which occurs earlier in this contribution.

The abbreviations used in regards to the deposition of type material are also

the same.

The methods by which specimens are measured or described are the

Same as those published in my revision of Sinophorus (Sanborne, 1984) and

are not repeated here.

The generic description which follows is essentially that of Townes,

1970, with appropriate additions and modifications.

Cymodusopsis Viereck

Cymodusopsis Viereck, 1912. U. S. Nat. Mus., Proc. 43: 588.

Type-species: Cymodusopsis aristoteliae Viereck. Examined in 1984.

Fore wing 3.2 - 6.5 mm. long. Body slender to moderately robust.

2 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

Eyes moderately convergent ventrad, with lower facets larger than upper

facets, not distinctly hairy. Genal carina joining oral carina near base of

mandible. Tentorial pit (clypeal fovea) separated from inner margin of cmpd.

eye by distance 2.0-4.0 x dia. of single pit. Mandible short, its lower margin

with narrow lamella near base, its teeth equal. Mesopleuron matte, with

fine, sparse punctures (except brevicauda which has rather large, dense

punctures). Posterior mesosternal carina complete. Propodeal spiracle

round. Areola usually completely confluent with petiolar area, rarely part-

ially or completely separated by transverse carina. Propodeal trough not or

weakly excavated. Hind basitarsus without a mid ventral row of closely spaced

hairs. Tarsal claws short, with weak to moderately strong teeth at base.

Fore wing with areolet open (usually) or closed. Hind wing with nervellus often

intercepted by discoidella. Abdominal petiole weakly decurving to straight,

and lacking a glymma. Ovipositor 1.3-2.0 x as long as hind femur, slender

to stout, straight to weakly upcurving.

The genus is distributed from the low arctic regions of Canada, Alaska,

south to Central America and parts of South America. Its hosts are presumed

to be microlepidoptera as is indicated by the meagre host records available.

Relationships. The relationship of Cymodusopsis to other Porizontinae is a

matter of speculation at this point. Larval studies presently being conducted

by David Wahl on the entire subfamily may shed more light on the matter.

In my opinion, the similarities between Cymodusopsis and Cymodusa are

superficial (e. g.,convergent eyes, shape and form of abdominal petiole) and

the result of convergence. There is a group of species within Diadegma

(apparently all undescribed) that are even more similar and can only be

separated from Cymodusopsis by the presence of a glymma on the abdominal

petiole. If Cymodusopsis is related to this group of species it opens up the

possibility that Diadegma or Cymodusopsis is a paraphyletic group. I choose

to treat Cymodusopsis as a separate genus for the time being as it can easily

be separated from Cymodusa ( eyes not as strongly convergent in females

and not distinctly hairy, tentorial pit much farther away from inner margin of

cmpd. eye in Cymodusopsis) and from Diadegma which has distinct glymmae.

Key to the Nearctic Species Groups and Species of Cymodusopsis Viereck

1. Antennal scape and pedicel with frontal surfaces pale or white. Areola

completely confluent with petiolar area, not indented at junction with

petiolar area, the combined areas forming a broad trough (Figs. 3,4).

re wena ores ecole a ccc cccccccccocccrcoccseesseAristoteliae Group. 2.

Antennal scape and pedicel uniformly black. Areola often partially, rarely

completely separated from petiolar area by transverse carina, indented

at junction with petiolar area (FigS. 5-12).ccceccrcccecersceresessce

2. Abdominal tergites 2-6 with apical ferrugineous bands. Southwestern U.S.

eS 0b SA EO ee biwild Mees apa scics Se evelbisoapay aucBp. (p. 6).

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 3

Abdominal tergites with antero-lateral corners ferrugineous, rarely with

some tergites entirely ferrugineous, never with a banding pattern....

east of Rocky Mts........-e-sscerceeseee aristoteliae Viereck. (p. 4).

Second abdominal tergite approx. 1.0-1.2 x as long as wide in females,

1.4-1.5 x in males. Propodeal trough broad (Fig.5), not coarsely

wrinkled. Hind femur extensively to entirely black. Melanosoma Group

Colorado..... ye ger ae Os Bienes’ ese. melanosoma, n. sp. (p. 8).

Second abdominal tergite approx. 1.4-1.8 x as long as wide in females,

1.8-2.2 x in males. Propodeal trough not as broad (Fig. 6), usually

coarsely wrinkled. Hind femur variously coloured. Nigricincta Group.

@eeee0aeeeeeezVn2rweeeverveexseeen2xeeessts e868 ee @eee#ecssecscsececseeeee4u25uoegesc@e@geee#eesceeevs2v0ee8e0e8e8e8e0e0e86e (p- 10).

Areolet of fore wing closed (variabilis can be traced through both halves

OF COUDIEU hie tal ee CPEELEAES EES OES O88 OW 5 Sains wie 4 HES ERES e

Areolet of fore wing open......... Pe es Fe ne eA PER prey hae ee eR a

Abdominal tergites extensively rufous. Ovipositor approx. 1.3 x as long

as hind femur. Oregon....... :6 kA pe »--.ebrevicauda, n. sp. (p- 10)

Abdominal tergites black and ferrugineous banded or entirely black.

Ovipositor approx. 1.6-1.8 x as long as hind femur........ecseseele

Abdominal tergites entirely black. British Columbia south to Colorado...

«eR ae ee 6 AE Ak 9 WORE & chaik’ de hes, Se ene --. nigricincta, n. sp. (p-11)

Abdominal tergites with banding pattern. Widespread in U.S.....cccceee

cee c cece rcoreeeccccercesercerescorosecese Variavilis, Ne sp. (p. 13)

Abdominal tergites with banding pattern........ variabilis, n. sp. (p. 13)

Abdominal tergites uniformly black or fuUSCOUS....cscrccesscseccceesd:

Abdominal petiole with a dorso-lateral carina running from base to spir-

acle, the sides of petiole weakly concave. Ovipositor slender, its

depth at mid length less than width of hind basitarsus. Boreal forests

of Alaska and Canada. .....css-scsorccerce- + -asperata, 1. Sp. (Pel

Abdominal petiole without a dorso-lateral carina, its sides weakly convex.

Ovipositor Tamer BlOOl. 2s. een acesazece pd aa da ek pee ao Dine eae ae 9.

Cmpd. eyes weakly notched opposite antennal sockets. Face not unusually

wide. Clypeus strongly convex. eastern North America.......... itis

5 ees eg atk Re ace cree eee ee cores eeee- macdunnoughi, n. sp. (p. 18)

Cmpd. eyes distinctly notched opposite antennal sockets. Face unusually

wide. Clypeus weakly convex. western North America.......c.c-eee.

oP asic st aS Sk ms FL OE he Mesionsriy ate ttin -latifacies, n. sp. (p. 20)

A Contrib. Amer. Ent. Inst. , vol. 22, mo. 11, 1986

Aristoteliae Group

This group is characterized by a broad propodeal trough which is not

indented at the junction of the areolar and petiolar areas and by the antennal

scape and pedicel which are coloured white on their frontal surfaces. The

group comprises two species, aristoteliae Viereck and albiscapa, n. sp.

Cymodusopsis aristoteliae Viereck

Fig. 3

Cymodusopsis aristoteliae Viereck, 1912. U. S. Nat. Mus., Proc. 43: 588

male, female. des. Holotype: Female. Kirkwood, Missouri (U. S. N. M.).

Examined in 1984.

Cmpd. eyes strongly converging ventrad in female. Tentorial pit sep-

arated from inner margin of cmpd. eye by distance 2.0-2.5 x dia. of single

pit in female . Second abdominal tergite approx. 1.6-1.8 x as long as wide in

female. Propodeal trough very broad (Fig. 3). Fore wing with areolet usually

closed. Antennal scape with frontal surface white.

Female. Antennal flagellum with 24-27 articles, with basal article approx.

4,.5-4.7x as long as wide. Antennal scape with inner margin approx. 1.4.x

as long as outer mergin. Cmpd. eyes strongly converging ventrad, moder-

ately hairy, with lower facets approx. 1.8 x as wide as upper facets. Tent-

orial pit separated from inner margin of cmpd. eye by distance approx. 2.0-

2.5 xX dia. of pit. Face strongly shagreened, matte, with moderately dense,

silvery pubescence, its punctures minute. Clypeus moderately convex, with

anterior margin moderately convex. Mesopleuron with lower 0.5 weakly to

moderately shagreened, weakly polished to matte, sometimes with distinct

rugosities anteriorly, its punctures medium sized and separated by distance

1.0-1.5 x dia. of single puncture. Propodeum short, with apex not projecting

between bases of hind coxae. Areola completely confluent with petiolar area,

the combined areas forming a broad trough with surface strongly rugoso-

punctate. Hind femur approx. 6.0 x as long as deep. Hind tarsal claw with

weak teeth at base. Petiolar segment of abdomen weakly to moderately de-

curving, with petiole approx. 1.5-1.7 x as long as postpetiole. Second abd.

tergite approx. 1.6-1.7 x as long as wide, with thyridium near basal 0.2.

Ovipositor approx. 1.7-1.9 x as long as hind femur, nearly straight, with

depth at mid length slightly greater than width of hind basitarsus. Fore wing

approx. 3.1-6.5 mm. long, with areolet usually closed.

Antennal flagellum, head, thorax, petiolar segment, 2ndabd. tergite,

median dorsal areas of térgites 3 to apex, hind coxa, black. Front of scape

white. Palpi white to fulvous. Fore and middle coxae black to fulvous dorsally,

yellow ventrally. Fore and middle trochanters yellow. Hind trochanter with

basal segment fuscous, apical segment yellow. Femora, fore and middle tibiae,

lateral areas of tergites 3 to apex, ferrugineous. Hind tibia with sub-basal

and apical fuscous bands, the median band yellow-ferrugineous. Hind tarsal

segments fuscous.

Male. Similar to female except that cmpd. eyes not as strongly convergent

ventrad, hind femur slightly more stout, colour of tergites 3 to apex more

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 5)

extensively ferrugineous.

Material Examined. 4 males, 18 females

Distribution. CANADA. Alberta: Lethbridge. UNITED STATES. Arizona:

Portal. Arkansas: Fayetteville; Little Rock. Florida: Hialeah. Georgia: St.

Catherine Is., Liberty Co. Kansas. Douglas Co. Missouri: Columbia; Kirk-

wood. Texas: Carswell Airforce Base; Kerrville. Unknown State: Milford.

Host and Biology. This is a prairie grassland species. It has been reared

from Aristotelia pudibundella (Gelechiidae) and a species of Loxostege

(Pyralidae) on cotton. Peak abundance appears to be in June and July but

adults are active from April to November indicating two or more generations.

Remarks. This species is very closely related to albiscapa but differs from

it in the absence of a regular banding pattern on the tergites.

C. aristoteliae

r 500

6 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

Cymodusopsis albiscapa, new species

Fig. 4

Cmpd. eyes moderately converging ventrad in females, with hairs

slightly longer than normal. Fore wing with areolet open. Areola unusually

short, broad. Ovipositor approx. 1.5 x as long as hind femur. Frontal sur-

faces of scape and pedicel marked with white. Abdominal tergites with regular

banding pattern.

HOLOTYPE. Female. Antennal flagellum with 26 articles, the basal article

approx. 4.8 x as long as wide. Antennal scape with inner margin approx. 1.4 x

as long as outer margin. Cmpd. eyes moderately converging ventrad, with

hairs slightly longer than normal (relative to other Porizontinae), with lower

facets approx. 1.4 x as wide as upper facets. Face strongly shagreened,

matte, with dense silvery pubescence arising from minute punctures. Tent-

orial pit separated from inner margin of cmpd. eye by distance approx. equal

to 2.0 x dia. of pit. Clypeus weakly convex, its anterior margin moderately

convex. Mesopleuron with lower 0.5 strongly shagreened, weakly polished ,

its punctures small, separated by distance 1.0-2.0 x dia. of single puncture.

Propodeum moderately long, with apex projecting between bases of hind coxae

approx. 0.3 x the length of coxa. Propodeal carinae weak, with costula absent.

Areola completely confluent with petiolar area, approx. 0.7 x as wide and

0.2 x as long as petiolar area. Surface of petiolar area with numerous weak

transverse wrinkles. Hind femur approx. 5.9 x as long as deep. Hind tarsal

claw pectinate for over half its length. Petiolar segment of abdomen more or

less straight, with petiole approx. 1.4 x as long as postpetiole, the sides of

petiole smooth, polished. Second abdominal tergite approx. 1.8 x as long as

wide. Ovipositor approx. 1.5 x as long as hind femur, nearly straight, with

depth at mid length greater than width of hind basitarsus. Fore wing approx.

3.8 mm. long, with areolet open. Hind wing with nervellus weakly intercepted.

Antennal flagellum, head, thorax, abd. petiole, bases of abd. tergites

2-6, hind coxa (except extreme apex), basal segment of hind trochanter,

black. Front of scape and pedicel white. Palpi, tegula, ivory. Fore and

middle coxae, fore and middle trochanters, apical segment of hind trochanter,

external surface of fore tibia, externo-median surface of middle tibia, white.

Fore femur ferrugineous; middle femur ferrugineous with weak baso-ventral

infuscated patch; hind femur ferrugineous with baso-ventral and apico-dorsal

fuscous patches. Front tibia with internal surface ferrugineous; middle tibia

weakly fuscous internally, with weakly fuscous sub-basal and apical bands ;

hind tibia with internal surface, sub-basal and apical bands, fuscous, the

externo~median surface fulvous. Apical bands of abdominal tergites 2-4

ferrugineous, 5-6 fusco-ferrugineous.

Range and Variation. No significant variation in specimens at hand except

size. Fore wing 3.3-4.1 mm. long.

Male. Unknown

Holotype. Female. Denio, Nevada, May 31 1978, H. & M. Townes. Deposited

in H. & M. Townes collection, Gainesville, Florida.

Paratypes. California: Davis, VIII-20, 1955 (1 female), A.T. McClay (UCB);

Mint Canyon, Solemint 28-VI-1955 (1 female), W. R. Richards (CNC).

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae)

Colorado. Boulder, 4.5 mi. w., 5500', June 20, 1961 (i female), W. R. M.

Mason (CNC).Nevada: One female, same data as for holotype.

Etymology. The species epithet refers to the white markings on the front of |

the antennal scape and pedicel.

Host and Biology. There are no rearing records for this species. There

appear to be two generations per year.

Remarks. See remarks under aristoteliae, p. 5.

C. albiscapa

f 500

8 Contrib. Amer. Ent. Inst. , vol. 22, no. 10, 1986

Melanosoma Group

This group is characterized by the unusually robust body, broad petiolar

trough which is indented at the junction of the areola and petiolar region, and

by the second abdominal tergite which is approx. as long as wide in females.

The group comprises a single species, melanosoma, n. sp.

Cymodusopsis melanosoma, new species

Fig. 5

Cmpd. eyes moderately converging ventrad in females, not distinctly

hairy. Lower 0.5 of mesopleuron with rugosities. Propodeal trough broad,

with petiolar area lacking distinct transverse wrinkling. Petiolar segment of

abdomen moderately decurving, the petiole with a weak carina from base to

spiracle. Second abdominal tergite of female approx. as long as wide. Ovi-

positor approx. 1.7-1.8 x as long as hind femur. Body extensively black.

HOLOTYPE. Female. Antennal flagellum with 23 articles, the basal article

approx. 4.3 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes moderately converging ventrad,

with hairs slightly longer than normal, with lower facets approx. 1.4 x as

wide as upper facets. Face strongly shagreened, matte, with minute punctures.

Tentorial pit separated from inner margin of cmpd. eye by distance approx.

equal to1.5 x the dia. of pit. Clypeus strongly convex, its anterior margin

weakly convex. Mesopleuron with lower 0.5 strongly shagreened, matte, with

distinct rugosities, the punctures small. Propodeum moderately long, with

apex projecting between bases of hind coxae approx. 0.3 x length of hind

coxa. Propodeal carinae strong, complete. Areola approx. 0.5 x as wide

and 0.5 x as long as petiolar area, separated from petiolar area by weak

transverse carina. Petiolar area very broad without distinct transverse

wrinkling. Hind femur approx. 4.9 x as long as deep. Hind tarsal claw with

teeth extending over half the length of claw. Petiolar segment of abdomen

moderately decurving, with petiole approx. 1.4 x as long as postpetiole, with

a weak dorso-lateral carina running from base to spiracle. Second abdominal

tergite approx. 1.1 x as long as wide. Ovipositor approx. 1.7 x as long as hind

femur, weakly upcurving, with depth at mid length approx. equal to width of

hind basitarsus. Fore wing approx. 3.7 mm. long, with areolet open. Hind

wing with nervellus weakly intercepted.

Antennal flagellum, scape, pedicel, head, thorax, abdomen, coxae,

trochanters, basal 0.4 of middle femur, hind femur, black. Palpi, tegula,

fulvous. Fore femur ferrugineous; middle femur with apical 0.6 ferrugineous.

Fore tibia ferrugineous internally, whitish externally; middle tibia fuscous

internally, with fuscous sub-basal and apical bands, the externo-median surface

whitish; hind tibia coloured as middle tibia except that externo-median surface

pale ferrugineous. Hind tarsal segments fuscous.

Range and Variation. Antennal flagellum with 23-25 articles, the basal article

4,2-4.5 x as long as wide. Propodeum with areola partially to entirely sep-

arated from petiolar area by incomplete to complete transverse carina.

Ovipositor approx. 1.7-1.8 x as long as hind femur. Fore wing 3.2-3.8 mm.

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 9

long. Palpi fulvous to fuscous. Tegula fulvous to fuscous. Apical segment of

fore trochanter yellow to black. fore femur often infuscated basally.

Male. Hind femur 4.5-4.8 x as long as deep. Second abdominal tergite approx.

1.4-1.5 x as long as wide. Coloured as female except that infuscation of fem-

ora often greater.

Holotype. Female. Independence Pass, Lake Co., 12,100', Colorado, 31-VII-

1961, tundra, B. H. Poole. Deposited in Canadian National Insect Collection,

Ottawa.

Paratypes. 18 males, 16 females, same data as for holotype except that 6

specimens captured 8-VIII-1961. Colorado. Hoosier Pass, 12,000', 8-VilI-

1961, (1 male, 2 females), B. H. Poole (CNC); Loveland Pass, 12,000' summit,

¢(~VII-1961 (3 females), B. H. Poole (CNC).

Ktymology. The species epithet refers to the highly melanic colouration of

this species.

Host and Biology. There are no rearing records for this species. All spec-

imens have been taken at an altitude of approx. 12, 000' in tundra conditions.

Remarks. The very broad petiolar area, nearly square 2nd abd. tergite and

extensive black colouration distinguishes this species from all other neartic

Cymodusopsis.

C. melanosoma

10 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

Nigricincta Group

This group is characterized by the elongate second abdominal tergite

(approx. 2.0 x as long as wide), and narrow, coarsely wrinkled propodeal

trough which is distinctly constricted at the junction of the areolar and pet-

iolar areas. The group comprises six species: brevicauda n. sp., nigricincta

n. Sp., variabilis n. sp., asperata n. sp., macdunnoughi n. sp., and

latifacies n. sp. There are some features of brevicauda which may indicate

that this species is incorrectly placed, specifically, the weakly convergent

cmpd. eyes whose upper and lower facets are approx. equal sized, the very

short ovipositor, and the extensively red abdomen which is unique to the genus.

Cymodusopsis brevicauda, new species

Fig. 7

Cmpd. eyes weakly converging ventrad in female, with hairs slightly

longer than normal, with lower facets slightly larger than upper facets. Fore

wing with areolet closed. Ovipositor approx. 1.3 x as long as hind femur.

Abdominal tergites 3 to apex entirely rufous.

HOLOTYPE. Female. Antennal flagellum with 38 articles, the basal article

approx. 4.0 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes weakly coverging ventrad, with

hairs only slightly longer than normal, with lower facets approx. 1.2 x as

wide as upper facets. Face with dense silvery pubescence arising from min-

ute punctures. Tentorial pit separated from inner margin of cmpd. eye by

distance approx. equal to 3.0x dia. of pit. Clypeus unusually broad, nearly

flat, with anterior margin weakly convex. Mesopleuron with lower 0.5 coarse-

ly, densely punctate. Propodeum moderately long, with apex projecting be-

tween bases of hind coxae approx. 0.3 x the length of hind coxa. Propodeal

carinae strong, complete. Areola and petiolar area completely confluent.

Petiolar area with coarse, reticulate pattern of wrinkling. Hind femur approx.

5.6 x as long as deep. Hind tarsal claw nearly straight, with weak teeth at

base. Petiolar segment of abdomen moderately decurving, the sides of petiole

weakly excavated, the petiole approx.1.5 x as long as postpetiole. Second

abdominal tergite approx. 1.4 x as long as wide, with thyridium near basal

0.2. Ovipositor approx. 1.3 x as long as hind femur, weakly upcurving, with

depth at mid length approx. equal to width of hind basitarsus. Fore wing

approx. 6.5 mm. long, with areolet closed. Hind wing with nervellus inter-

cepted at base.

Antenna, head, thorax, petiolar segment of abdomen, basal 0.8 of 2nd

abdominal tergite, mid dorsal triangular area at base of 3rd abdominal ter-

gite, dorsal surfaces of fore and middle coxae, entire hind coxa, basal seg-

ment of hind trochanter, hind tarsal segments 4,5, black. Median area of

mandible, palpi, tegula, ventral surfaces of fore and middle coxae, most of

fore and middle trochanters, apical segment of hind trochanter, yellow. Fem-

ora, tibiae, tarsal segments 1-3 (all legs), ferrugineous. Apical 0.2 of 2nd

abdominal tergite, most of 3rd tergite, tergites 4 to apex, rufous. Wings

weakly infuscate.

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) a

Range and Variation. Known only from holotype.

Male. Unknown

Holotype. Female. French Glen, Oregon, VII-10-1978, H. & M. Townes.

Deposited in Townes collection, Gainesville, Florida.

Etymology. The species epithet refers to the unusually short ovipositor.

Host and Biology. Unknown

Remarks. The combination of weakly converging eyes, broad clypeus, short

ovipositor, and extensively red abdomen clearly distinguishes this species

from all other nearctic Cymodusopsis.

Cymodusopsis nigricincta, new species

Cmpd. eyes moderately converging ventrad in female, with hairs of

normal length. Basal article of antennal flagellum 3.7-3.8 x as long as wide.

Fore wing with areolet closed. Ovipositor approx. 1.6-1.7 x as long as hind

femur, straight, with depth at mid length greater than width of hind basitarsus.

Abdomen black.

HOLOTYPE. Female. Antennal flagellum with 34 articles, the basal article

approx. 3.8 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes moderately converging ventrad,

not distinctly hairy, with lower facets approx. 1.4 x as wide as upper facets.

Tentorial pit separated from inner margin of cmpd. eye by distance approx.

equal to 2.0 x dia. of pit. Face strongly shagreened, matte, with small punc-

tures which are separated by distance less than or equal to dia. of single punc-

ture. Clypeus moderately convex, with anterior margin moderately convex.

Mesopleuron with lower 0.5 shagreened, weakly polished, with medium sized

punctures which are separated by distance less than or equal to dia. of single

puncture, with some rugosities present medially. Propodeum moderately long,

with apex projecting between bases of hind coxae approx. 0.3 x the length of

hind coxa. Areola completely confluent with petiolar area, polished, with

weak wrinkling. Petiolar area with numerous, coarse transverse wrinkles.

Hind femur approx. 5.5 x as long as deep. Hind tarsal claw with weak teeth

at base. Petiolar segment of abdomen moderately decurving, the sides of

petiole weakly excavated apically. Petiole approx. 1.5 x as long as postpetiole.

Second abdominal tergite approx. 1.8 x as long as wide, with thyridium near

basal 0.2. Ovipositor approx. 1.8 x as long as hindfemur, straight, with

depth at mid length greater than width of hind basitarsus. Fore wing approx.

0-09 mm. long. Hind wing with nervellus not intercepted.

Antenna, head, thorax, abdomen, coxae, basal segment of all trochanters,

black. Palpi fulvous. Tegula white. Apical segment of fore trochanter yellow;

apical segment of middle trochanter pale fulvous; apical segment of hind tro-

chanter fuscous basally, ferrugineous apically. Femora, fore and middle tib-

iae, ferrugineous. Hind tibia with base and apex weakly infuscated, the med~

ian area ferrugineous. Hind tarsus fuscous. Ovipositor unusually dark. Wings

weakly infuscated.

12 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

Range and Variation. Antennal flagellum with 25-34 articles, the basal art-

icle 3.8-3.9 x as long as wide. Antennal scape with inner margin 1.3-1.4 x

as long as outer margin. Petiole approx. 1.4-1.5 x as long as postpetiole.

Second abdominal tergite approx. 1.6-1.8 x as long as wide. Ovipositor approx.

1.7-1.8 x as long as hind femur. Hind femur approx. 5.3-5.6 x as long as deep.

Fore wing approx. 4.5-5.5 mm. long.

Basal segment of fore trochanter yellow to black. Fore coxa black

dorsally, yellow ventrally , to entirely black. Hind femur often with some

infuscation ventrally.

Male. Hind femur approx. 5.2-5.4 x as long as deep. Petiolar segment of

abdomen weakly decurving. Colour as in female.

Holotype. Female. Stone Mt. Pk., British Columbia, 3800', VI-19-1973, H.

& M. Townes. Deposited in Townes collection, Gainesville, Florida.

Paratypes. CANADA. British Columbia: Stone Mt. Pk., VII-17-22-1973,

(4males, 4 females), H. & M. Townes (HMT). UNITED STATES. Colorado:

Doolittie Ranch, Mt. Evans, 9800', VII-4, VII-18, 1964 (2 males, 1 female),

C. Dasch (CDC) and VII-3-1961 (1 female), W. R. M. Mason (CNC); Echo

Lake, Mt. Evans, 10,600', VIII-4-1961 (1 female), S. M. Clark (CNC); Fort

Collins, May, (18)94, on alfalfa (1 male, 2 females), coll. ?, (USNM); Gould,

VITI-6-1974 (2 females), H. & M. Townes (HMT); Loveland Pass, 1990', VII-

18-1964 (1 female), C. Dasch (CDC); Phantom Valley, R.M.N.P., 9400',

June 16, 1948 (1 male), H.,M., G., D., & J. Townes (HMT); West Chicago

Cr., 9800', VII-25- 1964 (1 female), C. Dasch (CDC). Idaho: Galena Summit,

nr. Stanley, 8700', VIU-4-1978 (1 female), H. & M. Townes (HMT); Stanley,

Aug. 2-9, 1978 (9 females), H. & M. Townes (HMT). Washington: Mt. Ranier,

VU-8-1940 (2 males, 3 females), 5700', H. & M. Townes (HMT). Wyoming:

Battle Lake Rd., Sierra Madre, 8,000', VII-18-1961 (1 female), J. G. Chilcott

(CNC).

Etymology. The species epithet is in reference to the black colouration of the

ence

Host and Biology. There are no rearing records for theis species although

a number of speciemens have been collected on alfalfa. Adults are active

from May to August indicating two or more generations per year in some

areas.

Remarks. This species is most closely related to the asperata-variabilis

sister pair but can be distinguished by the larger size, fore wing with areolet

closed, and black colouration of the abdomen.

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 13

C. nigricincta

e 500

Cymodusopsis variabilis, new species

Fig. 8

Cmpd. eyes moderately covergent ventrad in females, with hairs

slightly longer than normal, with lower facets approx. 1.4x as wide as upper

facets. Propodeum long, with apex projecting between bases of hind coxae

a distance approx. equal to 0.4 x the length of hind coxa. Petiolar segment

of abdomen weakly decurving. Ovipositor with depth at mid length greater

than width of hind basitarsus. Hind wing with nervellus intercepted.

14 Contrib. Amer. Ent. Inst. , vol. 22, no. 10, 1986

HOLOTYPE. Female. Antennal flagellum with 26 articles, the basal article

approx. 5.0 x as long as wide. Antennal scape with inner margin approx.

1.5 x as long as outer margin. Cmpd. eyes moderately covergent ventrad ,

with hairs slightly longer than normal, with lower facets approx. 1.4 x as

wide as upper facets. Width of frons along line intersecting posterior margins

of antennal sockets approx. 1.5 x width of face at level of posterior margin of

clypeus. Face strongly shagreened, matte, with dense silvery pubescence

arising from minute punctures. Clypeus strongly convex, its anterior margin

moderately convex. Tentorial pit separated from inner margin of cmpd. eye

by distance approx. equal to 3.0 x dia. of pit. Mesopleuron with lower 0.5

strongly shagreened, matte, with minute punctures which are separated by

distance approx. equal to dia. of single puncture. Propodeum long, its apex

projecting between bases of hind coxae a distance approx. equal to 0.4.x

length of hind coxa. Propodeal carinae strong, complete. Areola approx.

0.5 x as wide as petiolar area, completely confluent with petiolar area.

Petiolar area with numerous coarse, transverse wrinkles. Hind femur approx.

6.4 x as long as deep. Petiolar segment of abdomen weakly decurving, the

petiole approx. 1.5 x as long as postpetiole, with sides smooth, polished.

Second abdominal tergite approx. 1.8 x as long as wide. Ovipositor approx.

1.6 x as long as hind femur, weakly upcurving, with depth at mid length

greater than width of hind basitarsus. Fore wing approx. 3.5 mm. long,

with areolet open. Hind wing with nervellus intercepted.

Antenna, head, thorax, petiolar segment of abdomen, bases of abdom-

inal tergites 2-6, hind coxa, black. Palpi pale fulvous. Tegula white. Fore

coxa fuscous dorsally, yellowish ventrally; middle coxa ferrugineous dor-

sally, yellowish ventrally. Fore and middle trochanters mostly yellowish;

hind trochanter with basal segment fuscous, apical segment yellow. Femora

ferrugineous. Fore tibia whitish externally, ferrugineous internally; middle

tibia with extreme base and externo-median surface white, the sub-basal and

apical bands and internal surface, ferrugineous; hind tibia fusco—ferrugineous,

with very weak sub-basal and apical bands. Hind tarsus fuscous. Apical

bands of abdominal tergites 2-6 more or less ferrugineous.

Range and Variation. Antennal flagellum with 24-28 articles, the basal art-

icle 4.8-5.0 x as long as wide. Antennal scape with inner margin approx.

1.4-1.5 x as long as outer margin. Areola approx. 0.4-0.5 x as wide as pet-

iolar area. Hind femur approx. 6.2-6.5 x as long as deep. Petiolar segment

of abdomen straight to weakly decurving. Ovipositor approx. 1.5-1.7 x as

long as hind femur. Fore wing with areolet open (usually) or closed. Hind

femur ferrugineous but showing varying degrees of infuscation, particularly

basally and ventrally. Hind tibia ferrugineous with weak sub-basal and apical

bands and showing varying degrees of infuscation with resulting loss of band-

ing pattern . Abdominal tergites 2 to apex with ferrugineous apical bands to

entirely black, with many intermediates.

Male. Hind femur approx. 5.8-6.0 x as long as deep. Coloured as female

but with much greater tendency towards melanism so that abdominal tergites

very commonly uniformly dark.

Holotype. Female. Iron Co., Michigan, July 1, 1970, Nemah Hussain.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. CANADA. Alberta: Coleman, VI-12-1962 (1 male, 1 female), K

C. Hermann (CNC); Elkwater, VI-4-1952 (1 female), A. R. Brooks (CNC);

Kananaskis, For. Exp. Stn. Seebe, June 29, 1962 (1 female), W.R.M. Mason

(CNC); Lavine, VI-11-1952 (1 male), A.R. Brooks (CNC); Morley, 8 mi. e.

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 15

VI-22-1962 (1 male), W.R.M. Mason (CNC); Onefour, VI-2-1956 (1 female),

O.Peck (CNC); Waterton, June 11, 1962 (1 female), K. C. Hermann and June

11-1962 (1 male), W.R.M. Mason (CNC). British Columbia: Kelowna, VII-14-

1950 (1 female), B. P. Beirne (CNC); Kamloops, VI-1-1938 (1 female), G. S.

Walley (CNC); Oliver, V-17-1959 (1 male), L. A. Kelton (CNC); Robson, VI-

13-1952 (1 male), H. R. Foxlee (CNC); Trinity Valley, V-21-1959 (1 female),

L. A. Kelton (CNC).New Brunswick: Kouchibouguac Nat. Pk., (1 female),

VIII-30-1977, S. J. Miller (CNC). Prince Edward Island: Alberton, VIII-4-

1940 (1 male), ex. lep. on fireweed, J. McDunnoughi (CNC). Saskatchewan:

Elbow, VI-17-1960 (1 female), A. R. Brooks (CNC); Sakatoon, V-9-1949 (1

female), A. R. Brooks (CNC). UNITED STATES. California: nr. Glacier Pt.,

Yosemite Pk., July 16, 1948 (1 male), H.,M.,G.,D., & J. Townes (HMT)s;

Lee Vining, June 25, 1948 (1 male), H.,M.,G., & D. Townes (HMT); May L.,

Yosemite Pk., 10,500', July 27, 1948 (1 female), H.,M.,G., D., & J. Townes

(HMT); nr. Mono Pass, Inyo Co., 12,000', VIII-15-1957 (1 female), C. D.

MacNeill (CAS); Nevada Co., Whitecloud campground, V-30-31-1964 (14 males,

2 females), P. H. Arnaud (CAS); Mint Canyon, Solemint, IV-28-1955 (1 female),

W. R. Richards (CNC); Oakland, V-3-1937 (1 male), E. S. Ross (HMT); Ruby

Lake, Inyo Co., 11, 500', VIII-13-1957 (1 female), C. D. MacNeill (CAS); nr.

Sonora Pass, 8,000', July 6, 1948 (3 males) and 8500', July 4 (1 female), July

7 (1 male), H., M., G., & D. Townes (HMT). Colorado: Doolittle Ranch,

Mt. Evans, 9,800', VIII-3-1961 (1 male), W.R.M.Mason (CNC) and July 20,

1964 (1 female), C.D. Dasch (CDC); Echo Lk., Mt Evans, July tl, 1978 (1

female), B. & R. Dasch (CDC); Gould, VIII-5-1974 (1 male), H. & M. Townes

(HMT); Loveland Pass, 9850', VIN-8-1961 (1 male), B. H. Poole (CNC); Mt.

Evans, 11,700', July 29, 1961 (2 males), C. H. Mann, S. M. Clark (CNC);

Muddy Pass, Jackson Co., 8,800', VII-15-1961 (2 males), B. Poole (CNC);

Nederland, Science Lodge, 9,500', VI-28-1961 (1 male), B. H. Poole and

VH-4-1961 (2 males, 1 female), J. R. Stainer, and VI[-3- 1961 (1 male), B. H.

Poole (CNC); Niwot Ridge, nr. Ward, July 4, 1961 (5 females) C. H. Mann

(CNC).Idaho: Lowman, Aug. 9, 1978 (Imale), H. & M. Townes (HMT); nr.

Stanley, Aug. 8, 1978 (Imale), H. & M. Townes (HMT). Michigan: Iron Co.,

July 1, 1970 (1 female), Nemah Hussein (HMT). Montana: Glacier Nt. Pk.,

July 4, 1973 (1 female), B. & C. Dasch (CDC). Nevada: Angel Lake, 12 mi.

sw. Wells, 8,400', VI[-11-1961 (1 female), B. H. Poole (CNC); Denio, May

31, 1978 (2 females), H. & M. Townes (HMT); Tuscarora, June 5, 1978

(1 male, 3 females), H. & M. Townes (HMT). New Mexico: Las Cruces (1

female), coll. ?, (USNM). Oregon: Brothers, July 9, 1978 (2 males), H. & M.

Townes (HMT). Texas: 10 mi. w. Ft. Davis, 5,000', May 29, 1959 (1 female)

W.R.M.Mason (CNC); unknown locality, Apr. 7, 1934, ex. Plutella maculi-

pennis, coll. ? (CNC).Utah: Henry's Fork Pk., Summit Co., VII-1-10-1979

(3 females), S. & J. Peck (CNC). Wyoming: Lusk, VII-6-1961 (1 female),

H. Howden (CNC). MEXICO. Toluca, 8800', 17-VIII-1954 (1 female), J. G.

Chillcott (CNC).

Etymology: The species epithet is in reference to the variable nature of the

colouration of the abdomen (with or without a banding pattern) and the areolet

of the fore wing which can be open or closed.

Host and Biology. The species has been reared from an unidentified lepid-

opteran on fireweed and from Plutella maculipennis on cabbage. There is

probably a single generation per year and occurs later in areas of high elev-

ation.

Remarks. This species is closely related to asperata but differs from the

latter in having a much stouter ovipositor and in the petiole which is sm ooth

sided.

16 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

C. variabilis

t 500

Cymodusopsis asperata, new species

Fig. 9

Cmpd. eyes moderately convergent ventrad in females, not distinctly

hairy. Fore wing with areolet open. Abdominal petiole with sides rough and

usually grooved dorso-laterally at base. Ovipositor with depth at mid length

less than width of hind basitarsus. Antenna uniformly black. Abdominal ter-

gites uniformly black.

HOLOTYPE. Female. Antennal flagellum with 29 articles, the basal article

approx. 3.7 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes moderately conver gent ventrad,

with lower facets approx. 1.4 x as wide as upper facets. Face shagreened,

matte, with dense silvery pubescence arising from minute punctures. Tent-

orial pit separated from inner margin of cmpd. eye by distance approx. equal

to 4.0 x dia. of pit. Clypeus moderately convex, its anterior margin moder-

ately convex. Mesopleuron with lower 0.5 shagreened, matte, with widely

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 1é

spaced, minute punctures. Propodeum moderately long, with apex projecting

between bases of hind coxae a distance approx. equal to 0.3 x the length of

hind coxa. Propodeal carinae strong, complete. Areola completely confluent

with petiolar area. Hind femur approx. 5.2x as long as deep. Hind tarsal

claw with weak teeth at base. Petiolar segment of abdomen weakly decurving,

with petiole approx. 1.6 x as long as postpetiole. Sides of petiole rough.

Second abdominal tergite approx. 1.6 x as long as wide, with thyridium at

basal 0.3. Ovipositor approx. 1.9x as long as hind femur, weakly upcurving,

with depth at mid length less than width of hind basitarsus. Fore wing approx.

4.2mm. long, with areolet open. Hind wing with nervellus not intercepted.

Antenna, head, thorax, abdomen, dorsal surface of fore and middle

coxae, entire hind coxa, basal segment of hind trochanter, black. Palpi ful-

vous. Tegula white. Ventral surfaces of fore and middle coxae yellowish.

Fore trochanter with basal segment yellow dorsally, fuscous ventrally; mid-

dle trochanter with basal segment fuscous, apical segment yellow; hind tro-

chanter with apical segment fuscous. Fore and middle femora and tibiae

fusco-ferrugineous. Hind femur dark fuscous basally, paler apically. Hind

tibia fuscous basally internally, and apically, with externo-median surface

fulvous. Hind tarsus fuscous.

Range and Variation. Antennal flagellum with 24-28 articles. Basal article

of flagellum approx. 3.7-3.8 x as long as wide. Lower 0.5 of mesopleuron

with fine to small punctures. Petiolar segment of abdomen weakly to mod-

erately decurving. Second abdominal tergite approx. 1.6-1.8 x as long as

wide. Hind femur approx. 5.2-5.4 x as long as deep. Ovipositor approx. 1.8-

1.9 x as long as hind femur. Fore wing approx. 4.0-5.0 mm. long. Femora

commonly more ferrugineous than in holotype, with fuscous areas confined

mainly to baso-ventral surface.

Male. Hind femur approx. 4.6-5.0 x as long as deep. Petiolar segment of

abdomen nearly straight. Coloured as female but with greater tendency to-

wards melanism particularly the legs . Second abdominal tergite approx.

1.8-2.0 x as long as wide. Hind tibia long, very slender.

Holotype. Female. Tangle Lakes, Alaska, VIJI-6-1973, H. & M. Townes.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. CANADA, British Columbia: Williams Lake, VIII-20-1960 (1 male)

B. Heming (CNC). Labrador: Hebron, VII-28-1954 (1 male), E.E. Sterns (CNC).

Northwest 'Territorieg:Kovaluk R., 69 11N131 W, July 6-10, 1971 (1 female),

W.R.M.Mason (CNC); 20 mi. e. Tuktoyaktuk, 13-16 July, 1971 (1 male), W.

R.M. Mason (CNC). Quebec: Payne Bay, 21-VII (1 male), 26-VII (1 male),

27-VIT (2 males), 11-VIII (1 female), 12-VIII (1 female), 19-VIII (1 female),

1958, W.R.M. Mason (CNC). Yukon: British Mts., Firth R., 24-VII-1956

(1 male), R. E. Leech (CNC). UNITED STATES. Alaska: Tangle Lakes

(1 male, 2 females), same data as for holotype (HMT).

Etymology: The species epithet is translated as 'rough' in reference to the

sides of the abdominal petiole.

Host and Biology. There are no rearing records for this species. Adults

are active during July and August and ther appears to be a single generation

per year.

Remarks. This species appears to be very closely related to variabilis but

18 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

it is easily distinguished from the latter on the basis of the following charac-

ters: sides of abdominal petiole rough; hind femur stouter; ovipositor longer

and more slender; abdomen never with a banding pattern.

C. asperata

f 500

Cymodusopsis macdunnoughi, new species

Fig. 10

Cmpd. eyes weakly notched opposite antennal sockets, weakly converg-

ent ventrad in females, with hairs of normal length. Propodeal carinae very

strong. Fore wing with areolet open. Ovipositor approx. 1.7 x aslong as hind

femur, weakly upcurving, with depth at mid length approx. equal to width of

hind basitarsus. Hind femur rather stout. Abdomen uniformly dark fuscous.

HOLOTYPE. Female. Antennal flagellum with 28 articles, the basal article

approx. 3.9 x as long as wide. Antennal scape with inner margin approx. 1.4 x

as long as outer margin. Cmpd. eyes weakly notched opposite antennal sock-

ets weakly convergent ventrad with hairs of normal length. Face strongly

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 19

shagreened, matte, with minute punctures. Tentorial pit separated from

inner margin of cmpd. eye by distance approx. equal to 2.0 x dia. of pit.

Clypeus strongly convex, its anterior margin moderately convex. Mesopleuron

with lower 0.5 strongly shagreened, matte, with small punctures which are |

separated by distance 1.0-2.0 x dia. of single puncture. Propodeum moder-

ately long, with apex projecting between bases of hind coxae a distance approx.

equal to 0.3 x the length of hind coxa. Propodeal carinae very strong. Areola

approx. 0.8 x as wide, 0.6 x as long as petiolar area, with surface granular.

Petiolar area with moderately coarse transverse wrinkles. Hind femur approx.

4.5 x as long as deep. Hind tarsal claw pectinate along basal 0.5. Petiolar

segment of abdomen straight, with petiole approx. 1.5 x as long as postpet-

iole, its sides smooth, polished. Ovipositor approx. 1.7 x as long as hind

femur, weakly upcurving, with depth at mid length approx. equal to width of

hind basitarsus. Fore wing approx. 3.6 mm. long, with areolet open. Hind

wing with nervellus weakly intercepted.

Antenna, head, thorax, abdomen, dorsal surface of fore coxa, entire

middle (except extreme apex) and hind coxae, black. Palpi, tegula white.

Ventral surface of fore coxa, fore trochanter, apex of middle coxa, middle

trochanter, apical segment of hind trochanter, yellow. Basal segment of hind

trochanter fuscous. Fore and middle femora ferrugineous; hind femur ferr-

ugineous with basal and apical infuscation. Fore tibia ferrugineous; middle

tibia ferrugineous, with weak apical infuscation; hind tibia with basal and apical

fuscous bands, the median band pale ferrugineous. Hind tarsus pale fuscous.

Range and Variation. Known only from holotype specimen .

Male. Hind femur approx. 4.0-4.5 x as long as deep. Hind tibia rather short

and stout. Coloured as female except that banding of hind tibia generally more

distinct.

Holotype. Female. Petite Riviere, Nova Scotia, 5-IX-1935, J. Macdunnoughi.

Deposited in Canadian National Insect Collection Ottawa.

Paratypes. Quebec: Park Reserve, Kam. Co., 950', 24-VIII-1957 (6 males),

W.R.M. Mason (CNC).

Etymology.This species is named in honour of J. Macdunnoughi, the collector

of the holotype specimen.

Host and Biology. Reared from unidentified microlepidoptera on cherry

(pupal case missing). This appears to be a late summer species.

Remarks. The weakly notched, weakly convergent cmpd. eyes, stout hind

femur, unusually short and stout hind tibia of males, in combination with the

uniformly coloured abdomen will distinguish this species from other nearctic

Cymodusopsis .

20 Contrib. Amer. Ent. Inst., vol. 22, no. 10, 1986

C. macdunnoughi

\ $00

Cymodusopsis latifacies, new species

Fig. i

Cmpd.eyes weakly convergent ventrad in female with hairs of normal

length. Face unusually broad, particularly in male. Clypeus unusually broad.

Tentorial pit separated from inner margin of cmpd. eye by distance approx.

equal to 4.0 x dia. of pit. Petiolar segment of abdomen weakly decurving.

Ovipositor approx. 1.8 xas long as hind femur, more or less straight, with

depth at mid length greater than width of hind basitarsus. Abdomen black.

HOLOTYPE. Female. Antennal flagellum with 26 articles, the basal article

approx. 4.2 x as long as wide. Antennal scape with inner margin approx.

1.4 x as long as outer margin. Cmpd. eyes weakly convergent ventrad , with

hairs of normal length, with lower facets approx. 1.4 x as wide as upper

facets. Face shagreened, matte, unusually broad, with minute punctures.

Tentorial pit separated from inner margin of cmpd. eye by distance approx.

equal to 4.0 x dia. of pit. Clypeus broad, moderately convex, its anterior

Sanborne: Revision of Nearctic Cymodusopsis (Ichneumonidae) 21

margin weakly convex. Mesopleuron with lower 0.5 strongly shagreened,

matte, with small punctures which are separated by distance less than or

equal to dia. of single puncture. Propodeum short, with apex projecting

between bases of hind coxae a distance approx. equal to 0.2 x length of hind —

coxa. Propodeal carinae very strong. Areola completely confluent with pet-

iolar area, approx. 0.4 x as wide, 0.5 x as long as petiolar area. Surface of

petiolar area polished, with moderately coarse transverse wrinkling. Hind

femur approx. 5.3 x as long as deep. Hind tarsal claw pectinate for slightly

more than half its length. Petiolar segment of abdomen weakly decurving,

the petiole approx. 1.6 x as long as postpetiole, its sides slightly rough,

polished. Second abdominal tergite approx. 1.5 x as long as wide. Ovipositor

approx. 1.8 x as long as hind femur, more or less straight, with depth at

mid length greater than width of hind basitarsus. Fore wing approx. 4.1 mm.

long, with areolet open. Hind wing with nervellus weakly intercepted.

Antenna head, thorax abdomen, dorsal surfaces of fore and middle

coxae, hind coxa, basal segment of middle and hind trochanters, black. Palpi

and tegula white. Fore , middle coxae with ventral surfaces yellow. Fore

trochanter yellow, with some infuscation on basal segment; middle trochanter

with basal segment mostly fuscous, apical segment yellow; hind trochanter with

apical segment fuscous basally, yellow apically. Fore femur pale ferrugineous;

middle femur ferrugineous, with small baso-ventral infuscated patch; hind

femur ferrugineous, with weak baso-ventral and apico-dorsal infuscated

areas. Fore tibia with sub-basal and apical bands, its inner surface ferrug-

ineous, its externo-median surface white; middle tibia coloured as fore tibia;

hind tibia with weak basal and apical fuscous bands, the median band yellow-

ferrugineous. Hind tarsus fuscous. Wings weakly infuscated.

Range and Variation. Antennal flagellum with 26-28 articles, the basal art-

icle approx. 4.2-4.4 x as long as wide. Hind femur approx. 5.3-6.1 x as long

as deep. Second abdominal tergite approx. 1.3-1.5 x as long as wide. Fore

Wing approx. 3.5-4.5 mm. long. Lower 0.5 of mesopleuron sometimes with

weak rugosities. No significant colour variation in specimens at hand.

Male. Face very broad. Hind femur approx. 5.6-5.8 x as long as deep.

Second abdominal tergite approx. 1.7-1.8 x as long as wide. Coloured as

female.

Holotype. Female. Ochoco Cr., Oregon, VII-11-1978, H. & M. Townes.

Deposited in Townes collection, Gainesville, Florida.

Paratypes. CANADA. Saskatchewan: Saskatoon, 9-V-1949 (2 females), A. R.

Brooks (CNC). UNITED STATES. California: Nevada Co., White Cloud

Campground, 30-31-V-1964 (13 males, 3 females), P. H. Arnaud Jr. (CAS).

Colorado: Boulder,4.5 mi. n., 5500' , June 20, 1961 (1 female), W. R. M.

Mason (CNC).

Etymology. The species epithet is in reference to the unusually broad face.

Host and Biology. There are no rearing records for this species. Adults are

active in spring and early summer.

Remarks. The broad face will easily distinguish this species from all nearctic

Cymodusopsis except brevicauda but is distinguished from the latter by its

(latifacies) small size , longer ovipositor (relative to hind femur), and black

abdomen.

22 Contrib. Amer. Ent. Inst. , vol. 22, no. 10, 1986

C. latifacies

500

ACKNOWLEDGMENTS

This project was funded in part by an NSERC (Natural Sciences and

Engineering Research Council) operating grant held by Dr. S. B. Peck,

Department of Biology, Carleton University, Ottawa. The remainder of funds

came from an NSERC post-doctoral fellowship held by the author. Mr. Luis

Ling produced the scanning electron photomicrographs.

REFERENCES

Sanborne, Michael. 1984. A revision of the world species of Sinophorus

Foerster (Hymenoptera: Ichneumonidae). Mem. Amer. Ent. Inst., no.

38. 403 pp.

Townes, Henry. 1970. The genera of Ichneumonidae, Part 3. Mem. Amer.

Ent. st, +) nes 19% 307 Dp.

Viereck, H. L. 1912. Descriptions of one new family, eight new genera and

thirty three new species of ichneumon- flies. U. S. Nat. Mus., Proc.

43: 575-593.

Cym odusopsis spp.: Figs. 1-2, frontal view of head: 1, melanosoma; 2, nigricincta.

Figs. 3-6, propodea: 3, aristoteliae; 4, albiscapa; 5, melanosoma; 6, nigricincta.

Cymodusopsis spp.: Figs. 7-11, propodea: 7, brevicauda; 8, variabilis; 9, asperata; 10,

macdunnoughi; 11, latifacies.

A REVISION OF THE WORLD SPECIES OF SINOPHORUS FOERSTER

(HYMENOPTERA: ICHNEUMONIDAE) : SUPPLEMENT 1

Michael Sanborne

Department of Biology, Carleton University

Ottawa, Ontario, Canada, K1S 5B6

ABSTRACT

Four new species of Sinophorus Foerster (Hymenoptera: Ichneumonidae)

are described from the palearctic region: albotibialis, amplificatus, centun-

culus, katoensis. New distributional and host data are included for tumidus

Sanborne and ventosus Sanborne.

This paper represents the first supplement to my recent revision of the

world species of Sinophorus (Sanborne, 1984). Additional supplements will

follow as new species are discovered. Four new species from the palearctic

region are described below (albotibialis, amplificatus, centunculus, katoensis)

and run through my key with explanations on how to differentiate them from

the species identified in the couplets. The last part of the paper deals with

new distribution and host records for two nearctic species (tumidus Sanborne,

ventosus Sanborne).

Sinophorus albotibialis, new sp.

DIAGNOSIS. Head approx. 2.1 x as wide as height of face and clypeus. Face

with medium sized punctures which are separated by distance less than the

dia. of single puncture. Lower 0.5 of mesopleuron with medium sized punc-

tures which are separated by distance less than or equal to dia. of single

puncture. Hind femur approx. 4.3-4.9 x as long as deep. First abdominal

tergite without a lateral, longitudinal groove. Suture separating 1st abd. ter-

gite from sternite approx. 0.5 below midheight of petiole in females, 0.7

below in males. Hind tarsal claw approx. 1.8 x as long as empodium, with

teeth confined to basal 0.5 of claw. Ovipositor approx. 2.0-2.1x as long as

hind femur. Tergites 2, 3, 4 of male with narrow apical white bands. Hind

tibia extensively white. Hind wing with nervellus intercepted.

ETYMOLOGY. The species epithet refers to the hind tibia which is extensive-

ly white.

HOLOTYPE. Female. Head approx. 2.1 x as wide as height of face and cly-

peus, broadly triangular in frontal view. Antennal flagellum with 33 articles.

Face with medium sized punctures which are separated by distance less than

dia. of single puncture. Clypeus polished, with medium sized punctures,

2 Contrib. Amer. Ent. Inst., vol. 22, no. 11, 1986

weakly convex, weakly separated from face, with anterior margin moderate-

ly convex. Anterior 0.4 of clypeus produced below lower margins of cmpd.

eyes. Mandible with upper tooth about as long as lower tooth. Gena approx.

as wide as basal width of mandible. Temple, in lateral view, approx. 0.9 x

as long as cmpd. eye, in dorsal view, moderately (approaching strongly)

sloping inwardly. Mesopleuron with coarse wrinkling between antero-dorsal

lobe and speculum which extends to mesopleural fovea. Speculum conspic-

uously punctate above. Mesopleural fovea elongate-oval, moderately imp-

ressed. Lower 0.5 of mesopleuron with medium sized punctures which are

separated by distance less than or equal to dia. of single puncture. Propod-

eum with basal transverse carina forming round-bottomed 'V' shaped border

around areola. Costula complete. Median longitudinal carina strong, regular

medially, moderately strong, regular at junction with basal transverse

carina. Lateral longitudinal carina absent. Propodeal trough narrow, deeply

excavated. Areola polished, approx. 0.3 x as long, 0.7 x as wide as petiolar

area, and 0.3 x as wide as distance between propodeal spiracles. Petiolar

area polished, with approx. 13 moderately coarse transverse wrinkles. Hind

femur approx. 4.5 x as long as deep. Hind tarsal claw approx. 1.8 x as long

as empodium, with 5 moderately strong, weakly slanting teeth which are

confined to basal 0.5 of claw. First abdominal tergite without a lateral long-

itudinal groove. Suture separating 1st abd. tergite from sternite approx. 0.5

below midheight of petiole. Petiole approx. 1.2 x as long as postpetiole.

Postpetiole with moderately dense pubescence. Ovipositor approx. 2.1.x as

long as hind femur, 1.4 x as long as thorax, weakly upcurving, not tapering,

with depth at midlength less than width of hind basitarsus. Tip of dorsal valve

of ovipositor approx. 1.1 x as long as 5th hind tarsal segment. Fore wing

approx. 5.8 mm. long, with 2nd recurrent vein straight , perpendicular.

Hind wing with nervellus intercepted near basal 0.2

Colour. Head, base and apex of mandible, thorax, abdomen, coxae, troch-

anters, black. Median area of mandible, maxillary palpus, tegula, external

surfaces of fore and middle tibiae, base and externo-median surface of hind

tibia, basal 0.6 of hind basitarsus, whitish. All tibiae, internal surfaces of

fore and middle tibiae, interno-median surface of hind tibia, ferrugineous.

Hind tibia with sub-basal band pale brown and apical band dark brown.

Labial palpus, apical 0.4 of hind basitarsus, brown.

Variation. Based upon one other female. Antennal flagellum with 33-34

articles. Costula complete or incomplete. Median longitudinal carina mod-

erately strong to strong, regular to irregular at junction with basal trans-

verse carina. Hind femur 4.3-4.5 x as long as deep.Ovipositor approx. 2.0-

2.1x as long as hind femur. No significant colour variation observed.

MALE. Based upon single specimen. As in female except that suture separ-

ating Ist abd. tergite from sternite approx. 0.7 below mid height of petiole,

hind basitarsus with basal 0.9 white, and abdomen with white apical bands on

tergites 2, 3, and 4.

Holotype. Female. U.S.S.R. Tashkent region, Parkent, V-20-1980, Kaspary-

an. Deposited in Zoological Institute, Academy of Sciences, Leningrad.

Paratypes. U.S.S.R. Tashkent region, same data as for holotype (ASL);

Turkmenistan, Krasno Znamya, Murgab River, 4-V-1980 (1 male), Kasparyan,

(ASL).

Sanborne: World species of Sinophorus: Suppl. 1 (Ichneumonidae) 3

Host and Biology. There are no rearing records for this species. The three

known specimens were all captured in the month of May.

REMARKS. This species belongs to the Pleuralis Group and is closely relate-

ed to punctifrons Sanborne (palearctic). This species traces to couplet 12

in my key (Sanborne, 1984) to palearctic females where it will key out to

punctifrons. It differs from this species in having small punctures on the face

and mesopleuron and in the extensively white hind tibia. The male traces to

couplet 14 where it keys to sticticalaefrom the palearctic region. It differs

from this species in the presence of apical white bands on tergites 2, 3, and

4 and in having a much less swollen hind femur. Some males may also trace

to couplet 15 where it will key to juniperinus. It differs from this species in

having a much narrower propodeal trough, much less swollen hind femur,

and much more extensively white hind tibia.

Sinophorus amplificatus, new sp.

DIAGNOSIS. Known only from female holotype. Head approx. 2.0 x as wide

as height of face and clypeus. Face with punctures which are separated by

distance less than dia. of single puncture. Mandible with upper tooth longer

than lower tooth. First abd. tergite with a very weakly impressed lateral,

longitudinal groove. Suture separating ist abd. tergite fron sternite approx.

0.4 below mid height of petiole. Hind femur approx. 3.9 x as long as deep.

Ovipositor approx. 2.0 x as long as hind femur, strongly upcurving, not

tapering. Hind wing with nervellus not intercepted. Hind femur without an

apical black band. Hind tibia black and ferrugineous banded.

ETYMOLOGY. The species epithet is fromthe latin word meaning enlarged,

in reference to the swollen hind femur. |

HOLOTYPE. Female. Head approx. 2.0 x as wide as height of face and clyp-

eus, triangular in frontal view. Face moderately rugoso-punctate, the punc-

tures of medium size and separated by distance less than dia. of single punc-

ture. Clypeus strongly shagreened, matte, moderately convex, with medium

sized punctures which are separated by distance less than dia. of single punc-

ture. Anterior 0.4 of clypeus produced below lower margins of cmpd. eyes.

Mandible with upper tooth longer than lower tooth. Gena approx. 1.2 x as

wide as basal width of mandible. Temple, in lateral view, approx. as long

as cmpd. eye, in dorsal view, moderately sloping inwardly. Mesopleuron

with moderately coarse wrinkling between antero-lateral lobe and speculum

which extends to mesopleural fovea. Speculum densely punctate above.

Mesopleural fovea elongate-oval, weakly impressed. Lower 0.5 of meso-

pleuron strongly shagreened, matte, with medium sized punctures which are

separated by distance less than or equal to dia. of single puncture. Propodeum

with basal transverse carina forming flat-bottomed 'V' shaped border around

areola. Costula present laterally. Median longitudinal carina strong medial-

ly, weal and highly irregular at junction with basal transverse carina. Lat-

eral longitudinal carina absent. Propodeal trough moderately broad, deeply

excavated. Areola strongly shagreened, with several transverse wrinkles,

approx. 0.3 x as long and 0.6 x as wide as petiolar area, and 0.4 x as wide

as distance between propodeal spiracles. Hind femur approx. 3.9 x as long

4. Contrib. Amer. Ent. Inst., vol. 22, no. 11, 1986

as deep. Hind tarsal claw approx. 1.5 x as long as empodium, with 5 mod-

erately strong teeth which are confined to basal 0.5 of claw. First abd. ter-

gite with a very weakly impressed lateral, longitudinal groove. Suture sep-

arating 1st abd. tergite from sternite approx. 0.4 below mid height of petiole.

Petiole approx. 1.2 x as long as postpetiole. Postpetiole with sparse dorsal

pubescence. Ovipositor approx. 2.0 x as long as hind femur, strongly up-

curving, not tapering, with depth at mid length slightly less than width of

hind basitarsus. Tip of dorsal valve of ovipositor approx. 0.8 x as long as

5th hind tarsal segment. Fore wing approx. 6.0 mm. Hind wing with nerv-

ellus not intercepted.

Colour. Head, base and apex of mandible, thorax, abdomen, coxae, middle

and hind trochanters, apical 0.6 of hind basitarsus, black. Tegula, basal 0.4

of hind basitarsus, whitish. Maxillary palpus pale ferrugineous, labial palpus

brown basally, ferrugineous apically. Fore trochanter with basal segment

dark ferrugineous internally, yellowish externally. Femora, internal surface

of fore tibia, middle tibia, median band of hind tibia, ferrugineous. Hind

tibia with sub-basal and apical bands brown.

Holotype. Female. U.S.S.R. Primorskij Kraj, Khasan, Golubin, Utes, 7-8.

9.1982, Tobias. Deposited in zoological Institute, Academy of Sciences,

Leningrad.

Host and Biology. Unknown

REMARKS, This species is very closely related to the nearctic species

constrictus but is easily distinguished by the longer, more slender, much

more strongly upcurving ovipositor. This species traces to fuscicarpus

(coupiet 7) in my key to palearctic females and differs from this species in

having black and ferrugineous hind legs and hind wing with nervellus not

intercepted.

Sinophorus centunculus, new sp.

DIAGNOSIS. Known only from female holotype. Head approx. 2.2 x as wide

as height of face and clypeus. Face with small punctures which are separated

by distance less than dia. of single puncture. Mesopleural fovea sub-round,

deeply impressed. Lower 0.5 of mesopleuron with medium sized punctures

which are separated by distance less than dia. of single puncture. Hind femur

approx. 4.4.x as long as deep. First abd. tergite with a moderately impress-

ed lateral, longitudinal groove. Suture separating ist abd. tergite from ster-

nit approx. 0.2 below mid height of petiole. Ovipositor approx. 1.8 x as long

as hind femur, weakly tapering. Hind femur with baso-ventral area and apex

black. Hind tibia black and ferrugineous banded.

ETYMOLOGY. The species epithet is from the latin word meaning 'small

patch' , in reference to the baso-ventral black patch on the hind femur.

HOLOTYPE. Female. Head approx. 2.2 x as wide as height of face and cly-

peus, triangular in frontal view. Antennal flagellum with 32 articles. Face

with small punctures which are separated by distance less than dia. of single

puncture. Clypeus shagreened, matte, weakly but evenly convex, weakly

Sanborne: World species of Sinophorus: Suppl. 1 (Ichneumonidae) 5

separated from face, with anterior margin nearly straight. Anterior 0.3 of

clypeus produced below lower margins of cmpd. eyes. Mandible with upper

tooth longer than lower tooth. Gena approx. as wide as basal width of mandible.

Temple, in lateral view, approx. 1.1x as long as cmpd. eye, in dorsal view,

moderately sloping inwardly. Mesopleuron with moderately coarse wrinkling

between antero-lateral lobe and speculum which does not extend to mesopleural

fovea. Speculum conspicuously punctate above. Mesopleural fovea sub-round,

deeply impressed. Lower 0.5 of mesopleur on with medium sized punctures _

which are separated by distance less than dia. of single puncture. Propodeum

with basal transverse carina forming 'V' shaped border around areola. Costula

absent. Median longitudinal carina moderately strong medially, weak and

highly irregular basally at junction with basal transverse carina. Lateral

longitudinal carina absent. Propodeal trough broad, deeply excavated. Areola

approx. 0.4 x as long, 0.7 x as wide as petiolar area, and 0.4 x as wide as

distance between propodeal spiracles, with several weak transverse wrinkles.

Petiolar area with approx. 15 moderately strong transverse wrinkles. Hind

femur approx. 4.4 x as long as deep. Hind tarsal claw approx. 1.4 x as long

as empodium, with 5 moderately strong teeth which do not extend to base of

tip. First abd. tergite with a moderately impressed lateral, longitudinal

groove. Suture separating 1st abd. tergite from sternite approx. 0.2 below

mid height of petiole. Petiole approx. 1.4 x as long as postpetiole. Postpet-

iole with very sparse dorsal pubescence. Ovipositor appr ox. 1.8 x as long as

hind femur, 1.4 x as long as thorax, moderately upcurving, very weakly taper-

ing towards tip, with depth at mid length approx. equal to width of hind basi-

tarsus. Tip of dorsal valve of ovipositor approx. 1.2 x as long as 5th hind

tarsal segment. Fore wing approx. 5.0 mm. Hind wing with nervellus not

intercepted.

Colour. Head, base and apex of mandible, thorax, abdomen, coxae, troch-

anters, base of middle femur, baso-ventral patch and apex of hind femur,

internal surface of middle tibia, sub-basal and apical bands of hind tibia,

apical 0.7 of hind basitarsus, black. Tegula, labial palpus, brown. Median

area of mandible, maxillary palpus, fore femur, middle femur, hind femur

(except black areas noted), fore tibia, external surface of middle tibia, med-

ian band of hind tibia, ferrugineous. Basal 0.3 of hind basitarsus, whitish.

Holotype. Female. U. S. S. R. Alma-ata, 2500-2800 m., 7-VII-1979, Kasp-

aryan. Deposited in Zoological Institute, Academy of Sciences, Leningrad.

Host and Biology . Unknown.

REMARKS, This species is related to nigritellus and villosus (Albidus Group).

It differs from both species in the ferrugineous colour of the median band of

the hind tibia and from villosus in the longer, more strongly upcurving ovi-

positor. The species will trace to couplet 18 in my key to palearctic females

where it would best fit the description given for 18a. This will lead to couplet

19 where it would best fit the description of turionus. The much shorter ovi-

positor will easily separate centunculus from turionus.

6 Contrib. Amer. Ent. Inst., vol. 22, no. 11, 1986

Sinophorus katoensis, new sp.

DIAGNOSIS. Head 2.0-2.1 x as wide as height of face and clypeus. Face and

lower 0.5 of mesopleuron with small punctures which are separated by dis-

tance less than or equal to dia. of single puncture. Hind femur approx. 4.0-

4.3 x as long as deep in males, 4.3-4.5 x in females. First abd. tergite with

a weakly impressed lateral longitudinal groove. Suture separating ist abd.

tergite from sternite approx. 0.2-0.3 below mid height of petiole in females,

0.3-0.4 below in males. Ovipositor approx. 2.0-2.1x as long as hind femur,

not tapering, moderately upcurving. Hind wing with nervellus intercepted.

Abdomen of females with postpetiole and abd. tergites 2,3 rufous but some-

times abdomen entirely black. Male abdomen with narrow apical white bands

on tergites 2-5. Hind femur entirely ferrugineous in females. Hind tibia

black and white banded.

ETYMOLOGY. The species is named for the holotype collecting locality of

Kato, Manchuria.

HOLOTYPE. Female. Head approx. 2.0 x as wide as height of face and cly-

peus, triangular in frontal view. Antennal flagellum with 35 articles. Face

with small punctures which are separated by distance less than or equal to

dia. of single puncture. Clypeus strongly shagreened, matte, with small

punctures which are separated by distance less than or equal to dia. of

single puncture, weakly separated from face, with anterior margin weakly

convex, the anterior 0.3 of clypeus produced below lower margins of cmpd.

eyes. Mandible with upper tooth longer than lower tooth. Gena approx. as

wide as basal width of mandible. Temple, in lateral view, approx. 0.6 x as

long as cmpd. eye, in dorsal view, strongly sloping inwardly. Mesopleuron

with weak wrinkling between antero-dorsal lobe and speculum which extends

to mesopleural fovea. Speculum conspicuously punctate above. Mesopleural

fovea sub-round, deeply impressed. Lower 0.5 of mesopleuron strongly

shagreened, matte, with small punctures which are separated by distance

approx. equal to dia. of single puncture. Propodeum with basal transverse

carina forming round-bottomed 'V' shaped border around areola. Costula

complete. Median longitudinal carina moderately strong. Lateral longitudinal

carina moderately strong, highly irregular. Propodeal trough narrow basal-

ly, moderately broad apically, moderately excavated. Areola shagreened,

weakly polished, approx. 0.3 x as wide, 0.6 x as long as petiolar area, and

0.3 x as wide as distance between propodeal spiracles. Petiolar area with

approx. 18 moderately coarse transverse wrinkles. Hind femur approx. 4.2.x

as long as deep. Hind tarsal claw approx. 1.3 x as long as empodium, with

6 moderately strong, erect teeth which extend slightly beyond basal 0.5 of

claw. First abd. tergite with a moderately impressed lateral, longitudinal

groove. Suture separating 1st abd. tergite from sternite approx. 0.3 below

mid height of petiole. Petiole approx. 1.4 x as long as postpetiole. Post-

petiole with sparse dorsal pubescence. Ovipositor aprox. 2.2 x as long as

hind femur, not tapering, moderately upcurving, with depth at mid length

approx. equal to width of hind basitarsus. Tip of dorsal valve of ovipositor

approx. 1.1 x as long as 5th hind tarsal segment. Fore wing approx. 6.0 mm.

long. Second recurrent vein moderately sloping. Hind wing with nervellus

intercepted near basal 0.2.

Sanborne: World species of Sinophorus: Suppl. 1 (Ichneumonidae) 7

Colour. Head, base and apex of mandible, abdominal petiole, abdominal

tergites 4 to apex, sub-basal and apical bands of hind tibia, apical 0.3 of

hind basitarsus, black. Antennae fuscous. Median area of mandible, palpi,

tegula, external surfaces of fore and middle tibiae, externo-median surface

of hind tibia, basal 0.7 of hind basitarsus, whitish. Trochanters, interno-

median surface of hind tibia, pale ferrugineous. Coxae, femora, internal

surfaces of fore and middle tibiae, ferrugineous. Postpetiole, tergites 1 and

2, rufous.

Variation. Head 2.0-2.1 x as wide as height of face and clypeus. Antennal

flagellum with 34-37 articles. Propodeum with areola weakly polished to

matte. Hind femur 4.2-4.5 x as long as deep. Ovipositor approx. 2.0-2.1x

as long as hind femur. Fore, middle coxae sometimes replaced dorsally by

black. Hind femur sometimes with weak apical dark band. Postpetiole rufous

to black. Abdomen sometimes entirely black.

MALE, Head approx. 2.0 x as wide as height of face and clypeus. Hind femur

approx. 4.0-4.3 x as long as deep. Suture separating Ist abd. tergite from

sternite approx. 0.3-0.4 below mid height of petiole. Colour generally more

melanistic than female. Palpi white to ferugineous. Hind coxa extensively to

entirely black Hind femur usually with distinct apical dark band. Sub-basal

and apical bands of hind tibia wider than in female. Petiolar segment of abd.

black. Abdominal tergites 2-5 often with narrow apical white bands or abd-

omen entirely black.

Holotype. Female. China: Manchuria, Kato, XII.2. ?, ex. Pyrausta nubilalis .

Deposited in Canadian National Insect Collection, Ottawa, Canada.

Paratypes. 78 females, 30 males. Same data as for holotype except that em.

dates range from Dec. to Feb. Two females, three males deposited in P.M.

Sanborne collection, the remainder in the United States National Museum,

Washington.

Host and Biology. All known specimens were reared from Pyrausta nubilalis

REMARKS. This species belongs in the Xanthostomus Group and is very

closely related to exartemae (palearctic). It will trace to exartemae (couplet

37) in my key to the palearctic females. It differs from this species primarily

in colour (abdomens of katoensis often extensively rufous, abdomens of

exartemae always black). Females of katoensis with entirely black abdomens

can be distinguished from exartemae by the entirely ferrugineous hind femur

(sometimes a very weak dark band present, this band always distinct in

exartemae). Males of katoensis will key out as exartemae and I have not

been able to find characters which will distinguish between the two.

New Distribution and Host Records

Sinophorus tumidus Sanborne: Host. Homeosoma electellum (Lepidoptera)

on wild sunflower. New Distribution Records. Ontario: Cumberland, Stitts-

ville. Saskatchewan: Bienfait, Trossachs, Weyburn.

Sinophorus ventosus Sanborne: Host. Stibadium spumosum (lepidoptera) on

Helianthus annuus (sunflower). New Distribution Record. Texas: Bushland.

8 Contrib. Amer. Ent. Inst., vo]. 22, no. 11, 1986

ACKNOWLEDGEMENTS

This research was funded in part by an NSERC (Natural Sciences and

Engineering Research Council) operating grant held by Dr. S. B. Peck,

Department of Biology, Carleton University, Ottawa. The remainder of funds

came from an NSERC post-doctoral fellowship held by the author. Special

thanks go to Dr. D. R. Kasparyan for providing me with specimens of albo-

tibialis, amplificatus, and centunculus, to Dr. C. E. Rogers for providing

reared specimens of ventosus, and to Mr. Gene Bisdee (Biosystematics

Research Institute, Ottawa) who provided me with reared specimens of tum-

idus .Specimens of katoensis came from the Canadian National Insect Coll-

ection (Dr. John Barron) and the United States National Museum.

REFERENCES

Sanborne, Michael. 1984. A revision of the world species of Sinophorus

Foerster (Hymenoptera: Ichneumonidae). Mem. Amer. Ent. Inst., no.

38. 403 pp.

ERRATA

World Species of Sinophorus. Mem. Amer. Ent. Inst., no. 38.

P. 33. Para. 2, line 11. 'makes' should read 'makers' (2 x).

P. 39. Para. 3, line 4. Underiine Sinophorus and Campoplex.

P. 40. Couplet la. Place brackets around Fig. 148 (i.e., (Fig. 148)).

Couplet 5b. Fibs. should read Figs.

P. 45. Couplet 38b. (50) should read (51).

P. 48. Couplet 57b. 2.1-2.0 should read 2.1-3.0

P. 61. Couplet 2b. 1.9-2.0 should read 2.1-2.7

P. 68. Couplet 9b. 2.1-1.7 should read 2.1-2.7

P. 224. Para. 1, line 4. After first ' distance ' stroke out 'approx. distance’

| line 6. Remove second 'than lower'

P. 305. Para. 2, line 2. Remove 'face' at end of line.

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