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Contributions ,

of the

American Entomological Institute

_ ot.

Volume 18, 1981

“Tr

Howden, H. F. & O. P. Young. Panamanian

Scarabaeinae: Taxonomy, distribution, and habits

(Coleoptera, Scarabaeidae). 204 pages. Feb. 25, 1981.

Hopper, H. Pearson. A revision of the genus

Aphanistes of America north of Mexico (Hymenoptera:

Ichneumonidae). 56 pages. September 24, 1981.

Michener, Charles D. Classification of the bee family

Melittidae with a review of species of Meganomiinae.

135 pages. December 23, 1981.

Reinert, John F. Medical Entomology Studies - XV.

A revision of the subgenus Paraedes of the genus Aedes

(Diptera: Culicidae). 91 pages. November 10, 1981.

Contributions

of the

American Entomological Institute

Volume 18, Number 1, 1981

PANAMANIAN SCARABAEINAE:

TAXONOMY, DISTRIBUTION, AND HABITS

(COLEOPTERA, SCARABAEIDAE)

H. F. Howden and O. P. Young

PANAMANIAN SCARABAEINAE:

TAXONOMY, DISTRIBUTION, AND HABLTS

(COLEOPTERA, SCARABAEIDAE)

Hee. Howden! and O.P. Young”

ABSTRACT

Twenty-one genera of Scarabaeinae that occur in Panama are

briefly reviewed. In addition the genus Cryptocanthon is included

sluice it occurs Om eilher Gide of Panama. A total of 113 species

are recognized with twenty-eight described as new. The old and

new taxa ’are listed below in the’Contents. © Zoogeoscraphic and

ecological data were compiled from a total of 7113 Panamanian speci-

mens bearing accurate data.

CONTENTS

Introduction . i

Nomenclatorial Limitations 1

Materia! Pxamined i). . (ate. ia Ge se ee

Collections Studied 2

Gazetteer of Localities 3

Oe em rk ek ee ea ee ee ecg NL 3 8

Denny (OO. (ie OCA ataetide Fo ek eee ee ee ee. 8 Ed

Ney 4@ Genera, Of Sear apaeitiae Occurring in Panama... . .. 11

Dey eee Ue ee On we a Gee a 9

Crave ane: Hirsg oe. oe Momitliiatus Bales ss so... 22

Drove iG Hats: ae a cee ae 5 septemmaculatus (Latr.) . 23

Vemiads Fates 2 le. ae dewroUer Wat.) ek a ce 24

Memon Mars Gop a ce eG LG ae Cle vrclals War. so. s . 24

cearloeete Chest.) wn tw OLY Piuraius (Germ..)> . 0... .<,25

Peers Pape ee ete oe 1S fie pllie Lucas, ... . 4 26

eee Taisen ow ws, ZO

pegs eos ltd ~ Mequinecriatis Har, . 2°. 627

Candee (Har) fie. . trees LS SENSE Dn ae 3

lDepartment Of Biolocy, Carteton University, Ottawa, Ontario,

KIS SB6, Canada:

2Department of Entomology. University of Maryland, College

Park, Maryland 20742. U.G.A..

Ateuchus

seminudus

aeneomicans

Contrib. Amer.

morsel group.

angustatus Har.

subhyalinus Har.

euryscelis Bates .

femoralis (Chev. )

lamprimus Bates .

caelius Bates

v. meridionalis (M.,

Pseudocanthon

perplexus (LeC.)

Deltochilum

@. panamensis Howd.

mexicanum Burm.

Darive Bates.

pseudoparile Paul.

V. acropyge Bates .

Cryptocanthon

Agamopus

lampros Bates

Pedaridium ~,

pilosum (Rob.)

previsetosum MO. sp.

boitimeri n. sp.

Bdelyrus

(Bates }

Onthocharis

panamensis Paul.

Scatimus

quadridentatus Balth.

ovatus Har. ;

Uroxys

gorgon Arrow

Mela COrgon 1. SDs...

gepressiirons 1. 6D. 4

micros Bates

microcularis n. sp.

platypyga n. sp.

macrocularis n. sp.

bidentis n. sp.

Balunensis 1. 8p.

dvybasl Nn. Sp,;

(Mae)

(Hav)

candezei

Brnt.

ano

frist. ,: vol. 10,¢moer 11981

29 guatemalensis (Bates) ,

oO - Canthidtum i 7: :

centrale “houc.::

32 tuberifrons m1. sp: .

33 haroldi ee :

angusticeps Bates

elegantulum Balth.

bituberifrons mn. sp...

3D discopygidiale n. sp.

35 subdopuncticolle n.

lene Drosum 1. sp...

36 planovultum n. sp.

37 DELCEDLIDIIe 1. Sp,

37 vVesperiinum 1. sp.

38 pallidoalatum n. sp.

38 variolosum n. sp.

39 leucopterum 1. 6p.

ardens Bates

39 aurifex Bates

40 hespenheidei n. “sp.

a Onthophagus

incensus oay...

7 Davest th. aC.

43 Cc. panamensis Bates.

44 stockwelli n. sp.

45 nyctopus Bates

i acuminatus Har.

transisthmius n. sp.

46 marginicollis Har.

lebasi, Bouc.

oy chryses Bates

48 coscineus Bates

48 dicranius Bates

landolti Har. ‘

49 atrosericeus Bouc. .

50 cyanellus Bates

50 mirabilis Bates

Sharpi Har.

a2 praecellens Bates

54 belorhinus Bates.

= Ontherus .

ze brevipennis Har.

0) didymus Er.

62 Sirius, M., SDs

Pe Dichotomius

66 C., eolonicus (Say),

68 nevermanni (Lued.)

satanas (Har.)

69 agenor (Har.)

69 inachus (Er.)

Howden and Young: Panamanian Scarabaeinae Ti

POMOPAIUS 1. Gis Gye Wee we ee pe See eg 128

CAMVOGCISVE Ws SPa sc ee eye ene ome cc. Bete alee 129

CES ee eke oa Pe ee CO ea ee Ne a Gece 130

TW DTS ROU a eee col ee Ee NS arta ba gc eels 132

TG GUS ee ee See ye a RN sa £33

POSPACTOCCMSTS Gaia cc cee ey Mey it 2 a aed pueeeuh ina ta 153

DO ee Pe ee alee awa eek ees 134

Phanaeus Orde Nie oct ae pene UC ESS es Wen Coro uO Ng Ney te. @ Ova cw 6 A

heemee a a ee ee a Ga, 135

Deltans Bares fee er ee te tee ee 15

amore Bet reS i oe Oa. ee ed Gee ae BO

UME OM MATS Wie gy Wi Get St ee bug ae ede ek ew ee es as we caer 13/

Sy CU RG a ie a 138

UCL ere UNS NR er ee ae 18

ye Ve ise ee e189

OU a ee a a a ee 140

Ro Cor hte he Oe eG ea AE

See ee OR

Oa TENG EE gel tac aah ho ESRD LN uO as Ate igen PVA aE ge enertny

COCO aT OTC PY LING I Ge Sag citar ice iol GRRE He eg heh ee ee ae Ore 144

OPUS OM GO en a a We a na i eho roe dO

One Or VAG Wee et a a oO

Biocon Per... Aiea das ae Se ak gs 147

HOpe@rapnic Peatures ond. faunal Ratierns. 6. (sake) bel sles 148

Habits): Comments, on the behavior Ola Few Species oo. .ier 152

MTC W MOCO MCT EG Rl ayia ue ee Sg a week aa BD

Literature Cited : Se ake aca he

POGOe 2 FOO Oe oe en BURR ore le ea eas 1684 204

PANAMANIAN SCARABAEINAE:

TAXONOMY; (DUSTRIBUTION., (RAND HABITS

(COLEOPTERA, SCARABAEIDAE,)

Hw PF. sHowden andwdol.. wove

INTRODUCTION

In the last decade interest in the biology and Dehavior of

dung beetles belonging to the subfamily Scarabaeinae has increased

greatly.: This -has been due, in parlivtomi ne tmicrest yin. tneie wiility

both as a natural sanitation system and as an agent for pasture

improvement (e.g., Weeda, 1967). From the ecological point of view,

the various methods utilized by the beetles for partitioning, the dune

resource are extremely interesting: and. cemplex,.., However, im order

to adequately study and document the habits of the Scarabaeinae

for any geographic area the species must be taxonomically well

known. The inadequate taxonomic status of the Panamanian

Scarabaeinae stimulated the initiation of this review. The only

previous publication that has attempted to enumerate and describe

the Panamanian species of Scarabaeinae is the Scarabaeidae volume

by H.W. Bates (1886-1890) of the Biologia Centrali-Americana. In

this volume Bates lists 58 species from Panama. The present survey

reviews 1135 species in 21 genera, elving Drier deccrio tons. Keys,

and biological information for the Panamanian species.

NOMENCLATORIAL LIMITATIONS

Since this work is intended as a review and not as a revi-

sion, the reader should recognize the inherent limitations in the

application of the scientific names used. Of primary importance

is the recognition that nomenclature and taxonomy are not in-

separable. For Panama we hope that our taxonomy is reasonably

good although our nomenclature, in some cases, is doubtful.

Taxonomically we have attempted to define the species, including

variation, from specimens taken in Panama. We have not included

in the descriptions non—-Panamanian material (as should be done

in a revision). In many cases specimens from Costa Rica or

Colombia differ slightly and consistently from Panamanian speci-

mens. Frequently we have briefly mentioned these differences.

However, when describing a new species we have excluded most

non-Panamanian material from the type series because of these

small but consistent differences. From the nomenclatorial stand-

point we have attempted to apply names as broadly as possible.

If a species was described from Colombia and a slieht wariant

occurs in Panama, we have used the available name and avoided

proposing a new name.

Correct nomenclature often depends on the examination of

types, and here again we lack the degree of accuracy that one

would expect to find in a revision. However, a number of the

types have been seen as noted in the descriptive section. The

2 Contrib: “Amer. Ent. insic, vol. 10, mos. 15° 1961

Biologia material has been carefully ctudied...but where no type

has been designated for a cotypic series, we have refrained from

doing ‘so. Lectotype selection should await. proper revisionary

studies.

if our nomenclature sliould prove to Be inaccurate, it can be

changed. This shoulda be a Yelatively favre necessity since a

reasonavle eliort has been: made to verify the names used herein.

MATERIAL EXAMINED

The specimens séen were derived largely from two sources.

The first source was museum holdings accumulated over a period

of “years. “hese accumulations, “while oftem lacking ecological

data Othér than date and locality, were valuable both taxonomi-—

Cally and historically. in some €ases, due to clearing of land

or-other types of habitat “disturbance, a species’ may no longer

occur in “an “area. Thus.° some of the older material was’ of ‘con-

siderabie historical importance. ~The sécond: source of specimens

consisted of recent Intensive: collecting efforts: by* both ecologists

and systematists. The authors alone totalled nearly two man

years of collécting and ‘observations’ in Panama, and’ numerous

individuals, as mentioned in the Acknowledgments, assisted us.

Much of the biological information discussed was accumulated by

this, recent effort.

COLLECTIONS STUDIED

In. citing. the collections studied we utilize the name of the

Cily if which the museum is situated, and for a private. collection,

we cite the individual's last name. The museum collections

studied and the curators assisting are as follows:

Cambridge Museum of Comparative Zoology, Harvard

University. As. Newton;

Chicago Field Museum of Natural history. 7? Hoc Dybas.

London British Museum (Natural History). R. Pope.

Ottawa Canadian National Collection, Biosystematics

Research Institute. A. Smetana.

Paris Museum National d'Histoire Naturelle. A.

Descarpentries.

Washington United States National Museum. R. Gordon.

Types cited as: "Howden". are deposited on? l6an ‘in the

Canadian National Collection, Biosystematics Research Institute,

Canadian’ Department’ of “Agriculture, Ottawa, Canada.

Howden and Young: Panamanian Scarabaeinae &

GAZETTEER OF (LOCALILE GES

Because one of the objectives of this survey is te encourage

biological investigations in Panama, we have devoted considerable

effort to organizing and presentine the 1204 toeeliiies tnat are

represented inwour examined material, A few tocaliiy mames..were

added after the manuscript was essentially compiete: these are

not included™iw Table 1.7, lf nothing else, Tabje |= combined wit

Figure 1 should indicate -how unevenly entomolegists have collected

ins Panama.

LOCALITY “DATA

Most of the place names used in this review are slandarca

names approved by. the Board on Geographic Names; U.S. Depari—

ment of the Interior, as published in 1969 as Official Standard

Names Gazetteer No. 110, Panama and the Canal Zone. The

Panama Gazetteer provided latitude and longitude coordinates,

which, according to. its. procedures, ‘were read: at the map

symbol for populated places and other entities occupying limited

sites, at the mouths or lower ends of streams, at the summits of

mountains and hills, at the extremities of capes and poinis, and

near the centers or midpoints of other features''. Several names

not found in the Panama Gazetteer were located in The Gazetteer

of Selander’ and Vaurie (1962). ° Localities that could not be

authenticated in a gazetteer were usually located on one of several

maps, such as the’ Esso Standard’ Oil road map of | La hepulbaca

de Panama", 1964. To determine each locality's elevation above

sea level and its relation to nearby major features, several other

Maps ‘were, utilized, These included U.o.) frmy Corps, ot Enoaineers,

Army Map Service, Series: #1301-- 1:1,000,000 - Panama and

Central America; #EZ6)l——1-500,000 = Panama: @rbol-— 1-100 7000. —

Canal Zone; #E761-- 1:50,000 - Panama, Topographical.

The Army Map Series #E761 maps provided exact or contour

elevation data, but unfortunately only central Panama has been

mapped at a. scalée.’of.):50,000." “Hence, for much or Panama,

Army Map Series #E461 was used. This map, however, indicated

elevations in zones of 0-200 meters, 200-500 meters, 500-1000 meters,

1000-2000 meters, and »2000 meters. Although mountain elevations

are for the summit and river elevations are for the mouth, it is

understood that specimens collected at mountains or rivers may not

have been’ at the elevation indicated in the locality list. Distance

méasurements given imuthe locality list (but not on label’ data)

are straightline measurements obtained from maps and are not

road distances. Locality names found on specimen labels that can

not be authenticated and located by coordinates are not listed in

the main table, but are given separately. When necessary, local-

ity names on specimen labels were corrected for spelling before

listing.

Contrib. Amer. Ent. Inst., vol. 18, no. 1, 1981

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Howden and Young: Panamanian Scarabaeinae he

TAXONOMY OF THE SCARABAEINAE

Panamanian species belonging to the subfamily may be dis-

tinguished from other Scarabaeidae by the following combination of

characters: antenna nine or ten segmented with a three segmented

club; mandible and maxilla partly membranous; mandibles not

visible from above, clypeus large;, pygidium exposed, not covered

by elytra; mesosternum short; middle coxae widely separated,

obliquely placed: spiracles presént in’ lateral membranes o1

abdomen; posterior tibia with a single apical spur; tarsal claws,

when present, simple. |

The species are primarily dung and/or carrion feeders, at

least 19 the adult" stage,

Key to Genera of Scarabacinae

occurring in Panama

ds ocutéellum clearly wisible, between ace Ol Siviras . . 2

Scutellum not visible between bases of elytra . ... 3

Zl). Clypeus shallowly, broadly emarginate; body

rectangularly elongate, flattened dorsally . eee

CA Pee ae ae . Eurysternus Dalm. (p. 14)

Clypeus anteriorly bidentate; body shape oval,

dorsally*convex:) 4, Meee ieposom: eal (p. 18)

Be). Middle and hind tibiae dilated apically or broad

and ‘flattened; hind tibia’ if not; dilated’ ai apex

then at basal third ‘distinctly vectamawilar in

CVOSS: SCCUIOI. 4 OR ge eee eas 4

Middle and hind tibiae slender; “frequently curved,

scarcely: enlarged apically; hind tibia at basal

third trapezoidal in ¢ross Gection’ <9) 7 4 secre. 19

ENO) Fore tibia with four “teeth on outer mergin: hind

tarsus with basal segment not expanded

apically, pie ay as long as tibial spur Peete

: Onthophagus: Latr. tp. 93)

Fore tibia with ‘three or four teeth on outer” maroin,

if with four teeth hind basal tarsal segment

expanded apically, moteleader sala. Bre ga ek 8

Aa, Anterior tarsus absent, or if present without claws;

basal segment of antennal club hollowed,

receiving other segmonits ity. * BCE Te 7 I ae 6

Anterior tarsus present and with claws: ‘basal

segment or antennal-iclub not distally COncCavete a ws 9

(5). Antero-median angle of metasternum prolonged as

a long, ‘sharp spine “extending between “apices of

the ‘front’ coxae? postero-median ‘edée of pronotum,

distinctly aneulate between’ pases ‘of ‘elytra’ Shu...)

Oxysternon Lap. (oe 146) }

19).

LAO)

Lett).

13Ct2).

Contribs (Amer) ints «dusty. woh ike, vino. bs .1961

Antero-median angle of metasternum not spiniform;

postero-median edge of pronotum not distinctly

NOTA G iy pies aha Van ts ee Pw petA Cw DUG War Eo uis made aul gh Sle oe oi agi 7

Anterior clypeal margin with three indentations

delimiting two acute teeth ae aoe

Coprophanaeus roe pe 40)

Anterior clypeal margin with.a single median

indentation, sometimes obsolete; margin lacking

USGL ING ly late Ne Om ue i I og 8

Outer pronotal margin just posterior to anterior

angle notched or abruptly sinuate; anterior

pronotal marginal bead not broken behind eye;

promotum Of malo” NOL Tlattenea, not triangular

LY SAPs hate, oo oo. oulcophanaeus Ols.. <(p. 137)

Outer pronotal margin posterior to anterior angle

not notched nor abruptly sinuate; anterior

pronotal marginal bead broken just behind eye;

pronotum- of imale. usually flattened, triangular

(Shee oe oe oe OR haneeus, Nac lL, (poe baa)

Elytron with 7 or 8 discal striae (a 9th sometimes

present on epipleuron), or striae obscured by

dense punctures and/or setae; median longitudinal

promolal “oroeve weakly developed or absent... . . 10

Elytron with 9 discal striae (10th on epipleuron);

median longitudinal pronotal groove usually

sirongly developed 4) o4.\,i:7 0... Copris.Geof. hoe weed)

Entire dorsal surface moderately to densely

punctate, or with dorsal setae on head, pronotum

ane -elyiras uc). oe Wee el tas eer ame ea a LL

Dorsal surface not as above Sr aise Gk iee an 6. dhe

Body shape rectangular; elytral intervals not

distinctly convex; anterior. margin of clypeus

NOL stawply dentate 4). a0. Bdelyrus Har. fo. G6)

Body shape oval; elytral intervais may be

Gistinctly convex;,.anterior margin’ of clypeus

dentate, teeth acute . . .Pedaridium Har. Soe ae

First segment Of hind tarsus triangular; clypeus

eflen rucose, sometimes with coarse. punctures... . 13

First segment,.of, Wind; tarsusi(slender jor, broadly

rectangular) and wit troneate..or-roynded apex;

clypeus ‘notudistinctly rugose, often, smooth,or

Linein, punctabe,) S0metings coarsely «punctate. va... 14

Hind coxa in extreme).postermior., position, . with

strone. median.wentral: reduction.in width .of

abdominal segments; middle coxae, separated

by distance of less than width of middle

POM Reena a ee ed ORES. Er, CBatcb 21)

1A( 12).

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VOCS):

ZOULON

Howden and Young: Panamanian Scarabaeinae

Hind coxa more anterior, with six abdominal

segments easily identifiable; middle coxae

separated by distance greater than width

of middle femur. i... 4 Dichotomius Hoge... ip.

Lateral pronotal fovea round, without depression

or .row of; punctures, immediately .amterion to

fovea; usually seventh elytral stria, excluding

scattered punctures, not Bi ebes to. anterior

margin.

Lateral pronotal pag ee OF oblone, With a row

of coarse punctures and/or a longitudinal

groove in line with fovea? seventh elytral eiria

extending “to anterior margin.

1 23)

Body shape rectangular, elongate; dorsally flattened;

fifth ventral abdominal segment. medially

shorter than fourth or Sixth; “Une S1xin segment

fourito dive: dimes: longer tian iui:

Onthocharis Westw. (De

Body shape not as above; fifth abdominal segment

hot. as, above

Mesosternum distinct; first segment of posterior

tarsus gradually ‘enlarged toward the apex;

first five abdominal segments narrowed medially,

sixth longer

Mesosternum very short: basal segment of posterior

tarsus very elongate, scarcely enlarged: median

ventral abdominal segments of approximately

equal lengths) e7Fe twee he wCamt nid) be ph

Pygidium: with a’ deep, nearly (median transverse

Sulcus; body ean | convex

Pygidium lacking a “median transverse sulcus;

body ‘strongly ‘convex’... GAtenehus® Weber <(p.

Mesosternum smooth or nearly so; lateral pronotal

groove pronounced, ee tovleebiy

punctate init) ler Sd roxyen Westwie, (ps

Mesosternum distinctly punctate; lateral pronotal

punctures delimiting or forming irregular

SLO E ly ee ha tant ee uted an ah eae eh Pn ae ig areeal a Vl

Posterior edge ofyvertex (netvabruptiy: declivous,

at»most only slightly so) behind ver between the

Ones, ;

Posterior edge ‘of vertex “clearly declivous

Eyes not visible: from) above’ without protracting

the head: elvirgiiidise Cartinate laterally

. Cryptocanthon Balth. Gee

Agamopus Bates (ie

47)

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14 Contrib. Amer. Ent. Inst: . vel 46,7 ne. bp 1961

Eyes visible Trom above with head Tully retracted)

elytral: disc dacking tateral or apical carinde .

Pseudocanthon Bates mp. 35)

21(19). Lateral margin of elytral disc at or behind

umbone with strong carina(e); third to seventh

intervals often briefly carinate near apices . :

; Delicehwum Esen.- (p. 36)

ELyira! Nea ae. and seica prominences.

ae ee : Canthon Hoff.) (p. 19)

Eurysternus Dalman

Burysternus Dalman, 1624, p. 8.

Type-species: Scarabaeus caribaeus Herbst (= Eurysternus

planus Dalm.), listed by Lucas, 1920.

Aeschrotes Serville, 1828, De Gor.

Eurysternus, as presently delimited; may be recognized by

the following combination of characters: body shape elongate,

clypeus lacking teeth, labial palpus with two segments, elongate

scutellum visible, elytral disc often flattened and with a sharp

lateral carina delimiting epipleuron, elytron with eight striae, and

mesosternum short.

The genus is in need of revision, hence some of the names

used herein may be incorrectly assigned.

Key: to’ species of Burysternus

i. Posterior margin of hind: femur a ee tooth «(if

peesent) Dacal’.: ee 8 : 2

Posterior margin: of Hind femur curved or sinuate,

tooth (if present} median or in’ distalhalf 3: . . 3

2 Anterior margin of fore femur feebly arcuate; length

14 toc) mime -. Is ?@laudi¢ans Kirs.

Anterior margin of fore femur ‘angulate; leneth |/ to

eee 2 Sp bepe is: © Har.

Ba Posterior margin’ of hind femur with tooth ‘or

tubercle; lenoth 1/-mmeorl less 2 ies. POS 4

Posterior ‘margin of ‘hind femur lacking tooth or

tubercle; length 1/ to 22°mm...°. 3. velutinus Bates

A. Posterior margin of prosternum straight or very

ae arcuate st Ay; leneth Ll to: 14 0mm

: 4. mexicanus Har.

Posterior margin of prosternum angular or sinvate: . 5

S¢ Posterior portion of prosternum smooth, margin

angulate (sometimes feebly) medially (Fig. 3);

posterior femur eradually expanded in. apical

halt; length 14 to lO mm... eo) eearibacis Hbst.

Howden and Young: Panamanian Scarabaeinae 15

Posterior portion of prosternum with tubercle on

either side of midline, margin. sinuate: posterior

femur abruptly es in apical half; length

5:50 (Oo amm oh as eee i ee

I. Eurysternus claudicans Kirsch

Fig. 6

Euryéternus clautlicans Kirsch, 1870 (18/1),-p. 360.

Type, Dresden (?). Not seen.

Length 14 to 17 mm. Dull brown to brownish black dorsally.

Dorsal setae short, inconspicuous. Pronotum trregularly tumid

either side of midline, often with four shining, impunctate spots

Al are across anterior: third;:. Elytra’ metymottica: “striae aretinct,

represented by depressed double lines widened by shallow, indis-—

tinct punctures. Metasternum mot depressed: in center, lencituding!

median third smooth except for scattered coarse punctures.

Hind femur (Fig. 8) usually toothed at basal sixth, posterior

margin otherwise straight, hind tibia of females (in dorsal view)

ben? and sinuate in .apical third.

MATERIAL EXAMINED. PANAMA, 58 specimens from: BOCAS DEL

TORO PROV. - Rio Changuinola. CANAL ZONE —- Barro Colorado

Island; Frijoles; Madden Forest; Rio Trinidad. PANAMA PROV. -

Cerro.Campana; Las: Cumbres;)c COLLECTION DATES: February,

April, May, June, July, August, September, October, November,

December.

OTHER COUNTRIES, 29 specimens from: MEXICO, NICARAGUA,

Ooty RICA, COLOMBIA, PERU.

REMARKS. This species has been collected at agouti, rat,

tamandua, and unknown carrion; on feces of the howler monkey,

man, spider monkey, and tapir; and on leat Surfaces, “On

Barro Colorado Island, claudicans is active nocturnally, particu-

larly in the period from 1800 to 2000 hours. ~ The species is° most

abundant during the wet season, and is equally abundant on both

carrion and. feces... In Panama clavudteans “occurs primarily in

lowland forests.

Z. Eurysternus plebejus Harold

Figey 2. 9

Furysternus: plepeyuse: Marold. 1050, 0.) a.

lype or. cotype, Paris. Nor seen,

Length: 7 10. 11 mm.” Dorsally) brewn mottled with black or

dark brownish black mottled with brown; dorsal setae frequently

conspicuous, particularly on: elytral apices, © Pronotal’ punctures

ocellate, surface dull between punctures, usually lacking conspicu-

ous, impunctate shining spots; surface slightly irregular, frequently

feebly depressed medially in posterior fifth. Elytral striae (Fig.

2) feebly impressed. Metasternum centrally with shining shallow

depression. Hind femur (Fig. 9) lacking tooth on posterior margin

in males, a small tooth present at basal sixth in females. Fore

16 Contrib, Amer: Bats (basins vole. 08) ones: ia) 1981

femur with distinct median obtuse angle on ventral (tibial) edge.

Hind tibia not sinuate or strongly bent.

MATERIAL EXAMINED. PANAMA, 191 specimens from: CANAL

ZONE — Ancon:: Barro Colorado Island: Cabima: Gamp Chagres;

Fort Kobbe; Madden Forest; Margarita; Paraiso; Pipeline Road;

Rio Trinidad. -CHIRIOQUL PROV. = Bugaba,: S00-E500"ft: “COCLE

PROV. — El Valle. | CODON (PROVE «=; Coco" Solo; Portobélo: ‘Santa

Rita Ridge. PANAMA PROV. - Bayano Bridge; Cerro Campana.

COLLECTION “DATES: Every month of the year.

OTHER COUNTRIES, 32 specimens from: MEXICO, COSTA RICA,

COLOMBIA, VENEZUELA. |

REMARKS. This species has been obtained from Berlese

Separagion Ol fores, (igor liiler,~ irom roiting fruit of Gustavia sp.,

from, eavrriom, of agouti, mat,; Sloth, and tamandua, and.from feces

of horse, howler monkey, spider monkey, Manivanad tapir. Lt

has also Deen taken, perching on deaf surfaces.’ On. Barro Colorado

Island, plebejus is diurnally active, particularly from 1100 to

1300 hours, and appears to be equally abundant in all months of

the year, It does; exhibit a, “seasonal shift’ in food utilization,

however, occurring primarily on feces in the dry season and on

carrion in the wet season.

In Panama plebejus is seemingly restricted to lowland forest

below 1000 m in elevation.

3. Eurysternus velutinus Bates

Pig a 10

Eurysternus velutinus Bates, 1887, p. 39; Halffter and Halffter,

TO) Dei.

Cotypes, London and Paris. Examined.

Length 1/7 to,22 mm... Dorsally brownish black; setae fine,

numerous, similar in size on pronotum and elytron, less closely

spaced on elytron. Pronotal surface except at midline evenly

convex, midline in posterior fourth indented, in anterior fourth

often feebly raised and shining; surface otherwise closely,

shallowly ocellately punctate. Elytral striae represented by

scarcely depressed, paired shining lines and obsolete punctures;

elytral intervals dull, most with two to three irregular rows of

fine setae. Metasternum with median area shallowly concave, more

bO posteriorly; latcrabsuriace to edae.of middle coxa minutely

punctate, finely Setose.. Hind femur lacking posterior tooth (Fig.

10). Hind tibia feebly sinuate on ventral edge (in lateral view).

MATERIAL EXAMINED. PANAMA, 2 specimens from: CANAL ZONF

~i Madden: Horesi. CHIRIOUL PROV, Volcan de’ Chiriaui, 2000-3000

Py | COLLEC PION DATES: December,

OLE RY COUNTRIES, lb specimen from: ECUADOR,

REMARKS. This species has only been collected at human

léeces..

Howden and Young: Panamanian Scarabaeinae As}

4. Eurysternus mexicanus Harold

PLO Sache id

Eurysternus mexicanus Harold, 1869b, p. 505.

TVPG. OF, COLY Deg toQrkss, | i NOR Geen,

Length 1110: 14.mm.:. Dorsally, very dark ;OYown lo Dlack,

frequently mottled with..brown, more, conspicuously, so,,on head and

pronotum;. dorsal setae small. Pronotum andemved ani midline. in

posterior fourth, on ‘either side ianterior tho imdentation' a.iemooth,

feebly raised swelling extending, obliquely toward, lateral, fovea;

three. small polished swellings: present).al\ anterior Sixth, one on

midline anda proximal one ,on each, side) remainder or pronoium

moderately punctate, setose.” Elytral striae shallow, vepresented

Dy a narrow .double, line, punctures mbsolete.)Foavertor, margin (of

prosternum (Fig. 4) nearly straight. Metasternum with smooth

circular indentation medially,"area dull. (hore Temir abruptly

anoulate at;basal ventral third.) Hind) femur (hia. Ly warn

posterior tooth at distal third, distal half of femur swollen, some-

what oval. Hind tibia moderately ‘arcuate, mov sinuata,

MATERIAL EXAMINED. PANAMA, 65 specimens from: BOCAS DEL

ORO? PROV,..— Almirante, ‘Changuinolal sDistrictm (CANAL ZONE. ain p

Chacgres, .. COCLE: PROV. El. Valle, oRig Guava bee | (OP ANT Ay TON sx

Cerro Campana., COLLECTION: DAT EO: lanwary february. Nadie OT.

meplember, October,

OTHER. COUNTRIES, 27, specimens trom: MEATCO), NICARAGUA,

RAO RGA

REMARKS... This..species ‘has, been collected on Nerse manure

and on.human feces.

5. Eurysternus caribaeus (Herbst)

PAS wn Hog i dae

Scarabaeus caribaeus Herbst, 1789, p. 300.

Type, Berlin (?). Not seen,

Eurysternus caribaeus (Herbst), Halffter and Halffter, 1978, p. 57.

Length 14 to 18 mm. Dorsally brown mottled with black;

setae inconspicuous except along lateral margins of, pronotum and

near .elytral. apices. Pronotum with mumerous) moderate.punctures,

surface not noticeably (inregular, a) cistinet,) Shining, dark spot

present on. midline, at; anterior, Tourth, : Elvival striae »moderately

indented, narrow: elytral @dise longitudinally feebly) concave

laterally. Metasternum irregularly, shallowly impressed near mid-

line, mostly dull. Prosternum.(Figo. 3) with posterior margin

medially obtusely ‘angulate. Hind femur) (Pig. 12)..with posterior

margin, toothed near apical fourth: in male, near.middie’ in: female.

Hind tibia only moderately arcuate.

MATERIAL EXAMINED. PANAMA, 50 specimens from: CANAL ZONE

— Barro Colorado Island; Madden Forest; Pipeline Road; Rio

Trinidad, PANAMA RRON «i. Baveno: Bridge: Cerro Campanas, Rio

Pequeni.( COLLECT ION CORT Ee enrol Ce. AOU SE .

september, October, December.

CIHER COUNTRIES 20 amnecimene, from: MEATCO.) CONT A REC

COLOMBIA, ECUADOR,

18 Contribs ‘Amer? Minty ast. | vol 13, me,v iy 1961

REMARKS. Specimens have been obtained at rat, tamandua,

toad, and unknown carrion, and on feces of coati, horse, man,

howler monkey, spider monkey, and tapir.. lt is more common on

Carrion ane on Barro Colovado. leland) ie active ai night. lt is

most abundant during the wet season. In, Fandma -the spectes is

restricted to lowland forests. Halffter and Halffter (1978) have

discussed some aspects of the biology of caribaeus.

6. Eurysternus magnus Laporte

PIS, 15

Eurysternus magnus Laporte, 1840, p. 93.

ivpe, erobably iW Paris... Nou! seem,

Lenoth 15 to [9 mm: Worsally very dark brownish black,

elyira mottled with brown; dorsal Setae minute except near apices

Of elytra, Pronotum, with surtace dul), finely punctate; ‘fairly

evenly convex except for an irregular circle of six slightly raised,

usually shining, small tumosities; two tumosities at anterior fourth,

two more widely separated near posterior third, and two proximate

Ones) Near midline alt-=posterior fourth. Elytral striae consisting

of two slightly impressed shining lines. Metasternum with small

central, dull depression; median surface posteriorly flat in females,

cariniform in males. Hind femur (Fig. 13) with posterior margin

toothed, .at,apicaliihird in both sexes. Hind tibid. not sinuate,

ventrally arcuate.

MATERIAL EXAMINED. PANAMA, 47 specimens from: BOCAS DEL

TORO PROV, = Rio Changuinola, 5000 ft. CHIRIOQUL PROV. — Baldwin

Forest; Boquete;: Cerro Punta (2. km E,) 2200 moc Harimann'’s Finca,

1200 We Las Lacunas; Rio Chiriqui, 5600 ft.; Viejo, Volcan de

Chiriqui, 04000 41.) COULECTION DATES: january, February, March,

May, June, August, September, October, November.

GIHER: COUNTRIES, 12 spectmens from: MEXICO. COSTA “RICA.

REMARKS. Specimens have been collected at carrion and on

human and horse dung. The species is active nocturnally and is

restricted to forests above 900 m in elevation.

Megathoposoma Balthasar

Megathoposoma Balthasar, 1939a, p. 18; Halffter and Martinez,

1906, 9,141. (cemeric synonymy and detailed generic des-—

éription).

Type-species: Megathopa candezei Harold, 1873, by monotypy.

Mecathona Wechschoitz, 1022, 9, 34: Bates, 160/, p. 25; Howden,

1966.0. 726:

Body shape (Fig. 5) elongate-oval. Much of dorsum punctate,

many punctures annular; glabrous. Head somewhat triangular in

shape; clypeus closely bidentate, a transverse band of setae

present dorsally on each tooth near base; head lacking tubercles

or carina; eyes dorsally moderate in width, not narrowed poster-—

iorly. Pronotum fairly evenly convex, lateral fovea small and

circular; marginal bead lacking posteriorly. Scutellum small,

acutely angled posteriorly. Elytra each with eight discal striae

Howden and Young: Panamanian Scarabaeinae Lo

and an additional two lateral striae in anterior third below

carinate-eighth strra;: each. discal: stria, represented: by) mapressed

line -with scattered elongate punctures; conspicuous, shining, nearly

square black markings irregularly spaced along striae; intervals

nearly flat, moderately punctate, the punctures separated by

approximately one diameter. Pygidium feebly «convex, closely

punctate. Proepisternum and proepimeron not separated by carina.

Mesosternum very narrow; middle coxae widely separated by

metasternum,

This monotypic genus is restricted to Mexico and Central

America.

1. Megathoposoma candezei (Harold)

FignaiS

Megathopa candezei Harold, 1873, p. 102; Bates, 1887, p. ‘26.

Hy CeyUr arises (Mets Seen.

Megathoposoma deltochiloides Balthasar, 1939a, p. 19.

Megathoposoma candezei (Harold), Balthasar, 1942, p. 43; Halffter

and Martinez, 1966, p. 150.

Length 18 to 24 mm; greatest width 11 to 13 mm. Dorsally

with underlying black background, pronotum mottled cupreous and

green, head: nearly unifonm .cupresus;celytra eupreous, on conecn

with rows of squarish black markings (Fig. 5) along striae:

pygidium: cupreous. Legs and: ventnal surface «mostly: black,

frequently. with -greenish »castsn9 Pore: tibia \withathres teeth on

outer ‘margin vim apical: thinrd;: tibial»spur.ot. male, deeply emaremate,

each half of bifurcation sharply acute and nearly as long as base

below emargination. Middle tibia with small tooth on outer margin

at-apical third. Hind (tibia not, cistinetiy exmanced near apex,

slender; apex. with. single tibial spur. Ventral euxiaces (of, temora

closely punctate, ‘most punctures ‘each, withe minute vseta.

MATERIAL EXAMINED. PANAMA, 1 specimen from: CHIRIQUI

PROV’.

OTHER. COUNTRIES, 20 specimens, fronts MBAICO, GUATEMALA,

NWECARAGUA,. COSTA RICA,

REMARKS.” This> species: is (easily. recboniged: Dbynits orelatively

elongate-oval shape, punctate dorsum with a greenish cast and

elytra with rows of shining, almost square, black spots. It ranges

from: southern..Mexico to. Panama.

Wille (1973, 1974) has described the biology of candezei in

some detail.:;.Thexspecies)'has: been’ taken outside.of: Panama in

tropical forested areas in traps baited with human feces.

Canthon Hoffmannsegg

Canthon Hoffmannsegg, 1817, p. 38; Vulcano and Pereira, 1964

(Catalogue); Halffter and Martinez, 1977 (Revision).

designation, Paulian, 1938.

The genus Canthon Hoff. and its close. relatives are currently

under study by G. Halffter and A. Martinez. Recently Canthon,

20 Contrvps Amer, hatiwbiste ly voli Te. ne, sty) £981

sensu lato, was divided into a number of genera and subgenera ,

and in some cases there are differences of opinion on the exact

status — genus, subgenus or species group — desirable for some

of the proposed divisions. Much of this should be resolved when

Halffter and Martinez complete their studies. Since Canthon is

still under revision we have decided to adopt a conservative

approach - recognizing only Pseudocanthon and Canthon as distinct

genera. (Thus, for convenience) we have considered Scybalocanthon,

Glaphyrocanthon, etc., as subgenera of Canthon and have not

included them herein. Our key to species hopefully is adequate

and placement of the species into groups can be determined by

consulting Halffter and Martinez, 1977 (and subsequent work).

Canthon, as delimited here, can be recognized by the

following combination of characters. Moderate size of 4.0 to 14.0

mm. Clypeus ‘antertorly emarginate or with, two to four teeth.

Frons and vertex lacking tubercles. Dorsal ocular areas delimited

laterally by a nearly complete canthus. Posterior of vertex with

complete, sometimes fine, marginal bead. Pronotum lacking marginal

bead basally; lateral pronotal margins angular or arcuate, their

basal portions not appearing subparallel when viewed from above;

anterior angles quadrate or acute. Mesosternum short and trans-—

verse, usually more than 10 times as wide as long. Meso-meta-

sternal suture evenly and ivery feebly arcuate. Elytron, maximally

with nine striae, these often obsolescent or lacking. Fore tibia

obliquely truncate, with three outer teeth. Hind tibia slender, not

noticeably expanded apically nor flattened near apex, nearly

quadrate in cross section. Tarsal segments flattened, triangular

Ori trapezoidal, Tarsal claws feebly to, moderately curved, not

dentate:

Key to species of Canthon

ne Hind femur ventrally with fine line or carina

PESSCOU AM IACOME ING aNLeMIOnRVedGe By ea lel ole 2

Hind, femur ventrally lacking) line. or carina

Me AOR ECG Lae ts A Ce i LL

Ze Hind femur ventrally with fine line or carina

adjacent (toposterior ‘edge. .).. A 3

Hind femurventratly ‘lacking fine line Or carina

SOA CeM CG wOOSTerion ed meat Gh elu a Cine 4 5

Ok Pygidium with basal carina a. aes size

Comin or wore 0.) lie ae 4

Pygidium lacking distinct Veal. Carina: ‘dorsal

color @reenish gold or ereen, or bluish green;

AVERAOS SIZ e OOM ee ae Cue Ka. 4 LOL. morsel group

A Dorsal color Not ‘umitorm, pronotum brown. with

black ee. COTES Cea

Wi Ba ee a Ns oan Cia part)

Donsal es tes black, sometimes with purple

TOC LECCHOM SA OND a koa ea Oa auinotiia lis’ Har.

10,

Nee

14.

Howden and Young: Panamanian Scarabaeinae 21

Clypeus anteriorly bidentate .

Clypeus anteriorly quadridentate .

CO OD

Dorsum lacking distinc! setac; size over > mm. 4 yi

Dorsum with scattered, distinct setae; size under 6

mm; color dorsally black, usually, with) feeble

metallic) Tuster |.) 1 ee a ae EL air g

Pygidium with basal carina’ present; color dorsally

uniform, Diluish’ black orwereen : :

alta indigaceus chevrolati Han.

Pygidium lacking “basal carina; color dorsaliy

black with yellow spots. .2. septemmaculatus (Latr.)

Thorax ventrally with proepimeral carina incomplete

(extending from coxa to thoracic margin) or

absent; dorsally bicolored or pronotum mottled . i,

Thorax ventrally with proepimeral carina complete;

color uniform, black with blue reflections Nae!

Oe COV TOLLE. Har.

Pronotum black, often with’ Dlue or @veen Cast,

Suze sally (UNGer im ee EA Aa es 10

Pronotum brown with black erat) ike usually

7. mm or more... 7. Cyanelius sallel tar. (in. Danis

Two inner clypeal teeth V-shaped, distinctly widened

basally; elytra and metasternum alutaceous, dull,

Brown QM DWlaCK a er ie eas (Germ. )

Two /inner clypeal teeth very slender, cloncare;

elytra and metasternum smooth between punctures,

shining Drown and black . 027) 207 miemiiie | lucas

CIV DEUS, ANLEPIOMIN GUAGE Uae eee ee lie es

CIVPCUS BMTOTIOR IY DCO Tae he ieee ciel ell in ol aliadals LG

Thorax ventrally with proepimeral carina complete

(extending from coxa to thoracic margin); at

least head and pronotum black, often with bluish

rel lections 200, Meas aE ORNS A) AURA EH Goffe s ME NN A Te

Thorax. ventrally with proepimeral carina incomplete

or: laterally obsolete: pronotum, yellowish, elytra

DEOW Te lei wc le oe aie ide anita, Cel Ones CAINS 10) Alaa

Pyeidium, With Dasal carina: thorax ventrally with

COMDIELE DRGEDIMeT AL Cama nei i moan mime ig iil ad ig 14

Pyeidium lacking Dasal Carina: Thorax ventrally

lacking: Proepimeral Caring so. ol moniiatue Bates

Head and pronotum bluish to bluish black ‘or black,

elvyira similar om reddish, Drown wwitn black

Lae aE MLE ERE CONE MEE WAN Umi MGIC

22 Contrib. Amer: (Ents lest: . vole’ 16 4 no. 1961

Dorsally untformly green, often with gold or

metallic reflections (see also note on 15a.

ecaelius Bates! 2 Po ee eae. 2 15. lamprimus’ Bates

be Antero-lateral margin of prothorax with deéenticle

On Wentral edges. elytra smooth, shining <;-. 16

Antero-lateral margin of prothorax lacking denticic

On Veniral @dee, elyvira Sstiacreened, > dul!

2716)" viridis: mericdionalis Wiehe oH.

16. Elytra black with bluish reflections Sh Sane

. 13. euryscelis Bates

Wivire oe or owe ele with black

WOGCErS 2k et fe ee en te Suplyalinus’ Har.

1. Canthon moniliatus Bates

Figs. .0,.20;,.21

Canthon moniliatus Bates, 1887, p. 27.

Cotypes,. Londom and Paris. ..London specimens examined.

Length 6.0 to 8.0 mm. Dorsally moderately shining, head

and elytra of varying shades of brown, head usually darker.

Pronotum yellowish, medially at basal margin with blackish

Marking, occasionally with larger -black marking. medially near

ANGELO: Marcin... Pyoidiunm brown,..approximately same. shade as

elytron. Clypeus (Fig. 20) anteriorly bidentate. Gena laterally

at, Glypeal junction feeply produced, very slightly oblusely angled;

eye dorsally ‘approximately three times as lone as wide. .-Pronotum

(Fig. 6) with anterior lateral margin straight, ventral edge

tacking 2 tubercle. Hlvyiron with eioht discal striae represented

by shallowly imdented lines, each stria with relatively large,

Shallow punctures separated from adjacent punctures by approxi-

Mately one diameter; iniervals.smooth, impunctate. Pyosidium with

Dasal Canina absent, pyeidial surface smooth, dull, shagréened.

Anvenwa- Drown, club: erey. Proesternum with, proepimeral carina

absent. . Metasiernum. between coxae. smooth and shinifio. Fore

tibia (Fig. 21) similar for both sexes, spur slender, apex acute.

Hind femur VvVenivally lacking, lines. adjacent.to margins.

MATERIAL EXAMINED. PANAMA, 249 specimens from: CANAL

ZUNE = Barro Colorado, Island; Coco Solo; Madden Forest; Pipeline.

Koad: Bio. Civicito, PANAMA PROV. — Bayano Bridge (9. km SE);

Cerro Canipamas, COLLECTION DATES: Every month of the’ year.

ODRE COUNTRIES, '9. specimens. from: COSTA, RICA,

REMARK. Canton moniliatus, fas..been, collected flying; in

léal Iitter and. citting on. leat suriaces;. on fallen. Entada..sp.

peds covered With’ white fungi and on Agaricaceae fungi; ‘at

Opossum, peccary,.ral, Snake, and sloth Carrion; and on howler

monkey, Human, jaguarund),..tapir, and rodent feces.

On Barro. Colorado Island, this, spectes is equally -abundant

atl Any time curing the day. .Albeough the species. is relatively

uncommon,.. beth teneral and mature adults occur throughout. the

year. Canthon moniliatus (as well as Onthophagus sharpi) utilizes

Howden and Young: Panamanian Scarabaeinae Zo

the widest range of food types of the 56 species of Scarabaeinae

found on Barro Colorado Island. «1L is ene of only Six species

that is found at carrion in the dry season ane one ol four species

found on fungi at any time. It does not aggregate in the manner

of C, angustatus. Usually only one or two individuals are found

at one time on the same food source. Panamanian specimens of

moniliatus are found only in lowland moist forests, usually below

200°.

2. Canthon septemmaculatus (Latreille)

Pie sso 22; 20

Ateuchus septemmaculatus Latreille, 1811, p. 180.

Lype, 21 aris.> Not seen.

Canthon septemmaculatus (Latr.); Harold, 18608, p. 51.

Length 6.5. to 12:0°mm.:~ Dorsally dull: head oreenish Diack,

pronotum, elytra and pygidium very dark brown to black; pronotum

with five yellowish spots, a spot adjacent to each pronotal angle

and one posterior median spot; two additional yellowish spots in

basal vhalf of pygidium; onevon eithen side! of midline. Clypeus

(Fig, 22): anteriorly bidentate. Margin: of cena adjacent to cltypeus

very obtusely angulate, only feebly produced; eye dorsally moderate

im ‘size, approximately three times as) lone a5 wide. 9 Pronotum

with antero-lateral margin nearly straight, ventral edge lacking

tuberelé. | Elytron’ with allebut marainal stria: obsolete. > Pygidium

with basal carina absent; pygidial surface dull, finely granular.

Antenna dark brown to black. Prosternum lacking proepimeral

carina. Metasternum between coxae feebly shining, surface very

finely granular or shagreened. Fore tibia (Fig. 23) of male with

Spur scarcely expanded apically, shallowly cleft; spur of female

slender and acute at apex. © Hind femur ventrally with only

anterior’ line present adjacent 10) maveoin.

MATERIAL EXAMINED. PANAMA, 182 specimens from: CANAL

ZONE =) Barro Colorado Island: Cocco) Selo; @er0z4a1 > Curundu:

Farfan; Fort Clayton; Fort Dawie Port Kebpe: Prijoelest:Gamboa;

Madden Dam; Margarita; Pedro Miguel; Pipeline Road; Rodman;

Summit Parks Taberntllaw COCLE PROV pa aAguadulice; bi Varie.

COLON: PROV .>= .Santav Rita 7Ridee. «LOS SANTOS PROV Gi-¢fonost,

PANAMA “PROV. i Arraijans” Camp Chacres;: Cerro: iaul4 Cerro

Campana; Cermeno;: Panama’ City; Puntar Vacamonte... COLLECTION

DATES: Every month tol the: vear except Fapruary.

OTHER COUNTRIES, i7iepecimens frome’ COLOMBIA, VENEZUELA.

REMARKS. This species has been found sitting on -leaf

surfaces; at rat carrion; and: on feces of coati, dog. horse, nowler

monkey, and man.:, OnsBarro: Colorado Island septemmaculatus

is active throughout the day, ‘primarily iinvthe wet) seasoned: In

the. early .wet: Season itis most: abundant’ on feces butaby: the: late

wet season is. most>sabundant on’ carrion.» -lt:is one, of the few

species. of -Scarabaeinae able to successfully; and:routinely pirate

food balls from other ball-rolling scarabs. The species apparently

is moderately eurytopic,; oceurrine An: moist and dry forest, forest+

pasture’ ecotones, shrub-land,:'and im small: cleared areas in the

preceding habitats. it is mainly limited, however, to, areas: below

24 Contrit: (Amer) oRnts Tish): wel loo mo. ay dS)

1000 m in elevation.

Su) Canthon Geyrelter Harold

Ries 20.) 125

Canthon deyrotlei Harold,| 1863," p. 132,

i Type or cotype, Paris. Not seen. |

| Length, 6s5 to 12.0 mim.) Dersally shinims black, usually with

blue or purple reflections. . Clypeus (Pig. 24) anteriorly quadri-

dentate, suriace granubate. Gena laterally at elypeal junction

ODLUSsely aneled Torming small tumercie: eve dorsally moderate,

approximately three times as lone as wide. Pronotum with: antero—

lateral margin slightly arcuate, ventral edge with small tubercle.

Elytron With (discal striae marked by fine Iindented limes, intervals

between mostly smooth. Pydigium with basal carina complete,

pyeidial surface smooth, very timely shagreened.. Antenna: black.

Prosternum «with proepimeral carina complete. Meéetasternum between

comae mostly smooth wand shining. Fore tibia (fio. 25) of male

with spur widened ‘andioutwardly bent at middle; thence tapered

to: rounded ‘apex; Spuriol female: similar to that of ‘male, more

Sstender. and with apex acute. () Hind femur’ ventrally with line

present adjacent (lo \anterior .marcin;. (posterior: line absent.

MATERTAL OE AAMINED.:. PANAMA, “None Listed “by Vulcano and

Pereira (1964) as occurring in Central America and Colombia.

OTHER COUNTRIES, 8 specimens from: BELIZE, EL SALVADOR,

NICARAGUA, COSTA RIGA, “COLOMBIA:

REMARKS. Seemingly “a rare species: that: ‘should occur in

Panama.

4 Canthnon ingioaceus .chevrolatt Harold

Pigs i263 27

Canthonchevrolati) Harold, > 1868, .p...119.

ype OR Cory pe, Paris. \ NOU iseen:

Canthon indigaceous chevrolati Harold; Halffter, 1961, p. 288.

Length fsa to 1isO\mm. “dorsally pronotum:and| elytra shining,

head “and. pyeidium (dullito feebly shining) Cotoriranging ‘from

bright green to bluish black. Clypeus (Fig. 26) anteriorly

bidentiates Gena laterally: aticlypeal (junctions scarcely or not

produceds eye, moderate approximately three: times: longer than ‘wide.

Pronotum with antero-lateral margin slightly outwardly arcuate,

Vetira ledge usually lacktie tubencies: Df tubercle: (present then

Near Wanterion angle... Hlyiron with discal striae obsolete. © Pygidium

with basal carina ‘complete, pygidial, surface smooth, dull,. very

finely shagreened.. Antenna black:.or: brownish black. Prosternum

with | proeepimeral Carina (‘absent ‘or spresent: only in) basal. fourth

adjacent to coxae. Metasternum: between coxae (smooth, feebly

shining, finely shagreened. Fore tibia (Fig. 27) of male with

spur: Gradually (expanded to moderately “notched. apex; spur of

female slender, \acuteto abruptly rounded! atvapex.))) Hind femur

venivally with iwell defined Wine present adjacent: to anterior margin,

posterion line! absent.

Howden and Young: Panamanian Scarabaeinae 2)

MATERIAL EXAMINED. PANAMA, 1 specimen from: CANAL ZONE -

Pipeline (/Roads s/GOLLECT LON oDAT EO: .obiann

OTHER COUNTRIES, 169 specimens from: MEXICO, EL SALVADOR,

COST AxoRIGA.

REMARKS. The species has been taken in horse and cow dung

and in traps baited with human feces.

5. Canthon lituratus (Germar)

Pio Gai cto sa) ne

Ateuchus lituratus Germar, 1613, .p. 117.

“Type, Berlin. Not seen.

Canthon quadripustulatus. Guerin, )1355;p¥ 507.

| Type, location uncertain.

Canthon lituratus (Germar); Harold, 1868. p. 95. eee

Canthon lituratus quadripustulatus (Guérin); Bates, 1887, p. 33

(Bates considered that var. quadripustulatus included

Panama specimens).

Length) 4.5/t0 6.5 mm. Dersally heed and i pronotum, shining

green; elytra dull, black mottled with brown or with brown spots

near humeri and apices; pygidium dull, very dark brown to black,

sometimes with metallic sheen. Clypeus (Figs. 7, 28) anteriorly

strongly quadridentate, inner teeth broadly V-shaped. , Gena

laterally near clypeal junction produced so that side of gena

behind lobe or angle is nearly parallel to that of opposite gena;

eye dorsally narrow, approximately four times as long as wide.

Pronotum with antero-lateral margin straight or feebly arcuate,

edge, ventrally lacking tubercle}, pronotal isuriace with. numerous

small punctures, area between finely shagreened. Elytron with

all striae except marginal one feebly indicated by wague lines;

intervals flat, distinctly granulate. Pygidium with basal carina

complete; pygidial surface granulate, with scattered small shining

spots. |Antenna, particularly ‘club, tan... (Prosternum with

proepimeral carina incomplete, present only in fourth adjacent to

coxa. Metasternum between coxae only moderately shining, finely

granular, moderately punctate, most punctures (in clean unworn

specimens) with a minute seta. Fore tibia (Fig. 29) with apical

spur long and slender, similar in both sexes. Hind femur ventrally

with line adjacent to, anterion, mareinw present, pesterior -line

lacking.

MATERIAL EXAMINED. PANAMA, 41 specimens from: CHIRIQUI

PROV. ~ Galera’ de: Chorcha:: Rovira. Volcan (‘de Chirtqul.) COCUE

PROV. = Aguadul¢es) 4 CODLEOT LON DATES wie pmiars,) July, ‘September ,

OTHER. COUNTRIES, 50.,epecimens. ¢rom: COLOMBIA, VENEZUELA.

REMARKS. This species has been recorded only from dog

feces. It appears to be restricted) in Panama to elevations above

800 m.

26 Contrib. “Amer. Ent. inet. wel. 18> mo. 1, 1981

6:.°Canthon mutabilis. Lucas

Fies. 30, 31

_Canthon mutabile Lucas, 1857, p. 100.

Type, Paris (7)... Not seen.

Canthon mutabilis Lucas; Harold, 1808, 5. 126. |

Canthon mutabilis transversalis Harold, 1868, p. 126 (close to

Panama form according to Bates, but Harold's ''variety a"

Gescribes the elytral pattern better. Schmidt, 1922, named

a number of forms of mutabilis; however these have not been

listed as valid subspecies by Halffter and Martinez, 1977).

Length 5.0 to 6.5 mm. Dorsally shining; head and pronotum

green, pronotum sometimes with brownish cast along lateral margins;

surface of head and pronotum with distinctly separated, small

punctures, elytra reddish brown, the Sutural intervals and a

transverse band at apical third black or elytral disc black-with

margins reddish brown; pygidium black with greenish cast.

Clypeus (Fig. 30) anteriorly quadridentate, the inner teeth slender.

Gena laterally adjacent to clypeus briefly, angulately produced;

eye dorsally narrow, approximately four times as long as wide.

Pronotum with antero-lateral margin slightly angulate between

pronounced anterior apd median lateral angles, ventral edge with

Simall tubercle beneath, slight angulation. Elytron with nine

striaie, the elont diccal Ssiriae represented by tive Slightly indented

dines; intervals minutely punctate, Pycidium with basal carina

complete, pycidial suriace dull, Granular. Antenna, particularly

club, tan. Prosternum with proepimeral carina incomplete, approx-

imately half of carina present. Metasternum between coxae

Shining, surface with numerous small punctures. Fore tibia (Vic.

31) with apical spur acutely tapered in both sexes, base of spur

Sligntly broader in male. Hind femur ventrally with anterior line

present adjacent tO margin, posterior line absent.

MATERIAL EXAMINED. PANAMA, 12 Specimens from: CANAL ZONE -—

Ameon; Batto (olorado lclanda: Coco Solo: Corozal? CHIRIOUI PROV. -

GUdpela.. COCLE PROV. = Acuaduice.. LOS SANIOS PROV. — Cerro

Lea Vela “PANAN PROV. = La’ Chorrera,... COLLECTION DATES:

February, May, June, August, September, December.

OTHER COUNTRIES, 2/7 specimens trom: COLOMBIA, VENEZUELA,

BRAZIL, ARGENTINA.

REMARKS. Canthon mutabiirs has been recorded from dung

and taken sitting on leaf surfaces in lowland forests.

7. Canthon cyanellus sallei Harold

P1g6) 02, OO

Canthon sallei Harold, 1863, p. 174.

Cotypes, London and Paris. London specimens examined.

Caninon sallet trianguiatus Gchmiat, 1920, p.. 124 (a Panamanian-—

Costa Kican color form).

Canthon cyanellus sallei Harold; Halffter and Martinez, 1977, p. 90.

Howden and Young: Panamanian Scarabaeinae Zi

Length 6.0 to 9.0 mm. Color dorsally variable; head black

usually with greenish cast, pronotum yellowish to tan with margins

and midline at center blackish; elytra tan to brown to dark brown

with margins sometimes black; pygidium yellowish often with dark

apical spot (this type of coloration typical of the vManriely

triangulatus found in Panama and Costa Rica). Clypeus (Hig. no2)

anteriorly quadridentate. Gena feebly produced or dentate adjacent

to clypeal. junction; eye narrow, three or more times longer than

wide. Pronotum with antero-lateral margin straight or feebly

arcuate, ventral edge lacking tubercle. Elytron with nine striae,

all but: lateral. stria. obsoletey. Pygidium with fine basal carina

complete, pygidial surface finely shagreened, dull. Antenna tan. —

Prosternum with proepimeral carina indicated only near coxa or

absent. Metasternum between coxae minutely punctate, surface

mostly shining. Fore tibia (Fig..33). with spur.ot male not greatly

expanded, apex shallowly notched, female spur slender, acute tip

bent downward. Hind femur ventrally with fine line -present near

anterior margin; posterior line present in male, absent or only

feebly indicated in female.

MATERIAL EXAMINED. PANAMA, 145 specimens from: BOCAS DEL

TORO PROV. - Almirante;. -Changuinola’ District,. CANAL ZONE: -

Barro. Colorado. Island; Canoa, Saddles: -Corozal: Curundus fort

Kobbe; Frijoles (6 km NNW); Madden Forest; Pipeline Road; Rio

Trinidad; Skunk Hollow;.Summit Park. COLON PROV. -— Alhajuela.

DARIEN PROV. - Santa Fe. PANAMA PROV. - Juan Mina; La Campana.

GOLLECTION: DATES::January; February,; April, May.:lune, july,

August, September, October, November, December.

OTHER COUNTRIES, 93 specimens from: MEXICO, NICARAGUA,

COSTA- RIGA;.. COLOMBIA; « ECUADOR, PERU, ~ERINLDAD.

REMARKS... Canthon c. sallei has. been. collected in forest

leaf. litter: dJlying;. at human amd-tapir feces son agoull, peccary,

rat, and sloth carrion; at a refuse heap of Eciton burchelli; and

on the carcasses of wood roaches and passalids. This species is

active throughout the day on Barro Colorado Island, where it is

the only Scarabaeinae to search for odor trails (and food) by

flying. low over the forest floor, for extended periods of time (up

to 15 minutes). Other species use various modifications of the

sit-and—wait technique. for feed detection. ©. sallei. is primarily

a carrion consumer, only rarely occurring on fecal material. It

is restricted to the wet season, with worn and damaged specimens

very rarely occurring im the dry season. .In Panama c., sallei is

found in lowland moist forest, usually below 200 m elevation.

8. Canthon aequinoctialis Harold

Pigs: 46s O55 o0s..00

Canthon. aequinoctialis Harold, 186e,..p. 79.

Type, .Pavisu, o NOL ceam.

Leneth. 410 to,l3.mm. .-Color: dorsally shining: black,

occasionally with purple reflections. Clypeus (Figs. 34, 36)

anteriorly bidentate. Gena laterally at clypeal junction obtusely

angulate, appearing feebly toothed; eye dorsally relatively wide,

28 Gontvib. Amer. Baro odnsty Sve 16, me. creo 196)

approximately twice as long as wide. Pronotum with antero-lateral

margin nearly Straight, ventral edge lacking distinet tubercle.

Elytron with nine’ striae, all but’ lateral stria obsolete. »Pygidium

with basal carina complete, pygidial surface smooth and shining,

impunctate. Antenna reddish or reddish tan. |©Prosternum. with

proepimeral carina complete. Metasternum between coxae smooth

and shining. (Fore tibia: (Pigs. 35, 37) with epur greatly expanded

apically in male; spur slender and acute apically in female.

Hind femur ventrally with both an anterior and a posterior line

adjacent to respective ‘margin.

MATERIAL EXAMINED. PANAMA, 500 specimens from: CANAL

ZONE — “Barro: Colorado Tstand: “Fort Kobbe;: Madden: Forest;

Pipeline’ Road; Rio’ Chagres. ‘CHIRTOUL PROV. =: Progresso. )')‘COLON

PROV. 2° Santa “Rita Ridge.’ COLLECTION DATES: “Every: month of

tie vear.

OTHER COUNTRIES, 64) specimens: from:) BELIZE, COSTA RICA,

COLOMBIA (Amazonas).

REMARKS. Canthon aequinoctialis has been collected from

heat Vitter: in Wight traps: a1 agoull, peccary, and ral. carrion;

and: at coati, “Atiman, j@euarundi, and tapir feces.

Otvthe: }o especies) of ‘Canthon oceurring in? Panama, this is

the only one that is nocturnally active, and on Barro Colorado

Island is most active during the period from 1700 to 2100 hours.

Reproductive activity may occur throughout the year on Barro

Colorado island; as both teneral and mature adults have been

oblained in every month of the years. INhis species is one of. the

most abundant Scarabaeinae on Barro Colorado Island and appears

to have a stable population’ density during the entire year.

During the wet season it is equally abundant on carrion and dung,

but ts found ‘only on) dungin the dry season." The species

occurs. primarily in towland. wet forests,

Oo. Canthon. juwencus ‘Harold

Rigs o VE] 385) 39

Canthon juvencus Harold, 1868, p. 111; Halffter and Martinez,

1977p. 00)

Type cor -cotypes Parisi (Not seen.

Canthon varipilis' Bates;*° 1067. p. 30.

“Cotypes, ‘Hondon and Paris,” Examined.

Length 3.0 162.5 “mm Doreally. black, oceastonally with

cupreous or metallic reflections; surface of head, pronotum and

elytra with scattered distinct punctures, each puncture with an

erect pale scta. Clypeus (Pics. tG;1/. SO) anteriorly bidentate.

Gena laterally at clypeal junction slightly produced, obtusely

angled. Eye dorsally approximately three times as long as wide.

Pronotum with antero-lateral margin nearly straight except at

middle, there margin offset by small ventral tubercle. Elytron

with all but marginal stria obsolete. Pygidium with basal carina

complete, pygidial surface finely shagreened with scattered setose

punctures being more pronounced near apex. Antenna dark brown,

club greyish brown. Prosternum with proepimeral carina complete.

Howden and Young: Panamanian Scarabaeinae 20

Metasternum between coxae shining, the surface with scattered

punctures, usually each puncture: with minute Seta... Fore tibia

(Fig. 39) with spur of male of uniform width to apex, apex

notched; slender and tapered to acute apex in female. Hind femur

ventrally with line adjacent to anterior margin present, posterior

lime absent.

MATERIAL EXAMINED. PANAMA, 20 specimens from: CANAL

ZONE - Barro Colorado Island; Coco Solo; Fort Kobbe; France

Pield;: Madden Porest: Pipeline Road. COUON PROV) =| Santa kita

Ridge. . PANAMA PROV. + Pearl Islands:. Tabega Island. COULECT ION

DATES: January, April, May, June, December.

OTHER COUNTRIES, 12 specimens from: ‘COLOMBIA, VENEZUELA,

BRAZ TL

REMARKS. This species has only been recorded from human

feces. Canthon juvencus is active diurnally and occurs at the

beginning and end of the wet season in lowland forests.

10. Canthon morsei group (Panama form)

Piees 145 15 us0 yp al

Canthon morsei Howden, 1966, p. 728; group currently under study

by Halffter and Martinez.

Type, Otlawa i.° Examined,

Length) 4.59 10°60 mms Colon dorsally’ shining: oreen,

frequently with golden reflections, occasional specimens black with

blue reflections; pronotum and elytra minutely punctate, surface

of head finely punctate, the punctures at least twice the size of

those on pronotum. Clypeus (Figs. 14, 40) anteriorly bidentate.

Gena laterally at clypeal junction feebly produced, obtusely

angulate; eye dorsally relatively narrow, approximately four times

as long as wide. Pronotum with antero-lateral margin nearly

straight, slightly inwardly arcuate posteriorly, small tubercle

usually present at middle of ‘wentral edge. Elvyiron with @ight

moderately distinct discal striae, each stria slightly but distinctly

impressed, the impression shagreened and with occasional widely

spaced punctures; intervals between striae shining, surface mainly

smooth, minutely punctate. Pygidium lacking distinct basal carina.

(in this respect differing from Mexican morsei), pygidial surface

dull, shagreened; shallowly, closely punctate. Antenna brown to

greyish brown. Prosternum with proepimeral carina present only

on inner half adjacent to coxae. Metasternum between coxae feebly

shining, finely shagreened and minutely punctate. Fore tibia

(Fig. 41) with spur of uniform width to narrowly notched apex,

spur of female slender and acute “at apex.) Hind femur ventrally

with both anterior and posterior line present adjacent to respective

margin.) Male’ genitalia “asin? fig) #15,

MATERIAL EXAMINED. PANAMA, 40 specimens from: CANAL

ZONE -— Barro Colorado Island; Gamboa (1 mile S); Madden Forest.

CHIRTQOUL PROV. =) Hartmann’s Finca.’ COLLECTION. DATES: May,

June, July, September. |

OTHER COUNTRIES, 15 specimens from: MEXICO, EL SALVADOR,

ECUADOR.

30 Contrib. -Amer, Ents. inst.) vole 16,-ne. hb, 1981

REMAKKS. This:species has been captured at carrion:.and:.on

human, feces. Canthon morsei is active diurnally, occurs only: in

the wet season and is foumd in moist forest over a broad, range -of

elevations.

il. Canthon angustatus Harold

bigs.- 42, 43

Canthon aneustatus Harold, 1867a, p. 79.

Type, Paris. » Not «seen.

Length 4.5 10 6.5 mm. Dorsally shining; head and: pronotum

largely yellowish tan to reddish brown with margins darker, often

nearly black; pronotum with median black marking present at

anterior and posterior margins, the small elongate anterior marking

occasionally obsolete; larger posterior marking always present,

often hastiform; elytra dark reddish brown, often semitransparent

with metathoracic wings vaguely visible. Legs usually yellowish,

apices and bases of femora and tibiae blackish. Clypeus (Fig.

42) anteriorly quadridentate, gena not produced at clypeal junction;

eye moderate in size, approximately 2 1/2 times as long as wide.

Pronotum with antero-lateral margin irregular; small tubercle

present On Margin in Unworn Specimens. Blytron with discal

Siridae Sepresenied only by extremely faint lines. .Pyoidium: with

fine Dasal carina complete, surface finely shagreened,, dull.

Antenna with club yellowish brown. Prosternum with proepimeral

Carina incomplete, the carina. broken in: lateral, fifth... Metasternum

between «coxae shining; very finely closely punctate. Fore ‘tibia

(Fig. 43) with spur of male apically cleft; tibia moderately

expanded, elongate; spur of female. acute apically, very feebly

Ssimuous.. hind femur ventrally lacking anterior and posterior

lines.

MATERIAL EXAMINED. PANAMA, 320 specimens from: CANAL

ZONE = Barro Colorado Island: Fort San Lorenzo; Pipeline Road.

COLON FPROV;.- Alhajuela,. (COLLECTION DATES: Every month of the

year. |

OTHER COUNTRIES, 19) specimens. from: BELIZE,<-NICARAGUA,

COSTA: RICA; cECUADOR, PERU.

REMARKS. This species has been collected: sitting on» leaf

surfaces and. at howler monkey and.human feces. One specimen

Dearsoan “at licht label. -Canthonangustatus:is active,diurnally,

and-on Barro Colorado, Island. is most, active during the. periods

from 0600 to 1000 hours and from 1500 to 1800 hours. These

intervals: coincide with the defecation periods of the: howler monkey,

the .feges of which angustatus appears fo prefer...This species. is

the most abundant scarabaeine on Barro Colorado Island, with

hioh’ population’ levels indintained in.all months of. the, year:

Teneral adults also occur. throughout the year, suggesting continu-

ous reproductive activity.

This is one of only two species of Scarabaeinae (the other

is Canthon subhyalinus) that occurs in the forest canopy (> 20 m),

where individuals form: and roll primate feces that is adhering. to

leat surfaces. °° The beetle remains attached to the ball: as it) falls

Howden and Young: Panamanian Scarabaeinae ot

to the. ground, landing on top ofthe’ ball... Once on the ground

the. beetle reshapes the: fecal. material into’ @)pal) and then rolls

it away. The species is known to: occur only in lowland moist

forest areas where howler monkeys are found.

12. Canthon subhyalinus Harold

Bigs ekg 5

Canthon subhyalinus Harold, 1867a, p. 79; Halffter and Martinez,

1 1, Ban, Cn

Lype vor \cotype,, Paris... Not seen,

Length 4.5. to..5.5 mm... Dorsaliy: with .jbead,*pronetum, base

and «suture. of elytra, and pygidium,; /shimineybiack. Disc of each

elytron translucent tan to reddish brown, the folded metathoracic

wings often partly visible through elytron. Clypeus (Fig. 44)

anteriorly: bidentate. «Gena laterally .at elypeal junction with

margin continuous with that of clypeus or very slightly produced;

eye dorsally approximately three times as long as wide. Pronotum

with antero-lateral margin interrupted at middle by ventral

tubercle, the margin anterior to tubercle straight, posteriorly

from tubercle to median angle inwardly arcuate. Elytron with

discal striae obsolete or absent. Pygidium with basal carina

complete, pygidial surface finely transversely shagreened. Antenna

brown. Prosternum with proepimeral carina complete. Metasternum

between coxae shining, mostly smooth with scattered minute

punctures. Fore tibia (Fig. 45) with spur in male expanded to

unevenly divided apex, one lobe of division slender and sharply

acute, the other lobe four or more times as broad, tapered

abruptly and often irregularly to sharp apex. Spur of female

slender, tapering to acute apex. Hind femur ventrally lacking

lines adjacent to margins.

MATERIAL EXAMINED. PANAMA, 106 specimens from: CANAL

ZONE. - Barro Colorado Island: Rio, Trinidad: Road to 1.4 Pita.

DARIEN PROV. -— Santa Fe. , COLLECTION DATES: January, February,

March, April, May, June, July, September.

OTHER COUNTRIES, 23 specimens jfromu, MEXICO, COSTA RIGA,

COLOMBIA, PERU.

REMARKS. This species has been found in forest leaf litter

and perching on leaf surfaces; and at howler monkey, human,

jaguarundi, and tapir feces. Canthon subhyalinus is diurnally

active on Barro Colorado Island and, like angustatus, is particu-—

larly active during the periods of 0600 to fO00 hours and 1500. to

1800 hours. This species also seems to prefer howler monkey feces

and is only rarely found on other fecal material. Competition

with angustatus appears to be reduced, however, by a preference

of subhyalinus for flat terrain with a high canopy, areas.in which

angustatus occurs in relatively low densities. C. subhyalinus can

also obtain primate fecal material adhering to leaf surfaces in

the forest canopy. Most teneral and mature adults occur in the

dry season, with only worn and damaged individuals appearing in

the mid to late wet season. The preferred habitat is level lowland

moist forest.

32 Contribs Amer. Ent. cinstes wok: 16) uno. 945 0981

13) Canthon euryscelis Bates

Figs. 46, 47

Canthon euryscelis Bates, 1887, p. 28: Halffter and Martinez, 1977,

: p. 79-80.

Cotypes, London and Paris. London specimens examined.

Length 3.8 to 5.2 mm. Dorsally with head and pygidium

feebly shining or dull (granular), pronotum and elytra smooth and

shining; uniformly black with bluish reflections. Clypeus (Pig.

46) anteriorly bidentate. Gena laterally at clypeal junction

feebly, obtusely angulate; eye dorsally approximately three times

as lone as wide.” Pronotum with antero-lateral margin sinuate,

ventral edge with tubercle. Elytron smooth, striae obsolete except

for lateral one. ‘Pysidium with fine, complete basal carina;

pygidial surface shagreened and minutely punctate. Antenna

brown to tan. Prosternum with proepimeral carina complete.

Metasternum between coxae shagreened and very minutely punctate.

Fore tibia (Fig. 47) with spur of male slightly expanded apically,

shallowly notched; spur of female tapered to acute apex. Hind

femur ventrally lacking lines adjacent to margins.

MATERIAL EXAMINED. PANAMA, 1 specimen from: CHIRIQUI

PROV. — Bugaba.

OTHER COUNTRIES, 12 specimens from: MEXICO, BELIZE,

GUATEMALA, COSTA’ RICA.

REMARKS. Some of the non-Panamanian specimens were taken

in pit-fall traps baited with human feces.

14. Canthon femoralis (Chevrolat)

Figs. 46, 49

Caprobius femoralis Chevrolat, 1834, p. 110.

Type or cotype, London. Examined.

Canthon femoralis (Chevrolat); Harold, 1868, p. 62.

“Tenens 70 to 6.5 mm. | Dorsally ‘shining black with bluish

reflections, elytra frequently partially brown, particularly near

apices. Clypeus (Fig. 48) anteriorly quadridentate. Gena laterally

at clypeal junction not notched or produced, the margin continuous

with clypeus; eye dorsally relatively wide, approximately 2 to

2 1/2 times as long as wide. Pronotum with antero-lateral margin

inwardly arcuate behind tubercle on ventral edge. Elytron with

discal striae obsolete. Pygidium with basal carina complete,

pygidial surface feebly shining, finely shagreened and occasion-—

ally with minute punctures. Antenna brown, club greyish brown.

Prosternum with proepimeral carina complete. Metasternum between

coxae smooth and shining. Fore tibia (Fig. 49) with spur of

male slightly expanded to notched apex, spur of female slender,

apex acute. Hind femur ventrally lacking lines adjacent to margins.

MATERIAL EXAMINED. PANAMA, None.

OTHER COUNTRIES, 54 specimens from: MEXICO, GUATEMALA,

COSTA RICA, COLOMBIA, ECUADOR.

REMARKS. Specimens from outside Panama have been collected

during the day flying to human feces. Since the species has been

Howden and Young: Panamanian Scarabaeinae ou

taken in countries on either side of Panama, there is a distinct

possibility: that;it will also| be) found: in Panama,

15. Canthon lamprimus Bates

Figss:' SOs hcok

Canthon lamprimus Bates, 1887, p. 29; Halffter and Martinez, 1977,

Dei CUE

Cotypes, London and Paris... London specimens examined.

Length 4.0, to.535 mm. »Dorsally, shining, green), often, with

golden reflections; all..dorsal, -suriaces often, with scattered «minute

punctures. Clypeus (Fig. 50) anteriorly bidentate. Gena laterally

at .clypeal,.junction,, slightly, produced,..obtusely angled: eye, dorsally

approximately three times as long as wide. Pronotum with antero-

lateral margin,.interrupted.at.middle bysventral sbubéercie. the margin

anterior..to,tubercle. straight, posteriorly .to; median, angle inwardly

arcuate... Elytron, with.,alb.but, lateral .6teia’ ebsoltere.er (absent.

Pygidium with basal carina complete, pygidial surface dull,

closely: shagreened.) Antenna |browil; (clap ereyn wa Prosternum with

proepimeral carina complete. Metasternum between coxae shining,

smooth except for scattered minute punctures. . fore tibia (Fig. 51)

with, spur gradually expanded ‘to apex, apex moderately notched:

spur’ of female tapered to ,acute .apex.. | Mindy temun vente tly

lacking distinct lines adtacent To \marotines

MATERIAL EXAMINED. PANAMA, 225 specimens from: CANAL

ZONE. = Ancon;::. Barro, Colorade island: Coeo: Boies Fart. Kopbe*

Frijoles; Madden Forest; Margarita (4 km NE); Paraiso; Pipeline

Roads. Rio, Ciricito;. Rio Trinidad.) Skier OMe ou alee mee ey ae gr ve

El Valle. (Las Minas) Trail, 2400. ft, jp) COLON. PROVs j=) Alhajuela:

yanta Rita Ridge... .PANAMAU PROV. Arrat tans CeO Ree gO 3

Cerro,,Campana.,, COLLECTION "DATMSc january ovrebraary. Narch,

April, May, June, July, September, October, November, December.

OVHER. COUNTRIES) No. records.

REMARKS. Canthon lamprimus has been found sitting on leaf

surfaces; at, mouse, peccary,,) rat, tamandua,,.and.cnake:. carrion;

and from coati, howler monkey, human, margay, and tapir feces.

This species on ,Barro,Colorado Island. is active, throughout

the day and is most abundant. during the dry season. Teneral

adults occur only in the dry season and adults captured in the

mid - wet season or later are very worn. Canthon lamprimus is

one of only six species of Scarabaeinae that occur on carrion in

the dry season, whereas it is one of 24 species that utilizes

carrion..in) the wet,season.:, lts primary food..in» both seasons,

however, appears to be vertebrate feces. Although this species

has been,collected at elevations of 700 to 900 m,)it occurs” primarily

in lowland moist forest.

isase Canthon ‘caelius Bates

Canthon caelius Bates, 1887, p. 29 (possibly = lamprimus Bates).

type, ‘London. "Examined,

Bates described this species from a unique female. No

34 Contrib. Amer. nate list... voli 213,) mo, 15-1981

subsequent material assignable to this species has been discovered

to the authors’ knowledge: “There are’ three possibilities: 1,

caelius is a rare species or one occupying some unusual niche;

2, the specimen described as caelius is a very atypical example

of lamprimus; 3, specimens that we have assigned to morsei should

be called caelius, but if so Bates' description of the bicarinate

hind tibia is incorrect which seems unlikely.

Halffter and Martinez (1977) have been the most recent in-

vestigators tO examine. the type and, they. relate it to lamprimus,

separating it as follows: eee

Color .and microsculoture equdl-on ihe pronotumxand elytra;

green with copper retlections; leas brown. lamprimus Bates

Pronotum brilliant @reen; élytra Subopaque,: bluish, black;

tibia and tars? dull brown, ventral, surface of Temora

With mcrae. orcen, vellecrions 1.7% 4 i) 91 eadelius-~ Bates

In addition to the above differences, Bates mentions that

caelius has a more distinctly punctured pronotum. Unfortunately

the size and density or: the pronotdl punctures, the color, etc.,

all vary in the séries of Jamprimus.we have examined and we

suspect that the name caelius is based on an atypical lamprimus.

It as for his reason that we have tot includéd@it in ‘the key or

included’ a anore formal: description.

1G. “Canthon vitidis meridionalis (Martinez, Halfiter. and. Halffter)

Pies (525° 53

Glaphyrocanthon viridis meridionalis Martinez, Halffter and

Ha lifter, 1964p. 32"

Canthonm Wiridis meridionalic (M., BH. & H.):°> Halffter and Martinez,

LDL Pp OU:

Type, Washington... Examined.

benoth 5.010.575 mm. Dorsally dull, finely: granulate;

dark blue. to bluish black,. Clypeus (Figo. 52): antertorly Didentate.

Gena lateraliy ‘at clypeal junction obtusely anoulate, appearing

feebly toothed; eye dorsally approximately three times as long as

wide. Pronotum with antero-lateral margin slightly inwardly

arcuate, ventra) -edee “lacking.distinct tubercles: Elytren with

discal striae obsolete, indicated by faint depressions. Pygidium

with basal Carina’ complete; pyeidial surface dull, granular.

Antenna brown to dark brown. Prosternum with proepimeral carina

complete. Metasternum between coxae feebly shining, finely granular.

Fore tibia. (Fie. 53) Sith imner margin sinuate tear middle, spur

of male expanded and slightly cleft apically, spur of female

Sinuoushy Papered. Hing: .femur ventrally lackine lines near margins.

MATERIAL EXAMINED. PANAMA, 10 specimens from: CANAL

ZONE = Ancon;. Barro Colorado: Island ;. Pipeline Road; -Tabernilla.

PANAMA “PROV, = Belle Viste: Cerro Aguli. « COLLECTION -DATES:: April,

May, June, july.

OTHER, COUNTRIES, 3 specimens from: EL SALVADOR, NICARAGUA,

COSTA RIGA.

REMARKS. This ispecies. has. been collected sitting on leaf

surfaces; froma, nottine-log,. from. carrion, and from.human-.and

tapiy feces. .Canthon viridis meridionalis is: diurnally active,

Howden and Young: Panamanian Scarabaeinae 35

occurs during the early wet season, and is found in moist forest

below 1000 m in elevation.

Pseudocanthon Bates

Pseudocanthon Bates, 1887,1p. 35; Matthews, 71966, p. 83.

Type-species: Canthon perpiexus LeConie, by monotypy.

The following combination of characters will separate this

gcnus; from. related genera “in, the; Canthonina:(Haliiter: and

Martinez, 1977): size small, length less than 6 mm; clypeus

quadridentate; eye dorsally large; vertex with hind edge un-

margined; pronotum parallel sided or nearly so in basal two-thirds;

excavated propleura with proepimeral carina complete; elytra with

scattered. minute, setae, stria 7.obsolete anteriorly, , stria 8

obsolete posteriorly; fore tibia with inner apex sharply angulate.

This is a small genus containing nine species, most of which

are Antillean. One species occurs in Panama.

1. Pseudocanthon perplexus (LeConte)

Figs, 165'.19

Canthon perplexus LeConte, 1847, p. 85.

Type, Cambridge. Not seen.

Length 3.6 ‘to''5.0° mm, e@redtest width 705 "4,0 min:

Oblong-oval’ (Fig. 16), black, shiny, legs rurous. "Head (Fic, 19)

finely alutaceous with dense, shallow punctures. Dorsal portion

of eye twice as long as wide. Pronotum with shallow, dense,

evenly distributed punctures; areas between punctures smooth.

Elytral striae distinct, impunctate and superficial; intervals flat,

smooth, with scattered, very short, sparse setae. Pygidium with-

out transverse basal carina, shallowly punctate. Median lobe of

metasternum densely punctate. Last abdominal sternite shortened

medially in male. sternite of female not shortened medially.

Fore tibia with three prominent teeth on outer margin, proximal

tooth somewhat reduced and separated from distal pair; fore tibial

spur of male spatulate, bifurcate apically; fore tibial spur of

female tapering to acute apex, reflexed toward outer margin of

tibia; hind femur margined on anterior border of ventral surface.

Tarsal segments trapezoidal, with first’ segment of hind tarsus

longer than second.

MATERIAL EXAMINED. PANAMA, 12 specimens from: CANAL

ZONE = Balboa; Madden’ Vorest. COULECTION’ DATES: May, ‘lune,

OTHER COUNTRIES, 62 specimens from: UNITED STATES; MEXICO,

EL SALVADOR, “COSTA "RICA, COLOMBIA TRIN EDAD.

REMARKS. Pseudocanthon perplexus (LeC.) and P. chlorizans

Bates may be geographic races of a single species (Howden, 1966).

Specimens of chlorizans Bates as redescribed by Matthews (1966),

are seemingly indistinguishable from perplexus (LeC.) from the

United States. If only one species is represented, then perplexus

(LeC.) (= chlorizans Bates) has one of the most extensive distri-

butions of any Scarabaeinae occurring in Panama, ranging from

the United States to South America and to the Antilles and Trinidad.

36 Contrib. Amer. Vem. oinst) oles, me. ol ge 1

Specimens ‘have ‘Deen collected’ at light,’ in) forest floor: litter,

in carrion traps, and at human feces. he’ data suggest ‘nocturnal

aclivity \and «al wide range. ot.food utilization... [ae species occurs

primarily in lowland forest. situations,.in, Panama.

Deltochilum Eschscholtz

Deltoctilum” Eschscholtz, 1622,°"o. 37: (Howden, “1966, -p. 733.

Type-species: Deltochilum dentipes Eschscholtz, by monotypy.

Moderate to large species, 12 to 28 mm in length. Anterior

edoe ofvelypeus with two or four teeth. “Anterior angles of prono-

itm ‘acute. “Elytron with humeral umbone’ usually with short’ longi-

tudinal carina and with at least one, often two, additional

carinae between umbone and lateral margin of elytron; third to

seventh intervals near elytral apex briefly, longitudinally

carinate. “(Middle and “hind ‘tibiae Not ‘expanded: near “apices;

middle tibia with two apical spines; hind tibia with one apical

Spine: Anterior tarsus oréeatly reduced or absent.

ppectes an the “oes ‘Are typically bali— rollers, the balls

Herne composed of either dung "Or Carrion.

Key to. species of Deltochilum

Li. Head (in normal position) wider than long: clypeal

tecthuesepateted medially by iwo,or mene Limes

their, own bength; pyeidium, apically jrounded.. not

Strongly. netlexeds ata pe) oo. saligs 2

Head (in normal position) approximately, as "lene

asywides clypeal. teeth, separated. medially by. .no

mone than own lengih: pygidium. apically. acute,

reflexed (possibly a male character); length 11

PO ee i a On Malou acropyge:... Bates

ue Median clypedl geeth) separated by more than .toree

pines), (hein tengtho. pyeidium, nearly tlat:. second

hing tarsal ceoment as large as third; one

lenotO lO) mm on oredtem, 6 a... a >

Civpeal teeth separated iby. no..more than ee

Their. benoths. pyoidinm: comvex;, second hind

tarsal Seament.as large jas third and fourth

COULD INSCUS eM ET Ws Oe Oil VLE SS mi eum ienee 1h eye 4

ce Antertor, half of, lateral pronotal. margin: straight

Of With avusltentucurve: dorsal: surface iDlack

with CeRRcky reflections; length 26 to 28 mm

1. gibbosum panamensis Howd.

eetene: bale. or ae pronotal margin sharply

Simuate; dorsal surface, with purple reflections;

LOR EO ee ee ee eC a RUIN Bar Mm.

A Dorsal surface with bluish reflections; length 14

to LO mms mangin,of lind trochanter. feebly

COGTHGG: DA OIA a a parile Bates

Howden and Young: Panamanian Scarabaeinae of

Dorsal surface with brownish reflections; length

{ie to 13 mm} manginwof hind) trochanter evenly

APCUG CS AT DOU Gewese) ne An pseudoparrle Paul.

1. Deltochilum gibbosum panamensis Howden

Fig. 54

Deltochilum (Hybomidium) gibbosum panamensis Howden, 1966, p.

MDS eee PRRRRM isced NT eOmOn On Cie a Naan

Type) Ottawa, examined

Length 26 to 28 mm. .Color black with coppery reflections.

Anterior edge of clypeus (Fie. 54) quadridentate, ithe outer teeth

sometimes: ‘feebly,developed,..«Pronotum with lateral. margin, anterior

and posterior to rounded median angle nearly straight, often

feebly arcuate; either. inwardlyor, outwardly... Blytral) sitiae, repre—

sented by rows of poorly delimited shallow punctures; umbone

longitudinally carinate; between umbone and complete epipleural

carina’, (a. humeral, carina extends) posteriorly slightly dese, than

half. length, of elytrons five small carinag (\preseqt nears apices ot

third to seventh iintenvals.. Pycidiuim \veny ireebix convex, curlace

with contiguous ocellate punctures, Hind trochanter with tuft of

setae.

MATERIAL EXAMINED. PANAMA, 11. specimens from: BOCAS .DEL

TORQ) PROV... =. Rio. Changuinola,, 2400) fi.) GaN AL OND et Pd pe lane

Koad. COLON. PROV. ~) Santa’ Rita Rid@eiy (Pia MAA BO iin A earn

Campana. COLLECTTON. DATES: sJune,) Septenper si) etobern.

OTHER COUNTRIES, 4 specimens (of g. panamensis) from:

COSTA RICA; 150 specimens (of gibbosum subspecies) from: UNITED

STATES, MEXICO, BELIZE, GUATEMALA, EL SALVADOR (closely

related forms are also in South America).

REMARKS. This’ species has been recorded from human feces

and rat carrion. Iti seemingly vrs active) only at might auring’ the

wet season, occurring \in ‘moist forests below! 1000 “mim ‘elevation.

Zi, Deltochilum mexicanum Burmeister

Pigidi55

Deltochilum mexicanum Burmeister, 1848, p. 135.

TC “pe py Hal lel (TG tine: Mies) vit Noe eeeny

Length, 19.10.24) Mme wiC Color) Diack eatin: Diuaieihy Wor alin le

reflections. Clypeal margin (Fig. 55) anteriorly bidentate,

laterally margin on each side distinctly sinuate. . Lateral, pronotal

margin anterior to median angulation abruptly sinuate (almost

angulate) near middle. Elytral striae represented by rows of

small, indistinct punctures; umbone with well-developed longitudinal

carina; epipleural carina’ complete: five, short) longitudinal carinae

present) near apices, of :third. to, seventh intenvals,. «“Pyeidium: flat:

closely, ocellately punctate. Hind trochanter lacking setae,

posterior margin obtusely angulate in both sexes.

MATERIAL... EXAMINED, .PANAMA,..13.specimens from: BOCAS DEL

TORO, 2 ROY coin Rie Aer winole AAO Cty ODO Ae i CR LOM ED BRCM oe

Hartmann’ s. Finca,,...0500/i0ns.Repr,., ba. Portunay:3200 )ft.3: Moloam de

38 Contribs Amer. obnt. 1nst. “wolle bo, no. 1571961

Chiriqui, 4000-5000) ft "“DARTEN PROV. = Cerro: Tacarcuna,’ 4800 ft.

COLLECTION (patES> May; july, September.

OTHER COUNTRIES, 85 specimens from: MEXICO, GUATEMALA,

El SALVADOR, ‘COSTA RIGA, COLOMBIA, PERU:

REMARKS... his species has. been collected.at. human -feces

and unknown carrion. il 15 active mainly at night and during

the wet season. It is restricted to middle elevation moist forest

habitats.

8. Deltochilum parile Bates

Fig. 56

Deliochibum parite Bates, YOO), 6.55. Patan, 1930, p. 231.

Lecrely pe. 1 artis... NOE Seen.

Leneth 14 to’ 16 mm. Color black, often with bluish’ reflec-

tions. Anterior edge of clypeus (Fig. 56) bidentate, distance

DeLWeen Teeth slisnily less than twice lénoth OF one tooth.

Lateral prendial Margin nor apruptiy anoulate, arcuate ‘medially’.

Elyivar striae opsolcre,” represented by rows of Teéblée depressions

(punctures); humeral umbone feebly carinate, humeral carina

extenaine fall the éeih ot elyiron, “epipleurdl carina complete;

third to seventh @lytral intervals cach with short carina or

tubercle Wear‘ apex. “F yoldtum convex, surface with fine scattered

punctUves. | Hind trochanter or mate with posterior margin’ angulate,

margin Nearly Straient im ‘female:

MATERIAL EXAMINED. PANAMA, 5 specimens from: CHIRIOQUI

PROV. ="Harimann 6 Einca, ‘COLEBCTION DATES: May.

OTHER COUNTER IFo. 20 Specimens from: “COSTA RICA, ECUADOR:

REMARKS. This Gpecies hasbeen ‘collected on carrion, and‘is

seemingly active nocturnally. Distribution is restricted to middle

and high elevation moist forest habitats.

4. Deltochilum pseudoparile Paulian

Pig«.5/

Deltochilum (Deltochilum) pseudoparile Paulian, 1938, p. 283.

lype, Paris... Not seen.

Leneth 11 4e.13ymm. = Color Black, ‘often with brownish’ -reflec-

tions. Anterior edoe of clypeus (Pig. 57) bidentate, ‘distance

belween*“ceth only. one third more than lenoth of one tooth.

Lateral pronotal marein arcuate’ medially, not abruptly angulate-

Elytra and associated Carinae resembling’ parile except punctures

of intervals more pronounced and carina on humeral umbone some-

times doupled. —Pyoidium only moderately convex, ‘surface with

fine scattered ‘punctures. * tind: (trochanter im both sexes with

posierior marein nearly straroht.

MATERIAL EXAMINED. PANAMA, 98 specimens from: CANAL

ZONE — Barro Colorado Tsland: Madden Forest:'Pipeline® Road.

COLON PROV. = Santa kita Ridge, ~“COLEECTION DATES:: February,

April. “me, tuly, Wuacust, “September, Octener, “November, December.

OTIES “COUNTRIES, 49 specimens from: MrEAICO. COSTA -RICA.

KEMAKKO. This species “nas “been recorded from Human ‘Teces;

Howden and Young: Panamanian Scarabaeinae 30

and from agouti, peccary, rat, and tamandua carrion. On Barro

Colorado Island, D. pseudoparile is nocturnally active, has only

been collected from carrion, and is most abundant during the late

wet season, when vertebrate carcasses are also most abundant.

The species is limited in Panama to lowland moist forests.

5. Deltochilum valgum acropyge Bates

FA gS .2 bea: 99

Deltochilum acropyge Bates, 1887, p. 36.

Lype,- Londen i.i-.Examineds

Deltochilum valgum acropyge Bates; Paulian, 1938, p. 273; Howden,

19665: Dias 18

Length 11 to 13 mm. Black, head and pronotum distinctly

punctate. Head (Fig. 58) approximately as long as wide; clypeus

with two distinct anterior, narrowly separated teeth, and on each

side at clypeal-genal margin with low, obtuse tooth; vertex closely

punctate, nearly. flat;. dorsal. eyes, large... Pronetum, with) dateral

margin angulate at middle, otherwise straight to slightly sinuate

before and behind; pronotal surface punctate, closely so laterally

and, posteriorly. Elytral striae indicated by fine double line,

line expanded by widely separated, shallow strial punctures;

discal intervals opaque, with numerous shallow, approximate

punctures; humeral umbone and apices of third to seventh intervals

carinate. Pygidium (Fig. 59) with apex thickened, acutely angled

and outwardly produced; disc nearly flat, shallowly punctate.

Fore femur with margin unmodified; fore tibia with three apical

teeth (the middle one smaller) and numerous serrations on outer

margin. Apical half of hind tibia bent inwardly (probably true

of males only).

MATERIAL EXAMINED. PANAMA, 1 specimen from: CANAL ZONE -

Pipeline Road, 27.V.1977 (Ratcliffe).

OTHER COUNTRIES, 1 specimen (type) from: BELIZE.

REMARKS. The unusual pygidium will readily identify the

species. Considering the fact that over 100,000 Scarabaeinae have

been examined from Central America and northern South America,

with only two specimens of acropyge being seen, it would appear

that either the species is exceedingly rare or has some unusual

habit. The exact status of acropyge (= valgum or subspecies or

valid species) can only be decided when more material becomes

available.

Cryptocanthon Balthasar

Cryptocanthon Balthasar, 1942, p. 36; Howden, 1973a, p. 39.

Type-species: Cryptocanthon paradoxus Balthasar, by mono-

typy-

While this genus has not been recorded from Panama there

is a strong possibility that it will be found there, since the

genus is known to occur in Costa Rica and Colombia. Because of

this the generic description is included.

40 Contribs (AMer Ene last. wooly Oro ios]

Size).small, 2.5 toa75: mm, oval (Pig. 60)... Clypeus with

two.téeth ‘on .anterior margin; ‘clypeal—frontal suture obsolete.

Bye: nov extending onto dorsal surface of vertex, "Visible. from

above only af head) extended. Sides: of. pronotum obtusely ancgulate;

pronotal disc ‘moderately convex except for shallow concavity on

each side pear lateral posterior marein.,. Surface of head and

pronotum, at least laterally, coarsely punctate. Scutellum absent.

Blytrom moderately to-stronely ‘convex, Six or’ seven obsolete dorsal

Sitlae,\ each indicated by two wavy dines. Intervals flat to

Sligntly. convex} Most Species wilh a) row Ol setose punctures on

each side of each interval, (Side on elytron “at seventh stria

ridged and sharply inflexed (as in some Deltochilum), the inflexed

portion’ at same anole bul wider than (epipleuron @€xcept near apex;

epipleuron distinctly raised: above infiléexed portion of elytron.

Pygidium unevenly convex, with a distinct swelling or longitudinal

ridge, medially, base near midline abruptly. transversely concave;

basaltedge \cariniform medially. «/Prosternum. laterally excavated

for’ reception of antennal clube ) Mesosternum broad, separated

from “méetastéernum by curved suture or ridge. Metasternum broadly,

evenly convex, lacking any distinct excavations; broad between

middle coxaeé.. Abdomen six’ segmented, terminal segment medially

about three times as jong as penultimate segment; terminal segment

laterally narrower in ‘female than ih’ male . Fore: tibia anteriorly

truncate. Fore tarsus short, aboutas long as: truncated’ portion

of tibia. Middle and hind tibiae narrow basally, broadened

medially ‘and (or) toward apex) “Hind tibia often longer and’ ‘bent

inward in male. Antenna: Nine seemented, «club three: segmented,

similar in Shape to club. of some, Canthon.

Cryptocanthon is not closely related to: other New World

Canthonini. “Balthasar related the genus to Ipselissus.: Olsoufieff

(= Ipsepilissus Paulian) and Zonocopris Arrow, but we believe it

is “perhaps more closely related to Bdelyropsis. Pereira, Vulcano

and Martinez. It resembles this latter genus closely in the shape

of the thorax (including the elytra) and abdomen. However,

Cryptocanthon can be immediately separated from any related New

World genus" by the shape of the eyes, ‘whichido‘ not extend

dorsailty “onto! the. vertex.

REMARKS. Most rof the: species have been ‘collected’ in very

wel ‘situations, both in lowland forest”.and “in cloud’ forest... Speci-+

mens” have: been taken sifting wet litter or in traps baited: with

human téces, . Lhe genus ranges from-Mexico to ‘Brazil.

Agamopus Bates

Agamopus Bates, 1887, p. 42; Halffter and Martinez, 1968, p. 226.

Type-species: Agamopus lampros Bates, by monotypy.

Size ‘small to’ moderate, 3.5 to. 6.5 mm; ‘oval,’ moderately

convex. “Head with clypeus.fteebly bidentate; frons and “vertex

lacking horns or carina, one (South American) species with small

tubercle. Pronotim ‘lacking protuberances ‘or grooves, lateral

fovea “obsolete; posterior maroainal bead lacking): at least medially.

Scutellum not visible. Elytron with eight striae (nine including

epipleural groove), eighth ventro-lateral in position, the seven

Howden and Young: Panamanian Scarabaeinae 41

discal $triae ‘represented by ‘slightly indented, punctare dines;

intervals nearly flat. Pygidium (Fig. 63) with basal groove and

a second, Teebly arcuate,’ deep transverse groove extending across

pygidium at apical third.» ,Prosternum ‘declhivous- ‘anterior To coxae,

nearly flat and relatively broad behind. Mesosternum moderately

wide, transverse. Metasternum broadly separating middle coxae.

Abdominal sternites distinct, narrowed: medially. © Fore tibia. with

apical two teeth: on: outer, margin d@istinel, thind moot ,erten

obsolete or poorly developed. Middle tibia) grnadually expanded” to

apex; hind tibia only feebly expanded to apex. Hind tarsus with

two basal segments subequal, basal segment approximately twice

as long as wide, terminal. segments. long and slender; all ‘tarsi

with claws.

Halffter and Martinez (1968) place Agamopus in the

Canthonina... Previously ‘the genus was, Incorrectly, placed, in, the

Coprini.,. ihe genus, containing, four or sive species, to eine rily

south, American..in. distribution. . All) of ine members .or ihe -cenus

can: be. recognized by the ‘relatively slender, hind tibia. and the

very characteristic pygidium with the deep transverse sulcus near

the’ middie.

1. Agamopus lampros Bates

Figs. 614702. °G3

Agamopus lampros Bates, 1887, p. 42; Halffter and Martinez,

1968, p. 236.

Cotypes, London and Paris. London specimens examined.

Length 3.2 to 4.8 mm. Dorsally shining brown to very dark

brown; pronotum finely punctate, elytral intervals (Fig. 61)

impunctate. Clypeus (Fig. 62) moderately reflexed anteriorly,

margin broadly, shallowly emarginate, feebly bidentate; clypeal

disc with transverse row of large punctures with scattered adjacent

smaller punctures. Frons and vertex smooth to finely punctate

posteriorly, protuberances lacking. Eye dorsally wide,. approxi-

mately 15 facets wide opposite posterior end of gena. Pronotum

lacking grooves or indentation, foveae obsolete. Elytral striae

represented by fine, shallowly indented punctate lines; intervals

nearly flat, shining. Pygidium (Fig. 63) with strong transverse

sulcus at. apical’ third. © Fore tibia tridentate, basal. tooth often

poorly developed, inner apical angle acutely produced in male,

not produced in female. Hind tibia relatively slender, not

expanded apically.

MATERIAL EXAMINED. PANAMA, 26 specimens from: CANAL

ZONE. - Fort Clayton. CHIRIQUL PROV. -— Boqueron; David; Dolega;

Potrera. PANAMA PROV,.= La CammanayiLa.Chorrera:, Rie Las

Lajas. COLLECTION DATES: May, September, December.

OTHER COUNTRIES, 46 specimens from: MEXICO, GUATEMALA,

EL SALVADOR, COSTAO RICA. COLOMBIA.

REMARKS. The species has been taken in traps baited with

human feces, and on ‘cow dune, carrion, “and at ‘light)*'/Agamopus

lampros is active at night and is found primarily on fecal material

during the wet season. Distribution of the species is seemingly

limited to low and mid-elevation areas with scattered forest.

42 ContribsvAmers Ent. imast., vol. 18, no..4d, 1961

The single Central American species, lampros, is easily

recognized by its odd, transversely sulcate pygidium (Fig. 63),

relatively slender hind tibia, and by the absence of pronotal

grooves or indentations.

Pedaridium Harold

Pedaridtum Harold, 1000, p.° 54; Paulian, 1936, p. 20/7.

Type-species: Pedaria hirsuta Harold, by monotypy.

Size small, body shape oval, sometimes elongate oval;

dorsally setose, sometimes sparsely so. Clypeal margin bi- or

quadri-dentate; head lacking horns or tubercles, some species with

vertex feebly carinate; eyes dorsally narrow to moderate in width,

Harrowed posteriorly. Pronotum convex, lacking horns, protuber—

ances or grooves; lateral fovea obsolete; posterior marginal bead

absent; lateral pronotal bead with close, ventral parallel carina

at edoe of episternum, the ventral carina obsolete or absent in

anterior third. Scutellum abeent or not visible.” Elytral disc

With @1gOnt Striae, each sirta represented by either a narrow,

shallowly indented, sometimes punctate, line or by a depressed

band of confused punctures; elytral intervals flat or raised, each

side of interval with a row of setose punctures (the related genus

Triehillum Hare has only 4a Single row of punctures on each

interval). Pygidium variable in size and convexity. Mesosternum

wide medially, separated from metasternum by wide U- or V-shaped

suture. Metasternum between posterior half of middle coxae broad,

surface punctate, setose, at least minutely so. Abdominal

sternites usually narrowed and fused along midline. Fore tibia

with two or three small teeth on outer margin in apical third.

Hind tibia gradually expanded toward apex, oval or flattened in

eross section at apical third. Hind tarsus with basal segment

shorter than adjacent segment, usually not slender and elongate.

Pedaridium and its close relative Trichillum are badly in

need of revision as the characters distinguishing the genera are

poorly defined and variable. Both genera contain a number of

small (under 10 mm) species. All have some dorsal setae and are

distinctly convex. Pedaridium is distinguished from Trichillum

(Paulian, 1936, Martinez, 1967) mainly on the basis of the length

of the basal segment of the hind tarsus. In Pedaridium this

segment is shorter than the next, while in trichillum if is longer.

Thevdiiiicutty ia wtilizing tais Character iS illustrated by the

eriginal placement Of the two Central American Species In

Trichillum when, according to. the characters utilized by Martinez,

1967, and others they should be placed in the genus Pedaridium.

Based only on present usage the three Panamanian species in this

complex should all be assigned to Pedaridium.

Key to species of Pedaridium

1 Clypeal margin quadridentate; vertex nearly

flat, evenly. feably, CONVEX york SoG TA. Meee ots Z

Clypeal margin broadly bidentate; vertex with

feeble transverse carina; length 3.5 mm.

Howden and Young: Panamanian Scarabaeinae 43

3; Dottimert vn. sp.

Zs Elytra with longitudinal median third: of intervals

convex and bare; thé aréa’ between tmiciuding

striae composed of broad band of confused

punctures, a row. of conspicuous setae on each

Side; length.3:0 to' 3.5. mm on %.. 1. pilosumr (Rob.

Elytral intervals flat, each with row of small

punctures on each side, most punctures each

with short, inconspicuous setae; elytral striae

répresentéed Dy line and vow of punctures:

length 3.3 Wim 3s eee 2 rev icelooul i. Sy,

1. Pedaridium pilosum (Robinson) New Combination

Figs, 66; 67°68

Trichillum pilosus Robinson, 1948, p.. 149.

Type, Washington. Examined.

Length 3.0 to. 4.) mm. .-Dorsally, @arvk brown to black: Nead,

pronotum and elytra (Fig. 66) with numerous, conspicuous, erect

setae; elytral setae approximately as lene a5 widih of one interval.

Clypeus (Fig. 67) quadridentate, inner teeth slender, separated by

a broad U-shaped emargination; ‘lateral teeth approximately halt

length’ of inner ones: “Entire surface ‘or nead, elypeal disc, “trons

and vertex even, very slichtly convex; evenly, coarsely “punctate;

each puncture with erect seta, surface between punctures smooth

and shining; eye dorsally narrowed posteriorly, at posterior of

gena “only one to three facets wide. § Pronotum with Ttovea repre—

sented by a/small, shining, rounded swelling,’ surface of pronotum

closely, coarsely punctate; punctures usually separated by less

than one diameter, each” puncture with central’ seta oLlytra with

inconspicuous, crenately punctate striaey each interval on each

side adjacent to striae with row of coarse punctures, middle

longitudinal half of interval tmpunctate, shining, convexly elevated;

seventh interval Carinate, €iehnin interval feebly so; apical sixth

of other intervals offen carinate. Pydidium and ventral surfaces

including femora’ ‘coarsely punctate; punctures: often shallow, most

setose. Pore tibia?’ with ‘three small teeth on outer*maregin ix

apical half. Sexual differences slieht; malice with sliohtly less

convex pygidium and with apical tibial spur wider. Male

genitalia as in’ fig. 60: :

MATERIAL EXAMINED. PANAMA, 89 specimens from: BOCAS DEL

TORO ‘PROV. — Almirante: >: CANAL ZONE>’= Barro Colorado Island:

Rio Ciricito; Madden Forest; Pipeline Road; Skunk Hollow.

CHIRIOUL PROVY (— Havimanm's* Fincay 1200 im, 1500 mo YCOCLE

PROV. = El Malle, ©2200 (1.99 COLON “PROV. —*Santa’ Rita’ Ridge.

PANAMA PROV. - Cerro Campana. COLLECTION DATES: January,

February, March, ‘April, May,” june,” jury.

OTHER COUNTRIES 39 specimens trom: “COSTA RICA, ECUADOR,

REMARKS. This ‘species has been collected in light traps;’ in

forest floor litter; in the fermented crown of a felled palm; at

horse, human, peccary, sloth, and tapir feces; and on unidentified

44 Contrib. wmes! inte nists, woh. oS gnd. 1. 1981

carrion. .Pedaridium, pilosum is necturnally active, and most

abundant in the dry season on Barro Colorado Island, where it

seems to be found almost exclusively on dung. The species occurs

in. both low. and middle elevation. moist forest.

There is one species, P. bradyporum-Bouc. that. occurs in

Costa Rica that might be confused with pilosum. The species was

originally described in the genus. Trichillum but as in the case of

pilosum should be placed in Pedaridium if one is to follow current

usage. If bradyporum is found in Panama, which would appear

likely, it may be distinguished from pilosum by the following

combination of characters: size smaller, approximately 3 mm, less

oval, setae slightly reduced in number and length, seventh

elytral interval moderately carinate but with apical sixth of

remaining intervals not distinctly carinate. In addition, the

punctures in the elytral intervals are smaller and distinctly

separated while in pilosum they are contiguous or nearly so. The

characters separating pilosum from the other two Panamanian

species are wiven in the key.

2. Pedaridium brevisetosum «n.' sp.

Figs. 69; (/0,.71

HOLOIYP RE. Male, leneth G.3. mim, ¢ereatest width 2:0. mm.

Dorsally very dark brown with scattered short erect setae (Fig.

69) on head, pronotum and elytra. Head and pronotum closely,

similarly punctate; punctures separated by approximately one

diameter; punctures of elytral intervals irregular, frequently small

to minute, Clypeus (Fis. ./0): broadly quadridentate, the inner

two teeth slender, widely separated by shallow emargination; two

lateral teeth with bases in nearly straight line with bases of

inner teeth; lateral teeth small, approximately half length of inner

teeth. Clypeus on each side laterad to outer teeth with a slight

obtuse angle present indicating that, in some specimens, six teeth

might be present. ,Clypeal disc, frons, and vertex essentially in

same plane, feebly convex. Eye dorsally moderate in size,

narrowed posteriorly, two facets wide at posterior edge of gena.

Pronotum almost evenly convex, lateral fovea indicated by a

slightly raised irregular area. Elytra with striae indicated by

faintly impressed lines, crenately punctate; intervals flat to

feebly convex laterally, not carinate, surface mostly smooth and

shining, a few coarse punctures present on either side near striae.

Placement of punctures irregular and in some areas intervals

completely impunctate; most coarse punctures each with short erect

seta. Pygidium convex, punctate-setose. All ventral surfaces

closely punctate, setose; in some areas punctures shallow; abdominal

sternites one to five constricted and fused at coxal edge near mid-

line, apical (sixth sternite) the only obvious segment extending

across midline. Fore tibia with two distinct teeth on outer margin

near apex; tibial spur small, nearly as broad as long. Male

cenitalia as: an.digw fla

FEMALE... Unknown.

TYPE: MATERIAL: Holotype, made, ae. Canal Zone, Barro

Colorado Ishand, \OC10'N,. J9°50'W, 16.V1L.19//, Hea, Hespenheide

Howden and Young: Panamanian Scarabaeinae 45

(Howden).

REMARKS. The non-carinate head and very feebly punctate

elytral intervals will distinguish this species from the others

known: to oecunsim Panama.’ The males genial ere: also. very

distinctive.

3+ Pedaridium bottimer? n. ‘sp.

Figs.uWey ch die. ie

HOLOTYPE. «Male, length 3:5: mm, .ereatest width 2:4° mm.

Dorsally very dark brown. Head, pronotum (Fig. 72) and,<10 a

lesser degree, elytral intervals coarsely punctate; pronotal punc-—

tures separated by one to three diameters, each with an upright

seta. Clypeus (Fig. 73), bidentate, teeth widely separated, acutely

angled; emargination between teeth shallow, approximately one

quarter as wide as head. Clypeal disc with widely separated

punctures, surface between very finely shagreened, shining. Frons

between anterior margins of eyes with low transverse carina;

carina depressed at midline, extending approximately two-thirds

of distance between inner margins of eyes. Vertex behind carina

with punctures similar to those on pronotum. Eye dorsally moder-

ately wide, narrowed posteriorly; four facets wide at posterior

end of gena. Pronotum evenly convex, lateral fovea indicated by

faint irregular swelling. Elytra with striae slightly indented;

strial lines interrupted by well-spaced crenate punctures separated

by three to four diameters. Elytral intervals slightly convex with

scattered punctures on each side near striae; placement and size

of punctures both being highly irregular, but with punctures

generally more numerous laterally and in apical half; seventh

interval rounded, not carinate. Pygidium moderately convex;

basal groove very distinct, nearly twice as wide as in other

Panamanian species; pygidial surface with scattered coarse punc-

tures, punctures medially separated by one to two diameters.

Ventral surfaces moderately punctate, femora feebly so; punctures

often with short setae. Sternites one’ to tive ‘becoming Tused hear

median third and very narrow, the narrow fused sternite extending

across the midline; width of this sternite one tenth or less the

median width of sixth sternite; sixth sternite approximately twice

as wide medially as laterally. Fore tibia with three distinct

teeth on outer margin in apical half; tibial spur distinct, broad

at base, narrowing to acute apex. Male genitalia as in’ fig. 74.

FEMALE. Unknown.

TYPE MATERIAL: Molotype. mate. Pranama, Canal Zone, Barro

Colorado Island, 13. VEIT 4963, "Li." Bottimer: (Ottawa). Paratype,

Lo male, "Panama, Canal Zone. Coco Selo’ Hospital, 7. Vib. 19/5

(Ratcliffe).

REMARKS. "The bidentate clypeus and’ the ‘transverse carina

on the frons will immediately distinguish this species from any

others known lO Occur in Fanama or melanporing ‘Countmes. Itoi

interesting to note that, while the external, morphology ‘differs

radically trom’ pilgsum, the oenttalla are quite similar. "This: is

the reverse of the situation found in brevisetosum which morpho-

46 Contribs Aimer. “Bint. Inst. vot. 15,>mo..1751981

logically appears: more closely related to pilosum. except for the

characters of the cenitalia which show radical differences.

3 The species described above is named: in honor of. Mr... Ls).

Bottimer who has collected a number of the species mentioned in

this paper.

Bdelyrus Harold

Bdelyrus Harold, 10094. p. 9/7; Pereira, Vulcano and Martinez,

1960, bp. 155.

Type-species: Bdelyrus lagopus Harold, by monotypy.

DiZe. Moderate, fo $0 mm. Body shape elongate oval;

feebly 16 moderately longitudinally convex dorsally. .Most of

dorsal Surfaces distinctly punctate. Head, lacking lhorns,.tubercles

or transverse carinae; clypeal margin obsoletely emarginate and

wery slignily, rellexed medially, or moderately, emarginate. or

produced and retlexea ait and méar midline. Pronotum. lacking

protuberances, fovea or abrupt indentations. Scutellum not visible.

Elytta. Gach with ¢éyen diccal striae. ihe ouwler Siria: adjacent. to

ADTUDL carina, delimiting sharply iInilexed epipleuron;. discal striae

representea by broad, Shining, sometimes punciate tines; lines, at

most, feebly impressed. ..Pyeidium varying in convexily, oiten

nearly horizontal, in approximately same plane as terminal

sternite... Prostermum declivous anterior to coxae, broad and nearly

flat posteriorly... Mesosternum broadly transverse. .Metasternum

anterior]y.widely. separaling middie, coxae, inner .coxal margins

very slightly convergent posteriorly,. Abdominal sternites not

fused medially, sutural lines distinct. Fore tibia relatively

broad ana flattened.in apical.hali, apically truncate, outer. margin

in apical half with three distinct teeth. . Anterior tarsus small,

length approximately equal to apical width of tibia; tarsal claws

small, Giten concealed. by long setae. Middle and hind tarsi with

basal segments approximately as broad as long, long ventral

setae often obscuring segments; width of segments diminishing to

apex.

The closely related Mexican-Central American genus Bdelyropsis

Pereira, Vulcano and Martinez, is easily separated from Bdelyrus

by its distinctly bidentate clypeus with a deep V- or U-shaped

Median cmaraination.. .for. other, generic, diiferences and, the

descriptions of the Mexican—Central American species of Bdelyropsis,

see Howden, 1971.

1. Bdelyrus seminudus (Bates)

Figs. 645. 05

Aphengium seminudum Bates, 1887, p. 42.

Cotyoes, London. and Paris... London. specimen. examined.

Bdelyrus seminudus (Bates); Pereira, Vulcano and Martinez, 1960,

Das 2.

Leneth 6.10.3 mm. . Dark brown to black; dorsum (Fig. 64)

closely .punciais.. less. 60 in elyiral siide;, punetures in most areas

Howden and Young: Panamanian Scarabaeinae 47

separated by one to three diameters; most punctures each with

minute central seta, some punctures annular; pronotum medially

with fine secondary punctures. Clypeal margin (Fig. 65) very

shallowly, broadly emarginate medially, not distinctly reflexed.

Eyes dorsally 8 to 12 facets wide near posterior edge of gena.

Pronotum with marginal bead poorly developed, absent posteriorly.

Elytral striae each represented by slightly impressed line inter-

rupted by moderate sized non-setose punctures. Pygidium with

posterior groove; surface punctate, shining between punctures.

Obvious sexual modification lacking.

MATERIAL EXAMINED. PANAMA, 9 specimens from: CHIRIQUI

PROV. -— Volcan de Chirigqui (cotypes).. .PANAMA PROV, — Cerro

Campana, 3000 {t.), Vill; Certo cjete, OF 17,0, O72) i, FOO mm, VI.

OTHER COUNTRIES, 42 specimens from: NICARAGUA, COSTA

RICA, COLOMBIA.

REMARKS. Only two recently (1970 to date) collected speci-

mens were seen from Panama. While Bdelyrus seminudus does

come to traps baited with human feces, closely related South

American species have been taken in the bracts of Heliconia and

from bromeliads. Others have been collected under wood chips

from a freshly felled tree in heavy Amazonian forest.

There are apparently several undescribed species in the

genus which is seemingly best represented in the Amazonian region.

Onthocharis Westwood

Onthocharis Westwood, 1847, p. 230.

Type-species: Onthocharis mirmidon Westwood, listed by Lucas,

1920.

Body shape elongate; dorsally glabrous, shining. Clypeus

bidentate, quadridentate or sexdentate; frons and vertex smooth,

tumid, slightly carinate or feebly biluperculate. —Bye donsaily

narrow, usually less than six facets in width. Pronotum relatively

evenly..convex,. dacking horns, tubercles, ,icges of, erooves: lateral

fovea small, shallow or obsolete. Scutellum minute or invisible.

Elytra nearly parallel sided; striae distinct, depressed; intervals

feebly to distinctly. convex... Pycidium evenly convex, relatively

large. Mesosternum, short to moderate in tenoth. , Metasternum

anteriorly broadly separating “middie coxac: inner edocs of coxae

slightly convergeni posteriorly, (Pore, [ibia with three teeth on

outer apical half. Middle and hind tibiae broad and distinctly

flattened. Middle and hina tarsi with at least basal segment

greatly expanded and flattened in species 5 mm and Over in size:

in species. 5 mm,and under basal tarsal segment oiten. cloneate,

only slightly tlatiened.

This genus has not been carefully studied since Westwood

defined it in 1847. The above description defines the genus

following current usage. However, as noted by Paulian (1939) the

one Panamanian species is atypical and actually belongs. to a group

of small species, usually less than 5 mm in length, that has a

shortened mesosternum and a narrow, elongate basal segment of

the, hind tarsus... Seemingly the closest relative to the Panamanian

epecies is: one OCcuUrring In the Upper Amazon Dasin near Leticia,

Colombia.

48 Conterb .Amer. dnt. clnses, wolus 18 n0.4d), F981

1. Onthocharis panamensis Paulian

Bias, go, 10

Onthocharis panamensis Paulian, 1939, p. 145.

Type, Cambridee. lb xamined.

Length 3.5 to 4:0 mm, @regtest width 1.0 mm; body shape

(Fig. 75) elongate, elytra distinctly longer than wide, almost

parallel sided. Dorsally shining, brownish black with cupreous

reflection or shining green; pronotal suriace with mumerous fairly

uniformly but widely-spaced small punctures. Clypeus (Fig. 76)

with two: slender, .clongate teeth, one on each side of U-shaped

Medidn emareinalion:Jateral.to eaen elongate: tooth, an obtuse

toolh and beyond a, second tecble or obsolete tooth; i.e. clypeal

marcin with four distinct teeth and two feeble or obsolete. ones.

Posterior clypeal disc and frons medially somewhat tumid, at

posterior edge on each side of swelling a feeble tubercle indicated.

ouriace of head closely, finely punctate; eyes dorsally very

Narrows Pronotum, lackime protuberances or orooves, lateral fovea

Shallow, small; anterior and lateral marginal Bead, present, bead

lackime@ posteriorly, “Blyiral striae distinctly imdented, obsoletely

punctate; intervals convex, some with minute, scattered punctures;

otherwise surtace smooth and. shining. .Pygidium relatively large,

smooth, shiming, with, scattered: fine punctures. Pore tibia

tridentate,

MATERIAL EXAMINED... PANAMA, 2 specimens from: CANAL

ZONE - Barro Colorado Island. PANAMA PROV. - Cerro Campana,

S60 my 2beIV 1978,

REMARKS. The small size,;shining dorsum, four or six, toothed

clypeus,.and elongate body, shape should be sufficient to, recognize

the species. [his te the only species in the genus recorded) from

Centralv.America.. The habits of \panamensis are unknown; one

specimen was taken sitting on a. leaf.

Scatimus Prichson

Scatimus, Erighson, (o47,. pp. 110; Balthasar,’ 1939, pp. 3/.

Type-species;: Scatimus cucullatus Erichson, by monotypy.

Size moderate, oO tO. J mm. Color, dark brown to black.

Anterior clypedl Marain dentate or distinctly sSinuate;: clypeal

Surface Not rugose.. Frons with transverse carina extending

approximately five-sixth distance between anterior margins of eyes.

Pronotum evenly convex; lateral fovea distinct, a line of punctures

extending anteriorly, line Sometimes very abbreviated. Elytron

with eight striae, six between suture and umbone, intervals on

disc flat, polished. Pygidium transverse, basal marginal groove

pronounced. Mesosternum closely, distinctly punctate. Middle

coxae widely separated, inner margins parallel. Abdomen with

six sternites, basal four often narrowed near midline particularly

in meles. .Middie and hind tibiae expanded .toward apex; outer

tibial surface with one, sometimes two transverse carinae.

The genus may be confused with some Uroxys or Ateuchus,

but the transverse carina of the frons and the distinctly punctate

mesosternum will distinguish Scatimus.

Howden and Young: Panamanian Scarabaeinae 49

Key (lo Species of) Scatimus

L Carina,,on,frons often at, least) sivehtbhy

elevated’ laterally, the .capina: medially

posteriorly arcuate.) oie die QuaGrVoontatue ina Lun.

Carina) on, frons mot, elevated laterally, ine

canina, medially anteriorly arouate . 02s oMaluie,. ilar.

1. Scatimus quadridentatus Balthasar

Bigee 1s Oy. eo

Scatimus quadridentatus Balthasar, 1939, p. 89.

Type, Vrague: Net ysceenk

Lenoth 4.5,.to. 7.0mm.) Cly pens ((Pigs..) 17.) 0/0) “anteriorly

Sharply bidentate,- margin: laterally lobed,’ sinuate orvarcuate.

Eye dorsally, in, typical, specimens, large, not’ narrowed, posteriorly,

about 1,3. times longer, than wide to: posterior edee of oenay | 1n

atypical specimens (usually from lower elevations) eye dorsally

distinctly narrowed. posteriorly, about twice as lone’ as wide. to

posterior edge of gena.,, Frons with carina extending three-quarters

the distance between anterior .margins,of eyes, carina highest

laterally; carina. posteriorly arcuate.’ Pronotum), either with

lateral line of four, or more punctures: extending ‘anteriorly beiore

lateral fovea or with clump of punctures in an elongate depression

anterior to fovea. | Fore \femur with: posterion,.( ventral) surface

with scattered to numerous coarse punctures; numerous coarse

punctures often present on specimens with narrowed eyes. Male

fore: tibial spur: not modified. Genitalia as (im tie.) (0).

MATERIAL EXAMINED. PANAMA, 28 specimens from: CHIRIQUI

PROV. -—.2 km/N Sta. ‘Clare, Hartmann Ss, bimncay) 1200) mi COLON

PROV. = 10. mi.. SE Colon, Gamtay Rites Ridge O70 maa CC Ti Me oe

El Valle.’ PANAMA PROV, =: Cerro Campana, O00 m. COLLECTION

DATES: May, june, July, December,

OTHER. COUNTRIES, 6 specimens from: COSTA RICA.

REMARKS. Specimens from Costa Rica:.and Chiriqul Prov.,

Panama, all have the wide dorsal eyes. Two females from Cerro

Campana, Panama Prov., vary slightly in eye width, being

slightly wider than those from the Santa Rita Ridge. Specimens

from Santa Rita Ridge, Colon Prov., all have the eyes narrowed

dorsally. Initially we believed that two close species were repre-

sented. Subsequently, after examining the male genitalia and being

unable to detect any differences and noting a seeming convergence

in eye size, we now consider the variation in eye size as possibly

intraspecific. Sinée eye Size’ often is related to time of activity

(nocturnal or diurnal) it is possible that we have combined two

species under the name quadridentatus. However, some proof is

needed to ‘show if the ‘difference in eye size’ is’ or is not’ coupled

with behaviorial differences.

Specimens have ‘been’ taken) at light, “at carrion and in

traps baited with human feces.

50 Contvib. Amer. Ent. dost. vole 10. no 1.446198!

2., Scatimus..ovatus. Harold

Figs 700,01

ocalimus ovatus Harold, 1262, a. 401.

on Py oe or COnype, Paris. «Net seen!

Length, > to 7 mm. Clypeus (Pie. 60) anteriorly feebly

bidentate, margin laterally arcuate or somewhat sinuate. Eye

dorsally narrow, nearly twice as long as wide. Frons anterior to

eyes. wiih low tvansveree carina. ol, uniform-height; carina anteriorly

arcuate (median portion Of Carina dnierior in position). .Pronotum

with longitudinal line of punctures extending from lateral fovea,

two or three punctures usually present posterior to fovea and a

lime anteriorly composed of “10 or more punctures, some contiguous.

Fore. femur with at least half of posterior (ventral) surface

cCOarsely punctate. Male tore tibial spur not. modified. Genitalia

as, in (ig. “or.

MATERIAL EXAMINED. PANAMA, 131 specimens from: CANAL

ZONE = Barre Coloradc island: Cabima: Camaron Arm: “Coco Solo

Hospital; Fort Clayton; Fort Gulick; Madden Forest; Tabernilla.

CHIRTOUT PROV. Aleadrrope; Bueaba, 600-1500 i175, David; Dolesa:

Gualaca: Potera; Kio Vabasara. COCEER PROV. — 1 Vaile: Penonome.

COLON PROV. — Santal Rita Ridge. LOS SANTOS PROV. — Macaracas.

PANAMA PROV. - Cerro Campana; Cermeno; La Chorrera; Tocumen.

VERAGUAS PROV, — Parailon. COLLECTION DATES: January, May,

june, july,. September, October, November; December.

OLHER COUNTRIES, 29 specimens from: MEXICO, EL SALVADOR,

COSTA RICH, COLOMBIA,

REMARKS. The species S. ovatus has either an unusually

wide range or, quite possibly, several species are included under

the name. There is geographical variation in the size of the eye

dorsally, shape of the clypeal teeth, dorsal punctation, etc., but

in Panama the specimens seen exhibit little of this variation.

This species has been captured on the feces of horse, cow,

howler monkey, man, Spicer monkey, ard tapir.

o. OValus 19 active ai night, occurs only on fecal material,

amd 16 resiticied [oO the wel season. lis distribution is one’ of

the “pDroadest Of the Scarabacinae in. Panama, occurring in’ the

lowland forests of all but two provinces.

Uroxys Westwood

Uroxys Westwood, i842, p.59;. 184/, pe 229.

Type-species: Uroxys cuprescens Westwood, by monotypy.

Leneih,.3 to 41mm... Golor red-brown.to. black: . Anterior

clypeal margin dentate; clypeal-frontal area sometimes with trans-

Verse; Carfina....Pronotum.with Jongitudinal groove near lateral

edge; posterior margin with adjacent row of shallow, longitudinal

punctures. \Elytron with eight striae; intervals on. dise flat,

polished. Pygidium wider than long, with disc smooth and with

basal marginal sulcus or groove pronounced. Underside glabrous,

punctate. Fore tibia usually with three distal teeth on outer

margin, ©6St Of margin not. serrate... Middle coxae widely separated.

Howden and Young: Panamanian Scarabaeinae a

Sexual dimorphism often evident in clypeal surfaces, fore tibiae,

pronotal edges, sixth ventral abdominal segment, and pygidium.

The genus’ 'Uroxys isnot’ clearly defined” and "to badly if

néed of a révision! “As currently “constituted 1) secins “close to

Scatimus and Agamopus, but the pronotal groove, lack of a

distinct transversé carina on the middle tibia’ and, in’ many

species, the sexual dimorphism will distinguish Uroxys from

related: genera.

Key. to species of. Uroxys

Ds Anterior of frons with transverse ridge(s) or

groove (sometimes feeble); pronotum often

stroma ly. convex behind ‘anterior nee sides

near middie mearly .paraliel .) 2.4). slike te 2

Anterior ot. trons, evenly convex, wath carina

or groove, sometimes .feebly, iransyersel,

rugose,. becoming. monre.stronely .so on .clypeus;

pronotum: relatively .éevenly convex,. sides jareuate

or angled near, middle’ sci ie eee el a

L Frons with one distinct transverse carina;

pronotum with lateral, longitudinal, groove net

extending, to anterior and posterior manoins:

length greater than..6 mm... . eee O

Frons. with two.slight,. parallel transverse elena.

tions with depression between; pronotum with

lateral longitudinal, groove extendine [oO

anterior. and posterior margins; length less

than: ©. i0Misl ey on edule Gare Oe ee se Cone Hie ep.

ce Meso-metasternal suture broadly arcuate anter-

iorly; distance,» between, dorsal ocular areas

approximately..seven times. their width; posterior

margin of hind trochanter continuous with

posterior ey? of aie moe 9 ho) J 1 mm

1. COrgor w Arr.

Metoemetastern at Siti Ao ae aheulate anter-

iorly; distance, betweenidorsal ccular areas

approximately 8.5 times their width; hind

trochanter Ee inte lengthpb.20 f«9

mm . - ree MOLAGOR GON (Tsui)

4. Dorsal ocular area approximately as long as

wide; distance between De two, torthree limes

One vena nla an, ANE WE eS

Dorsal ocular area at eer ete as ane as

wide:.distance, between ‘evyes-at lLeast.four’ times

VO) LUE Nias | a in ba can Aleut ea) Mang ae A Cs a alee, Wigs ac @ px pi bine

ay Male pyeidium -flat; female with. sixth ventral

abdominal segment as long medially as segments

two. to five combined: leneth A to 5 mm

HZ Condrip. hmem. Bnd. Pasties iwol. tO sno. 11, | 1981,

Se eu Sen Chiu car vegan ive de Loe wplaly Dy Ga.) a Sp.

Male, pygidium strongly convex;;: female with sixth

ventral abdominal segment no longer medially

than, segments, four;and five combined; léeneth

SRO Cadet uta s saniea a de Mine tiviemu li CamiH Bn ke ener ase on tl 6

6. Anterior margin of clypeus bidentate, slightly

indented immediately ‘lateral to teeth, which

are weakly reflexed; male with two strong

teeth on fore tibia, basal tooth much reduced

ae Oo. bidentis nh. Sp.

Anterior margin of ely pens WLih Three strong

indentations, ‘teeth and’ maroin. abruptly reflex-—

ed; male’ with three; strong teeth on fore tibia

pe Macrocularis’ nN. Sp.

Ts Basal sulcus of pygidium ea sinuate;

Ten Oth bo Aes Stam tae mi tats 8

Basal sulcus of pygidium feebly sinuate or

shharoht, on each Side ior mide: tenoth

ee Oe a ee eto eo Berea tag: | 9

8. Dorsal ocular areas twice as long as wide;

distance between ocular areas five times

their width ooo i aa I MICros: . Bates

Dorsal ocular areas three ‘times | as ; Vane as

Wide; distance ‘belween ocular areas “e1eht

Limes Tete Wry ae vO meer ogularis We SD

oe Males with fore tibial spur short, spatulate and

obliquely truncate; clypeal margin straight

adjacent to clypeogenal suture. 6.9. gatunensis n. sp.

Males wath’ fore tDial\ spur slender and

detlexed "distally; ‘clypealmarein indented

At UROon with Cy MeO ena RV SULUTE soe. eases)

ATE Uae te BO ay Oeste Sp,

1. Uroxys gorgon Arrow

Pigs OZ O38. O22" 103

Uroxys gorgon Arrow,’ 19333° p. 397%

ne Type, London. Examined.

Males oitator form so tenoch Oto imme s Oblono, shining

black .s Clypeusi fies. 62, 102) “bidentate, “teeth elongate and

strongly retlexed.( "Head dorsally (very finely punctate.’ Genal

Margins obtusely ianoulate anduretlexeds clypedosendal suture distinct

and depressed...“ Transverse carina “on froms moderately elevated,

weakly arcuate anteriorly. Dorsal ocular areas approximately

1.5 times as long as width at posterior edge of canthus (+17 facets

wide at that point), distance between ocular areas approximately

seven, times their widths Pronorim para kel sided for! most ‘of

Howden and Young: Panamanian Scarabaeinae 53

leneth, as wide as elytra; transverse’ tumesity) present) al anterior

third; a small depression on each side of midline adjacent to

anterior margin; lateral posterior marginal area nearly vertical,

lateral anterior area almost horizontal; surface faintly punctate;

median longitudinal sulcus feebly present in posterior fourth;

lateral fovea in form of deep longitudinal sulcus extending over

half length of; pronotum:but not-reachine anterior Or (posterior

margin, sulcus faintly crenulate; pronotum distinctly margined

basally with row of large longitudinal punctures immediately

anterior to’ margin. . Elytron’ ‘strongly convex, fainvly punctate,

with slight humeral umbone; striae distinct but shallow, with

distinct’ punctures: evenly spaced for most of length’ ofeach stria;

posterior fifth of striae one and two deeply furrowed; intervals

flat, faintly punctate; “apices flattened : but not produced:

Mesosternum faintly punctate; meso-metasternal suture medially

broadly anteriorly arcuate, angulatée laterally, valmost touching

anterior margin of mesosternum, as close to margin as to mesocoxal

cavity; metasternum swollen but not keeled and with median

posterior depression. Ventral abdominal segments five and six of

equal median length andeach longer, than segments) 2.03, or 4;

sixth segment narrowed medially. Pygidium (Fig. 103) moderately

convex, \faintly’ punctate, “twice aavwidewwe lone; deep sulcus

surrounding disc, margin: formed, beina twice as wide apically as

laterally; sulcus basally straight or) slightly arcuate toward’ apex

on each sidé' of midline? “Fore tibia” with thiec! ouner aiolal veer

equidistant from each other, proximal tooth somewhat reduced;

bases’ of all ‘teeth occupying less) (ham hall jor venoth or tourer

margin; Inner ‘marogin curved; distal endw with tarce' deourwed

rounded” projection” at inner corner, projection” as long as tibial

spur; tibial spur stout, ibDlunmt, Doth (decurved ana vecurved. Middle

and hind femora strongly widened medially; posterior mareoin

stronoly élevated, ‘with a flattened! proyection ‘medimiiyv), oestertor

margin of hind trochanter even with posterior margin OL femur.

_ Genitalia asin fia. 08.) Minor: forms dittering irom major Torn

in the following’ characters: ‘length’ 9 to 10 mm; clypeal dentition

reduced, margins of geendad only weakly petlexed, carina ‘on: irons

reduced ‘to:a slight elevation; Tore tibia less eroncared ‘and

Curved .

FEMALE. Dilferine’ from: mate im the Tollowmechavacters:

transverse carina’ on) Troms’ strongly elevated) tore tiple shorter

and broader; sixth ‘abdominal ‘segment; ner narrowed: ‘medially;

pycidium wider’ and shorter.

MATERTAL: EXAMINE Dun PANAMA SA) @pecimens trom’ BOCAS) DEE

TORO PROV. =—‘Almirante. \ CANAL ZONE =' Barro Colorado Island:

Rie Aeoue ‘Saludy COLON «PROV. (Ape hia ie vie vel ae. Cri My

PROV. - Cerro Campana. « COLLECTION DATES Mivery month or tne

Vea,

OTHER COUN TRIG Le; snecimens trom: COSTA RICA. COLOMBIA,

ECUADOR.

KEMAKKS. Uroxys corgon, on’ the basis or 'sizeialone, cannot

be confused with any other species of Uroxys known to occur in

Panama.

54 Comtvit.. Amer, dnt. .lnsts. vol. 16... mo... 14,,1961

Tis. mocturne) species has, been, vecorded, ai. iioht,. perching

On leat suriaces,.jn.iur of,tive three-toed, siolas,: on. pellets. of

Dynastor darius (a large Lepidoptera) in. rearing cage, and at

sloth and, toad feces... It, apparently shares with U.. metagorgon

the unique. abit of. agtaching itself, to the, fur of,living .sloths

which live, in, lowland .moist, forests. Closely..related species. in

South America are also associated with sloths.

On Barro, Colorado Island,. laboratory,.experiments .suggest

that @Orgom.can survive on, the, feces ofa wide variety. of

vertebrates, but .in the field specimens are much: more. restricted

in their range of fecal utilization. Sloth feces occurring on the

cround. 1s. normally quite hard, and dry, and. in. the laboratory

gorgon will stanve to.death, in. the presence, of.this. material.

Freshly produced sloth feces (5 min.) is normally soft enough

to allow consumption by dung beetles, and perhaps only consumers

Fiding.on the .sloth,can efiectively utilize. this. potential. food

source.

Cr ULOxy>s Meta gordon ns sp.

Bigs, 100, .101,, 106

HOLOTYPE. Male, jength./.5 mm; elytra only, 4.0 mm;. width

4.4 mm. + Oblong, shiny,:-reddish black (teneral)... Clypeus (Fic.

100) bidentate, teeth strongly reflexed, triangular in shape.

Margin, of gena obtusely angulate. Surface of head finely punctate.

Clypeogenal. suture distinct .but superficial. Transverse carina .on

frons ,moderately. elevated, weakly arcuate, anteriorly. . Dorsal

ocular areas approximately twice as long as width at posterior

edge of canthus (+12 facets wide at that point), distance between

ocular areas approximately 8.5 times their width; frons posteriorly

with faint. wrinkles... Pronotum parallel sided. for.most.of length,

wider, than elytra; disc with transverse tumosity at anterior third,

small depression, adjacent to.anterior margin on,each side of

midline; lateral. posterior area. meariy.vertical, lateral anterior

area almost horizontal; surface finely pumctate; median longitudinal

sulcus feebly expressed in posterior fourth: lateral fovea in form

of deep longitudinal sulcus extending over half length of pronotum

but nol reaching anterior, or ,posterion maroin; sulcus distinctly

crenulate with distinct punctures anteriorly; pronotum distinctly

margined basally with row of large longitudinal punctures immedi-

ately anterior to.mangin.. Elytron strongly convex, finely punctate,

with slight umbone; each stria distinct but shallow, with distinct

punctures, evenly spaced for most of. leneth; posterior .fifth.of. first

and second striae furrowed; intervals flat; apices flattened on

intervals one and two, but not produced. Mesosternum finely

punctate; meso-metasternal. suture sharply anteriorly arcuate,

angulate laterally, almost touching anterior margin of mesosternum,

ae close 10 margin .aS.t0 mesocoxal cavily;.distance between anterior

margin of mesocoxal cavity and anterior margin of mesosternum

equal to (in other species 0.5x, except depressifrons and dybasi)

length of pleurosternal suture posterior to forecoxa; metasternum

Howden and Young: Panamanian Scarabaeinae 55

slightly swollen, with median posterior ‘depression. Ventral

abdominal seements 2,°°3,. 4, and 5 °er equal meciam tengr, “each

shorter than’ Sixth; sixth segment narrowed medially. ~ Tyoidium

(Fig. 101) moderately convex, faintly punctate, twice as wide as

lone; Sulcus surroundine dise déép, forming margin twice as’ wide

apically as basally; sulcus basally straight either side of mid-

line, Pore Tibia with three outer distal’ teen camila ant iron

each otner, proximal: toot somewhat reduced: ail wee teem

occupying approximately ‘halt Of Ouler margin “OF tila: iiner

margin evenly curved; distal end with large decurved scalpriform

PLOjection at inner Corner,’ projection os lene’ ao tioidttsour |

tibial’ spur ‘stout with rounded apex, outline sinuete.. Micdle’ ana

hing: femora’ stronoly widened» medially; posterior maroin elevated ;

hind trochanter projecting laterally. Genitalia as in fig. 106.

ALLOTYPE. Female, length 7.0 mm elytra oniy,’ 3.6: mur

width 3.8 mm. Differing from male in the following characters:

anterior clypeal teeth * shorter, proader,* and lees: rel tewea, toate

verse Carina on irons only slionily’ €levared: promotum without

antertor depressions at margin, NOL as wide as Clytra; tore mi p12

snorter ‘and broader, "sixth abdominal “seement Nor Narrowed

medially .

IVP MATERIAL: “Hototype,’ “male. 1 anava. * anal Zone. (Barre

Coloraao Island, 12-1X.1972.7 0." Younes, tien irae 1) conimeton).

milorype, femalé, Same data ‘as Nolotype, excepre 27. Vil lo/!

(Washington).

PARATYPES: 8° males, 4 females...” PANAMA®’ 2 males, 4 females,

same data as holotype’ except 16.X1:1971, 1201%019/2, 731 -%. 1972;

S1.V11.1973, (2. X. 1973, 1.19742" 3 males. Catal Zone Barro. Golorade

island, (1) 16,,1959,°H.S. D¥bas, “at om, ly Sovioy .P.

Young, on live three-toed sloth, (1) 8.V.1935, A. EY “Emerson, in

fur, oF sloth; 1 male, Colon“Prev.. Santa’ Rita idee, 200 1) *1o7N.

1976, A. Thurman, whitelicht. ‘COSTA RICA: 2 males, Turriatba,

13-17. 71121965, S.5) and W.t. Duckworth:

Paratypes are in? Chieago, Washington, Howden, Ratelifie:

REMARKS. “Uroxys meétagorgon 1s closely related to gorgon,

but can be distinguished from that species by the combination of

smaller size,’ dorsal ocular’ areas relatively more elongate and

farther apart, absence of reflexed genal margins, meso-metasternal

suture’ sharply angulate medially, shape of distal projection of

fore tibia, and projecting hind trochanter. Specimens examined

from Brazil and Ecuador, although similar, show consistent differ-

ences’ ‘that ‘seem TO “inditate specitic or’ supspeciiic differences.

They are therefore excluded from the type series.

itis species its ‘mecturnal,’ occurs orimarily in’ the: wet

season, and has. been recorded from ‘liont and irom the tur” of

live three-toed'sloths Vsee remarks under Ul soreon). - Distribu-

tion is apparently restricted in Panama to lowland moist forests.

3. Uroxys depressifrons n. sp.

Figs. 90. 99." 100 gs

HOLOTYPE. Mate, "length 3,0 "mim elytra only, 2.6 wm : width

2.2 mm. Oblong, faintly shining, reddish black. Anterior clypeal

56 Contribs camer. fonts dastion iwol, 8. ae. BO8T

margin quadridentate, medially strongly emarginate (Figs. 98,

104); margin reflexed, teeth more abruptly so. Genal margin

broadly rounded; clypeogenal, suture. indistinct... Surface of .head

distinctly punctate, Bromtal, area, (Fic. 104) with, two, slight trans-—

verse elevations and depression between. Dorsal ocular areas

approximately 1.5 times as long as width at posterior edge of

canthus (9 to di) facets wide at that. point), distance between

ocular areas approximately four times their width. Pronotum

moderately convex, without anterior tumosity; two-thirds of lateral

margin nearly vertical; surface minutely punctate; median longi-

tudinal sulcus feebly indicated in posterior fourth;>-lateral fovea

in form of deep and straight longitudinal sulcus, faintly crenulate,

extending to both anterior and posterior pronotal margins; pronotum

distinctly margined posteriorly with row of longitudinal punctures

immediately anterior to margin. Elytron moderately convex, faintly

punctate, with slight humeral umbone; striae depressed, with faint

punctures evenly spaced, posterior one-fifth of first and second

striae deeply furrowed, fourth extending farther posteriorly than

third, eighth terminating before joining second; intervals flat,

smooth; apices flattened but not produced. Mesosternum faintly

punctate; meso-metasternal suture medially sharply arcuate anter-

iorly, broadly rounded laterally, suture farther from mesocoxal

margins than from anterior margin of mesosternum; median longi-

tudinal depression of metasternum distinct on posterior two-thirds.

Ventral abdominal segments two to five of approximately equal

median length, segment six narrowed medially and twice as long

as segment five. Pygidium (Fig. 99) moderately convex, minutely

punctate, 1.5 times,as wide as. long; sulcus surrounding disc

deeper basally, forming margin of equal width laterally and at

apex; sulcus basally straight either side of.midline. ..Fore tibia

stout, with three outer distal teeth equidistant from each other,

proximal tooth somewhat reduced;:all three teeth occupying at

least two-thirds of outer margin of tibia; inner margin slightly

curved; distal end with small deflexed pointed projection at inner

commer, projection half jas) long .as,. tibial spur; tibial spur elongate,

tapering to point. Middle and hind femora moderately widened

medially, posterior margin weakly elevated; posterior margin of

hind trochanter even with posterior margin of femur. Genitalia as

in toey, LOGe

ALLOTVE i. Pemate, Jeneth 4.0; mms elytraconly, 2.3: mm:

width 2.1 mm. Differing from male in the following characters:

fore tibia relatively shorter and broader; sixth ventral abdominal

segment not narrowed medially.

TYRE MATERIAL Holotype, male.Panama, Chiriqui Prov.,

15 KmonW Hava del) Volcan, (Hartmann 's Pincay: 1500)m, 20431.V.1977,

S. Peck, dung trap (Howden). Allotype, female, same data as

holotype except 20-25.V.1977 (Howden).

PARATY PES: (40° males.(:57) females. BANAMA: (10 males; 18

females) Choariauis Prove 2 (ka WiCenre Punta, (Baldwin ‘Porest, 30.

VeROwWh. LOY 7) BAV 1. O77 S.0 Beek Hi Howden)! human dune traps:

a mMatesyoilitemale, Chirigad Prove 2 kmh Cerro Punta, 2200: m,

POV ee WOT iG . ' Peck) Human dung traps; 2 males, 2. females,

Chivigul Prove. 2k Nu Gtate Clara 100 om BS ai NN CQO%E"W.

Howden and Young: Panamanian Scarabaeinae mp

Hartmann's Finca, 30-31.V.1977, H. & A. Howden; 14 males, 15

females) "Chiriquiy Prov. ,° 15) km INW: Hate: del. Votcan, Hartmann's

Fincay) 1200 «im39/1500 bm) 1550 my) 20225) 01977, 20-0. Vwi 7 259 30,

31.V.19779) Sih Pecky human, dungiutraps,; tatrioniiwe pax oh tema le,

Chiriqui Prov!,’ Das Lagunas, 4) km. W) Hato del: Volcan, 1200" m,

27 VADTIS § Heom AD Howden, chumaniidunge traps, 41 males. 12

females, Chiriqui, Prov., Las. Lagunas, 5 km SW Hato del Volcan,

£300" myi622426 \ QU 197 A Ber eck plana Han es ea eae. ee

Peck, Berlese of carrion; 1 female; Cocle Prov.,,) Bl Valle, (trailte

Las Minas} 2400-2600! ft, 2b. 1 1959, Oo eo Dyiae, Benlese Or. Norse

manure. along trail, COSTA RICA sc males, 7 females, Punt.,

Monteverde, . 1400 °m,' 15007 m,, 23,274 Vi 19794 INV U.19079.: 24400. Pie 900,

H. & A. Howden, R.S. Anderson, in human dung trap, sitting on

leaves near trail at night.

Paratypes are in: Cambridge, Chicago, Ottawa, Washington,

Howden, Ratcliffe, Riley.

REMARKS. In body shape and clypeal elevations depressifrons

is closest to’ U.. gorgon ‘and. U.°> metagorgon, But it) cam be ‘easily

separated on the basis of size. Panamanian species of Uroxys of

similar size do not have clypeal elevations or the elongate

parallel-sided body shape.

This nocturnal species occurs in the higher elevations of

Panama and Costa Rica in areas where the ''dry" season is quite

moist.) It has*been captured perching (on) leaf. surfaces, at carrion,

and on horse and human feces.

4. Uroxys micros Bates

Figs! Foi Ole ae aoe ae

Uroxys micros Bates, -1837, p. 43,

Cotypes, London and Paris. London specimens examined.

MALE. Length 4:to (4.5, mm elyviranoniw 2.70 miy with a9

mm. Oval,. shining blacks:(* Clypeis/. (figs, 00.) 108) bidentate, not

indented immediately laterad of teeth, which are broadly triangu-

lar and strongly reflexed. Genal margin broadly rounded.

Clypeogenal suture distinct: clypeal margin indented where suture

intersects. Frons weakly convex, without elevations. Dorsal

ocular areas approximately twice as long as width at posterior

edge of canthus (seven to nine facets wide at that point), distance

between ocular areas approximately five times their width. Head

surface with fine “punctures. posteriorly.) coarse punctures” anter-—

iorby... Pronotume( Fig.) 106), casuwiderias* elytra; lateral edges of

pronotum extended into prominent angles, strongly sinuate in

lateral view, posterior third of margin nearly vertical; pronotum

weakly convex medially, surface finely punctate; median longitudinal

sulcus present in posterior fourth; lateral fovea in form of shallow,

non-crenulated longitudinal groove two-thirds as long as length of

pronotum and extending to posterior margin; pronotum indistinctly

margined basally with adjacent row of large longitudinal punc-

tures. Elytron moderately convex, faintly punctate, with very

slight humeral umbone; striae distinct but shallow, with distinct

punctures evenly spaced for most of length of each stria and with

58 Comtrito.: Ammer. dintus lnste) vols old, mo. bh) 190):

posterior fifth of first and second striae slightly furrowed,

seventh stria extending three-fifths length of elytron; intervals

flat; apices. somewhat flattened bul not. produced.) Mesosternum

faintly punctate; meso-metasternal suture medially moderately

angulate anteriorly, weakly. angulete laterally, three times. farther

from anterior margin of mesosternum than from mesocoxal cavity;

metasternum swollen, with faint or obsolete median posterior

depression; hind coxa. with portion .of posterior margin. in. form of

triangular projection directed posteriorly (usually under femur).

Ventral abdominal segments two to five of equal length medially

and each half as lome medially as sixth.’segment; sixth not

narrowed medially. Pygidium (Fig. 91) strongly convex, faintly

punctate, 1.5 times as wide as long; sulcus surrounding disc deep

basally, shallow elsewhere, margin formed being slightly wider

apically than laterally;. sulcus basally strongly trisinuate,

separating from disc two lateral rounded expansions, the median

basal area of disc narrowed. | Fore tibia elongate with inner

Mare@in Sinuate;, three cuter distal teeth with. proximal tooth. some-

what reduced and 1.5 times farther from middle tooth than middle

tooth is from distal tooth; bases of all teeth occupying approxi-

mately apical third of outer margin; apical end of tibia: with

small decurved, pointed projection at: tmner corner, less-than: one—

fourth -leneth cof tibial spur, . tibial spur elongate, thick, -blunt,

strongly deflexed on distal: third, extending to middle-of distal

segment of tarsus. Fore femur strongly constricted distally,

middle and hind femora slightly widened medially; posterior margin

of hind trochanter continuous with posterior margin.of.femur.

Genitalia as in fig. 110.

FEMALE. Differing from. male in the follewing characters:

clypeus transversely rugose, lateral pronotal fovea not extending

t© posterior margin, fore. tibia velatively. shorter and. breader,

middle coxa without posterior projection, sixth ventral abdominal

segment longest medially, and pygidium 2.5 times as wide as long.

MATERIAL EXAMINED. PANAMA, 88 specimens from: CANAL

ZONE = Barro Golorado Island; Fort Kobbe; Frijoles; Pipeline Road;

Skunk Hollows; .COLON «PROV: = Pertobello.... COLLECTION: DATESs

Every month of the year except August.

OTHER COUNTRIES, 10 speciunens from:.COSTA RICA, GUATEMALA.

REMARKS. There are six species of Uroxys in Panama that

are. 5.0 mm @ectess in body Jlenoti; oval in shape, lack elevations

on ihe -frons,, and that form a separate group from the.preceding

three species. Because micros is the only previously described

species in this group, the assemblage is referred to as the micros-

complex. Males of micros are distinguished from other species of

the -complex by the presence of a projection on: the posterior margin

of the metacoxa. Both males and females of micros have the most

strongly sinuate pygidial sulcus basally of any member of the

complex, the sinuation producing a narrowed median basal portion

of \the pycidial disc.

This nocturmal»species has been captured at light, from

forest floor leaf. Jitler;. perching on deal. surfaces, at..peccary

carrion; and on howler. monkey, human; .tapir,:.and»toad feces.

Howden and Young: Panamanian Scarabaeinae 50

5, sUnexys microcularis ns sp.

Figs, 92, 9-12, 13

HOLOTYPE. “Male; length 3.7 mm; elytra only, 2.2 mm. width

2,5 mm.’ Oval, Shining plack > Clypeus (Fie. 97) wideniace, not

indented immediately laterad to narrowly triangular, strongly

reflexed teeth. Head surface with fine punctures posteriorly,

coarse punctures anteriorly, and with slight rugosity along

lateral and anterior edges. Clypeogenal suture present but

indistinct; genal margin weakly angulate immediately posterior

to intersection of suture. Frons weakly convex, without elevations.

Dorsal ocular areas approximately three times as long as width at

posterior edge of canthus (three to five facets wide at that point),

distance between ocular areas approximately eight times their

width. Pronotum “slightly narrower than.:.¢lytra; (ateral edges. of

pronotum produced into prominent angles, strongly sinuate in

lateral view, posterior third of margin nearly vertical; pronotum

weakly convex medially, surface finely punctate; median longitu—

dinal sulcus present in posterior fourth; lateral fovea in form of

shallow, non-crenulated, longitudinal groove two-thirds as long as

pronotum and extending to posterior margin; pronotum indistinctly

margined basally with adjacent row of large, longitudinal punc-—

tures. Elytron moderately convex, faintly punctate, with very

slight humeral umbone; striae distinct but shallow, with distinct

punctures evenly spaced for most of length of each stria; posterior

fifth of first and second Striae sliehily, furrowed, sevenin siria

extending three-fifths length of elytron; intervals flat; apex some-—

what flattened but not produced. Mesosternum feebly punctate;

meso-metasternal suture medially moderately angulate anteriorly,

weakly angulate laterally, three times farther from anterior margin

of mesosternum than from mesocoxal cavity; metasternum swollen,

with very slight median posterior depression; hind coxa without

posterior projection. Ventral abdominal segments two to five of

equal length medially and each segment half as long medially as

sixth; sixth not narrowed medially. Pygidium (Fig. 93) strongly

convex, faintly punttate, 1.5 times as- wide as lone: sulcus

surrounding disc deep basally, shallow elsewhere, forming margin

slightly wider apically than laterally; sulcus basally trisinuate,

delimiting two lateral rounded expansions, the resultant three

curves of basal sulcus of equal magnitude. Fore tibia elongate

with inner Margin sSinuate;: three outer distal teeth equidistant

from each other, proximal tooth somewhat reduced; all three teeth

occupying approximately one-third of outer margin; distal end

with small deflexed pointed projection at inner corner, projection

less than one-fourth length of tibial spur. Tibial spur elongate,

thick, blunt, stronely ‘decurved on, distal third, extending to

middle of distal segment of tarsus. Fore femur strongly constricted

distally, middle and hind femora slightly widened medially;

posterior margin of hind trochanter’ continuous with posterior

marein of femur. Genivalia “as in ties. 112, "113.

ALLOTYPE. Female, length 3.6 mm; elytra only, 2.2 mm;

width Z2.4°mm.° Differing from the holotype im the following

60 Contribs: Amery (Ent Ens > Volvcl6y nol 61.1981

characters: clypeus transversely rugose, clypeal margin indented

immediately dateral, to teeth,.fore..trbia ‘relatively shorter and

broader, ifore tibial Gpurcvtihinner, oixth ventral abdominal segment

longest medially, and pygidium twice as wide as long.

TYPE MATER Tab! Helotype, male,.Panama; Colon Prov., Santa

Rita Ridge, 300 m, 10-11.V1.1977, H. & A. Howden (Howden).

Allotype,, female, Panama,:Canal, Zone,» 5 mi NW: Gamboa, 7.1V.1970,

M.A.R.U., light trap (Washington).

PARATYPES:: 70 males; .607: femalés. PANAMA: 4 males, 4

females). Camal ‘Zone. «Barro. Colorado Island, 20-23:V.1964, 31.V.

1970) Woe & So. Duckworth, HA. Hespenheide: | male, ‘Canal

Loney jeoce, Golo Yospiiak, oh) Ali IOy. Gtockwell, manure trap:. 1

female. Canal, Zone, Cerro, Campana, loots .1954,).0.L. Cartwright:

1 female, Canal Zone, Cristobal) 5.%.1965, Navy. collection, light

traps iomale, Canal Zone, Forty Olayton, \22..1%,1952: 3 males,

Canal Zone, (Fis Culek. ij. .4 993, .2) males, 4 temales, Canal Zone,

Hort, Robbe, MeViljlO7o..24.Vi 1976, BG. Ridey, blacklioht trap;

bk female, Canal Zone, 4 mi. W Gatun, 6.V.19/3, H:P.. Stockwell;

22 males, 29 females, Ganal Zone; Skunk Hollow, 7.V11).19/75, 13.

Ville 1O-21 Volvo me BO. Ratelifres 4d. (female, Canal: Zone,

Pacific area, Vill.19605, Navy collection, light trap; 19 males, 9

females, Cocle. Prov, hi Valles trarhto Las: Minas, 2400-2600 ft. ,

20.11.1059, 4.5. Dybas.in dunes. 3.males, 3 females,. Cocle. Prov.,

Ely Valle, 2200 fi. ol 15D. vow Dybac, Berlege, horse manure

On trail, 12 males, ./ hemates, Cocle:Prov., bb Valle;alone

iriputary Of hic Anton, 2200 ft... Dybas, in horse: manure; |

female, Panama Prov., Cermeno, 11.1X.1952; 1 male, 1 female,

Fanama, Prov, Ghacvres Bridge nr. Madden Dam, &.X.1952;. 1..male,

Panama Prov. Molinga Swamp (= Mojvinga Swamp’), 15.VIL.1952;. 3

females, Panama Prov.,:.Potera, 11.1X.1952; 1 male, 1 female,

PanamayProw., Rio Sajalices. near Villa. Real, 12.1V.1952; 1 female,

Panama. Prova) (Locumeny (25. %.1952.

Paratypes are in: Chicago, Ottawa, Washington, Howden,

Ratcliie,. Riley.

REMARKS. Males and femates of microcularis can. be. separated

from other members of the. micros complex in,.Panama by the

combination, Of the small dorsal ocular areas, three times as long

as wide, Separated by the, large relative distance of!eight times

their width.

Phis mocturnal epecies is usually collected in the: wet

Season, and) mas, been taken vat lioht, and on ‘horse, human, and

tapin.wteces.. tits seemingly distributed throughout the. moist

forests of Panama.

6. Uroxys platypyga n. sp.

Pigs, 84,85, 107

HOLOIYP ER. Male, length 4.7 mm: elyira: only, 2.6.mm;. width

3.0 mm. “Ovals shinane black, |.Clypeus’ (Big. 84). bidentate,

faintly: indented immediately. laterad of broadly. triangular and

weakly reflexed teeth. Head, surface. with fine punctures. poster—

lorly, cOarse punctures anteriorly. Clypecgenal suture distinct;

Howden and Young: Panamanian Scarabaeinae 61

genal margin rounded and weakly angulate immediately posterior

to intersection ‘of suture.’ Frons feebly convex, without elevations.

Dorsal ocular areas approximately as) long as width..at) postertror

edge of canthus (19 to 21 facets wide at that point), distance

between ooular,..areas: approximately twiceutheir width. «Pronotum

slightly narrower. than elytra; lateral edges’ of pronotum produced

into prominent angles, strongly sinuate in lateral view, posterior

third of margin: nearly Vertical; pronotum weakly convex: medially,

surface finely punctate; median longitudinal sulcus evident in

posterior: fourth;; lateral. Tovea, in) form: of shallow, non=-crenulate

longitudinal groove two-thirds length of pronotum; groove extending

nearly: to (posterior: smargin, .anterlor,, portion: of ;eroove terminating

near several large punctures; pronotum distinctly margined basally

and: with: adjacent row of large :longitudimal pumetures,) » Eyytron

moderately convex, faintly «punctate,with slight umbone;. sire

distinct but shallow, with distinct punctures evenly spaced for

most ofeach stria, ‘seventh striay extending threc+iitths, of Jengin

of: elytron; intervals flat, apices) flattened, not: produced. . Meso-—

sternum faintly punctate: meso-metasternal suture medially narrowly

arcuate. anteriorly; srounded’ laterally; | suture inrea iimes: tanther

from anterior margin of mesosternum than from mesocoxal cavity;

metasternum swollen, with median posterior Groove’ absent; sind

coxa without posterior projection. Ventral abdominal segments two

tovfive of-equal length medially, sixth séementiias: lone as 32°,

and 5 combined and longer medially than laterally. — Pygidium

(Fig. 85) flat medially, faintly punctate, twice as wide as long;

sulcus: surrounding disc, deep: basally, ‘shallow elsewhere; sulcus

basally trisinuate, separating from disc two lateral expansions;

resultant median basal area of disc narrowed. «ihore. tipiaivelongate

with, inner margin sinnate: outer imargine with thnree, teeth, basal

tooth slightly reduced and\’?.5 times ‘larther, from middietooth

than middle tooth is from apical: .one; all three teeth occupying

approximately one-third of outer margin; distal end with small

projection at. inner'.corner, projection less: than: one-fourth lemeth

of tibial spur; tiblalspureloneate;: thick) blunt, :strongly

deflexed in distal third, extending as far as base of distal seg-

ment of tarsus. Fore femur istrongly constricted ‘distally, middie

and hind femora slightly widened medially; posterior margin of

hind trochanter continuous with posterior margin of femur.

Genitalia as) iim fig. WO),

ALLOTYPE cremate yvlengih dado mmyielyirawoniin «Oi Ginn)

width) '2./:mm:. Differing from male (inthe following ‘characters:

clypeus transversely rugose, fore tibia relatively short and broad,

fore tibial spur thinner; sixth ventral abdominal segment as long

medially as segments two to five combined; pygidium three times

as wide as long and strongly convex.

TYPE MATERIAL: Holotype, male, Panama, Canal Zone, Barro

Colorado Island, 2) NO7Sy OLR Young) itapineifecesy trap

(Washington). Allotype, female, Panama, Canal Zone, Barro

Colorado Island, 13. Vil074, OP. Yourtey human feces trap

(Washington).

PARATY PES /:14 mabes, 24) femalesi’) (PANAMA2/) 110 males, (17

females, Canal: Zone, banne (Gelopadodlelandy ¢ SueVEt.IOGSs Mb dal

62 Contrib. Amer. Ent.c Imets, wol. 16. no. -1,01961

IX, 1941, VEI-VIILL.1942, Viex. 1943," [VEX 1947,) 17-19. 11959, 1.1970,

O.1VGiI9/Gy 1O7IW 1970, «GV Lb o72) 19 21. 1972) Our o7e, esd:

1975, 462171 141.1975, 15.10, 17, 1421975, kt. eile leaf vdebris,

J. Zetek, Berlese: Tunnel, O.P. Young, tapir, Human feces, rat

carrion, H.o, Dybas, By. Poster, Hiay Hespenheide, £.C. Williams;

1 male, Canal Zone, Fort. Kobbe, 24.V1.1976, £.G.. Riley; 3:females,

Canal Zone, Madden Porest. mile 2.5. 20.471.1971, 6.1.:1972; H.-P.

Stockwell, manure trap; 1 male, 3 females, Canal Zone, 5 mi. NW

Gamboa, 80°011 01970, 4. IVeIO70 MOA. Rau disht trap; 1 male; 1

female, Panama Prov., Aguacate, 300 m, 16.V1-1970,..B.. Chaniotis,

at light.

Paratypes are in: Chicago, Ottawa, Washington, Howden,

Ratcliffe, Riley.

REMARKS. Other. members of the micros complex in Panama

possess, in both sexes, a strongly convex pygidium. The. males

of platypyga, however, have the pygidium flat medially, and the

aedeaeus is Unique: within the complex with wery: Short, truncate

parameres. The female has the largest sixth abdominal segment

@l any member of the micros complex, the segment being as long

medially as Gegmenis’ two to five combined. Other species have

the sixth segment no longer medially than segments four and five

combined.

The “species is active at night im both. the wet and dry

Seasons and has been collected at light, under leaf debris, at rat

and unknown carrion, and on human and tapir feces. - ‘Ihe distri-

bution of platypyoa in Panama is restricted to moist lowland

foresis. |= (el ve

7. Uvoxys Macroculearis 1... Sp.

Fics. 06,. 07; 114

HOLOTYPE. Males lencthn 6:4 mm; elytra only; 2.0 mm; width

2.4 mm. Oval, shining red (teneral). Clypeus (Fig. 86) with

three indentations, median one most pronounced, indentations

delimiting ‘pair of retléxed, Droadly triangular teeth; margin

lateral to outer indéntation straight and reflexed. ‘Surface of

head with fine punctures posteriorly, coarse punctures anteriorly,

with slicohtrucosity alone lateral and anterior edges: Clypeogenal

suture distinct; genal margin angulate immediately posterior to

intersection of suture. Frons weakly convex, without elevations.

Dorsal ocular areas approximately as long as width at posterior

edge of canthus (14 to 16 facets wide at that point), distance

between ocular areas approximately twice their width. Pronotum

slightly narrower than elytra; lateral edge of pronotum produced

into prominent angle, strongly Simuate in. lateral view; posterior

third: of margin «nearly vertical: proneotum. weakly convex medially,

surface finely punctate; median longitudinal: sulcus absent;

lateral fovea in form of shallow, non-crenulated longitudinal

groove two-thirds length of pronotum and extending to posterior

margin; pronotum indistinctly margined basally, with adjacent row

ef large longitudinal’ punctures. Elytron moderately convex,

faintly punctate, with small humeral umbone; striae distinct but

Howden and Young: Panamanian Scarabaeinae 63

shallow, with distinct punctures evenly spaced for most of length

of each striaz: posterior fifth.of first and second. striae slightly,

furrowed, seventh extending three-fifths length of elytra; intervals

flat; apices flattened, slightly produced at ead. ol. fifi siria.

Mesosternum faintly punctate; meso-metasternal suture medially

moderately angulate anteriorly, weakly angulate laterally; three

times farther from anterior margin of mesosternum than it is from

mesocoxal cavity; metasternum swollen, with very slight median

posterior depression; hind. coxa without, posterior projection.

Ventral abdominal segments two to six of equal length medially;

sixth of equal length throughout. Pygidium (Fig. 87) strongly

convex, faintly punctate, twice as wide as long; sulcus surround-

ing disc deep basally, shallow elsewhere, margin formed wider

apically than laterally; sulcus basally trisinuate, separating from

disc two feeble lateral expansions; three curves of the basal

sulcus of equal magnitude.’ + Pore’ tibia elongate with immer: maroin

Simuate; outer margin with three equidistant teeth in apical thicd,

basal tooth somewhat reduced; distal end of tibia with small

deflexed pointed projection at inner apex, projection less than one-

eighth as lone: as tibial spur.. > Tibial spur elongate, thick,

blunt, strongly curved in distal third, extending as far as base

of apical tarsal segment. Fore femur narrowed distally, middle

and hind femora slightly widened medially; posterior margin of

hind trochanter continuous with posterior margin of femur.

Geniialia asin: fie. 14.

ALLOTYPE. Female, length 3.4 mnigi elytra enly, 2.07 mmy width

2.4 mm. Differing from male in the following characters: clypeus

tnwansversely rugose, fore tibia relatively short and “sroad, -fore

tibial spur thinner, sixth ventral abdominal segment longest

medially and twice as long as fifth segment.

TYPE MATERIAL: Holotype, male, Panama, Canal Zone, Gamboa,

26:1V.1975, O.P«. Young, tapir feces, trap (Washington): — Alletype,

female; Panama, Canal Zone; 5 mi: NW yGamboa, 13.4V.1970, MARU. ,

light trap (Washington).

PARATYPES: 6 males, 3 females. sPANAMA= 1 female; Canal

Zone; Barro Colorado -island,: X=X129P941,°Zetek: 6 males, 2° females;

Canal Zone, 5: min NW Gamboa; 723: 17) 1070; 420. 1970s 2bely. 1975)

M.AVR Us, O.P. Youngs, light trapyetapir feces: trap.

Paratypes are in: Washington, Howden.

REMARKS. Males of macrocularis can be distinguished from

males of other members of the micros complex in Panama by the

three indentations of the anterior clypeal margin. The median

indentation is most pronounced, but the lateral indentations are

also distinct. The entire clypeal margin is strongly reflexed.

These characteristics are found in females of several species but

not in other males. “Females of macrocularis appear to be most

closély related to: bidentis which’ i¢’ similar in. thei pycidium,.“sixth

ventral abdominal segment, and dorsal ocular areas. Females of

macrocularis may be distinguished by the three indentations of the

clypeal margin. The anterior clypeal margin of bidentis is only

medially indented and the teeth are only feebly reflexed.

Uroxys macrocularis is active at night in both the wet and

dry seasons. . lt has been collected only in the moist forest areas

64 Comprib.t Amer. -Baturinse,) aol do,. mo. 191.1981

of the Canal Zone where it has been taken at light and on tapir

feces.

8. Uroxys Didentis n. sp.

Figs: 66,609) 011

HOLOTYPE. Male: leneth) 3.7 mm; elyive tony, 72.3 mms. width

2.30. mm. oOvals shinine black, «Clypeus (Pies 66). bidentate, teeth

narrowly triangular. and: weakly reflexed. Surface of head: with

fine punctures posteriorly, coarse punctures anteriorly, without

rugosity.: Clypeogenal suture distinct; genal margin weakly

angulate immediately posterior to intersection of suture, otherwise

rounded. Frons weakly convex, without elevations. Dorsal ocular

areas approximately as long as width at posterior edge of canthus

(14 to 16 facets wide at that point), distance between ocular areas

approximately twice their width. Pronotum slightly narrower than

elytra; lateral edges of pronotum produced into prominent angles,

strongly sinuate im: lateral. view, posterior third of margin nearly

vertical; pronotum weakly convex medially, surface finely punctate;

median longitudinal sulcus evident in posterior fourth; lateral

fovea in form of shallow, non-crenulated longitudinal groove two-

thirds the: length of pronetum and extending to posterior margin;

pronotum indistinctly margined basally with adjacent row of large

longitudinal punctures. (Elytron moderately convex, finely punctate;

humeral umbone small; striae distinct but shallow, with indistinct

punctures evenly spaced for most of length, posterior tenth of

first and second striae slightly furrowed, seventh stria extending

three-fifths of length of elytron; intervals flat, apices slightly

flattened but not produced. Mesosternum faintly punctate; meso-

metasternal suture medially narrowly arcuate anteriorly, rounded

laterally, three times farther from anterior margin of mesosternum

than from mesocoxal cavity; metasternum swollen, with very slight

median posterior, groove; hind coxa without posterior. projection.

Ventral abdominal segments two to five each of equal median

length and, cacy 1.5 times sherter medially than sixth; sixth seg-

ment of uniform) width. Pygidium (Fig. 89) strongly convex,

faintly punctate, twice as wide as long; sulcus surrounding disc

deep basally, shallow elsewhere; margin formed slightly wider

apically tham laterally; sulcus basally trisinuate, separating

from disc two lateral expansions; the three curves of basal portion

of suleus of equal magnitude. Fore: tibia elongate: with inner

margin sinuate; outer margin with three teeth in apical third,

basal tooth reduced to small projection and 1.5 times farther from

middle tooth than middle tooth is from apical tooth; tibial apex

with very small decurved projection at inner corner,: projection

less than one-eighth length of tibial spur. Tibial spur elongate,

thiek, \blwat, strongly decurved: on, distal third; »extending «to base

of distal segment of tarsus. Fore femur strongly constricted

distally, middle and hind femora slightly widened medially;

posterior margin of hind trochanter continuous with posterior

margin Of femursewGenitallaywas; in tig. ill,

Howden and Young: Panamanian Scarabaeinae 65

ALLOIYPE.«.Memate, lengthy 3.39 mmpelytea only.) lad aim:

width 2vl:mms Differing from male in the; following, characters:

clypeus transversely rugose; fore tibia relatively short and broad,

with proximal, tooth, reduced: by half; fore: tibial spur thinner:

sixth ventral abdominal segment longest medially and twice as

long ias.difth: seoment,

TYRE MATERLALs Holotype. male: Panama, Chiriqui Prov.,.2

km)W Cerro Punta, Baldwin: Foresti. guNle1O7/, 4. Howden, cup

traps (Howden). Allotype, female, same data as holotype (Howden).

PARATYPES: 46 males, 22 females. PANAMA: 18 males, 8

females, Chiriqui: Prov:,, Las Lagunas; 4 km W.Hato del, Volcan,

1200 om, 1360),.m5.275 V1 977; bo 61977; SEO Ih] hy Oy bg i ) &

A. Howden, 4 human dung traps-3 day run, dung traps, evening

sweepings; 6 males, 6 females, Las Lagunas, 5 km SW Hato del

Voleam)/ 1360my).22:Vsd9774 19) 2m Jey Peek) (dung. raps, iittern: (4

males, ol: female, Chiriqui (Prow. ,e2vkmoN «Stal vClara,. fartmann >

Pineay, 1300)4m, 30-31.V 1977 5.) He eo A. Howdeny human, dune) traps:

l mate; 2) females, :Chiriqui::Prov.;- Hartmann 's Pinca) 20.V.1977.

25 Nl OFF, Se Si dauPeck, Ntter, ‘under bark, dung. tra pe; :9) mates.

4 females, Chiriqui: Prove,,2. km) W Cervo: Punta. Baldwin Forest;

3.V 1977... A. Howden, human, dung traps: male, Chiriqut Prov<,

Cerro: Punta, 4500.m,'23.V319/77, te dowden,) human dune traps:

2 ;‘mates, 1) female, -Chiriqui) Provu, 2¢km W, Certo Punta?) 1300. im:

242): 29.N.197/, WH. Howden, .human sdung traps; 4) males, Chiriqui

Proven 255) kite W (Cerro, Puntag 61720: m,.260 kel 9/5. a Stockwell,

manure, trap; l male, Chiriqui Prov. W.Finga Palo Sento, near

Nueva California, 4900 ft., 6.111.1959, Berlese —- floor litter in

Ya Lhe’.

Paratypes are in: Chicago, Ottawa, Washington, Howden.

REMARKS. Male members of the micros complex in Panama all

have the proximal tooth on the outer distal margin of the fore

tibia reduced to some degree. However, bidentis is the only

species in which the males have the basal tooth of the fore tibia

reduced by) more than. half, to the extent that. most of the paratypes

seemingly lack the basal tooth. See remarks under macrocularis

for differentiation of females.

Uroxys bidentis is nocturnal, occurs in both wet and dry

seasons, and has been found perching on leaf surfaces, under

bark, in forest floor litter, and on human feces. It is seemingly

restricted; tothe higher) elevations.(/91000 mm) in Chiriqul Province.

9. Uroxys gatunensis n. sp.

Fies. 94,195

HOLOTYPE. Male, length) 5.0 vam: elytraloniyy3i2) mmr) width

324 mm. “Oval, shining red (teneral). Clypeus (Fic. 94). bidentate,

teeth broadly triangular and strongly reflexed. Head surface

smooth on vertex; laterally and anteriorly with very fine punc-

tures. Clypeogenal suture distinct; clypeal margin angulate

immediately posterior to intersection of suture, margin of gena

rounded. Frons weakly convex, without elevations. Dorsal ocular

areas approximately twice as long as width at posterior edge of

66 Contribs Amer. “Pnty-nst.. volo 13s) noe 4, 61981

canthus (8 to 10 facets wide at that point), distance between

ocular areas’ approximately five times Théir width. Prenotum

slightly narrower than @lytra; lateral edges: ofypronotum produced

into prominent angles, ‘strongly sinuate in #ateéral view, posterior

half of margin vertical; pronotum weakly convex medially; “surface

with scattered, fine punctures; median longitudinal sulcus evident

in posterior third; lateral Tovea in form of ‘shallow, smooth longi-

tudinal groove one-third length of pronotum and not extending to

either’ anterior: or posterior margin; pronotum margined basally,

with adjacent row of large longitudinal punctures. ~<Elytron

moderately convex, faintly punctate, humeral umbone small; striae

distinct but shallow, with distinct punctures evenly spaced for

most ol length, seventh stria extending two-fitths of lencth. of

elytron, posterior fifth of first and second striae“slightly <furrowed;

intervals flat, apices slightly flattened but not produced. Meso-

sternum faintly punctate; meso-metasternal suture medially narrowly

arcuate antertorly, weakly “anculate laterally; suture three times

farther from anterior margin of mesosternum than from mesocoxal

cavity; metasternum swollen, with slight median posterior depres-—

sion; hind coxa without posterior projection. Ventral abdominal

segments two To tive of equal “lénoth medially and “each half as

long as sixth; sixth slightly longer medially than laterally.

Pygidium (Fig. 95) strongly convex, faintly punctate, twice as

wide as long; sulcus surrounding disc deep basally, shallow else-—

where; Margin’ formed of same width apically and laterally; sulcus

basally ‘slightly arcuate toward apex on each side of midline.

Pore tibia elonedte with innér margin sinudte: outer margin with

three teeth in apical two-fifths, teeth approximately equidistant

from each other, basal tooth somewhat reduced; apical end of fore

tibla with short, wide, blunt deflexed projection at inner corner

less than one-fourth *lenoth of tibial Spur.” Tibial spur ‘short,

Spatulate, “obliquely truncated, extending to fourth tarsal’ segment.

Fore femur narrowed distally, middle and hind femora slightly

widened médially; posterior marain of hind trochanter continuous

with posterior margin of femur. Genitalia collapsed (teneral).

FEMALE. Unknown.

TYPE MATERIALS Holotype, mate, Panama, Canal;Zone,* Barro

Colorado island. 10.17.1975, O.P. Young, tapir® feces’ ‘trap

(Washington).

REMARKS. The male of catunensis*is unique among members

of the micros complex in Panama due to the presence of a spatu-

Tate, “Opliquely truncated fore jibial spur;. in all other species

both: males and females possess elongate, tubular, distally

deflexed spurs.

This Species is waned for Gatun Lake, the body of water

surroundine Barro Colorado Island, Panama.

LO. Uroxys..dybasi. n..sp.

Figs. 96,07, 4150 16) 5117

HOLOTYPE. Mate, lenoth a.0%mms elytra only, 895° mm; width

3-5 mm.) Elongate oval, shining reddish: black. -Clypeus” (Figs.

Howden and Young: Panamanian Scarabaeinae 67

96, 115). bidentate, slightly: indented immediately laterad of teeth;

teeth broadly’ triangular and strongly .reflexedy. Head. suriace

with slight anterior transverse rugosity, distinct small punctures

throughout. . -<Glypeogenal suture;distinel; chypeal, margins distinctly

indented. at intersection of suture; eenal: margins Broadly wounded:

Frons weakly -convex,: (with very. slioht, broad: indentations.

Dorsal ocular .areas approximately. twice .as tome as-widih, at

posterior edge of canthus (12 to I4 facets wide at that point),

distance, between ocular areas approximately five times their width.

Pronotum (Fig. 115) at median angulation as wide as elytra;

lateral edges of. pronotum produced into prominent angles,

strongly sinuate in lateral view, posterior two-thirds of margin

nearly vertical; pronotum weakly convex medially, suriace idensely

covered’ with: fine, deep punctures; median longitucdingl sulcia

feebly .indicated: in. posterior, third; lateral fovea tn form oF

crenulated longitudinal deep groove three-fourths length of prono-

tim; not,extendine toveither anterior’ or “pos eller marcia, with

cluster of ‘Coarse’ punctures ‘In posterior third: pronotum marcined

basally, with adjacent row of large longitudinal punctures.

Elytron moderately convex, faintly punctate, humeral umbone small;

striae distinct but shallow, with distinct punctures evenly spaced

for most of length of each stria, seventh stria extending three-

fifths leneth of elytron:; ‘posterior tenth © irs) sirie iurrowed :

intervals flat, apices slightly flattened and constricted, not

produced. Mesosternum faintly punctate; meso-metasternal suture

medially moderately angulate anteriorly, moderately angulate

laterally, three times farther from anterior margin of mesosternum

than from mesocoxal cavity; distance between anterior margin of

mesocoxal cavity and anterior margin of mesosternum similar to

distance between pleurosternal suture posterior to fore coxa;

metasternum swollen, with distinct median posterior depression;

hind coxa without posterior projection. Ventral abdominal segments

two.to five of equal length medially, each only slightly shorter

medially than sixth; sixth slightly longer laterally than medially.

Pygidium (Fig. 97) strongly convex, faintly punctate, twice as

wide as long; sulcus surrounding disc deep basally, shallow

elsewhere; margin formed of same width apically and laterally;

sulcus basally very slightly arcuate toward apex on each side of

midline. Fore tibia elongate with inner margin broadly curved;

outer margin with three teeth in apical third, teeth: approximately

equidistant, basal tooth somewhat reduced and more broadly tri-

angular; apex of fore tibia with short, narrow, rounded, deflexed

projection at inner,conner, projection approximately half length of

tibial spur. Tibial spur elongate, ‘straight, pointed, extending to

fourth tarsal segment. Fore femur gradually tapering distally,

hind femur with slight median ventral posterior swelling; posterior

margin of hind trochanter continuous with posterior margin of femur.

Genitalia. ias in figs. 116,..41/.

FEMALE. dnknewn.

TYPE. MATERIAL: (Holotype, male, Panama, Camp’ Chaores,

Pornct iLO i Taylor. (Chieace) ;

PARAIYP ER: angle. COSTA: RICA: Punt... Monteverde, 1500 m,

214N 1979, He & A. Howden, im carrion. traps (Howden).

68 Contrib ‘Aner. Pati colaet.: vel vrs mo. 1) e196!

REMARKS. This species is most’ closely related in Panama to

gatunensis from which it can be separated by the combination of

indented lateral clypeal margin, shaft-like fore’ tibial spur,

angulate meso-metasternal suture, and very broad basal fore

tibial tooth.

This species is named for Henry Dybas in appreciation of

his: many contributions: to our studies.

Ateuchus Weber

Mteuchus, Weber,. 1901... 0. 10; Chapin, 1946, p..79.

Type-species: Ateuchus histeroides Weber, by monotypy.

Choeridium Serville, 1828, p. 356; Bates, 1887, p..44; Balthasar,

POted pa 44; Chamin. 1940, m9. 79.

Size small to moderate (4 to 11, mm,in Panama); oval (Fie.

118), strongly convex. Head with clypeus feebly to distinctly bi-

Genlate; [eons and. vertex tacking. Worns, distinct tubercles),.or

Carinae. (Fronotum teialively evenly conver, Jacking. posterior

marginal bead; lacking punctures or indented line before or

Denind Lateral Jovea... ocutellum absent. —Elytron with eight

Giriaen, ww OCS, COX tranaveroce,. mo. conical... Middle, coxae widely

separated...imner marging marailel. Pore tibia with three, or: four

teeth On, outer. marein, the margin, basally lacking distinct serra-—

tions...) Wares Of. most Suecies. with, fore tibial spur. expanded.

Ning tibia, lacking raneverse Carinae on Duter.surface;:hind tibia

strongly expanded in apical half.

OF the. Various Cenuera. treated. in this, work... fie mames used

fOr (he Panama species Ol. Ateuchus are “the least certain., In, his

key to the genus, Balthasar (1939d) discussed a number of problems

encountered, im his. Study. thatsindicates that.a thorough revision

LS (Needed,

Key to°species of Ateuchus

ie Elytral striae moderately impressed, atleast the

) inner four striae. ol each elytron moderately

to strongly impressed near apex; elytra usually

dark “Drown or*® black, lacking \distinct”’ green’ or

solder Cast ois A a rl nice 2

Elytral. striae feebly ipeges deus “only Slightly more

impressed near apices; elytra usually with dis-

tinct green or golden cast. . .2. aeneomicans (Har.)

2a Size moderate, usually 6 mm or more; elytral

intervals dull, opaque! 0: Si candezet (Har. )

Sivel-smally; usually under’ 5:5 mm; elytral inter-

Vals cnining, polished, . .3- @uatemalensis (Bates )

Howden and Young: Panamanian Scarabaeinae 69

1. Ateuchus candezei (Harold)

Pigs. iG. Tip

Choeridium candezél Harold, T1800, . 82,

Type. or cotype, Paris. Net seen. |

Length 5.9. to 7.1 mm. Dorsally very dark brown to. black:

head and pronotum shining, frequently with cupreous cast, elytra

dull, opaque. Clypeal margin anteriorly broadly (male) to moder-

ately (female) V-shaped, tooth on each side rounded to acute, at

least slightly more prominent than margin laterally; margin

laterally arcuate; clypeal dise¢ (hig. 119) finely punctate, punc-—

tures larger and more numerous near margins. Frons and vertex

feebly tumid, smooth to very finely punctate; eye dorsally relatively

narrow, about seven facets wide in basal half. Pronotum minutely

punctate or impunctate laterally, impunctate medially. Elytral

striae (Fig. 118) distinctly impressed and with widely separated

small punctures; intervals feebly convex, surface smooth, rinely

granular, giving dull appearance. Pygidium with indented groove

just anterior to basal marginal bead, surface at least in basal

half, dull. Fore tibia with three teeth on outer margin; fore

tibial spur of males expanded into a slightly irregular, transverse

oval shape; female spur slender, slightly bent near acute apex.

MATERIAL EXAMINED. PANAMA, 119 specimens from: BOCAS DEL

TORO PROV. — Almirante. CANAL” ZONE = Barro Colorado Island;

Black Tank Rd.; Cabima; Gatun (4 mi. W); Madden Forest; Mojinga

Swamp; Paraiso; Pipeline Road; Skunk Hollow. COLLECTION DATES:

Every month of the year.

OTHER COUNTRIES, 7 specimens from: TRINIDAD (same or very

close species).

REMARKS. The opaque or dull elytra will usually distinguish

the species’ from others occurring in Panama.

This species has been recorded from light traps, forest

floor leaf litter, fermented palm fruits, and polypore Tanel.izard,

rat, and opossum carrion; and from coati, howler monkey, human,

spider monkey, tapie) and toad feces.

On Barro Colorado Island, A. candezei is active nocturnally,

particularly during the period of 2200 to 0600 hours. It is most

abundant in the wet season, when it is equally common on both

carrion and dung; in the dry season it is found only on fecal

material.

Distribution is seemingly restricted to lowland moist forests.

2. Ateuchus aeneomicans (Harold)

lip eras a AY)

Choeridium aeneomicans Harold, 1868, p. 82.

Type or cotype; “Perisy,) Not eeen:

Length 4.5 to 5.6 mm. Dorsally shining, dark green to dark

brown or black with strong green or golden cast. Clypeal margin

anteriorly shallowly, broadly emarginate, tooth on each side

feebly developed to obsolete; margin lateral to each tooth evenly

arcuate; clypeal disc (Fig. 120) with closely spaced small to

70 Contrib. Amer. Ent. last... wolves, sire. 01) 3198!

moderate sized punctures. Frons and vertex feebly tumid, minutely

punctates eye dorsally. narrow. .Pronotum Jargely impunctate,

smooth and shining, scattered minute punctures present laterally.

Elytral striae feebly impréssed, punctures in. striae smail to

obsolete; intervals mostly smooth and shining, sometimes minutely

punctate. ..Pysidium. shining, with scattered minute punctures.

Fore tibia with three teeth on outer margin; fore tibial spur of

males broad, parallel sided, apical edge asymmetrically emarginate;

fore tibial spur of female slender, parallel sided to slightly

expanded, truncate, apex,

MATERIAL EXAMINED. PANAMA, 42 specimens from: CANAL

ZONE = Barro Colorado (sland: “Gamboa; Rio Civicito. . CHIRIOUI

PROV. = Bijgaba; Volean Ge Chiricui.. COCLE PROV. — El Valle,

COLON PROV. — Portobello... PANAMA PROV. — Cerro Campana.

COULECTION DALES: January, Februdry, March,. June,.October,

November, December.

OFHER COUNTRIES, None.

REMARKS. The scarcely impressed elytral striae and the

similarly colored green or golden pronotum and elytra will normally

distinguish the species.

Specimens have been taken on tapir and Ateles dung.

3. Ateuchus (near?) guatemalensis (Bates)

PPO O17 1

Choeridium guatemalence Bates, 1307, p. 45.

Cotypes, London and Paris. London.specimens,examined,

Length 4.1 to 5.6 mm. Dorsally shining dark brown to black,

head and pronotum frequently with cupreous or slight greenish

Cast, €lyira usually Shining black. Clypeal margin anteriorly V-

Shaped, (he emarcination slightly, Wider in maic, tooth on each

side (both sexes) obtusely rounded, margin laterally arcuate from

base of tooth; disc of clypeus (Fig. 121) punctate near margin

(male) or mostly punctate (female). Frons and vertex broadly

cOnVeX, SUDiace Smooth te, minutely punctate; Gye dorsally. narrow.

Pronotum shining, smooth or with widely scattered, minute punc-

tures. .Elyiral striae, distinctly. .impressed, with small, widely

Spaced or drregularly spaced, punctures; intervals feebly convex,

SmOOLh Q@nd shining; EPycidium with basal groove just anterior. to

posterior, marginal bead; pyegidial surface.smooth, and..shining.

Fore tibia with four teetm on, outer margin, the basal. tooth feebly

developed (may be lost in worn specimens); fore tibial spur in

male broadly, irregularly tobed> in female siender, elongate, almost

parallel sided to. truncate apex.

MATERIAL EXAMINED. PANAMA, 160 specimens from: CANAL

ZONE -— Barro Colorado Island; Madden Dam; Paraiso. COLON

PROV, =) €oce Sele: Ganta kita Fidee., COLLECTION DATES: Every

month of the year.

«. OVHER COUNTRIES, 2. specimens (of nominate, form), from:

GUATEMALA.

REMARKS. ASSie@ning the name. Guatemalensis fo..the Panama

population as described above iS questionable, since the Panamanian

Howden and Young: Panamanian Scarabaeinae Th

form is smaller and may differ significantly in other respects.

However, at present, introducing a new name in the genus, until

it is revised, seems an even worse procedure.

This species has been obtained at tree sap; in forest floor

leaf litter;.at..vat. carcasses:.and, on: coati, howler monkey, human,

peccary, spider monkey, and tapir feces.

Ateuchus guatemalensis on Barro Colorado Island is diurnally

active and most abundant in the dry season, when it is found

only on fecal material. Individuals captured in the wet season

are likely to. be found on: carrion. |

The favored. habitat seems to be the lowland moist forest of

central Panama.

Canthidium Erichson

Canthidium EFrichson, 1647." p. 100>" Bates, eo). 4 27.

Type-species: Canthidium lentum Erichson, designated by

Lucas, 1920.

Neocanthidium Martinez, Halffter and Pereira, 1964, D, 160, NEW

SYNONYMY.

Type-species: Neocanthidium bokermanni Martinez, Halffter

and Pereira, by original designation.

Small to moderate (3 to 11 mm) in size, oval to almost

circular in outline. Clypeus feebly.to distinctly bidentate, in

some species quadridentate. Frons flat or transversely carinate

between lateral tubercles or with two or three distinctly separate

conical tubercles, Pronotum lacking horns or anterior indentations,

evenly convex; lateral pronotal fovea small, often circular, some—

times obsolete. Scutellum not visible. Elytron with eight striae

on disc, the striae often feebly impressed or obsolete. Mesosternum

narrow, transverse. |(Fore ‘tibia with three distinct teeth ion outer

margin. Middle and hind tibiae apically expanded, tibiae lacking

transverse carinae. Basal segment of hind tarsus two or more

times as long as adjacent segment.

The genus Neocanthidium is treated herein as a synonym of

Canthidium. The major distinguishing character supposedly

separating Neocanthidium from Canthidium is a row of punctures

across the base of the pronotum. All degrees of variation from

large to small to no punctures are present in the various

Panamanian species and none of the characters mentioned by

Martinez, Halffter and Pereira (1964) seems to consistently

separate’ a group of species from the genus Canthidium as defined

by Harold. ie

The main characters distinguishing Canthidium as defined

herein are the very narrow (measured longitudinally) mesosternum

and the long basal segment of the hind tarsus. Other combinations

of characters could form a variety of groups, but if this were

done the groups formed would probably not represent monophyletic

lines. Some of the sexual characters of Canthidium illustrate

this. All males in the genus have the last abdominal sternite

adjacent to the pygidium narrowed medially. However, many males

also have the fore tibia modified in several ways. In some

species the fore tibial spur is slightly shorter than in the female,

TZ COontripi Amer. (ERE Tash? voll is me. Ths 198 1

but. In ceneral there: is little sexual modification. (Other: males

have ‘the fore: tibial spur Tlattened, “expanded: apically and often

with the, apex shallowly emarginate. “ Uhisi type of modification

also Occurs\.in Ateuchus sand some, other.ceneras Phe fore tibia of

Catithidium itself may be modified by “having the inner apical

anole produced into an .acute) lobe, Similar in this tovcome males

in. Onthophacus. (Other? males may have’ a distinct tuft of setae at

the base, or the tibial spun; iin) Thissease the spur is usually not

modified. A’ similar modification, “ise., tuit of setae, occurs in

some groups of Onthophagus. Within Canthidium the various

tibial, modifications can occur ain different combinations in the

males of different species, with seemingly closely related Species

having Quite dilierent Combinations, Thus; any @eneric groupings

based on the extermal modifications of, the male fore tibia. would

be, anvarbitrary ang artificial classification.

Tne genus is a large one, and, is mainly, if mot. .entirely,

Conlinied to areas of ieavy forest, im the, tropical and .subtropical

areas of the New Word,

Key (oO, species of Canthidium

t Clypeal-trontal region, with two or three ‘conical

tubercles, at Teast one ‘always with shining,

angulate apex; suriace ‘sometimes carinate

between tubercles: 6. ae 2

Clypeal-frontal region lacking: tubercles or ith

One to. three, feSbly elevated, broadly rounded

swellings, these not. distinctly angulate at

A a Re a er a keira! TO

2. Pyoidium with distinct, well defined’ tat least

in basal half) punctures, the punctures separated

Dy “approximately one diameter (fo. 3

Pyeidium impunctate or with minute, inconspicuous

UC MUS A AO Mee Oe ae 4

oe Clhypeal-ltrontal region with: two tubercles” connected

by “low ‘transverse carina; average body length

A OS Re ian anousticeps: "Bates

Clypeal- beer at region ‘Wak oe tubercles,

Sucrace ol Carinate between: average body

Peet emi Se a ea a rifes < Bates

is Pronotum narrowly impressed along posterior mar-

ein anterior to élytra, the impression with

single wow Of punciures,’ | punctures often

elongates average: Dodgy Teng over & mmoie |, 5

Pronotum not impressed adjacent to posterior mar-

ein, tacking transverse row ‘of punctures

adjacent tO °(marein; average body tength 4.0

CO Oe RR a IE Nh SOT LOS ?

10:

Howden and Young: Panamanian Scarabaeinae

Clypeus: with »anterior’ median é€dge with distinct

V-shaped emargination; frons not elevated

between clypeal-frontal tubercles; average body

length 6 mm {

Clypeus with anterior median edge ‘proadly,

shallowly emarginate; frons elevated between

the two clypeal-+frontal tubercles; avenage: body

length Gicimime heii ane cs el a en oldie Pei.

Two, frontal) tubercles strongly: developed: pointed;

pronotum tan,.head»and elytra vbrown to: black:

elyitralsirivae ieee, impressed (for genus)

04 DItUb eri irons. nm.) sp.

Two frontal tubercles feebly developed, scarcely

elevated (the species also keyed in second

portion) of first.couplet);head and pronotum

brown to dark brown, cupreous, elytra similar

or lighter imnecolor;:elytral striac mederavely

LMPrESSeG louie. Da eleg amie a Rabi (in part)

Elytral striae fine, scarcely impressed; feebly

punctate: or impunctates pyerdialyaurtace ion

males mostly smooth and shining, medially

Hat Ora ronver: (ih 0 He ii 8

Elytral striae impressed, distinctly punctate:

pygidial surface of males granular, dull,

medially often with shallow concavity; average

body ‘length 5:5 mmocke. (Gl yeomieeony aiaiale min. \Ssp.

Elytra lacking metallic sheen, usually of differ-

ent, color) thany preneliuin yay | CRN g ae a hitle -

Elytra with cupreous or greenish cast, ‘similar

in color ‘to pronotum; ,average . body length: 4.5

er ee aA arrow LO Chaco tari: ccm le simile ts 5 abl)

Clypeus medially with V-shaped emargination

between tooth on each side, a rounded, shallow

emargination at outerimargin oly each tooth,

the clypeal margin often appearing almost

quadridentate; elytra dark brown to black,

frequently with tan spots or rarely each elytron

tan except invantentonitaterat halt and ialone

margins; aera body: length: 4.5) mm eis

NRO ty variolosum me Se.

Clypeus with Hinodlenattiatyy broad V-shaped emargin-

ation between teeth, lateral emargination lack-

ing, the clypeus appearing simply bidentate,

the outer margin arcuate; elytron ivory to tan,

suture and lateral margin narrowly black;

avera ce body) demoted) mim of) Leo tepeepterame:) iin.) sp.

Pronotum posteriorly with transverse row of punc-—

tures immediately adjacent to margin anterior

TO API ceive tect Ma RUST IAN MITA ARO CKO CWS A Sh eat anal iele Te

iZ.

1A.

iD.

16,

Contribw fmer. inte inet.) solce lose. 14. 1981

Pronotum posteriorly lacking transverse row of

puncties, adjacent te Marcin okey ke. 13

Clypeal teeth obtuse, emargination between

broadly V-shaped; color dorsally uniform brown

or Diack; lacking ‘distinct metallic ‘reflections;

size normally more than /imm <.=.°> pei SG 2

Clypeal teeth, acute,. emarngination pee geen

narrowly V-shaped; color dorsally brown to

dark brown, elytra often. lighter; head and

pronotum: citenm with cupreous or green cast;

puna aie aa Onrmnay si6 vr

‘ a ae SR Balth. (in apart}

Eye dorsally at posterior inner edge of canthus

constricted,;. four to six facets wide; hind tibia

with inner narrow: edge straight to:arcuate,

not distinctly sinuate; frons distinctly tumid;

average body length 6 mm... .. 2. tuberifrons n. sp.

Eye dorsally al posterior inner edge of canthus

not constricted, more than 10 facets wide; hind

tibia with inner margin feebly (female) to

distinctly (male) sinuate; frons feebly tumid;

average Docy teneth 9.5 mm. . . - is centrale Bouc,

Eye dorsally at posterior edge of canthus

Constricted, ‘tfaree to six facets wide 1 6. ete. 5

Eye dorsally afl posterior edge of canthus not

COne elcted. FO OF More Tacers Wide. eee eee 14

‘Elytral striae distinctly punctate; pygidial

length to width*rdtio 2.5:4.5; average body

Tength 406 mina .9e sco. edz. vespertinum ne Sp.

Elytral striae impunctate or nearly so; pygidial

Tenet TO Widin ratio 3.5:5.5; average body

Tena oe. Shim aa . phanovultum: nn. sp.

Pyeidium- impunctate or minutely punctate;

surface usually shining, faintly shagreened,

finely sranulan, or: smooth between punctures. . 16

Pygidium with moderate sized punctures; surface

duils eranular; eee here 4.9 mm. ;

eee ee gel te tec ce dewunemeciie mW SD.

Elytra, at most only slightly different in color

to pronotum, brown or: Dlack, ee with

CUupreous or green cast ii... bis Ly

Elyiva distinctly diitghier im: color ann ee

elytra ivory to tan, pronotum: dark brown: or

black with slight metallic sheen; average body

(one eo i ee ee oe De licoalatuin / A. Sp»

Howden and Young: Panamanian Scarabaeinae 15

oe Elytra surface largely shining,” intervals’ often

minutely punctate, surface between gee oe

smooth, snotooranular<) 2%" Sa eeoeds

Elytca, dull tintervais® very fedbiy “punctate or

impunctate, surface distinctly granular; average

body: leweth 5 mm... ee a ere oe Date.

18. Strial lineson elytral’disc lacking distinct

punetures six 2. [os Ore

Strial lines on elyival dice dior, punctate:

averace body lengin 4.3 mm ; ae

ee an as eaacyen peak ta res er eee A Sp.

1. Metasternum between middle coxae smooth or at

: most minutely punctate; average body length

Le Nee Gd 15, 92) ibe Seamdens Bares (im part)

Metasternum ponvcen middle coxae with scattered

small but distinet: punctures; averace.-body

leneth 3567 mms ae yo ee) | Ey es penherdei n= sp.

1. Canthidium centrale Boucomont

Figs. 122-125

Canthidium centrale Boucomont, 1928, p. 203.

Corype;. Paris.) sNor seen.

Length 8.5 toel0:0 mms “Dorsal: color” black, cecasionally

brown (teneral specimens); pronotal disc and elytral intervals

very minutely punctate and slightly alutaceous, head dorsally

more distinctly, closely punctate... Clypeus* (Fig. 122): feebly ‘bi-

dentate, median emargination with shallow, very broad V-shaped

emargination; marginvlaterally on each side evenly arcuate to

middle:of gena. Dise of clypeusmearly dlats frome broadiy;

feebly tumid, tubercles or distinet swellings lacking. Eye dorsally

at posterior end ‘of gena’ broad, <1) orvmore facets wide. -Pronoimum

evenly,. broadly. convex, Jateral foveae obeolete, ipesterior margin

lacking adjacent transverse indentation but with contiguous row

of small, elongate ‘punctures, these absent at and near midline.

Elytra with ‘strial- lines distinctly indented and punctate, punctures

separated ‘by ‘three’to.. five diameters: intervals convex: Pyoidium

with scattered: fine*> puncturés, punctures closer in. basal third.

Fore. femur posteriorly with partly double, irregular longitudinal

row of setose punctures, remainder of surface smooth; middle and

hind femora with ventral surfaces smooth, impunctate. Fore tibial

spur of male’ (Fig. 123): slightly stouter and with more rounded

apex than in female; also in male a small basal tuft of setae

present adjacent to spur. Hind tibia of ‘maies with’ inner edge

strongly sinuate in basal half; in females the sinuate inner margin

feebly expressed or absent. Male genitalia as:in’ fies. 124.. 125:

MATERIAL: EXAMINED., PANAMA, 39 specimens from: CANAL

ZONE: = Barro Colorado Vsland: Madden. Forest. «PANAMA PROV. -

Cerro ‘Campana. -CCOLULECTION “DATES: january, February, March,

May, July, August, September, November, December.

76 Contrib. Amer. ointus dast.) wolwnt6.) mo. 6d) 9S]

OTHER COUNTRIES, 32 specimens frém: MEXICO, GUATEMALA,

ECUADOR. :

REMARKS... [Che elytral (intervals of the, Ecuadoriam specimens

are dull or opaque, whereas in the Central American and Mexican

specimens: the, intervals) are shining.

This species has been collected in light traps; in forest

floor leaf. litter; at rat. <arrion;, and son human, and tapir feces.

Canthidium centrale) is mocturnally, active’ and during’ the «day can

be found, in. lean litter.

2. Canthium Taverlirons ne sp.

Figs. 126-129

HOLOTYPE. Male, length 8.6 mm, greatest width 5.3 mm.

Dorsally dark brown, pronotal disc and elytral intervals very

minutely punctate.or impunctate, distinctly. alutaceous. -.Clypeus

(Fig. 126) bidentate, median emargination V-shaped, margin on

each side evenly arcuate. to middle of cena: ¢clypeal surface in

anterior bali closely, contrauously punctate; posterior half distinctly,

broadly tumid, «he tumid area minutely punctate; -frons: and vertex

Just posterior to tumosgilty shallowly concave, finely punctate. Eye

at posterior end of séena, narrow, fiven facets wide... Pronotum

broadly convex, lateral foveae obsolete, posterior margin (Fig.

127) with adjacent transverse depressed area, a line of distinctly

elongate punctures present in depression contiguous with margin,

punctures .obsolete:,at midline. ) Elytra with striae: distinctly

indented and punctate; punctures sin veach siria separated by Irom

two to five diameters; intervals convex. | Pygidium with. small

punctures “present in basal third. “fore femur with irregular

double row of setose punctures on posterior (ventral) surface;

middle andihind femora with ventral surfaces smooth, impunctate.

Fore tibial epur sliohtly thicker than: in’ female, otherwise. unmodi-

freddy. ca small basal tulisoh setae present. Hind: tibia: with inner

mMarcin-evenlyiy inwardly, curved; mot Sinuwate, (Genitalia \as in

figs. -hvoy B20.

ALLOPYPE.cPemale, leneth /.5 mm, @reatest width 4./.mm.

Dorsally black... Differing from holotype in) the: following major

characters: anterior half of .clypeal disc closely: punctate and

feebly transversely rugose; pygidium less punctate; fore tibial

spur mere slender, only three orcfour setae. present at base of

spur.

TV¥PE MATERIAL: Holotype; male, Panama, Chiriqui. Prov.,

1500\m, 15.°km NW) Hato del Volcan, Hartmann's Finca, .20+31.V.

19/77, Sa (Peck, dune traps 4-6 (Howden)s \Allotype) female, same

locality, as holotype,:6°51'N, 82040 W, 24-25.V.1977,. H. & A. Howden,

cup trap in ravine (Howden).

PARATYPES:. 7: males, 14 females. PANAMA: 5 males, 12

females, same data as allotype except 30-31.V.1977; 1 male, Cocle

Prov., trib.) Rio. Guewabo hl. Valle, 2200). (240 15,.1959,..HUS. Dybas,

in horse manure; 1 male, 2 females, Panama Prov., Cerro Campana,

S00 met SOOO Ds BOWE. LOFOy bh & CAL Howden, on feces.) 5ehX.1970;

H.P. Stockwell, human imeanure. trap, 22.V0h970,.B.C. Ratcliffe, at

Dlack light.

Howden and Young: Panamanian Scarabaeinae fe

Paratypes are in: Chicago, Ottawa, Washington, Howden,

Ratcliffe:

REMARKS. ‘Size waries from’7.0 to 9.5 mm also the degree of

convexity: of :the frons varies; often being’ more. "pronocunced in

larger specimens. Canthidium tuberifrons is seemingly related to

a complex ‘of species, \including C. convexifroms: Balthasar,” that

ranges from’ Mexico to Argentina.'+- In the: Case of Convexifrons

from Colombia, the species is smaller and dorsally distinctly

smoother and more shining than is tuberifrons.

3. Canthidium haroldi Preudhomme

Figs. 130-133

Canthidium haroldi Preudhomme de Borre, 1886, p. 111; Bates,

3 LEO Sw Var

Type,< Geneva 02). ' Net cseen:

Length 6.1 to 10.0 mm. Dorsally shining reddish brown to

black’ with: strong green or cupreous reflections; *pronotal idisc and

elytral intervals mostly smooth, very’ feebly,« minutely” punctate.

Clypeus (Fig. 130) feebly bidentate, median emargination shallow,

broadly V-shaped; margin on either side evenly arcuate to middle

of géna; clypeal frontal. juncture elevated, Tuberculate’ at each

end, the’ two: tubercles separated by approximately Same distarice

as.clypeal: teeth; surface of head, exchiding: tuberculate area,

moderately closely ‘punctate.!: Eye dorsally. at posterior ‘end Of

gena narrow, four to seven) facets ‘wide. sPrenotum ‘with lateral

fovea small, circular, sometimes obsolete; area immediately adjacent

to posterior margin indented and with single row of longitudinally

elongate “punctures: ¢ Elytral striae’ represented) by finely indented

lines, faint’ punctures present im (etrial lines; + punctures ‘Geparated

by 8 to 10 diameters or more; intervals almost flat. Pygidium

shining, finely ‘punctate and shaereéned, Posterior surface. of

fore femur? with. longitudinal, “irregular: row; or large, setose punc-

tures" in males, thevséetose (punctures: present but ‘small in females:

middle and hind femora’ with ventral’ surface minutely punctate

and shagreened. Fore tibial spur of male (Fig. 131) not greatly

modified, cylindrical, and with acutely pointed apex; tuft of setae

present ‘at inner: base “ol spur? duit lacking in feneales?’ Male

Cenitaliaras in figsa lah oe.

MATERIAL EXAMINED. PANAMA, 164 specimens from: CANAL

ZONE - Barro Colorado Island; Canoa Saddles; Curundu; Fort San

Lorenzo; Madden Porést.: oCHIRIOUD' PROV. = Bugeba.,: PANAMA

PROV, —' Cerro ‘Azul: (Cerro Campana, (Cerra jete. ) COLLECTION

DATES: “Every month ofthe year,

OTHER COUNTRIES, 24) snepimens' trom: (GUATEMALA, COSTA

RICA, ECUADOR.

REMARKS. Specimens from Ecuador have the pronotum consis-—

tently more distinctly punctate and the dorsal surface less shining.

This’ species has been: captured: perching on leat surfaces in

forest floor leaf litter; at rat carrion; and on coati, howler

monkey, ‘human, opossum, and tapir feces:

78 Contrib. Amer. “btw dnsi.) ovoki86,mo. +)..9961

On Barro Colorado Island harold) is; active: throughout the

day, particularly during the periods of 0600 to 1000 hours and

from 1500 to 1800 hours. The species is most abundant during the

late wet season.

4. Canthidium angusticeps Bates

Figs. 134-136

Canthidium angusticeps Bates, 1887, p. 49.

Cotypes, London and Paris. London specimens examined.

Length 6.9 to 8.1 mm. Head and pronotum reddish cupreous,

finely punctate; elytra brownish black to black with cupreous

reflections; intervals minutely punctate, smooth between punctures.

Clypeus (Fig. 134) distinctly bidentate, median emargination

deeply U-shaped, clypeus broadly, shallowly concave to margin;

Margin On each side im conjunctiom. with maroin of gena. unevenly

Sinuaté.« rons transversely caninate, the carina raised: at each

end, essentially bitubéerculate with carina connecting» the tubercles.

Eye dorsally at posterior end of gena narrow, five to seven facets

wide. Pronotum-evenly convex except for small, circular lateral

fovea on each side; posterior edge of pronotum with transverse

row. Of moderate: sized, cireular punctures; the row of punctures

not in. transverse. groove. -Elyltracwith striae represented: by

feebly indented lines interrupted by large shallow punctures

Separated by three to five diameters; intervals nearly. flat,

minutely punctate. Pygidium distinctly punctate, punctures

separated Dy (approximately ome diameter... Posterior surface: of

fore femur with scattered coarse, setose punctures; middle and

hind femora ventrally each with elongate; fine. punctures: parallel-

ing long axis of femoray:=Fore tibial: spur of male bhuntly acute.

Male genitalia as im figs. 135, 136.

MATERIAL EXAMINED. PANAMA, 3 specimens from: CANAL

ZONE: =~ meer Fort:/San korenzoy..201m:. PANAMA: PROV.<= Cerro

Campana, 650 m:.-COLLECTION DAVES: April, Aucust, October.

OTHER COUNTRIES, 5: specimens «from: .COSTA: RICA, COLOMBIA.

REMARKS. The Colombian specimen from Anchicaya,;: /O°km. E

of Buenaventura, has more heavily punctate elytral striae than

Panamanian specimens. This difference may indicate a sibling

species bul more material is needed. .A very closely related

species (7?) occurs at Limoncocha, Napo Prov., Ecuador. «There

appears to be in South “America ax widespread complex. of forms or

species related fo, angusticeps.

specimens were taken in traps baited with human» feces.

5. Canthidium elegantulum Balthasar

Figs. 137-140

Canthidium elegantulum Balthasar, 1939c,:p.. 12k

type, Pracue.: « Examined,

Lenoth 5.5 to 6.5 mm. Colom dorsally tan. on brown or

reddish brown, often with cupreous cast; elytra frequently lighter

Howden and Young: Panamanian Scarabaeinae 79

in color than’ prénotum; ‘surface sof head, pronotal disc and

elytral intervals smooth to minutely punctate, strongly shining.

Clypeus (Fie. 137). distinetly “bidentate, “sharply shaped between

teeth, shallowly emarginate on each side adjacent to tooth, margin

then evenly arcuate posteriorly to middle of gena; clypeal disc

shallowly concave. © Frons tumidy'a feeble, sometimes obsolete;

tubercle present on each side near inner junction of clypeus and

gena. Eye dorsally iat. posterior end of gena very narrow,’ two°or

three facets wide. Vertex nearly flat medially, surface smooth,

minutely punctate. Pronotum with shallow lateral fovea, a narrow

band contiguous to posterior margin feebly indented, indentation

(Fig. (438) with row of ‘shallow ovalito cireular puncturesy> -Elyira

with striae represented by feebly indented lines interrupted by

widely spaced shallow punctures, punctures usually separated by

more than six diameters; intervals feebly convex. Pygidial

surface minutely punctate, a few coarse punctures in basal groove.

Posterior surface of fore femur with a few scattered, coarse, setose

punctures; middle and hind femora ventrally with scattered very

minute punctures or surface impunctate. Fore tibial spur of male

tapered to acute apex, similar to female in this respect, but with

tuft: of setae at»base of spur’ that is lacking in females, Male

eenitaliaasin: figs., 139; «140.

MATERIAL EXAMINED. PANAMA, 99 specimens from: CANAL

ZONE - Barro Colorado Island; Gatun; Madden Forest; Pipeline

Road: Rio’ Ciricito: Skunk Hollow: Tabernilia. PANAMA PROV..5

Cerro;Campana; Rio Pequent.) : COLLECTION DATES: Every, month: of

pe .vear’.

OTHER COUNTRIES, None,

REMARKS. This. species: has been found-sitting on leaves; in

forest floor leaf litter; in and» onvrotting fe. [ruitson the ground;

under an old coati carcass; and. at bat, howler: monkey; tapin,

tayra, and human feces:

On Barro Colorado Island elegantulum is present throughout

the year but is. most common -durine-the-middie of the rainy

season. Diurnally active, it is most frequent from 0900 to 1500

hours.” The species 19: seemingly vestricted to the Jowland Torests

of central Panama:

5. Canthidium bituberiirons.n. sp:

Figs. 141-143

HOLOTYPE.” Male, length s5.3 mim, ereatest width 3.7 mm.

Head dorsally: largely shining black with greenish reflections;

pronotum shining yellowish tan; elytra shining dark reddish brown

to black; head; pronotal dise and elytral intervals smeoth or very

finely shagreened and minutely punctate. Clypeus (Fig. 141)

strongly bidentate, median emargination abruptly U-shaped; margin

laterally. evenly “arcuate: to middle of cena. Frons: near inner

anterior margin of -éach eye with strongly elevated; conical tubercle,

the area: between feebly concave: dorsal surface of head smooth;

eye dorsally at: posterior end of cena narrow, five to’ eight facets

wide. Pronotum evenly convex, lateral fovea small, indistinct;

80 Contiritba:: Amer. “aria best. : yokvers, mo. vb esi

narrow band immediately adjacent to posterior margin (essentially

marginal line) with row of small, circular or oval punctures,

these slightly larger and somewhat anterior to margin near

(posterior) midline basally. Elytra with strial lines distinctly |

impressed, each line distinctly punctate, the punctures separated

by four to six diameters; elytral intervals feebly convex, more

distinctly ‘so' near apices. :Pygidium mostly smooth and shining.

Posterior surface of fore femur with longitudinal irregular row of

small setigerous punctures, surface otherwise finely shagreened;

middle and hind femora ventrally minutely punctate and finely

shagreened. Fore tibia with spur slightly expanded, flattened and

curved downward in apical third, apex rounded; inner edge of

tibia at base of spine feebly produced and with small tuft of

setae, oMalecgenitalialias im ities. 142) 143.

ALEOIYPE. Female, tength 55 mms ereatestewidth 3.5 mm.

Differing from holotype in the following major characters: apical

sternite distinctly broader, of uniform width (as isonormal in

females); apical tibial spur narrower apically; tibia lacking

apical tuft of setae. |

TYPE MATERIALS: Hobotypey: mate, Panama, Panama: Prov .;

Cerro Canpana,® GO00 NT.) BOlV 11319703" He: eA. Howden, om ‘human

feces (Howden). Allotype, female, same locality as holotype, 27.

V.1974, O.P. Young, on howler monkey feces (USNM).

PARATYPES: 1 male, hk female: ’ PANAMA: 1 male, same

locality as holotype, 30.V1I1.1974, H. Stockwell (Howden); 1 female,

Panama Proview Cerro: Campana, 650) im, 6240 N,: “79956 Wye2le Aug.

Ode WeRe Bivin (EV Ge oRpley))

REMARKS. The paratypes do not vary noticeably. The

sharply bituberculate frons, yellowish tan pronotum, the nearly

black head and elytra, and the impressed elytral striae should

distinguish this species from other species occurring in Panama.

i. “Canthidiuim Giscepyeidiale n,. sp.

Figs. 144-146

HODOUYP Es Mate, lenetn 5.0 mm, oreatest width 3.7 mm.

Dorsally dark brownish black, pronotum light brown, head and

pronotum with cupreous reflections; pronotal disc and elytral

intervals shining, smooth to very minutely punctate. Clypeus

(Fig. 144) feebly bidentate, median emargination shallowly U-

shaped, clypeal margin laterally very slightly sinuate at junction

with gena; clypeal-frontal area trituberculate, median conical

tubercle anterior to and twice size of either lateral tubercle;

dorsal surface of head moderately, closely punctate, more densely

so on clypeus. Eye: dorsally almost: enclosed posteriorly by

posterior edge of gena, at most one or two facets wide at constric-

tion. Pronotum with circular lateral fovea shallow, small;

posterior pronotal margin lacking adjacent indentation or row of

punctures. Ebytra with ‘strial lines: moderately to strongly

impressed and coarsely punctate, punctures in most striae separated

by two to three diameters, intervals feebly convex, minutely

punctate. \Pygidial surface) dully granular, surface. ‘centrally at

Howden and Young: Panamanian Scarabaeinae 81

basal’ third, shallowly’ concave.’ Posterior surface ‘of fore femur

with three or four large setose punctures in apical half, remainder

nearly smooth; middle and hind femora ventrally very finely

punctate and shagreened. Fore tibia with apical spur broadly

expanded to almost truncate or feebly emarginate apex; tuft of

setae lacking at base of spur. Genitalia as in figs. 145, 146.

ALUOTYPE. Female, length 0.9 mm). enéatest) ridin oe7 mim.

Differing from holotype in the following major characters: clypeus

more distinctly punctate, partly transversely rugose; pygidium

shining, minutely punctate, lacking central shallow cavity, the

pygidium evenly convex; fore tibial spur relatively slender,

slightly tapered to, anteriorly) rounded \apex.

TYPE, MATERIAL?) Holotype,’ male, "Panama, Pananave rows,

Cérre Campana, G000"fts',: (BP40' NG 79C56" Wy Vii to 70;

Hee Ad Howden’ (Howden!) -Allotype,)femaleyscame locality: as

holotype, 10.11.1973, H. Stockwell (Washington).

PARATYPES: 6 males, 6 females. PANAMA: 1 male, 1 female,

same locality: as holotype;'28.1V.1973, (Hh Hespenheidas (hol 197 3,

Hi. Stockwell: 2’ males,’ 3° females) Canal Zone) PRort) Kop be; wo-24

June/1976j) EG.) Riley. “COSTA (RICA? 3) males) temale /F umtarenas

Prov) y/' 2400) 'm; 1500) «m', Monteverde (260240 Vins. Vtg oe Gea.

Howden, (males) cup traps, (female) sitting on leaf; 1 female,

Puntarenas \Prov., Osa Peninsula near Rincon) S742) Nien? 26" Ww

23.1.1968, H. Hespenheide.

Paratypes are, in: Washington, Howden.) Riley.

REMARKS. Variation occurs: primarily incolor,) the, pronotum

of ‘both the holotype and. allotype being reddish tan and’ the

elytra’ being reddish brown... Pheimale paratypes) have the head,

pronotum,’ and elytra) black witha distinct vcupreous,casty.4) In

other’ respects; except for) ithe sexual) difference as, noted, the

specimens are. quite) umiform.

8. Canthidium subdopuncticolle n. sp.

Figs. 147-149

HOLOTYPE. Male, length 4.9 mm,’ greatest ‘width 2.3 ‘mm.

Dorsally dark brownish black with cupreous cast; pronotal disc

shining, with numerous small to (moderate sized shallow punctures;

elytral intervals! impunctate,! alutaceous,’ feebly’ shining. -Clypeus

(Fig. 147) bidentate, each tooth rounded and separated medially

from its opposite by a broad U-shaped emargination; margin

laterally on ‘each: side’ sinuately) curved) to‘gena; | surface of,

clypeus closely punctate and transversely rugose. Frons with two

extremely vague swellings, one on each side midway between

anterior) edge’ of eye”-and.(midlindg, midline appearing) slightly ‘tumid

at certain angles, “Pye dorsally. an) posterior \end ot” cena’ 'moder—

ately narrowed, five to seven facets wide. Pronotum evenly convex;

fovea’ on each side. small; circular, shallows posterior pronotal

margin’ Tacking ‘adjacent (‘transverse eroove’ or row of ‘elongate

punctures. (\Elytral’ striae represented’ by slightly “indented | lines,

lines lacking “punctures ior? with) Vague punctures)present (in lateral

striae?) (Pygidium with ¢hallow,) somewhat irrecular,. scattered

82 Contribs (Amer: enti. bast: voleolS,.no. «l4.-1981

punctures, pygidial surface distinctly granular, dull; pygidium

one-sixth wider than lone.» Posterior surface. of fore femur.and

ventral suriaces.of middie and hind femora-impunmctate, alutaceous.

Fore tibial spur; not. greatly modified, slender, velatively. short

(worn”’), inner apical: angle of tibia adjacent. to.spur produced

and slightly bent downward, forming a broad, acutely angulate

lobe approximately as long as tibial spur, , Genitalia -as in. figs.

1A0, 109)

FEMALE. Unknown.

TYPE MALERIAL: sHoletype;, male.,Panama,. Canal Zone

(= Colon Prov.), Alhajucla, 28.V.1912, A. Busck.(Washington).

REMARKS. Initially we considered this species to be a small,

edd specimen of G. puncticolle Hary., but the two extremely vague

tubercles on the head, cupreous color, shallowly punctate pronotum,

obsolete, strial punctures, and cenitalic characters seem. to. preclude

the inclusion of the untque.male under puncticolle. |.Actually.

puncticolle of: authors seems tO peprésent a, complex. of forms,some

With two, come with: theee tubercles on the head... At. present the

closest form t0 subdopuncticolle is seemingly the blue variety

mentioned by Bates (1887, p. 49) that occurs from Costa Rica to

Mexi1Go.. ,ihis’ Dlue ‘form 1s distinctly larger im Size, the pronotum

and elytral striae are gnore sirongly. punctate and the male genitalia

differs! im. shape. :

y. Canthidium tenebprosum n. sp.

Figs. 150-152

MOLOTI PE. Mate, dleneth 5.3 am, ereatest width 8.2.mm,

Black, elypeus and. elytra stinged with brown; legs, brown; dorsally

icebly shining; promotum and, to lesser degree, elytral intervals

minutely punctate, surface finely granular. Clypeus (Fig. 150)

bidentate, median emargination broadly V-shaped, margin. laterally

evenly arcuate om ¢ach,cide. 16 maddie.o!f gena except slightly

notched at clypeal-cenal junction; surface of clypeus feebly,

transversely tugose. Prontal area feebly tumid, particularly just

anterioy, FO and) inside inner edge of eye, a vague median) tumosity

also present (indicating relationship to other trituberculate species);

frons and vertex moderately punctate, surface between punctures

Granulate. Eye dorsally al, posterior end:of e@ena: narrowed, three

10. five: facets, wide... -Pronolum moderately convex, lateral foveae

small, circular, shallow; pronotal surface finely punctate, . between

DuNnciUres \o@ranular, dull. .) Piytral, striae represented..by. feebly

impressed, limes, each line with, shallowly, impressed. punctures,

the punemires usually separated by four, to..six diameters; elytral

intenvals flat, obscletely« minutely punctate, .oranulate.. Pygidium

With -seatierec small to minute, punctures, surface finely granular

Ore shacreenecd, dull.) lostertonr surface f.fore, femur, with. two. or

Lhree coarse, .seloce. punctures: mean, apex, remainder of surface

shagreenedand with. scattered, very minute punctures.. Ventral

Surfaces of middle and, hind; femora. timely shagreened and. with

scaltered minute: punctures... Pore: tibial spur..not) obviously

modified, «slightly shorter than in: temale;: inner: anterior. angle. of

Howden and Young: Panamanian Scarabaeinae 83

tibia’ adjacent’ to spur produced into acutely angled: lobe; *tuit of

setae lacking. Genitalia as in figs. 151, 152.

ALLOTYP. Female, léenoth Si 4 mm) oreatest: width G72" mm.

Similar ‘to holotype ‘except in’ the following characters color

dorsally with distinct greenish cast; clypeal margin laterally less

strongly arcuate; pronotal surface only feebly granular, more

shining; pygidial’ punctures greatly reduced; posterior suriace’ of

fore femur with four or five setose -purictures: near apex; tore

tibla-with inner apical angle’ not. preduced, “nev aeute.

TYPE MATERIAL? Holotype, mate,” Panama; Giitvaut Prov.,

2 km’ W Cerro’ Punta, “Baldwin. Forest, 3. Vi. 197 7," Me poween, vcup

trap (Howden). Allotype, female, Panama, Chiriqui Prov., 2 km W

Cerro Punta, 6°5)'N; ~62936'W;°22.VA977, Ho Howden, cup trap

(Howden).

PARATYPES: 94 malés, 129 females. PANAMA? 50 males, 80

females, same data as holotype; 17 males, 17 females, same data

as allotype; 9 males,” 7 females, Chiriqui Prov.) Gevro- 7 onta,

1500 m,° 23-V.1977, Hs Howden; ‘cup traps; -4 males, 77 females,

Chiriqui Prov., 2 km. W Cerro Punta; 1760 mm, Baldwin’ Porest, 30.

V.-O.V1T.1977,°S:' Peck, ‘dune trap; 2 males, 4) féemdies, Chiriqui

Prov .,"/2-3: % 1975, OP. Youre, tapir feces; rat carcass tran; 2

malés, 2° females, ‘Chiriqui “Prov.; 2.5 km W Cerro Pinta; 6°51" N;

62°36'W, 1720 m, -287%1.1975,;° HV Pes Stockwell, “manure trap. — COSTA

RICAs’ 1 male, Alajuela Prov.;° Volean Poas,’ Poacite; 1040 m, 20:

VIT.-3.V111.1960,°S. Peck, carrion trap; 9 males, 12: femates,

Puntarenas Prov., Monteverde Reserve, 1500 m, 27.V-1.V1.1979, H.

one Howden, cup’ traps: and. larce duno trap.

Paratypes are in-vvall collections studied:

REMARKS.” The large: series. om hand ‘varies: in length’ trom

4.0 to 5.5 mms. The majority of specimens are’ dark brown to

black, but a’°number of ‘specimens; particularly females; may: have

a cupreous or’ ereenish' cast. “Alb specimens, unices worn. are

dorsally: granular, the surface, at most, feebly Shining? - The

. degree of rugosity’® of the -clypeus varies,:-in some specimens the

clypeal ‘surface’ is: more contiguously punctate than: rugese, The

tumosity of the frons’ also varies, but noné of the: specimens seen

have the frons distinctly trituberculate (or tuberculiate).” The

spacing ol the’ punctures on. the pronetum. om the elytral striae

and elsewhere, shows some variation, but does so within a relatively

narrow range.

The relatively dull pygidium might: cause some specimens ‘to

be keyed out as subdopuncticolle, but the rather elongate Dody

shape of ‘tenebrosum™ should separate it) trom the puncticolle group:

10... Canthidium planovultum nn... Sp.

Figs. 153-155

HOLOTYPE. Male, length 4.6 mm, greatest width 3.0 mm.

Color dorsally black with head and lateral margins of pronotum

brownish black; head and pronotum finely, distinctly punctate;

elytral intervals irregularly, + indistinctly “punctate: Clypeus’ (Fi¢.

153) feebly bidentate, median emargination shallow, U-shaped;

84 Contrib. pAmet. Ohintedsbneiue woly taboo. bb LOB TL

clypeal).margin).om either, side more) abruptly. curved. near .jumctions

Ol gend, the curve nearly unifonm, to.middle of cena. .Clypeal-—

frontal, area. wery slightly tumid,: entire surface of head closely

punctate, anterior half of clypeus contiguously punctate or trans-

versely rugose. Bye dorsally at posterior, end of .gena, broad. 12

to 13 facets wide. .Pronotum broadly convex, lateral: foveae

obsolete, posterior, edge of. pronotum lacking. adjacent elongate

punctures or transverse. depression... Elytral striae faint to

obsolete, lacking, distinct punctures:,,intervals flat, finely punctate,

the punctures) irregulac.in.size amd. shape,.diving surface

irregular appearance. Pygidium large for genus, approximately

1.2 mm wide, surface with widely scattered small punctures, the

area. between smooth, and shining. «Posterior surface of fore femur

and ventral surfaces of middle and hind femora with scattered

minute punctures and) finely, longitudinally. alutaceous.') Fore

tibial spur slightly thickened but not noticeably dissimilar from

female, inner apical margin of tibia at base of spur deflected

downward and extended to form a broad, acute lobe. Male

eenitalianvas, iin fos. c 150, i155.

ALLOTYPE. Pemale, length 4.4 mm,.greatest. width 2:9°mm.

Different from male in the following major characters: pygidium

with surface slightly irregular, less distinctly punctate; apex of

fore tibia not forming a broad, distinctly enlarged lobe but still

slightly produced, forming an acute angle, this character being

unusual in females of the genus.

IYPE MATERIAL: \soloiype, “male, Panama, Chirigur Prov.,. 2

km Wi Cérro Punta, [300m B25 Mia ee gotW 27. V. 1977, H...Howden,

cup trap (mowden). Allotype, female) Panama, Chiriqui. Prov., 2

kin Wy GCerrocPunta, Baldwin! Porest, .G.V 1.1977, Hew& A. Howden,

cup trap (Howden).

PARATY PRES: 09 males, 131 females... PANAMA: 60° males,: 70

females, same data as holotype; 14 males, 26 females, same

locality (as, alletype, 30.No-OuVial Ol, Ga. Peck» mega, dung .traps;.8

males,.16.females, same bocality as holotype, ,22.Vs-3.Vi.1977,. HH.

S Ay MOwWGehaa cup. traps: 2 males, Chiriaui Prov. «Cerro: Punta,

1500: 1m, 23. V.107/,. H. Howden, cup traps; 4, males,,.10.females,

ChiriquivProye, 20GvX019/5),\O0P,) Young, tapin.feces) trap, : rat

Carcass traps Ocfemaies, Chiviguin Prove, 2.5. kne.W.,Garro Punta,

mudubonm, cabin, 2o.41.19/5,. fH.) Stockwell, I9.Vil9/J5. Siok J. Peck.

COSTA RICK mate, Alajmela (Prov: Volcam Poas, Poacito,..1840..m,

20. V1 boo Vbh bs t0G0,, Sa Peck, carrion trap.

OTHER SPECIMENS: One. male, Mexico,., Oaxaca, 20 mi. S

Juchatengo, 6000 ft., 27-30.V.1971, S. Peck (apparently the same

Species, DUL dilfers slightly, in punctures on. fore tibia’ and

clypeus).

Paratypes are in all collections studied.

REMARKS. Considering the large series of planovultum on

hand, the species is remarkably uniform, varying moderately in

size, (rom. 3.9) to 419 mimes lhe omal form, <distinct convexity, and

dark brown to black color gives the species a fairly distinctive

appearance. However, for initial recognition, the characters as

outlined in the key should be. utilized.

Howden and Young: Panamanian Scarabaeinae 85

liv Canthidium perceptibile: tm. .' sip.

Figs. 156-158

HOLOTYPE. Male, length 4.3 mm, greatest width 2.6 mm.

Dorsally shining, dark brownish black with strong cupreous and

greenish reflections; pronotum and, to lesser degree, elytral

intervals minutely punctate. Clypeus (Fig. 156) bidentate,

median emargination V-shaped; margin laterally feebly arcuate

to slightly notched junction with gena; clypeal surface transversely

fugose.: Prons on either “side “and” anterior’ to "eye with “teable

tumosity indicatine tubercle, “however frons “definitely ‘not tapercu-

late; surface ‘closely purictate."| Eye’ dorsaily at) posterior, end’ or

gena narrow, three to five facets wide. Pronotum moderately

convex, ‘lateral foveae ‘small, “circular, conéave.; ilytron with

strial lines feebly impressed, each line distinctly punctate, the

punctures ‘separated “by approximately Tive”’ or six ‘diameters: inter~

vals nearly flat, ‘surface’ minutely” punctate on’ faintly tregular,

polished ‘and shining. Pygidium with°scattered: Small, punctures,

surface between mostly® smooth and ‘shining. Posterior surface: of

fore femar> with one or two’ ‘setose ‘punctures, near’ apex, remainder

of ‘surface minutely punctate and longitudinally shagreened.

Ventral surface of middle and hind femora minutely punctate and

lensitudinally “shaereened. Apical: spur of? fore tibia long and

slender, apex acute; adjacent inner angle of tibia narrowly,

anteriorly produced, forming ‘sharp, acute lobe a@oproximately nalr

the ‘léneth: of ‘spur; tibia Jacking: apical Tull ot /serae. |) Mate

genitalia as in figs. 157, 158.

ALTOTYPE.| Female; length’ 4.5) mm ereatest width 45.0 "mm.

Dorsally ‘shining, bluish: green’ to. black.’ (Dilterine trom holotype

in the following major characters? olypéus more rugose: elytral

striae léss strongly punctate: fore tibial spur slicntly tareoer,

more slender; fore (otal teeth lessiacure:

TYPE? MATERIAL? Holotype, male, Panama, (hiriqui ch rows. 2

km N Sta. Clara, V300°m, GO) "N, G2046 Wit terimana 6 Finca:

24-25.V.1977, H. & A. Howden (Howden). Allotype, female, Costa

Rica, Puntarenas, Monteverde, 1500°m, 23.V.19/9, H. ® A. Howden

(Howden).

PARATYPES: 8 males, '11 females.’ PANAMA: 1 male, same

locality :as. holotype, S03) VeTOP Tyne) Gy A roweden ) ins cup tre

baited with human feces; COSTA RICA: 7 males, 11 females,

Puntarenas, Monteverde, 1400 m, and Reserve, 1100 m, 27-31.V.

W711, He & A. Howdeny: cup Tragtseitiine on leaves:

Paratypes ‘are: ini Ottawa, “Washington, -lowden.

REMARKS. (‘The few ‘specimens exhibit’ no noteworthy variation.

The. species may. be'vcontused either: with C. aurifex” om Cy ‘ardens

because of its shining cupreous or greenish dorsum but can be _

separated from either of these species by the following combination

of characters: non-tuberculate frons, elytral strial lines distinctly

punctate, fore tibial spur of male slender and’ apex of male

genitalia ridged (Fig. 158). The ridged apex of the genitalia’ is

similar’ to that of: tenebrosum but perceptibile - Has’ ‘the

elytral intervals shining rather than opaque.

86 Contribs wmer. vente asin Vols io. no. Sly 1981

12. Canthidium vespertinum mm. sp.

Figs. 159-161

HOLOTYPE. Male, length 4.6 mm, greatest width 3.0 mm.

Dorsally very dark: brownish black, head and. pronoium: with slight

cupreous tinge; prometal disc. and elyival-intervals«closely, finely

punctate, —~pumctures frequently separated by one. diameter.

Clypeus. (Fig. 159) very feebly bidentate, median indentation

extremely shallow, very obtusely V-shaped; margin laterally

on each side evenly arcuate to middle of gena. Chypeal

disc, irons amd vertex.in. @.¢ingle plane, mearly flat; .tubercles

or swellings lacking; surface finely punctate, more conspicuously

so on clypeus. Eye dorsally at posterior end of gena broad, 15

er more facets wide. .Pronotum Broadly convex, lateral. toveae

obsolete; posterior pronotal margin lacking line. of elongate punc-

tures Or associated transverse groove... biytra with. strial lines

fteebly impressed, dictinct|y punctate, the. punctures iscparated, by

three to tour diameters;.¢elyiral intervals: flat..«Pygidium: minutely

punctate, feebly alutaceous, between’ punctures. kore femur with

posterior surface lacking setigerous punctures; only scattered

minute punctures. present, surface shining -and polished. . Middle

and hind femora with ventral surface similar to posterior surface

of fore femur. Fore dibiaiwith tarsal spur «wnmodified, inner

apical amegle ef fore’ tibia anteriorly forming. acute tooth,» Genitalia

as in figs. §160,- 161,

ALLOTYPE. Female, length 4.6 mm, greatest width 2.9 mm.

Differing from male in the following major characters: dorsal

surfaces 6lightly more, shining and more distinctly punctate;

pygidium less distinctly punctate; inner apical edge of. fore tibia

not produced, essentially right-angled.

PYPE MATERIAL: Molotyoe, male, Panama, Chiviqui Prov., 2

km N Sta. Clara: 1300 m,.6C5iN.. 62°46 Ws Hartmann'’s:. Finca,

24-25.V.1977, H. & A. Howden (Howden). Allotype, female, same

data as holotype (Howden).

PARATYPES: 5 males, 4 females. PANAMA: 1 female, same

locality jas nolctypew 20-314) 19/75: Peck,.dung trap, human

feces... COSIA RICA. al female, Cartago, Prov., 600 m, Turrialba,

CAT dd 19.0. 1979,0 be & A.) Howden, feces. cup trap; 5. males,

2 females) Puntarenas, Prov., 9 1400 m, Monteverde, 25;..27.V.1979,

He & A. Howden sitting jon Teaveo near trail.at night, and.in

cup traps.

Paratypes ate im: Ottawa; Washington, Howden.

REMAKKS. the one Panamanian paratype. measures 5.0 mm in

length but, otherwise. does not vary significantly from the allotype.

Within the Panamanian fauna this species is most likely to be

confused with tenebrosum; however, the complete tack. of any

tubercles'.en the ead, the weeny. broadadorsal portion of the eye,

andvthe relatively small’ size is sutficient to distinguish the

species wilmin the ceographie. area. being considered. .Costa- Rican

specimens: show jsome variation im. the sculpture ,of the: pronotum

and elytral interwals, «bul otherwise do mnot.vary significantly.

A elated species occurs. in: Belize,

Howden. and Young: Panamanian Scarabaeinde 87

13. Canthidium: pallidealatum mn. sp.

Figs. 162-164

HOLOTYPE... Male, lensth. 4.2>mm, @reavest width) 2./ om:

Head, pronotum, elytral margins and sutural intervals dark

brownish black’ to black, remainder oL elytra ivory to tight tan;

strial lines irregularly spotted with gray; dorsal surfaces shining,

pronotum and elytral intervals minutely punctate. Clypeus (Fig.

162) feebly bidentate, median emargination shallow, broadly V-

shaped, margin: laterally arcuate, to middle of cena. Surface of

clypeus moderately transversely rugose. Frons with two vague

swellings, one: on; each. side, near,.imner, anterior, edge of eye, area

between swellings feebly concave. Eye dorsally at posterior

edge. of..cena strongly. narrowed, three to four Jacets »wides . Frono-

tum evenly .convex;, lateral fovea small, oval, shallow; posterior

pronotal,margin lacking adjacent. transverse groove or row of

punctures. Elytral striae represented by faint lines interrupted

by large shallow punctures, the shallowly impressed punctures

delimiting striae and separated by approximately two to three

diameters, strial punctures frequently somewhat darker in color

than. surrounding portions of elytra; intervals neariy: tlat, closely,

finely .punctate, ..Pyeidium [inely punciate, surtace between qnuinc—

tures faintly granular... Posterior: surface of fore femur with two

coarse, setose punctures near margin at apex; remainder of sur-

face with vague minute punctures or finely shagreened. Ventral

surfaces and middle and hind femora minutely punctate and slightly

shagreened. Fore tibial spur, not) e@reatly modified, slender, , some-

what shorter. than-in, female, inner apical angle: ot tibia adjacent

to spur, produced anteriorly and slightly bent downward forming

a broad, abruptly rounded Jobe..approximately equal im lenoin to

tibial spur,.,.,Genitalia .as im, fties..103, 164.

ALLOTYPE. Female, lengui' 4.2.1, oveatest width 2.6 mm,

different; from holotype, in the. following major characters: clypeal

margin laterally. feebly, arcuate, clypeal suriace more strongly

rugosé; punctures in elytral striae cecasionally separated Dy tour

to. five, diameters.” frequently net conspicucusiy darker than

surrounding. surface... Fore tibial spur, one Unird. jonger.. than. in

male, inner apical angle of (iia 201, oroduced,

TYPE MATERIAL. Hotopioe, male, oranda.) Civiqui. brow, «

Las .Lagunas, 5 km. SW. Mate “del Volcan, 1360 m, 22-20.V.19/7:. 5.

Peck, dung trap (Howden). Allotype, female, same locality as

holotype, 22.V.197/, S..& ). Peck,. litter Berlese No. 371 (Howden).

PARATYPES:. 8 males, 2 females. PANAMA: 2. males, same

data as holotype; 6 males, 2 females, Chiriqui Prov., Las Lagunas,

Av kim W Hato del: Velean.. 1200-1360. mm 27 Nie 1077 eA

Howden, cup traps.

Paratypes are in: Ottawa, Washington, Howden.

REMARKS.).Variation im the type 7series is, slight, size raneing

from 4,0 .10,.5.0. mm. 7 Coler iol. ine elivira shows slight warlation

but this..seems primarily due to method. of preservation. ,.[hitially

the .elytral disc) was primarily ivory in color,, darkening after

death, vin. other: respects the type series is very uniiorm. [Lhe

species can .be readily. confused with. leuconterum. which also has

88 Contrib Amer. abn telnet. VolelOu vino ely hos!

mostly ivory colored. elytra and istatyleacgt partially sympatric.

Canthidium leucopterum Can be distinguished from the above

Species by the tollowime combination Or Characters: slightly smaller

average size, distinctly bidentate clypéus and a trituberculate

froms. Other distinguisnine ‘characters are ‘mentioned in the key.

The (Majority of eGpecimiens were taken in) traps baited with

human: Teces.

i4., Canthidtum vartolosum.n.\, Sp.

Figs. 165-167

HOLOTYPE Male. lenoth 200mm, ereatest width 2./° mm.

Dorsally very dark brown to black, humeral area light reddish

brown: promotum at elyital intervals minutely punctate, some areas

finely shagreened. Clypeus (Fig. 165) quadridentate, the inner

two teeth strongly developed, median emargination V-shaped;

Outer Teeth on veach Side \fecply: developed, obtusely rounded,

approximately one-quarter the length of adjacent inner teeth.

bateral maroin of clypeus slightly sinuate to small’ notch at

junction with Gena; ‘clypeal curtace shallowly closely punctate.

rons tridentarce, (ad three ‘conical protuberances feebly developed,

widely separated, wedian one distinctly anterior in’ position; .sur—

face between with scattered tine punctures and “granulate. Eye

dersally near the posterior end iof Gena narrow, two to three

faceto(wide.|) Pronotum evenly convex, lateral fovea on each side

Small) circular, Telatine y deep. posterior pronotal margin lacking

associated Groove. or punctures. “Elytral’ striae represented by

faintly impressed lines, rarely small punctures visible in strial

line but striae largely appearing impunctate; intervals largely

flat, minutely punctate, Shining.” Pyoidium smooth to minutely

PUNCtate, | shining. \|.Posterion or ventral surfaces ofall. femora

Smoot to minutely puriclate, Tinely»Siaoreened, “ Fore tibia with

apical spur not ebviously modified, Slightly shorter than in

female; vadjyacent immer sapical portion of tibia produced into

acutely "rounded lobe. slightly imflexed near tibia, the lobe

approximately Gquah iy lengen te: tibial’ spur.” ‘Genitalia as. in

figs. 160) 16).

RULOTYE EE. Pemale, leneth aol mm, oreatest width 2.5 mm.

Dittering, from male in the following major characters: clypeus

transversely rugose; pronotal fovea less deeply impressed, prono-

tum Slightly more obviously punctate; inner apex of fore tibia

nol ‘produced into an acutely anoled lobe. .

Pe a eR Toltetyipe, male, Panama, Chiriqui Prov...

bas Laconas, (360 mo 4 kmoW “Hato del. Volcan, 12Vl.1977, H. & A.

Howden, cup trap baited with human feces (Howden). Allotype,

female, same data as holotype (Howden).

PARATYPES: 7 males, 9 females. PANAMA: 3 males, 8 females,

Same TOCa lity as Nolonype, 22-27. Velo (oS. at. Peck, H. & A.

Howcens\) Watea I female Chintaur Prov.) (2 skim N Sta. Clara,

1000 OCR, See Ro WO artnann’ Ss Finca, 25.) GO-31.Vi1977, H.

SOR ower Numan feces. Cun trap, o, a ly Peck, litter ber:

Poy i, ber. #ay2 une@er bark: J male, Chiriqui Prov., 4900 ft., W of

Finga, Palo Santo, mre Neva California, O.111.1959, H.S. Dybas,

Howden and Young: Panamanian Scarabaeinae 89

Deriesen ravine: litter.

Paratypes are in: Chicago, Ottawa, Wastiinoron, Hower.

REMARKS. The (restricted ‘range, ‘small size amd quadridentate

clypeus owill usually: distinguish the species. (Size varies: from 3.4

to 4.6 mmand elytralvcolor varies trom (black to mostly savory.

15. Canthidium leucopterum n. sp.

Figs. 168-170

HOLOTYPE. Male, length 3.7 mm, creates width 7.0 ma.

Head, pronotum, and ‘pygidium “(dark Drown tor Bbiacky elytra ‘ivory

(darkening slightly when preserved), sutures and margins narrowly,

evenly brown or black. Clypeus (Fig. 168) bidentate, median

emargination narrowly U-shaped; margins on either side slightly

irregularly arcuate to middle of gena; clypeal surface feebly

retlexed in anterior half, “reflexed: portion finely, transversely

rugose. “Frontal “area ‘tritubéerculate, (median ty bercie anterior) 1

and more distinctly conical than either lateral tubercle; surface

near and between tubercles finely granular with small scattered

punciures.: Lye dorsally ial posterior end “ol saena wer marrow.

two-to four ‘facets wide. °. Pronotum, with lateral tovea appearing

as large »puncture, dise evenly convex mingtely punetate,, shining,

posterior margin lacking adjacent anterior row of punctures or

indentation. Elytral striae indicated as faint lines, strial: punc-—

tures obsolete; intervals nearly flat, surface slightly irregular.

Pygidial surface shining with small to moderate, relatively incon-

spicuous punctures.) Posterior surface, O17 fore: iemur tacring ‘Coarse

punctures, surface finely granular, with some scattered, minute

punctures; middle and hind femora ventrally minutely punctate

or-Tinely’ shagreened: or both.) Pore tibia with «spur apically

acute, unmodified; inner apex of tibia moderately produced, apex

acutely angled. “Gemitalia as) im: tos. hoo, 1170:

ALLOTYPE. Pemalée, Jengin 2,0 mm? orediest: widthe?. 3 mm.

Differing from holotype in the following major characters: clypeus

nearly flat, more stronely rugose? head and) promotum slightly

more densely and distinctly punctate; inner apical angle of fore

tibia not produced anteriorly, not acutely angled.

TYPE “MATERTAL: “Holotyoe, male, Panama, Chivriqui’ Prov. ,

(2\ km W)s Cerro: Punta, 1500) my 230Vl19779 "RL Howden,? cup traps

(baited with 2-3 cc human feces) (Howden). Allotype, female,

same data’ as’ holotype.

PARATYPES: 6. males,.7 females “PANAMAr 4 ?male; 2 females,

same data as holotype;, )+matle, 1 female, Chiniqui Prov., 2 km W

Cerro Punta, 6°51 N/020 360W. 225N 1977, (0. Howden,’ /cup traps; 1

female, same locality: as: holotype, 199.1977, Sie |... Peck,

Berlese sample No. 366; 5 males, 2 females, Chiriqui Prov., 4 km

W: Hato’ del Volean,) 1200-1660 mm (272 V il ONT LOFT Ue RA Howden:

cup traps; l omale, (Chiriquis Proy..,(Uas’ Lagunas, fh’ km SW: Hato

Gel Voleans 2220 1077) vos icePecky "latter berlese,) No. 471%

COSTA’ RICA: («1 female, Mélleza, 1300 m, ‘(near Candelas, Panama),

LaVoie Bek Howden,

Paratypes are in: Ottawa, Washington, Howden.

90 Contriy. Amer. “Enterinst>, wool. 1S) mo. 1371961

REMARKS. Canthidium leucoplerum may be confused with

variolosum and pallidoalatum, which frequently also have the

elytra mostly ivory im color. Une elytral margins evenly ‘and

narrowly marked with brown or black and the trituberculate frons

will distinguish teucopterum..> [hese markings do not show” signi-

ficant variation; size ranges from 3.0 to 4.2. mm,

16. Canthidium ardens Gates

Figs. 171-173

Canthidium ardens. Bates, 1887, p. 4&8.

Cotypes, tondon and Paris.:« London, specimens examined.

Length 3.7 to 5.6 mm. Dorsally brownish black to black

with distinct cupreous cast; head»and pronotum frequently with

@reenish buster im.certain. lights.. Promotal disc and-elytral

intervals appearing smooth and polished at low magnification.

Clypeus (Fig. 171) feebly bidentate, median emargination broadly

V-shaped; clypeal margin laterally arcuate to gena, there slightly

tO moderately notched; suriace of clypeus and) gena: moderately,

closely punctate. .Frens triftuberculate, tubercles. feebly developed,

median one approximately twice as. large as lateral ones and

anterior if position, «Surface of trons and vertex, moderately to

finely punctate. Eye. dorsally at posterior end. of gena narrow,

two €6.four facets wide. .Prenetum evenly convex except. for smal!

oval lateral: fovea on each, side, posterior: margin lacking adjacent

Sroove Or punctures.: -Elytra! siriae either lacking. or. indicated

by faint, offen obsolete lines, intervals when distinguishable

usually. smooth and flatjoshiminge. Pygidium minutely punctate,

shining. «Posterior suritace of fore femur with s.one or two coarse

Setose punctures im apical third; remainder impunctate,. finely

sShagneened; middle and Hind temora with ventral surface mostly

impunctate, finely shaoreened..- Fore tibial, spur.of .male.similar

tO thak of female. bul mene Dlunily rounded. Immer apical margin

ol. fore tibia. ofimale produced inte: distinct rounded anterior

projection approximately equal to hall the, lenoéth of adjacent spur:

Male; @enitatiavas, in) figs, (/2, 21/3:

MATERIAL EXAMINED. PANAMA, 25 specimens from: CANAL

ZONE = Barro Colorado. island: -[vinidad River... CHIRIOUL PROV. -

near Boquete on Rio Caldera, 6000 ft.; 2 km N Santa Clara,

Hartmann S$: Finca: Santa, Glara: Volcan de: Chiriqui, 2500-4000: ft.,

Champion (cotypes).. COCLE PROV. = EL Valle, utributany:of: Rio

Amtons) Hil Vale, tributary of Kio Guaybo.. COLLECTION DATES:

January) February; May, june; July, August, November.

OTHER COUNTRIES, G1 Specimens trom: COSTA RICA;

REMARKS. [he species varies considerably in«size but can

usually be Precoonized’ by -tSishining cupreous or greenish: dorsum;

feebly to distinctly triiuberculate frons,.<non-punctate elytral

striae, and by the oebsoletely punctate anterior midline of the

metasternum. «While: specimens that: we are calling ardens-vary

considerably. size and color, ‘a sertes Irom ~Turrvalba, ‘Costa

Rica, at 600 m, average 5 mm in length while a series. from

Monteverde, Cocta chica,..at 1400: ms average only, 4 mm. «in: the

Howden and Young: Panamanian Scarabaeinae gill

two seviesithere i¢ no overlap im size; the larger form. is more

cupreous;: ‘the: smatl ome greener: Panamanian specimens Mave mot

shown these consistent differences. At present we have no evidence

that:two or more. “species are: included under ardens, bul there as

thespossibility that’ we are confusing. two species under ome name.

The species (one or two) has been collected from fermented

palm: fruits; -rat-cdrrvon,:horse manmiure; and oman feces.

Canthidium ardens is diurnally active and has been most: frequently

collected am: the wel season,

The species is found in moist forests at both low and middle

elevations. |

17. Canthidium aurifex Bates

Figs. 174-178

Canthidium aurifex Bates, 1887, p. 48.

Cotypes, London and Paris. . London specimens examined.

bencth 3:33:00 5.5 mm. Dorsally. black with @istinct creenish

Or cupreous.reflections; dorsal surfaces shining, prenctal aise at

elVytral. intervals. with numerous extremely minute punctures.

Clypeus (Figs. 174, 175) very feebly bidentate, median emargina-

tion shallowly, broadly V-shaped.-..Clypeal margin laterally evenly

arcuate to gena, there. margin: feebly oliset, then evenly arcuate

to posterior of gena. Clypeal surface transversely rugose or

closely contiguously punctate; g@ena closely punctate; -fronms, tri-

tuberculate, each tubercle low, conical, the median tubercle

distinctly anterior. in: position>.«surface of vertex closely; moder

ately —punctate; less so posterioniys,, Hye dorsally at posterior end

of. gena marrow, three-to. five facets wide, Pronotum evenly convex

except. for. small-circular: lateral fovea: on each side;/ area vadjacent

te posterior: margin not. punctate, not depressed. <Elytral striae

consisting -of distinct: indented limes interrupted by fairly evenly

spaced. coarse punctures; intervals nearly flat,: shining, minutely

punctate. © Pygidium distinctly punctate, marticutarly in basal

halt, there, separated. by “a distance: approximately “equal to one:

diameter. . Posterior surface of fore femur..with scattered minute

punctures, the surface between finely shagreenead;, middle and hind

femora ventrally with scattered minute punctures, surface between

smooth to finely shagreened. Fore tibial spur of male (Fig. 176)

flattened and enlarged in apical half, the apex emarginate,

broadly. V-shaped; immer; edge of. tibia at base of spur slightly

produced anteriorly, forming an.acute angle. Male genitaliavas

in figs, 17751408.

MATERIAL EXAMINED. PANAMA, 91 specimens from: CANAL

ZONE —- Balboa; Barro Colorado Island; Canoa Saddle; Gatun Lake;

Fort Sherman; 5 mi. NW Gamboa; Paraiso; Pipeline Road; Rio

Trinidad. ), COLON PROV. =.Portebelo... PANAMA PROV. - Cerro

Campana, 850m. COLLECTION DATES: January, Feoruary ; March,

April, May, June, July, September, November, December.

OTHER COUNTRIES, 36. specimens from: COSTA RICA, ECUADOR.

REMARKS. Canthidium aurifex exhibits a great variation in

size and to a ‘lesser degree also varies considerably in color.

92 Contrib. Amer. nti inet.: voli wl6,) mo. 6b .i981

The sizecrange ‘given on the déscriplion: occurred in specimens: from

Barro Colorado Island. However, in some localities the size seems

to. ‘be fairly consistent. Specimens taken near Turriatba: (900 m),

Costa Rica, averaged about 4 mm in length while specimens from

Monteverde (1500 m), Costa Rica, averaged about 5 mm. Specimens

from Rio Palenque (200 m), Ecuador, varied between 4 and 5 mm

in length. (Color ranges: from black to ‘black with strong green

OY Cupreous mellections) the. color often uniform -incone ‘locality,

with Panamanian specimens often distinguishable from those from

Ecuador.

Panamanian specimens of aurifex have been collected from

forest floor leaf litter; leat surfaces; and at tapir, howler

monkey, and human feces.

On Barro Colorado Island this species. is active throughout

the daylight hours and is most abundant during the wet season.

Teneral individuals occur only in the early-mid wet season, with

dry season specimens usually damaged or well worn. A number

of specimens were collected along forest trails sitting on leaves

14 to 25 cm abeve the ground... The»species is primarily found in

lowland moist forests.

18. Canthidium hespenheidei n. sp.

Figs. 179-182

HOLOTYPE...Male, length: 3.6. mm, .ofeatest: width 2.3 mm.

Dorsally dark brown with strong cupreous reflections, shining;

pronotum and elytral intervals with scattered minute punctures,

those on elytra often vague. Clypeus (Fig. 179) bidentate, median

emargination U-shaped; margin laterally broadly, feebly arcuate

posteriorly to junction with gena; clypeal surface moderately

punctate, ‘the punctures: often contiguous. . Entire: frons: slightly

elevated above plane of clypeus, neariy flat, no tubercles “present:

Eye dorsally.at posterior end of cena narrow, two to. three) facets

wide. Pronotum evenly, moderately convex, lateral foveae

obsolete; pronotum lacking line of punctures adjacent to posterior

margin (Fig. 180). Elytral striae represented by feebly impressed

lines; the lines: impunctate or with small, poorly defined, .incon-

Spicuous punctures; punctures if present frequently separated by

10 or imore diameters. Elytral intervals nearly flat, with scattered,

minute punctures; most of surface smooth and polished. Pygidium

shining, polished, with scattered minute punctures. Posterior

surface of fore femur with one or two setose punctures near apex,

remainder of surface smooth and shining. Ventral surfaces of

middle and hind femora smooth, shining. Fore tibial spur slender,

apex ‘acute; Slightly shorter’ than: in female; inner ‘apical angle of

tibia strongly produced, abruptly bent downward, with apex

broadly oacute,. the projection. equal in: length to tibial.)spur;

tibial apex lackime distinct tuft! of setae, -Anterior.third of

metasternum between middle coxae distinctly punctate, the size. and/

or number of punctures not noticeably reduced medially (this

character separates the species from other small, cupreous species,

all of which have the metasternum impunctate or feebly punctate

Howden and Young: Panamanian Scarabaeinae oO

in anterior median third). Genitalia as in figs. 181, 182,

ALLOTYPE. Female, length 3.5 mm, greatest width 2.2 mm.

Differing from holotype in the following major characters: clypeal

disc transversely rugose; pronotum and elytral intervals with

minute punctures obsolete; inner apical angle of fore tibia not

produced; (tibial spur ‘longer.

TYPE MATERIAL: Holotype, ‘male, Panama, Panama: Prov.

Cerro iCampana, ) 850m, 8040"N,.79°56' W,'23.1,19/0,) Fae Mespenberde

(Howden). Allotype, female, same locality as holotype, 800 m,

30.V¥.1970,° HeP. Stockwell (Howden).

PARATYPES: 6 males, 24 females. PANAMA: 2 females, Canal

Zone, Barro’ Colorado’ Island; V11. 1969, /20.N 11519076, Nike Woodley,

]. Lawrence, Bo.) & Ta Hlavacs) 2) Temates, ‘Canal Zones mis NW

Gamboa, 9°09'N,: 79°43'W; 28.X1.1969,:H.A. Hespenheide;. 1 female,

Canal Zone, Madden: Forest,’ miley 2.5, 90057"). 79°37 We 24.17.1970,

H.A.: Hespenheide; 1 male; 2. females, Panama’ Prov., same locality

as holotype, 12: V1974) “YQiNG1973 2 30.N TOO Pe Stockwell; 1

male; PanamasiC.2.4, Camp Chaoreées,'\:26-27.X.1967,. 7. Vaylor;. 1

male, sPananiay Provi; Cerro Azils car 2000 fo 21176

Newton, in wet debris small forest stream. ECUADOR: 3 males,

15. females; Pichincha Prov., -4/ekmS Stow Domingo, 700: 11... Kio

Palenque, 22427 e1l1l.19/6).Ho-& Aor Howden, WV. (Nealia; (1. temale,

Piehincha Prov., 30kino Ee: Tandapin 40001. ey. V0 7541 Ga 8

Peck, wet’ ravine; 1 female, Pichinea Prov.) ima landiay 650" my,

15-20. NEeO75;) Sock leeches

Paratypes are in: Cambridge, Chicago, Ottawa, Washington,

Howden.

REMARKS. Canthidium hespenheidei is seemingly the smallest

species in the genus occurring in Panama, with the size ranging

from 2.6). to: 4.1 mae ly can occasionally: be. confused with small

specimens of C. ardens that lack distinc? tubercles, on the frons,

In these cases, the distinctly punctate anterior median third of

the metasternum and the shape of the male genitalia will distinguish

hespenheidei from ardens. The small size of hespenheidei, in

conjunction with the mearly flat trons, the errial lines On the

elytra lacking plinctures and the modification of the male tore

tibia will separate the species from other Canthidium, Fanamanian

specimens have strong coppery reflections while specimens from

Ecuador have strong @reen reflections and average Slightly larger

in. olze.

The species is named in’ honor of Dr. Henry Hespenheide in

recognition of the many interesting species of Panamanian

Scarabaeinae that he has collected’ and made available to us.

This ‘species’ has) most frequently) Deen taken Sitting’ on

leaves 10 to 15 cm above the @round, (nA few specimens were

collected at. human: feces,

Onthophagus Latreille

Onthophagus Latreille, 1802, p. 141; Bates, 1887, p. 66.

Type-species: Scarabaeus taurus Schreber, by monotypy.

The genus Onthophagus, with over 1500 species described, is

one of the largest senera of organisms. Even in Panama, where

94 Contrib. (Amer. “Entice... volocic. no.

1951

only 19 Species are. known (to o¢cur, There is a considerable

_ diversity.in the morphclogy. The genus, Decause of its large

size and diversity, is difficult to define morphologically without a

relatively lengthy description.

In Panama, species in the genus Onthophagus are small to

moderate im size, 3 to 13 mm im leneth, oblons oval.° with a

short thick compact body, convex above and: below, with middle

te@¢s Tat apart and hind leos far pack. “Males uswally have. the

head horned or tuberculate and (or) the pronotum with horns or

other protuberances; females have the neacd Carinate and sometimes

the pronotum with a carina orvother elevation. The mentum is

hairy, transverse, Guadrangular, emarginate; labial palpi hairy,

3 seginented, ficsi segmene slhigatly shorter than second,. third

very small and inconspicuous. . Labrum membranous, densely

€lligte,.comcealed by clypeus. . Mandible elongate, rounded,

Without (tect. membranous toward: apex, and ciliated on internal

edge. Maxilae termimate.in large membranous ciliated lobes;

Dalpl 4 secmented., first seament short and slender, second and

third short and thickened, fourth fusiform and truncate at the

end. .Ciypeus relatively smooth «in male, carinate in. female,

Variable in shape. Amtenna. 9 sésmented, with 3-ségmented club.

scutellaym: not vistble, «Elytrom short, somewhat. narrowed. and

rounded posteriorly) «with. Seven striae. «Pore tibia

quadridentate,

oftew Ssiender and selongate in wale; fore, tarsi present in Both

sexes; middle and hind, tibiae apically enlarged and: truncate:

tarsi Slender, ciliated on inner ecdae: first seament elongate. and

Subparallel: claws small, Metasternum long and parallel sided,

separated from mesosternum by rectilinear groove:

Pygidium

exposed. Males with last abdominal segment broadly emarginate

at middle: females with lasi> segment not constricted at. middle.

Key to species of Onthophagus

ee Pronotal surface with at least some scattered,

erect Selace, tie setae obvious im fresh. speci—

mens at 10x, loacer (han.one. puncture dia—

Meters ft sciae sileri, then clyira bicolored.. . . 2

Pronotal surface lacking setae longer than dia-

meter Of Duncture; elytra usually. but not

always uniform in color.

Dorsally brown to black or. Dbicolored;. pronotal

Suriace dull if uniformly: black, otherwise

Frequently chigiie: ayersee size under O mm. . 3

Dorsally uniformly Diack, often with green .or

Dlue. vellections;: pronotal surface. bétween

setose punctures smooth and shining; average

size’ 6 $0 9 Min = Je. © Gs eninitus panamensis .Bates

Pronetal punctures appearing annulate: rim

abrupt, depressed portion of punctures shallow,

that; color dorsally uniform .

Howden and Young: Panamanian Scarabaeinae 95

Pronotal punctures simply impressed, rim rounded;

dorsally» metthed or: bicoléred:. eign u 2 eiarae >

Pronotal punctures not contiguous, most separated

by distance equal: to,one-hall. diameter, om more;

size moderate, 5 to & mm 2.) 14. -almosericeus =. Bouc.

Pronotal punctures mostly contiguous; size small,

3.5°40 5.0: mm. ore i ae ee eee cincue Bares

Pronotum anteriorly with-bread protuberance,

either: with emalb: lateral tubercle on each side

or prominence feeble and, rounded; ei ypeus

anteriorly rounded, sometimes briefly truncate

but not shallowly emarginate; pronotum fre-

quently with sides broadly marked with brown,

longitudinal central third or more dark. brown

to black, sometimes with cupreous reflections .. . 6

Pronotum anteriorly with small V-shaped median

prominence often overhanging anterior marcin,

or prominence: very small ):rounded or lacking,

clypeus anteriorly shallowly emarginate on

truncate; pronotum usually mostly or entirely

black, frequently with ereenish cast gers

~ Sey oe Ie ea eapes . 13. landolti Har. (complex)

Pronotum in: beth sexes with well, detined truncate

or broadly convex protuberance; elyita usually

with intervals alternately tan and dark brown

or: biaek ; isi size.5.5, mm oaty.

es . 8. marginicollis ae (in part)

anne ue oe tumid ee head, elytra

uniformly -brown or drrecularly mottled, mot

striped) .awerage size 51mm @ «57 oe debast Bouc.

Pronotum and-elytra distinctly different in

color or elytimea: mottled. Gr bigeioredy) a) exioscous. 8

Pronotum and elytra similar in color (sometimes

pronotum with more pronounced greenish cast

but basic color uniformly dark: brown or black) . 10

Elytra uniform in color or mottled, neverm with

intervals: alnernately ditferine (im color sh .eca. 9

Elytra with: intervals alternately tan.and dark

brownor cblaick . c.cn78e mayeinicollis:; Hari (in part)

Pygidial surface basally between punctures dull,

finely @ranular or shagreened; hind femur

ventrally witha short row of -3:or 4 coarse

punctures, otherwise surface: sincoth cr with a

few minute punctures; male. with erect. horn

near anner posterior edge of each lieve. ;

cc Mae te aneeed mans Mn. SDs

10;

he.

14.

Contribs (Amer. Bat: imet:) velois, me. 2b, 4951

Pygidial surface basally between punctures

shining, smooth; hind femur ventrally with

widely scattered moderate-sized punctures;

male with, abruptly reflexed, nearly truncate,

anterior clypeal margin, lacking horns on

Hes ee a Be a a Op yises:. Bates

Pronotum punctate @r mot, evenly, conwex /or

with broad tumosities or with paired horns,

plates or ‘tubercles, never with a single

anterior median conical: hor. or ee

tubereher. ies a i

Pronotum heavily panciate with single ‘anterior

median horn or: tubercle; in males: horn pro-

jyecting’ anteriorly over bhbead, Dbifurcate

apically; in females forming anvacuteverti-—

cal stubercle at antéevion: margin; ‘average .size

Coe a, ee a a ko eho mirabilis: Bates

Color dark brown to black, frequently with

greenish-blue metallic reflections; middle

and hind femora not differing greatly in

color from. verntrat suriaces) brown to: black ve: . 12

Color brown to dark brown (pronotum and elytra

heavily punctate); middle and hind femora

(and often fore femur) mostly yellowish;

average size 5.5 mm wo. Se 12. dicranius: Bates

Pronotum smocth or. punctate). af punctate, the

punctures eee of uniform size in any

one area «.". ee ee eat Lae 13

Pronotum distinctly punctate; punctures centrally

of two sizes; medium punctures separated by

one to two diameters with small to minute

punctures interspersed (dorsal surface of

pronotum and elytra dull; color uniformly

Dlackiiwith; bluish cast); average size: 9 mmilo...

Jip. Cyanellus Bates

Pronotum bituberculate or with paired ‘swellings

either side of midline; some species with acute

tubercles deeply separated at midline athe ee ae

Pronotum evenly convex or with median protuber-

ance evenly rounded, anteriorly not impressed

OF Cr iGea at erieeior Midline A ee ke LY

Setae at apex of hind tibia mostly stout, with

some fine setae between, mostly of similar

lenoth; “pronotal aeons narrowly

Civica Ws, Ra ES

welac al apex vot hind tibia “alternating between

stout and slender, the slender setae often

considerably longer; pronotal protuberances

16.

12.

19.

Howden and Young: Panamanian Scarabaeinae oF

separated medially by'a distance: equaliite

about half width of vertex; average size 8 mm... 16

Elytralb«<intervals: finely::punctatey suniace teendy

shining; ;dorsal..color black: wsuallys with strong

a or cupreous cast; average size 6 to 7 mm

i wyan. DGae praeceblens, Bates

Elytral intervals obsoletely punctured or

impunctate, surface; dull, minutely: eranular;

dorsal color dark brown: to black; occasionally

with bluish reflections; average size 7 to 8 mm

(Se:belorhinus«: Bates

Pygidial surface distinctly shagreened or granu-

lar, ‘at; most feebly shining: if Merns| on vertex.

clypeal+frontal carina. absent..or Highest

medially; if; horns, lacking) then carina) on

vertex, nearly straight, and ace lower

medially. ig. Cia it ee ea BC

Pyeidial surtace between ‘punctures “at most

feebly Shaigreened, strongly shining. 11) horns

on, vertex, clypeal-frontal carina migher on

either side of midline; if no horns on vertex,

then carina on vertex ogival (brace-like)

AN SMAPS: abit wana ews ae cl OUP ern RIAN eee Renee LDN i ea)

Pronotal surface shining, at least medially;

color dorsally frequently with metallic coppery,

bluish or .eareenish east) go cig is ce NL ie

Pronotal surface dull, opaque, particularly in

median posterior half; color dorsally black,

occasionally with greenish cast on) pronotum:

average size. 0) 10) 7% Mimi. ig cin hal De me One) hates

Hind femur relatively stout, width to length

about 3:7; clypeal margin medially rounded or

with wide reflexed: area or: with broad). rounded

accumina lions! via Bi LO

Hind» femur relatively slender, width to Lengths

about 3:9; clypeal margin medially with

distinct, strongly reflexed horn (often

expanded apically) or with margin with sharp

V-shaped emargination; average size 7 to 8 mm

j Lie osharpin Har:

Metasternum between middle coxae finely,

shatlowly | punmetate Near: Comaliimaral mei ie die dann 20

Metasternum between middle coxae coarsely,

distinctly punctate near coxal margins,

punctures separated by approximately one

to two diameters; average size 8 to 10 mm

25) lute alte! ie Gy

98 Contrib, Amer. Enticinst.) vol. 18, ne. 1) 1981

20. Color dorsally black with blue or blue-green

reflections; “metasternum anteriorly distinctly

prow-shaped; well-developed males with |

anterior clypeal margin broadly truncate or

shallowly emarginate; size frequently over

Caimi. oe a ineensus Say

Color dorsally very dark brown to black with

feeble metallic luster, particularly on

protetum;: Melasternum anteriorly only feebly

tumid medially; well-developed males with

anterior clypeal margin obtusely angulate

medially; size usually under 8 mm

. 6. acuminatus Har.

1. Onthophagus incensus Say

fig. 183

Onthophagus incensus Say, 1835, p. 173; Howden and’ Cartwright,

1903, pe 2k:

Type, Cambridee. Examined. .

Length 7.0 40 11.0 mmo. Deorsally black with bluish or

greenish reflections, particularly on head and pronotum; head and

pronotal disc of large males impunctate, smooth and shining; in

small males and females head and pronotum moderately, closely

punctate, elyiral intervals feebly punctate, stmfiace dull; pygidium

Shallowly, closely punctate, dull. Metasternum between middle

coxae with midline briefly but distinctly carinate;. on either side

of carinate area metasternum with scattered, shallow, poorly

defined setose punctures. Middle femur ventrally with coarse

punctures in apical half; hind. demur ventrally with: surface

largely smooth or finely punctate (usually more distinctly punctate

in females).

MALE. Major form (fie. 163): clypeus in apical third

broadly, trumeately reflexedy the. reflexed: portion often feebly,

broadly emareinaie; clypeal surface smooth. Lateral’ margin of

clypeus and gena slightly sinuate, gena posteriorly inwardly

arcuate. . Clypeal-frontal carina absent; carina: of wertex: repre=+

sented by long; eylindrical, inwardly arcudié horn ‘arising from

inner posterior edge of eye. ~Pronctum with anterior protuberance

rising just Denind abterior margin, the protuberance’ delimited on

either Side by 4. concave groove for the reception of the horn,

each groove Geebly, longitudinally arcuate, not stronely convergent

posteriorly.s> Lateral anterior surfaces of pronotum ‘shallowly,

closely punctate, median posterior areas frequently impunctate,

snining. Pore tibia only: slightly eloncate, lacking apicaltuft of

setae, similar in.fovm to female. Minors formesclypeus arcuate or

only feebly truncate, scarcely reflexed, surface closely punctate.

Clypeal-frontal carina indicated by transverse swelling, carina of

vertex a low transverse ridge, median portion slightly posterior

in position; horns absent. Pronotal protuberance and related

grooves “obsolete to absent.

Howden and Young: Panamanian Scarabaeinae 99

FEMALE. Clypeus shallowly arcuate, evenly reflexed, surface

transversely rugose. Clypeal-frontal carina complete, transverse,

highest medially. Carina of vertex nearly straight, slightly

elevated on each side at termination near eye. Pronotum with

protuberance low, delimited on each side by a small rounded

tubercle, pronotal surface moderately punctate, closely so laterally,

less densely so medially.

MATERIAL EXAMINED. PANAMA, 103 specimens from: CHIRIQUI

PROV. = Baldwin: Forest:. Bambitos Cerre: Punta; 11 Valle de Nubes,

4500.ft;.: Finca Leridas Hartmann 's: fincas Las Lacunas, £360 -m:

Rovira: (12. miles. NW): Santa Glara; Volcan de Ghiriqui.-=PANAMA

PROV. #cCerro: Campana... VERAGUAS PROVe> + Potrerilios. COLEEC-

TION DATES: January, > March, May,: jume,;: October, December.

OTHER COUNTRLES, 104 «specimens from; MEXICO, EE SALVADOR,

COSTA RICA; VENEZUELA, COLOMBIA, ECUADOR.

REMARKS. The species has been obtained from horse, human,

and tapir feces. In Panama it is restricted to moist forest areas

at middle elevations,

2. Onthophagus batesi Howden and Cartwright

Fig. 184

Onthophagus batesi Howden and Cartwright, 1963, p. 21.

Type, Washington. ~¢ Examined.

Léernieth 6.0 t0°9°5 mm.” Doresally:dark brown to black;

occasionally with slight greenish luster; impunctate to finely

punctate, punctures of elytral intervals frequently with minute,

very inconspicuous setae. Pronotum varying from smooth and

polished (large males) to finely, distinctly, punctate (small males

and females), surface between punctures smooth and shining.

Elytra with striae feebly impressed, each stria represented by a

double line interrupted by shallow punctures, usually separated

by approximately two diameters. Intervals flat with irregularly

scattered minute punctures, surface between shagreened or finely

granulate, féebly shining. »~Pyaidium closely punctate, punctures

frequently irregular, sometimes becoming confluent, forming area

of transverse rugosity, surface shagreened, setae lacking or

minute. Metasternum between coxae with numerous large, distinct,

setose punctures present in two-thirds of area between coxae;

median area raised, midline shallowly impressed; median area,

particularly posteriorly, lacking large punctures). with’ only

scattered minute punctures, surface mostly smooth and shining.

Hind femur with a few scattered coarse punctures concentrated in

apical anterior half of ventral surface, remainder of surface with

a few minute punctures, the surface smooth and shining.

MALE. Major form (Fig. 184): clypeus with anterior margin

abruptly reflexed, the area almost truncate or very shallowly,

broadly emarginate; posterior portion of clypeus and frons lacking

a carina’ or with caring vacuely indicated laterally, ‘surface

nearly! flat, finely, distinetly; punctates(’ Lateral margin: of iclypeus

im conjunction with: cena slightly sinuate;’ifeebly indented at

junction between clypeus and gena. Vertex with two long, widely

100 Contpibe Aner: (Snr inst. , volute, me.) \ 1961

separated, slightly divergent ‘vertical horns; the horns nearly

straight, ‘teebly transversely Tlatvened, with a very distinct. U=

shaped carina extending from the middle of the horn downward

and across to other horn. Pronotum posterior to horns broadly,

shallowly, concave. Pronotal protuberance between horns broadly

rounded anteriorly, not sharply delimited laterally, remainder of

pronotum evenly convex. Fore femur slightly elongated, lacking

distinct tuft of setae but with slightly upturned tooth above apical

spur. Minor form: clypeus only moderately reflexed anteriorly,

the reflexed portion frequently truncate, surface of clypeus more

heavily punctate, occasionally transversely rugose, clypeal—frontal

carina often evident as low transverse ridge, horns on vertex

greatly reduced, but with U-shaped carina still evident. Pronotal

protuberance occasionally reduced to feeble tumosity. Fore tibia

may be unmodified, similar tovthat of female.

FEMALE. Clypeus with anterior margin truncate or feebly

emarginate, weakly reflexed, clypeal-frontal carina low, of nearly

Uniform heieht; clypéal eurface transversely. rugose.: Carina’ on

vertex transverse, very slightly arcuate with lateral termination

slightly posterior, not clearly defined. Pronotum with weak median

protuberance, often with very slight lateral concavity on each

side,

MATERIAL EXAMINED. PANAMA, 3 specimens from: CANAL

ZONE = Ancon; Camp Chagres;: Coco Solo Hospital. COLLECTION

DATES: July, August, ‘October.

OLHER COUNTRIES, 108 .specimens from: MEXICO, BELIZE, EL

SALVADOR, NICARAGUAG*COSTAX RICA.

REMARKS. One Panama specimen was captured in a light

trap. One Costa Rican specimen was obtained from the stomach of

a bat, Nyetidromus aibicollis. “Other. specimens from Nicaragua

were found “under cow ‘dune.. The ‘species is frequently common in

ClUt=OVer areas,

3., Onthophagus crinitus panamensis Bates

Pies 165

Onthophagus crinitus var. panamensis Bates, 1887, p. 68.

Cotypes, London and Paris. London specimens examined.

Length 7.0 to 1O0.0\mm: (Dorsally black; frequently with

greenish or metallic reflections, particularly on head and prono-

tums dorsum punctate ssetose, setae érect, pale yellowish Oni tan:

pronotal punctures separated by less than one diameter, surface

between smooth and shining. Elytral intervals distinctly, irregu-

larly punctate, the punctures slightly smaller than on pronotum,

separated by about one diameter, each stria represented by feebly

impressed double line frequently interrupted by punctures.

Pygidium with punctures similar to those of pronotum, surface

between smooth and shining, most punctures with long erect setae.

Metasternum and ventral surfaces of all femora with scattered

coarse setose punctures, most punctures separated by two to three

diameters.

Howden and Young: Panamanian Scarabaeinae 101

MALE. Major: form (Fig... 185):oclypeus with (sides converging

to broadly truncate reflexed apex, apex often very shallowly

emarginate, lateral margin of clypeus and gena forming nearly

straight line. Dorsum of head moderately to coarsely punctate.

Frontal carina absent, carina of vertex represented on each side

by. long, upright, inwardly arcuate ;horn)arising from posterior

portion of vertex behind eye. Pronotum with median protuberance

rising vertically above anterior margin, the protrusion limited on

each side by broad, shallow groove; groove on each side directly

beyond horn on head and extending inwardly behind anterior

margin: of protrusion, remainder of ;pronotum evenly convex... Fore

tibia distinctly elongated with a tuft of yellowish setae at base

of apical spur... Mimor' form: clypéeus. similar to;temale except

slightly. more broadly truncate anteriorly. + Clypeal-frontal carina

present, feebly developed. Carina on vertex present medially,

lateral horns feebly indicated or totally lacking in very small

specimens. Pronotum with minute swelling on either side of mid-

linesindicating vestige of: pronotal protuberance, otherwise pronotum

evenly conve Fore tibras aimmodi tied.

FEMALE. Clypeus arcuate, often more broadly so anteriorly.

Surface closely punctate to transversely rugose, both clypeal-

frontal carina and carina of vertex present, indicated by low,

nearly. straight, or: feebly “arcuate ridges; Righest mecially.:: Prono-

tum with small tuberculate swelling on either side of midline

posterior to anterior: margin, pronotum otherwise evenly convex.

MATERIAL EXAMINED. PANAMA, 126 specimens from: CANAL

ZONE -— Ancon; Barro Colorado Island; Cabima; Canoa Saddles;

Fort Kobbe; France Field; Madden Forest; Paraiso; Pipeline Road.

COLON: PROV. = Alhajuelas «PANAMA PROV g-omrravjare: COLLECTION

DATES: Every month wofmthe year .

OTHER COUNTRIES: Nome,

REMARKS «The: use of a subspecies: name tor Panama: speci-

mens’ is seemingly» valid: The mominate species from Mexicoinas

smaller ~punctures on the pronotum, the ‘head is usually less punc—

tate: and in large: males the horns. of the wertexmare more: arcuate

with -apices: flattened.” Whtlevthe differences “are: quite) small, they

seem to: beconsistent: and) hence the, Panamanian Specimens. are

treated: as’ distinct,

Onthophagus c. panamensis has been obtained at rat carrion

and on horse, howler monkey, human, and tapir feces. On Barro

Colorado Island, panamensis is active from 0700 to 1900 hours

with no! significant peaks of abumdance) during; that period. It.is

most abundant during the dry season and early wet season and

has only omee been recorded’ from carrion. iin: Panama the species

is seemingly restricted’ to) lowland: moist Torests.

4. Onthophagus stockwelli n. sp.

Figs. 186-189

HOLOTYPE. Male (major), length 9.0 mm, greatest width 4.5

mm. Color dorsally black with faint brownish-purple tinge.

Antennal club tanto reddish tan... Dorsal: surface shining, ‘setae

102 Contrib. Aimer. “Ent inst,, voli o6, mo.) 1961

mostly absent, a few very minute séelae. present on elytral intervals.

Clypeus (Fig. 186) with margin laterally arcuate and feebly

reflexed; “anteriorly more strongly reflexed, slightly thickened,

faintly emarginate, almost truncate; clypeal surtace closely —punc-—

tate; clypeal-frontal’ carina présent, low, “most distinctly elevated

on either side midway between midline and inner margin of .gena,

carina in profile bisinuate (almost biangulate). Gena not expanded

beyond line of clypeal: margin, junction with clypeus slightly

indented; cena and trons closely punctate. «Vertex: with: low: trans—

Verse Carina terminating om either cide meaqr inher edge “of eye in

vertical, nearly straight, cylindrical Norm; the two thorns very

siiohtly: diverging; height of shorns: approximately 1: mm}; vertex

behind smooth. Pronotum (Fig. 186) with anterior median pronotal

protrusion feebly developed, the pronotum behind head rising

nearly vertically to a: omail rounded tubercle on either side of

midline, protrusion thus bituberculate; remainder’ of pronotum

evenly convex; surface just posterior to-1ubercles: and near lateral

mareins finely; closely punctate, central posterior half of pronotum

impUnctate. Elyiral striae inipréessed; each “stria represented by

an impressed single line interrupted by punctures separated by a

distance equal to 3 to § diameters of one puncture; elytral intervals

feebly convex, with scattered, distinct small punctures, some with

minute setae; surface between punctures very smooth; shining.

Punctate area of pygidium enclosed by marginal bead, three-fifths

as long as wide, punctures separated by approximately one dia-

meter ,. sUtlace: Delween very Tinely Shagreened, shining. ~Fore

tibia (Fig. 187) unmodified, similar to female, lacking apical

tuft of setae and not clongated. Metasternum between’ middle

coxae. Obsolelelycarinate anteriorly, mear coxae with «a few

scattered lange punctures, otherwise metasternum centrally at most

minutely punctate, largely smooth and shining. Middle and: hind

femora ventrally each with a row of four to six coarse punctures

along median axis in apical third, mumerous coarse: punctures

present Near anterior mavoin, remainder of ventral surface minutely

punctate, mostly smooth.

ALLOTYPE. Female, length 8.2 mm, greatest: width, 4.3 am.

Differing from. mate in the following ¢haracters:: clypeal margin

(Fig. 188) very briefly truncate anteriorly, otherwise arcuate;

Margin eradually becoming more stroncly reflexed anteriorly, ‘much

less so than in‘ males clypeal—frontal carina ‘low, transversely

arcuate, highest medially;:: carina of vertex (Fig. 109): broadly

Ogival im Shape, horns absent; pronotum nearly as in :male but

with tubercles of protrusion more pronounced.

TYPROMATE RAL: Holotype. “male, Panama, Colon Prov., 270

m; 10 mis SE Colon; Ganta Rita Ridge, dO-127V1.1977 30S. Peck,

carrion trap, day 2 (Howden). Allotype, female, Canal Zone,

Barro Colorade Tsland, 25.7.19/5,, 0.7; Young, tapir feces. trap

(Howden).

PARATYPES- 107 males, 125 females: ~ PANAMA: 26 males, 27

females; same locality as allotype, taken on various days ‘in

January, February.) May, june, September, October, November,

Decembery.(by Laj. Bottimer,> P. ji Darlington, T.L.. Poulson, and 0:

Young, some: taken at Ateles: feces, .coati feces; howler monkey

Howden and Young: Panamanian Scarabaeinae 103

feces, human: feces, and tapir feces; 1 female, Canal Zone, Barro

Colorado ‘Island; 30. Viel9/65 E.Gu Rideyy i" female, “Panama Prov.’,

Cerro Azul, 24:¥ 19755 Ou Young; tapir! fecestivap. 1 ECUADOR. 74

males,.92” females;: Pichincha): 47: kim S Sto. Domingo, Ric: Palenque

Stave, 24225: 1b21976; Howdens* Nealisy (Peck, 49225 -V29]/5, 5. Peck,

26-31: V1I1 291976; °S. Peck;: all: numan’ dung traps, 17-25: FP51979... 7

Ling; /SviMarshally<) males 5 females: Esmeraida; 1) km SiaiSan

‘“berenzo,. (5 rm ps va oC ha quite: Stas; Galle LeVO7Sy Si heck .. dune trap

in forests: 3:males;"2-females, Manabi,: 7/3 km WE Chone,- 300 -m, -90

km (WStos Domingo; -6+9.V1.1976, S. 1 Peck, 1dune iraps.

Paratypes are in: Cambridge, Chicago, Ottawa, Washington,

Howden, Poulson, Ratcliffe, Riley, Stockwell.

REMARKS. Variation is average for the genus. Length ranges

from 5.7 to 8.7omm. «2Dorsal. color varies: from dark brown to

black with most specimens having slight blue or purple reflections;

a few however, have a distinctly green cast. In minor males the

pronotal protrusion may be greatly reduced, the tubercle on each

side poorly defined; likewise the horn on either side of the vertex

may be reduced to a small conical tubercle and in very small

males the carinae of the head may be similar to females. In

small females the two tubercles on the pronotum may be greatly

reduced. There is some variation in the density and size of the

pronotal punctures, but the species does not show the degree of

variation found in some of the more variable species of Onthophagus.

The closest relative of stockwelli is an undescribed species

from Oaxaca, Mexico, that has similar carinae and pronotal

characters, but differs in several small characters and is readily

separable by its grayish-black antennal club. Among species

sympatric with stockwelli, small females of acuminatus are the

most likely to be confused with it, but the ogival shape of the

carina’ion the vertex: of stockwelli' 4s: very distinctive, . Ihe

bituberculate pronotal protrusion of stockwelli is also usually

sufficient to: separate’ it from sympatric species. -lhe species. is

seemingly a wet-lowland forest form and is attracted to a variety

of different kinds of animal feces. On Barro Colorado Island,

stockwelli is mainly active from 700 to 1100 hours and 1400 to 1700

hours.

We are pleased to name this distinctive Onthophagus in

honor of Dr. Henry Stockwell who. assisted: both of us on.numerous

occasions, and, ,.whe. has,.collected. many-cl the.Scarapaecinae species

occurring in, Panama,

5. Onthophagus nyctopus Bates

big. oO

Onthophagus nyctopus Bates, 1887, p. 68.

Cotypes, Londen and Paris. London specimens examined.

beneth s,0 te° 7.2 mm. Dorgally Black, ‘dise of oronotum

opaque, imparting grayish sheen: .anterior face of pronotal protuber—

ance of some males greenish; dorsal setae absent... Head, »pronotum

and elytral intervals with-small, shallow, sometimes shining

punctures; punctures on pronotum Separated by one to three’: dia-

meters; punctures on elytral’ intervals trequently obsolete, ‘particu—

larly near suture... Elytral striae scarcely impressed, represented

104 ContribocAmer, Ente sls, wolsets sno. ol, 0981

by distinct double. line, «the lines: feebly interrupted by daint: ~

punctures separated by three to four diameters; intervals flat,

surface distinctly shagreened or finely granular, dull. Pygidium

closely: punctate, (punctures mearly contiguous.) surtace wetween

shining; setae very inconspicuous or absent. Metasternum between

middle coxae with scattered setose punctures near anterior lateral

edge, median portion smooth to very minutely punctate; elevation

of anterior midline rounded, not carinate. Middle and hind femora

with scattered, coarse, setose punctures on outer half or third of

ventral surface, remainder of surface smooth or minutely punctate.

MALE. Major form. (Pig. (290): combined clypeal and. genal

Marein evenly arcuate, narrowly retlexed, more strongly so

anteriorly; clypeal surface finely punctate, punctures separated

by approximately one diameter, surface between smooth and shining.

Clypeal-frontal carina very low but distinctly indicated. Frons,

anterior portion of vertex: and gena less closely punctate than

clypeus. Carina of vertex represented by two nearly straight,

cylindrical, somewhat posteriorly inclined horns on either side

near inner posterior margin of: eye; each horn connected: with. its

opposite by feeble ridge, inner edge of horns and connecting ridge

being broadly U-shaped. Pronotum posterior to horns longitudinally

concave, the pronotal protuberance between narrowly rounded

anteriorly; ‘remainder ‘of prenotum evenly convex. Fore tibia

unmodified, not elongated, lacking apical tuft of setae. Minor

form: head not differing greatly from major form except horns

reduced to small conical tubercles. Pronotal protuberance similarly

reduced, partioulariy: Weight of anterior face; concavity behind

reduced ‘horns virequently very small.

FEMALE. Differing from male in the following major characters:

clypeal margin less reflexed, carina of vertex low, arcuately

rounded on either side of depressed midline, the carina slightly

angulate on either side near lateral termination. Pronotum with

slight, rounded; anterior median protuberance; remainder of prono-

tum evenly convex, or with posterior midline faintly impressed,

the posterior median third of pronotal disc appearing flattened.

MATERIAL EXAMINED. PANAMA, 13 specimens from: CHIRIQUI

PROV> = Hartmann = Finca, 1200 m, 1300 mm... PANAMA PROV. —

Cerro Campatras CObUEC | TON DATES: “May, i Aucwst:.

OTHER VCOUNTRIES, (0 ‘specimens from: MEXICO,” COSTA RICA.

REMARKS. Females of the species may be confused with

females of acuminatus, but in acuminatus the pronotal surface is

shining between punctures whereas in nyctopus much of the

pronotal surface is dull, ‘opaque.

This species has been collected only at human feces.

6. Onthophagus acuminatus Harold

Pig. iol

Onthophagus acuminatus Harold, 1880, p. 30.

Cotypes,, Paris. . Not seen,

képoth 445 40 /,0 mm... ,Head\and pronotum, red-brown :to

Very dark’ brown or nearly black;..elytra..darker than, pronotum,

Howden and Young: Panamanian Scarabaeinae 105

dark brown to black; occasionally head and pronotum with metallic

Sheen. Dorsal setae lacking, pronotum with punctation variable,

frequently mostly impunctate in male major, minutely punctate in

either sex; if moderately punctate then usually female or male

minor. Elytra with striae shallowly but distinctly impressed and

with shallow strial punctures separated by approximately two dia-

meters; intervals feebly convex with scattered small to moderate

sized punctures, surface between smooth and shining. Pygidium

with close, moderate to large punctures, most with a small,

inconspicuous seta centrally; surface between punctures shining,

less so basally where surface finely shagreened. Metasternum

between coxae and ventral surfaces of middle and hind femora

with scattered small to minute punctures, surface between shining.

MALE. Major form (Fig. 191): clypeus broadly acuminate

anteriorly, apex reflexed with sides forming an obtuse angle,

lateral portions of margin only slightly reflexed, feebly arcuate,

feebly notched at junction with non-expanded gena. Clypeal-

frontal carina absent medially, poorly developed laterally adjacent

to and anterior to gena. Carina of vertex obsolete medially,

represented on either side near inner, posterior edge of eye by

nearly cylindrical, erect, slightly posteriorly inclined horn.

Surface of head almost uniformly, moderately punctate, most

punctures, separated by :less thanvone diameter... Prometum lonsi-

tudinally concave posterior to each horn; concavity on each side

delimiting anteriorly rounded pronotal protuberance, rarely pro-

trusion anteriorly flattened or slightly depressed along midline at

summit; disc of pronotum impunctate or minutely punctate, surface

very smooth and shining. Fore tibia similar to female, not un-—

usually elongated, lacking tuft of apical setae. Minor form:

clypeus with median apical margin scarcely reflexed, abruptly

rounded, sometimes only obsoletely acuminate; clypeal-frontal

carina low, complete, of nearly uniform height; carina of vertex

sometimes present medially, particularly when horns reduced to

small conical tubercles; pronotal protuberance reduced to slight

anterior median swelling.

FEMALE. Differing from male as follows: clypeus arcuate,

surface transversely rugose; clypeal-frontal carina complete, low,

arcuate either side of feeble median depression; pronotum with

slight, rounded, anterior median swelling or with pronotum evenly

convex, surface usually more distinctly punctate than in male.

MATERIAL EXAMINED. PANAMA, 314 specimens from: CANAL

ZONE, Ancon; Barro Colorado Island: Cabima;..Coco Solo: Corozal:

France Field; Gamboa; Madden Forest; Pipeline: Road: Rio :Frijoles:

Rio Trinidad; Skunk Hollow. CHIRIQUL: PROV. =) Bugaha,. 800-1500

{t.; Davids Rovira, COCLE PROV Eb lavadle. 2600.6. 4 PANAMA

PROV) «+: Cerro, Campana.’ COLLECTION DATES: Every” month, of) the

year:

OTHER COUNTRIES, 116 specimens from: COSTA RICA, COLOMBIA,

ECUADOR.

REMARKS. This species has been recorded at light; in

Virola sp. seed husks; on opossum, peccary, and rat carrion; and

at coati, howler monkey, human, margay, spider monkey, and

fapir feces.

106 Contrib. Amer. Ene. Tmst., wolv1d,-mo. 2; 1961

On Barro Colorado Island, acuminatus is active throughout

the period of 0600 to 2100 hours, particularly from 0600 to 0800

hours and 1600 to 1900 hours. It is one of the most abundant

Scarabaeinae and is equally common in all months of the year.

The preferred food appears to be feces, with a few individuals

being found on carrion in the late wet season. The species is

restricted to lowland moist forests, at least in Central Panama.

7. Onthophagus transisthmius n. sp.

Pras. 192, 103

HOLOTYPE. «Mate, ‘length 5.5 mm, eredtest width 3:2 mm.

Head and ‘pronotum’ (Pig. 192) very dark brown with cupreous

cast, elytra and pygidium reddish brown. Clypeal margin broadly

arcuate, anterior two-thirds harrowly reflexed: clypeal surface

finely punctate adjacent to margin, remainder of clypeus and

adjacent frons nearly impunctate and flat, clypeal-frontal carina

absent. Gena in same plane as clypeus, margin very slightly

flared to rounded median angulation. Vertex slightly uneven,

carina obsolete across vertex between horns, horn on each side at

inner edge of eye cylindrical, inclined posteriorly, feebly bowed.

Pronotum posterior to horns shallowly, broadly concave; anterior

median area between horns feebly elevated, face: only slightly

more convex than lateral areas of pronotum; concavities smooth,

remainder of pronotal surface with numerous, evenly spaced small

punctures, mostly separated by two to three diameters, obvious

setae lacking, surface between punctures smooth to finely shagreened,

shining. Elytral striae each represented by slightly impressed

double line, the line interrupted by shallow punctures separated

by two to four diameters; intervals flat, minutely punctate, sur-

face largely granular) punctures of marginal intervals: with small,

inconspicuous, pale setae. Pygidium with large, shallow, somewhat

irregular punctures, some lateral ones with setae; surface between

punctures shagreened, becoming granular basally, feebly shining

to dull. Fore tibia similar to female, not elongated, lacking

apical tuft of setae. Metasternum between middle coxae with

scattered, coarse, setose puUnCcIUreS Mear Ccoxae, minutely punctate

and shagreened medially, midline anteriorly elevated, broadly

rounded, not carinate. Middle and hind femora each with

scattered coarse punctures on outer half of ventral surface.

ALTOTYPS. Pemale, léeneth 5.9 mm, greatest width 3.1 mm.

Differing from holotype in the following major characters: clypeal

and genal margins (Fig. 193) evenly arcuate, clypeal margin only

slightly more arcuate anteriorly than laterally, surface trans-—

versely rugose. Clypeal-frontal carina low, arcuately elevated

to midline; frons, genae, and vertex moderately punctate; carina

of vertex low, slightly depressed medially, outer third on each

side slanted posteriorly toward posterior margin of vertex near

inner edge of eye; pronotum with feebly raised, rounded, anterior

median protrusion; posterior third of pronotal midline feebly,

broadly concave, opaque; pronotal surface with punctures slightly

deeper, more pronounced than in male, most with very minute,

Howden and Young: Panamanian Scarabaeinae 107

extremely short central seta, length of seta equal to approximately

half diameter of puncture.

TYPE .MATERIAL: Holotype, male, Panama, Canal Zone, Gamboa,

Limbo Hunt Club, «23624.% 1975.04? | Young .” tapim feces “irap

(Washington). Allotype, female, same data as holotype (Washington).

PARATYPES: 11>males,.7 females. «PANAMA: 3 males, 5 females,

same data as holotype; 1 male, 1 female, same data as holotype

except taken in human feces trap; 1 female, same locality .as

‘Holotype, IBALAG/59°O.P. Youngs wtapir fecés trap; 1) male, Canal

Zone, Frijoles, 2-3.X1.1975, Buena Vista P., 0.P. Young, tapir

feces trap; 1 male, Canal Zone, Skunk Hollow, 14.V11.1975, B.C.

Ratcliffe.

Paratypes are in: Ottawa, Washington, Howden, Ratcliffe.

REMARKS. Variation in the series is not great. Size ranges

from 4.3 to 5.5 mm. In females pronotal color usually has a

more greenish tinge and in both sexes the elytra vary from tan

to dark brown (in greasy specimens the metathoracic wings are

partly visible and tend to darken the elytra). In minor males

the clypeal margin is feebly reflexed, horns reduced to conical

tubercles and the pronotal protrusion is only feebly developed.

Pronotal punctures show little variation, as do most other charac-

Ters..

The species belongs to a complex of small species that occur

along the northern and western margins of South America from

Panama to Trinidad and Peru. Most, species im whe group have

conspicuous dorsal setae, at least on the elytra, and in major

males the majority of species have the horns on the vertex con-

nected by a reasonably distinct ridge or carina. Within this

complex transisthmius is most closely related to O. ophion confusus

Boucomont and to ©. nabelecki Balthasar, both occurring in

Ecuador and Colombia (these two may be synonymous). These

latter forms average at least 1 mm larger, the head of major

males is more distinctly punctate, the horns are bowed inwardly,

in some cases almost encircling the pronotal protuberance.

Pronotal color is usually a distinct oréen, the elytral: lines ave

less distinctly marked, and the pygidial punctures are more

feebly developed. All of the above characters seem to separate

the South American forms from the Panamanian transisthmius.

8. Onthophagus marginicollis Harold

Figs, 194, 195

Onthophagus marginicollis Harold, 1880, p. 31.

Type ior cotype,orerie.:.. Notuseen,

Length 4.5 toi6.0. mm. . Dorsal colors) head brown to: black,

when black frequently with metallic reflections. Pronotum with

margins, particularly lateral ones, tan to brown. Median third

to four-fifths of pronotum very dark brown to black, i black

frequently with faint greenish or metallic cast. Each elytron

with sutural third, fifth, and seventh discal intervals dark

brown to black, the intervening intervals (i.e., second, fourth,

etc.) ‘always ‘ltenter in color, tan to. brown.’ Pyeidium tan to

108 Contrib Aner. "DAtivlmsts )) veld, mes fy) 196):

brown) area near midline frequently darker. | Pronotal:suriace

moderately, closely punctate, the punctures separated by from one

to three diameters, more widely spaced in male; punctures smaller

medially) Pronotum with, lateral punctures ‘In male and ali

punctures in female each with a short inconspicuous, pale _ seta.

Elytral surface with stria represented by the usual double line

interrupted by shallow punctures separated by four to five dia-

meters; intervals flat, granular, minutely punctate in male or

coarsely punctate in female, the punctures of the latter each with

a fine seta. Pygidium coarsely punctate-setose, surface in basal

third between punctures dull. Metasternum between coxae with

Scattered Coarse puncliures only in lateral third adjacent to ¢oxde,

median third largely impunctate, smooth and shining, not carini-

form anteriorly. Ventral surfaces of middle and hind femora with

scattered coarse punctures mostly restricted to apical halves.

MALE. Major form (Pig...194): clypeus strongly reflexed) in

anterior third, margin finely arcuate. “Posterior hail of clypeus

nearly flat, transversely punctate-rugose; posterior margin of

clypeus delimited by a bow, nearly ‘straight transverse Carina.

Gena with margin slightly expanded beyond line of clypeal margin.

Frons and vertex moderately punctate; vertex with U-shaped carina

which forms long cylindrical horn above, and behind each eye

(Pig. 194)... Pronotum with median protrusion delimited. on each

side by Smooth dadentation for meception of ‘horm on vertex... Fore

tibia similar to that of female and noticeably elongate. Minor

form: clypeus less strongly reflexed, posterior clypeal carina more

elevated, horns of carina on vertex reduced or absent in extreme

cases, pronotal protrusion greatly reduced, in some cases obsolete.

FEMALE (hia. 105).) Head, similar to minor male, the: carina

of vertex broadly arcuate, of nearly uniform height, horns lacking.

Pronotum in large specimens with a distinct antero-median protru-

sion rising vertically beyond anterior margin, terminating laterally

on each side in a slightly produced rounded knob, the knob

almost directly beyond inner margin of eye. In small females

this protrusion may be almost totally lacking.

MATERIAL EXAMINED. PANAMA, 127 specimens from: CANAL

ZONES Noone Fort Kobe: s Prijoless Kio Civicito.. CHIKRIOUL PROV... —

David; (Galera de Chorcha; Remedios; Volcan de Chiriqui.. COCLE

PROV. | wweuadulce: Bl Valle, « PANAMA PROV..— Cerro Campana;

Maje; Panama Cily: Bio ba jagua. . VERAGUAS PROV..— Los

Algarrobos..., CODLEU TION. DATES: tanuary, March, May, june, july,

August, September, December.

OTHER COUNTRIES, 58 specimens from: MEXICO, GUATEMALA,

EL SALVADOR, NICARAGUA,.COSTA RICA, COLOMBIA, GUYANA, CUBA.

REMARKS. This species has been captured under bark, and

at Cows ‘horse; human, (and tapir feces. (bf occurs in.,open

pastures, scrub, and open secondary forest at low and middle

elevations.

Howden and Young: Panamanian Scarabaeinae 109

9. Onthophagus lebasi Boucomont

Pioss, WG. 107

Onthophacus. lebasi..Boucomont, 1932, pp..308, 326.

Ivpe, London. i xamined.,

Length..4.5 to.5.5,mm.....Dorsal color: (head. black with oneen

er cupreous, reflections... Pronotum.tan.to brown on) lateral thirds,

median longitudinal third darker, often greenish black; dark

median band often abruptly widened at middle. Elytra brown to

dark. brown, areas near striae sometimes slightly darker... Pygidium

same shade of brown as sides of pronotum. Surface of pronotum

coarsely punctate, punctures usually separated by less than one

diameter, most,.punctures bearing .a .distinct,, enect, tam seta; sur—

face between punctures very finely shagreened, moderately shining.

Elytral intervals, except for sutural one, with two to three rows

of small punctures, each puncture with a raised granule anteriorly,

an erect tan, seta arising, from..each) /DuUneciuUne. Fel Gium. Coarsely

punctate-setose,. suriace, between shining.....Metacternum and. ventral

surfaces of middle and hind femora with fairly evenly scattered,

coarse, SCTOse., punciures’.

MALE. Major form (Fig. 196): clypeal margin relatively

evenly, arcuate.and.only mederately reflexed;margin, of jcena, only

slightly irregular at junction. Clypeal surface flat with a few

scattered punctures; clypeal carina absent; frons and vertex

nearly flat, the posterior portion of vertex rising to transverse

plate-like carina slanted back against pronotal margin, the carina

anteriorly arcuate medially; on each side lateral terminus elevated,

forming small horn or tubercle. Pronotum unmodified (same shape

as female), slightly swollen near anterior margin. Fore tibia

elongated, inner apex conical and somewhat extended, lacking

distinct tuft of setae. Minor form not seen.

FEMALE (Fig. 197). Clypeus feebly reflexed laterally, more

so anteriorly. Median edge nearly straight with sides arcuate;

surface punctate-setose, sometimes transversely rugose; posterior

clypeal carina present, margin of gena continuing arcuate curvature

of clypeal margin. Vertex posteriorly with low arcuate carina,

carina highest, medially, surface before and behind carina with

scattered, coarse, setose punctures.

MATERIAL EXAMINED. PANAMA, 3 specimens from: CANAL

ZONE: — Barro, Colorado, Island: Palo. Seco..(0. 8) mi.) So. COLLECTION

DATES: February. :May,.lume,

OTHER COUNTRIES, None.

REMARKS. This species has been taken at howler monkey and

tapir feces. |

10. Onthophagus chryses Bates

Fig. 198

Onthophagus chryses Bates, 1887, p. 76.

Cotypes, London and Paris. London specimens examined.

Length 4.9 to 5.6 mm. Dorsally (Panamanian form) reddish,

pronotum dark brown to black with very strong greenish gold

110 Contrib. (Amer. Fri. mst. vol, 16, mo. 4: 21931

reflections. Elytra reddish tan to reddish brown with suture and

sometimes wide median transverse band greenish. Pygidium brown

to dark brown, usually with ereen or gold refiections... Head and

pronotum with small distinct punctures, frequently separated by

four to six diameters; each puncture centrally with minute brown

Seta, length of seta not exceeding diameter Of puncture; surface

between punctures very smooth, polished, shining. Elytra with

striae feebly indicated by scarcely depressed double line inter-—

rupted by shallow punctures two to three diameters apart; inter~

vals flat, most with one or two irregular rows ol minute punctures,

setae not evident, surface between punctures slightly irregular,

polished, shining. Pyoidium with’ well: separated, moderate sized

punctures, most separated by approximately one diameter; each

puncture usually with minute central seta. Metasternum between

middle coxae with scattered coarse punctures, punctures lacking

only me@ar faintly impressed midline, suriace between punctures

and alone midline smooth and: shining. Middle and hind” femora

with Scattered, distinctly separated, coarse, setose punctures dis-

trrputed over most Of veniral suriace, suriace between punctures

shining.

MALE. Major form (Fie. 190): clypeus anteriorly strongly

transversely reflexed; reflexed portion approximately two-thirds

width of clypeus, nearly vertical with edge very broadly, shallowly,

emar@inate.. “Sides of clypeus’ Dehind: retlexed portion distinctly

divergent, feebly arcuate, side of gena arcuately expanded beyond

line Of ciypeal maroin. Suriace ol clypeus, trons, and vertex

lacking dany indication ef carina; moderately punctate, punctures

Separated Dy a @istance Of One Or two diameters. Some punctures

on clypeus and eenae, particularly lateral ones, with fine setae,

surface between smooth. Pronotum anteriorly behind head tri-

tuberculate; tubercles directed anteriorly, delimited by longitudinal

concavities between (Fig. 198); summit of tubercles rounded, re-

mainder Gf. pronotum ‘evenly convex. ~fore tibia distinctly elongate

With large “apical lull of setae al base of spur. ©Minor form: not

seen but probably the reflexed portion of clypeus and pronotal

tubercles reduced in development and possibly carina, as described

for female, evident on head.

PEMALM. Clypeus weakly reilexea laterally, moderately so

anteriorly; anterior third of margin broadly. shallowly, emar-

@imate. ~ Surface of Clypeus (transversely rugose, clypeal—frontal

carina low, of nearly uniform height. Gena with margin flared

slightly beyona line of clypeal margin. Vertex with low, slightly

Sinuous transverse carina; carina in median two-thirds of nearly

Uniform Nelont, OulLer ‘portion on either side abruptly ‘sloping

gownward, tO posterior itimer edoe of eye... Pronotum: with anterior

portion’ immediately behind margin very feebly trituberculate,

swellings and indentations sometimes only visible with proper

lighting: Pere tibia unmodified, lacking Ttult of setae.

MATERIAL EXAMINED... PANAMA.© 4 specimens from: CHIRIQUI

PROV.-— Boquete; Buecaba: Cérro Punta, «VERAGUAS PROV. /-

Potreriliogs; COLEECTION DATES: January, March, (June.

OTHER COUNTRIPS, 2. specimens trom: COSTA RICA.

REMARKS, This’ species has been’ recorded’ from human feces.

Howden and Young: Panamanian Scarabaeinae pe

ll. Onthophagus coscineus Bates

Figsa boo, 4 200

Onthophagus coscineus Bates, 1867.05 go:

Cotypes, London (1 female) and Paris (pair). Examined.

Onthophagus digitifer Boucomont,’ 1932,. p.' 419. NEW SYNONYMY.

Type, Dresden (?). Not seen.

Teneth 275"to 4.1 mm. - Dorsallydull brown “to:-Diack: in

profile with conspicuous, erect tan setae; head (Fig. 200) with

close, simple, large punctures; pronotal punctures with contiguous,

raised, shining |margins; bottom of punctures flat, dull, sranuler

with céntral granule at base of seta shining.” Blytral etriae not

impressed; represented ‘by doubled shining .liné Interrupted Dy

shallow punctures; intervals flat, dull, shagreened, with scattered

setae, a small raised granule at base of each seia. Py eidium

obsoletely punctate-setose, surface dull, shagreened or very finely.

granular. Meétasternum: and ventral surfaces’ of atl femora with

numerous coarse punctures, surface between punctures smooth and

shining.

MALE. Major form (Fig. 199): clypeus broadly arcuate

laterally, reflexed and broadly emarginate anteriorly. Margin of

gena continuing arcuate line of ‘clypeal marain. . Clypea—irontal

carina..absent; carina of vertex straight, low, sor (uniiorm, hereht

between anterior fourth of eyes; vertex posterior to carina .as

heavily punctate as frons and genae. Pronotum with small,

anteriorly arcuate protuberance medially just posterior to anterior

margin; face and upper edge shining, remainder of pronotum dull,

contiguously' punctate. Fore. tibia slightly ‘clongated,. tacking

apical tuft of setae. Miner form: clypeal margin: less 'refrlexed:

pronotal protrusion very. feebly developed; forse tibia not elongated.

FEMALE. Clypeus as in male except straight, low -‘clypeai-

frontal carina present, other characters including carina of vertex

as described for male; pronotal protrusion feebly developed or

absent.

MATERIAL EXAMINED. PANAMA, 85 specimens from: CANAL

ZONE -. Ancon; Barro Colorado Island; Cocoll:; ‘Coco: Solo; Frijoles:

Gatun; Madden Forest; Pipeline. Road; Summit Park." CHIRITOUT

PROV. -. Rovira... PANAMA PROV. -— Cerro Campana. VERAGUAS

PROV. — Potrerillos. “COLLECTION DATES: Every month of the year.

OTHER COUNTRIES, 16° specimens from: ‘COSTA RICA)’ COLOMBIA,

ECUADOR.

REMARKS. Cotypes of coscineus Bates and specimens determined

by Boucomont as digitifer have been examined. All specimens

seemingly ‘represent one species; “hence digitifer is placed in

synonymy.

specimens of coscineus have been taken at dog, howler

monkey, human, peecary.,. and, tapir feces. .On Barro Colorado

Island the species is active during the period from 1000 to 1400

hours, and occurs in fairly uniform numbers throughout the year.

The species in Panama is limited to the central lowland moist

forests.

lig Contrib. Amer.) fntw lasts; volui?Be nov. dy), L961

12. Onthophagus dicranius Bates

Pag) 20)

Onthophagus dicranius Bates, 1887, p. 72.

Cotypes, London and Paris. London specimens examined.

Lenoth 4.5. %0 6.0, mm... Dorsally. uniform tan:to dark -brown;

middle and hind femora centrally pale, yellowish; remainder of

ventral surfaces similar to dorsum in color; dorsum heavily punc-

tate, smost punctures each with, very minute, inconspicuous seta,

setae on elytra most readily visible; all dorsal punctures, including

those on pygidium, shallow, bottoms flattened, usually feebly

ocellate; surface between punctures shining except on pygidium

where finely shagreened. Metasternum and ventral surfaces of

middle and hind femora with scattered small punctures, surface

between punctures smooth and shining.

MALE. Major, form (Fig. 201): .clypeus, with, anterior median

sixth or seventh of margin vertically reflexed into a transversely

flattened parallel-sided, apically truncate horn, the horn approxi-

mately as high as wide; remainder of clypeal margin evenly

arcuate and moderately reflexed, clypeal surface medially obsoletely

punctate, shining; remainder.of head) heavily, closely punctate.

Margin of gena continuing arcuate line of clypeal margin; head

lacking any imdication.of carinae,, Preonetum in jantertor, third

with an elongate longitudinal carina on either side close to mid-

line, summit of each carina obtusely angled at anterior third of

pronotum, thence extending downward and slightly converging

anteriorly toward midline; carinae impunctate, remainder of prono-—

tum closely punctate, evenly convex. Fore tibia slightly elongated,

lacking. distinct. tuft..of, setae, at.apex. Miner form: clypeal: horn

reduced to rounded elevation, width not greatly reduced; pronotal

carinae reduced in height, usually evident; fore tibia similar to

females.

FEMALE. Clypeus anteriorly sharply bidentate, emargination

V-shaped, teeth acute, moderately reflexed; remainder of margin

not or feebly reflexed. Head punctate, slightly convex medially,

lacking any indication of carinae. Pronotum evenly convex,

anterior seventh of midline impunctate, smooth and shining,

remainder of pronotal surface closely punctate.

MATERIAL EXAMINED. PANAMA, 8 specimens from: CANAL

ZONE - Barro Colorado Island. CHIRIQUI PROV. - Bugaba;

Hatimaniis Hince, (500. mai) COLLECTION DATES: (May, june,

September,

OTHER “GCOUNTRIES..4\ specimens from: MEXICO, BELIZE.

REMARKS. The species has been taken from rotting fruit of

Gustavia sp. and at human feces.

13. Onthophagus landolti Harold

Figen 202

Onthophagus landolti Harold, 1880, p. 34.

Type lor cotype, Paris. | Examined.

Length 4.2 >to. 6.0 mm, Head and pronotum dorsally black

with distinct greenish reflections, anterior pronotal margin often

Howden and Young: Panamanian Scarabaeinae 113

narrowly edged with brown. Elytra largely greenish black, fre-

quently with brown spots near basal and apical margins.

Pygidium occasionally entirely greenish black, more frequently

greenish black centrally, brown laterally and sometimes apically.

Dorsal surface of head and pronotum with numerous setose punc-—

tures, surface between shining. Elytral intervals shallowly punc-

tate with one row of punctures along outer edge of interval, most

punctures with pale, erect’ setae. Pygidial surface coarsely punc—

tate-setose. Metasternum with distinctly separated, coarse punc-

tures relatively uniformly distributed, separated by approximately

one diameter. Ventral surfaces of middle and hind femora with

similar coarse, setose punctures.

MALE. Major form (Fig. 202): clypeus anteriorly abruptly

reflexed, very shallowly, broadly emarginate medially; posterior

two-thirds of clypeus nearly flat with a few scattered coarse

punctures; ‘carina absent. Genai distinctly ‘expanded, Vertex

largely impunctate, carina vaguely indicated above and behind

each eye. Pronotum with median, anteriorly projecting, rounded

tubercle overhanging anterior margin; pronotum otherwise evenly

convex. Fore tibia considerably elongated with a tuft of yellowish

setae at the base’ of €onical apex. © Minor tormivelypeus searcely

reflexed anteriorly, carina on vertex complete; anterior pronotal

protuberance greatly reduced; fore tibia only slightly elongated.

FEMALE. Clypeus slightly reflexed laterally, more so anter-

iorly, median edge broadly emarginate; surface rugose, punctate-

setose; ‘posterior clypeal carina present. “Gena only slightly

expanded; vertex posteriorly with arcuate carina, highest medially.

Surface of frons and vertex coarsely punctate-setose. Pronotum

with anterior swelling obsolete or vaguely indicated and contigu-—

ous lo anterior’ mareain.

MATERIAL EXAMINED. PANAMA, 81 specimens from: CANAL

ZONE ~ Cocoli.: CHIRIOUL PROV -- 1Galeray de Chorcha; Hartmann’ s

Finca, 1200 m;Las“Lacunas: Gante Clara) COCLE PROV ee) a)

Valle. PANAMA PROV. - Cerro Campana; Rio La Jagua. VERAGUAS

PROV. = Potrertilos, OOLLEOCTION DATES: January) February ,

March, April, May, August, December.

OTHER COUNTRIES, 122 ¢specimens’ from: MEXICO, BELIZE, EL

~SALVADOR, COSTA RICK, COLOMBIA,

REMARKS. This species has been captured at horse, human,

and tapir. feces. [ti usuaiiy ie tound in forest and serub areas

below 1200 m.

14. Onthophagus atrosericeus Boucomont

Figw? 203

Onthophagus atrosericeus Boucomont, 1932, Op OLIg eh S20).

Type, Paris (?).7 Not seen,

Length’ 5.5° to’ 7.5 mm. Dorsally dull black, non-punctate

surfaces finely granular; head, elytra and pygidium impunctate

or only feebly punctate; pronotal disc distinctly punctate, punc-—

tures large, margins a conspicuous shining ring, bottom of punc-

ture flat, each puncture centrally with erect black seta; punctures

114 Comtvib«-Amer. Ent. anst., vol. @6;..n0...1,. 1981

nearly contiguous laterally, separated by one ito two diameters. at

center of disc, suriace between punctures Granular; anterior

pronotal angles on each side rounded, adjacent lateral margin

frequently. slightly.inwardly arcuate... Elytra with-..striae repre-

sented by distinct lines,. intervals oranular, flat, most.with two

slightly irregular rows. of erect. setae. Pycidium. with. scattered

erect sélae, surface granular. -Melasternum and. ventral. surfaces

Ol ali. femora with scattered. small to-moderate sized punctures,

surface, between mostly smooth and: shining. :

MALE. Major form (Fig. 203); clypeus abruptly. -reflexed in

anterior third, retlexed maroin-slightly, emarginate, remainder of

head flat to slightly convex, impunmctate;: margin of gena.arcuately.

produced beyond line of clypeal margin. —Pronotum with large,

anterior, median protuberance rising vertically behind anterior

margin, upper edge of protrusion, shitting, smooth, slightly depressed

Near Midline; Vertical face finely punctate, shining. . Fore. .tibra

Greatly. elongated, a tuil of selae present beside apical spur.

Minor form: clypeus teebly.retlexed, very broadly, shallowly

emarginate, gena only slightly expanded; carina of vertex feebly

Indigated On either side cf. midline between eyes; pronotal protu-—

Derance a fine, raised, shining, arcuate.carina just. behind

anterior «Margins fore tibia, not.unusually long, similar. to, female.

PEMALE. Clypeus feebly reflexed, broadly emarginate; -sur—

face mostly closely punctate and shining; only shining carina on

vertex present, the short, Straight catina positioned.,midway

between eyes: vertex impunctate;. pronetum with median. protrusion

Obsolete, represented by a slightly swollen, shining area adjacent

tO median. anterior margin.

MATERIAL EXAMINED. PANAMA, 68 specimens from: CHIRIQUI

PROV. — Baldwin Forest, 1/60 m: Cerro Punta (2 km E-~=—. 2000-2200

m); Ceres Punta (5 km ESE — 2600 m)... COLLECTION DATES: May,

June.

OLHER COUNTRIES, . described: from: COSTA :RICA.

REMARKS. This-species Nas been found at. horse dung and

human feees.-. [he distribution of atrosericeus’is seemingly

restricted in Panama to the forests avove 1/00 m elevation.

15. Onthophagus cyanellus Bates

Figs 204

Onthophagus cyanellus Bates, 1887, p. 81.

Cotypes, London and Paris. London specimens examined.

Pemetin. 7.5 to (0.5 mm. Color dorsally dull black, usually

with faint Dbluish tinge. Head and pronctum finely, closely punc-—

tate; pronolal punctures, particularly laterally, frequently with

minute erect Selaec, Setae scarcely longer than width of puncture;

slightly longer in females; pronotal surface between punctures

finely @ranular. Elyival intervals flat with two-or three irregular

rows Of minute tubercles, a: fine, short,,erect: seta .arising: from

base. of each. <ubercle,;surtace.. between, finely granular;.elytral

sitiac. represented by feebly impressed shining lines interrupted

by large shallow punctures. Pygidium opaque with scattered small

Howden and Young: Panamanian Scarabaeinae £5

punctures, many with minute setae, surface between granular.

Metasternum between middle coxae with small to large punctures,

midline finely impressed except in anterior fifth. All femora with

ventral surfaces with mixture of large and small punctures.

MALE. Major form.(Fig.: 204): clypeus. broadly transverse,

over half of total surface reflexed, laterally broadly sinuously

arcuate to rounded angles. Lateral clypeal margins short, gena

expanded, margin of gena. three to four times length of lateral

clypeal margin. Surface of head moderately to coarsely punctate,

occasionally rugose. Clypeal-frontal carina absent, carina of

vertex bow-shaped, vaguely indicated medially, terminating later-

ally on each side in a small rounded tubercle near inner posterior

margin of eye. Pronotum with anterior angles acute, laterally

with margin obtusely angled at anterior third, arcuate behind

angle, the area between angle and median pronotal protuberance

shallowly, “broadly concave... Proneotal.protuberdmee, truncate anter—

iorly, sides diverging arcuately to small tubercle on each side

above pronotal fossa; summit of protuberance often faintly indicated

on each side; surface anteriorly shining between punctures. Pos-

terior half of pronotum dull, surface between punctures granular.

Fore tibia distinctly elongated, a few apical setae present but

distinct tuft lacking. «Minor orm: clypeus with,entine mareain

feebly, nearly uniformly reflexed, surface transversely rugose.

Both clypeal-frontal carina and carina of vertex feebly indicated.

Frons and vertex closely punctate, often transversely rugose.

Pronotum. lacking abrupt angle laterally, central protuberance

feebly elevated, slightly more shining than surrounding convex

area of pronotum... Fore tibia: not elongated.

FEMALE. Clypeal margin arcuate, more broadly so anteriorly,

narrowly evenly reflexed, surface transversely rugose. Clypeal-

frontal. carina distinct; ‘low, of ainiform height, anteriorly arcuate.

Frons and gena closely, nearly contiguously punctate. Carina of

vertex with median third more anterior in position, the carina

slightly sinuous, of nearly uniform height, terminating on-each

side near posterior inner margin of eye. Pronotum with anterior

protrusion as described for male minor, feeble protuberance shining,

remainder.of pronotum opaque.

MATERIAL EXAMINED. PANAMA, 141 specimens from: CHIRIQUI

PROV.::< Baldwir, Hepests, Boguete: Bugaba: Genre..Punta,, 5/50..ft..

6250. ftic: Cerro. Punta: (2. ki 2000 mia ahineasweriadat.Hartnann. s

Finca, 1500 .ms7 1550.00 Moleam deo Chiriaui. COLLECTION DATES:

January, March, May, lume, Qclober,

OTHER COUNTRIES, 56 specimens from: MEXICO, NICARAGUA,

COSTA. RICA

REMARKS. This species has been captured at carrion, and at

horse, ‘hunian, and tapiy feces, « Tne species. occurs’ ins moist

forests at elevations between 1000 to 2500 m.

116 Contrib, /Amer, Bt. basics vol. 26.) mov .i: 1981

16. Onthophagus mirabilis Bates

Figs 205-207

Onthophnacgus mirabilis Bates, (00/5 op. 74; Howden, 19736, 'p. 334.

Type; Patis. "Ux anined,

Onthophagus orphnoides Bates, 1887, p. 75. NEW SYNONYMY.

Cotypes, London and Paris. Series examined.

Length 7 te t2°mm. Dorsally dark brown to ‘brownish Dlack,

glabrous; ‘head, pronotum, elytral intervals and pygidium with

close to contiguous, moderate-sized punctures, surface between

shining; elytral striae distinctly impressed. Metasternum between

middle coxae more’ strongly punctate laterally, number of punctures

variable, smaller specimens having more and larger punctures.

Ventral surfaces of middle and hind femora with scattered minute

LO Coarse” punctures, Gensity and size Variable, “but more numerous

im “small Specimens, particularly ‘small females. ~ Sexual ‘dimorphism

unusually pronounced; characters of each sex also strongly influ-

enced by allometric (heterogenic) growth.

MALE. Major: form (Pig. 205): clypeal margin converging to

anterior, median, almost vertical horn, the horn thickened near

apex and narrowly, deeply emarginate; clypeus otherwise almost

flat, feebly punctate. Junction of gena and clypeal margin obtusely

angled, sides of genae nearly parallel. Frons and vertex feebly

convex,’ not carinate or horned. Pronotum with conspicuous

conical, anteriorly projecting median horn (Fig. 9205), ‘the “horn

extending over head to middle of clypeus and apically bifurcate.

Fore tibia somewhat elongated and with conspicuous tuft of setae

al base (Of “apical spur. “Minor? torm (Pic. 206): both clypeal and

pronotal horns reduced to small tubercles; often some slight indi-

cation On ¢lypeal Norn of: an e@margination; fore tibiae not elongated,

similar to female.

FEMALE (Fig. 207). Large specimens (described by Bates,

1887, under the name O. orphnoides): clypeus anteriorly bidentate,

the two teeth reflexed and separated by U-shaped emargination;

clypeal surface otherwise flat and closely punctate; posterior

clypeal carina obsolete, represented by fine ridge. Vertex on

each side near inner edge of eye with low tubercle; ‘surface

punctate before, impunctate behind. Pronotum with acute conical

tubercle”or horn medially at anterior margin; pronotum medially

longitudinally concave, highest at obtuse angle either side of

CONnCaVIty ‘ab anterior’ third; concavity largely impunctate, “remainder

OF pronctum punctate. Somali! specimens have .clypeal: teeth,

tubercles on vertex, and pronotal horn all reduced in size; prono-

tal concavity very feebly indicated, entire surface heavily punc-

tate.

MATERIAL EXAMINED. PANAMA, 10 specimens from: CHIRIQUI

PROV. - Baldwin Poorest; Hartmann s*rinca, 1200'm; Las Lagunas;

canta Clara, Voleanm ce Cniregut, 2000" fi, COLLECTION ‘DATES:

Mays Junes July.

OTHER COUNTRIES, 1 specimen from: “ECUADOR.

REMARKS. It is not surprising that Bates described the two

sexes of this species’ under different names, although he did note

that the specimens of orphnoides were probably all females. Only

Howden and Young: Panamanian Scarabaeinae Ty

recently have both sexes been taken in one locality and enough

specimens have become available to indicate that the name

mirabilis represents the male and orphnoides the female of the

Same species.

The species has been taken at light, on carrion, and at

human feces. A few specimens have been collected by sweeping

over a forest trail at dusk in a wet valley forest 2 km west of

Cerro Parte’

17. Onthophagus sharpi Harold

Fig. 206

Onthophagus sharpi Harold, 1875, p. 138.

Cotypes,*: Paris: Not’ seen:

Length 6.5 to 8.8 mm. Dorsally shining brownish black to

black with faint purple or blue cast; pronotum minutely punctate,

lateral areas of pronotum and head more distinctly so; lateral

pronotal margins evenly arcuate. ‘Dorsal''setae ‘absent. - Elytron

with strial lines impressed, the lines regularly interrupted by

Shallow punctures; intervals flat to slightly convex, smooth

occasionally with scattered very minute punctures, surface shining.

Pygidium moderately punctate; the punctures separared: by ‘one’ 10

two diameters, most punctures with a minute seta; surface between

punctures smooth and shining. Metasternum and ventral surfaces

of all femora with scattered small to moderate punctures, surface

between punctures smooth and shining.

MALE. Major form: clypeus (Fig. 208) with erect horn

arising vertically at median anterior edge; expanding in “outer

two-thirds to form a wide arcuate T-shaped plate nearly truncate

dorsally and approximately equal in width to base of clypeus,

clypeus on ‘either sidé of ‘horn with edge mot reflexed, ‘almost

transverse to abruptly rounded lateral angle, posteriorly with

clypeal sides nearly parallel to junction with expanded gena.

Surface'‘of frons a@mdiivertex centrally teebly convex, tacking any

indication of carinae or “horns.” ) Pronotum. very slightly flattened

in anterior median third, posterior summit with flattened area

often with vague median! tubercle; flattened area more distinctly

punctate than remainder of disc, pronotum otherwise evenly

convex. © Fore tibia’ of approximately same Tenoth as’ female, Dut

with large distinct brush ‘of apical ‘setae. Minor form: clypea!

hern reduced to evenly rounded vertically reflexed anterior edge,

margin of clypéeus’ and eeéna otherwise arcuate to slightly sinuate,

gena: not’ noticeably produced.” "Fronotum’ lackine flattened” area,

evenly’ convex, ‘somewhat’ more’ distinctly’ punctate.

FEMALE.’ "Clypeus ‘anteriorly’ stronaly bidentate, median

emargination narrowly V-shaped. Tooth on either side acute,

moderately’ reflexed, remainder ‘of marein on’ each’ side evenly

arcuate, “arcuate ‘margin continuine “to posterior ed@e, orgena.

Clypeal-frontal carina feebly indicated’ in median half, occasion-

ally ‘obsolete, (vertex lacking earina. Pronotum evenly convex,

moderately” punctate.’ "Fore ‘tibial lacking ‘apical’ brush ‘of hair.

MATERIAL EXAMINED. PANAMA, 196 specimens from: BOCAS DEL

ii Contrib. Amer. Ent... Iist.. vol.-18. mo... 1, 1931

TORO: PROV. — Rio: Changuinola. CANAL ZONE — Barro Colorado

Island; Frijoless Madden. Forest; Pipeline Boad:. Rio Ciricito,

CHIRIQUL PROV..- Boquete; Bugaba; Hartmann's Pinca,.1300-.m.

FANAMA PROV. -— Cerro Campana... COLLECTION DAIES:- February,

April, May, june, july,. August, September,,. December.

OTHER COUNTRIES, 18. specimens from: GUATEMALA, COSTA

RICAy ECUADOR.

REMARKS. Onthophagus tapirus Sharp (1877, p. 130) was

described trom Nicaragua dnd is extremely close, if mot identical,

tO sharpl. ihe species tapirus supposedly difiers from sSharpi in

averaging slightly smalier, lacking the conspicuous flattened area

on the pronotum of ,tne males and baying a sreenish coppery cast.

There is also supposedly a difierence in clypeal shape but, this

Varies. sO much with heterogonic growlh that it isan extremely

Gifficull character to assess. We have seen specimens from Panama

that, meght be considered, to be tapirus but these were taken. at

the same time and place where typical sharpi were found. The

eroup should be caretully studied .but at, present,.all specimens

that.we have seen from, Panama.we would,.place.under the name

sharpi.

This. species. hac. been .recorded. from.fresh Scheelea-sp.

husks, fermented palm druits.. rotting truil. ef Gustavia sp.,

Maguita, sp. fruil, rotting bananas, spadix of Philodendron sp.,

and. fallen. flowers.of. Gustavia sp.; on.tree'.trunk, leaf .suriaces,

in. or under nest. of -[ricona.cupira, and at light; 4nm-or..on

Agaricaceae, Polyporus tenuis, Favolus sp., and fungus mold on

log; at rat and other carrion,.amd under skin..and.bones:.of -coati;

on howler.monkey.and human feces; and has been bred from

Heliconia mariae and UH. pendula.

On Barro Colorado Island sharpi is nocturnally active during

the period from 1600 to 0800 hours and is most abundant during

the early and mid-wet season... It probably has the broadest

feeding niche.of any of.the Scarabaeinae, in. Panama..

18. Onthophagus praecellens Bates

hie... 209

Onthophagus praecellens Bates, 1887, p. 70.

Cotypes, London and Paris. London specimens examined.

Lenetn 4.7. to: 7.54mm-.., Deorsally. shining. brownish black to

black, usually. with strong green or cupreous.cast.. Head. variably

punctate, pronotum with small, fairly uniformly distributed punc-

tures.separated by two.10 five diameters, .cach.puncture. frequently

with minute, central, ceta;, setal lensth,approximately equal. to

diameter.of punctures.scurface, between. punctures.shining, »smooth

to finely.shagreened. . Elytral.striae feebly impressed, with. double

line interrupted by shallow punctures separated by .one.to,two

diameters;. intervals mearly flat, with-scattered. minute. punctures,

surface. shagreened,. moderately. shining; setae on..intervals. present

only Near. marcins.«. Pycidium, with,poorly defined,. close: punctures,

surface in and. between. punctures. dull, shagreened: or. granular.

Metasternum between middle coxae with scattered coarse punctures

Howden and Young: Panamanian Scarabaeinae 119

except near midline, surface shagréened, feébly shining. Ventral

surfaces of middle and hind femora iareely impumectate, smooth

(extreme apex may have one or two punctures).

MALE. Major form (Fig. 209): anterior median clypeal horn

rising nearly vertically and. expanding to form wide, posteriorly

directed U-shaped apex, each arm of 7U'° directea toward adjacent

eye, terminating’ near posterior edge Of clypels: marcia of clypeus

on either side.of horn sinucusly expanded to upwardly flared,

angulate junction with margin of gena; margin of gena extending

directly toward posterior, turning inward opposite eye; shallowly

concave. surface of clypeus with. unevenly scattered punctures;

clypeal-frontal carina absent; carina of vertex represented by

small acutely pointed tubercle on,.either side near posterior inner

Margin of eye; vertex between tubercles with feeble depression and

ridge on. either .side of smooth midline... Pronetum in anterior

third with triangular, longitudinal upright plate on either side of

midline, outer angle of plate directed anteriorly and often over-

hanging vertex of head; apices of plates separated by approxi-

mately 0.5 mm. Fore tibia elongated, with apical tuft of setae

at pase.of spur. Minor Torm:, clypeus with sides, evenly .converaine

anteriorly to bluntly. acuminate, feebly reflexed, .apex; remainder

of head nearly flat, gena not flared; clypeal-frontal carina

absent; tubercles on.vertex. greatly reduced,.and. rounded but.still

evident; pronotal plates reduced to rounded tubercles; fore tibia

unmodified.

FEMALE. Differing from male as follows: clypeus arcuately

converging to emarginate, bidentate, leeply. reilexed iapex: clypeal

surface transversely rugose; clypeal-frontal carina very low,

usually indicated across entire width of ciypeus to margins;

vertex with broadly conical tubercle on either side nearly equi-

distant between midline and eye; pronotal plates greatly reduced,

but summit angulate, separated from opposite summit by approxi-

mately 0.3 mm, this varying slightly depending on size of female.

Fore tibia lacking apical tuft of setae.

MATERIAL EXAMINED. PANAMA, 209 specimens from: CANAL

ZONE .~.Barro Colorado Island; Cabima:. Coco Soloy Madden Forest:

Pipeline. Road..: COCLE PROV. ='0Fi Valle. 26000 f1.) COLON, PROV... -

Alhajuela. .. CHIRIOUI” PROV. = Corpo Pande: Las Lagunas... .COLLEC-

TION DATES? Every month of tne year.

OTHER COUNTRIES, 31 specimens from: GUATEMALA, COSTA

RICA, COLOMBIA, TRINIDAD,

REMARKS. This species has been captured at fermented palm

fruits, in. wet, debris of small stream, and in forest. floor leaf

litter;..at agouti, armadillo, bird, coati, howler. monkey, lizard,

peccary, rat, and tamandua carrion; and on coati, howler monkey,

human, opossum, spider monkey, and tapir feces.

On Barro Colorado Island, praecellens is active during the

period.from. 0700 19.1000 Bours. Jt occurs, in. all. months of the

year but is most abundant during the late wet season and early

dry season, and is equally abundant on both feces and carrion.

120 Contribs Amers Bote Insts, voli 16, sno. da: 1961

19. Onthophagus belorhinus Bates

Pyo2 210

Onthophagus belorhinus Bates, 1887, p. 69.

Cotyvpes, Eondon and Paris.” London specimens examined.

Length ©6.0°to'9.5 mm. ©" Dorsaliy dark brown, brownish

black or black; occasionally: with bluish reflections; “head ‘and

pronolam Shining, elyira dully doreqa! seiae’ absent except some—

times along margins of elytra. Head and pronotum finely to

moderately punctate, pronotal punctures small, on disc separated

Dy two to [Our diameters.” ( Blytrai striae represented by shining

double line interrupted by shallow punctures; elytral intervals

frequently impunctate or with Gxiremely minute punctures, surface

finely granular. Pygidium with moderate-sized punctures, the

punctures often poorly delimited, pygidial surface dull, opaque.

Metasternum between middle coxae with small to moderate-sized

punctures near coxae, minutely punctate and finely shagreened

medially. Middle and “hind temora with a tew Coarse punctures in

apical third on Vertical “surface, remainder of suriace Smooth or

finely punctate.

MALES Major form (Pig. 210): clypeus with anterior median

edge (about 0.3 mm wide) forming horn extending vertically for

0.5) to 0.7 mit, hence’ Difurcating mie two diverging, posteriorly

directed branches, each pointed toward adjacent eye, the branch

terminating in an acute polnl midway Detween anterior and poster-

tor moarein’’ Ol Heads anterior Clypeal marein on either side ‘of

horn truncate, “broadly tlared to abruptly arcuate sides; gena

elongate, side nearly straight, slightly divergent to abrupt angle

in line with eye; surface of clypeus and frons broadly, longitudin-

ally concave; clypeal-frontal carina absent; carina of vertex

represented by lone, cylindrical, ‘bowed horns laterally enclosing

an almost conical, median bituberculate pronotal prominence; fore

tibia ‘elongated with apical tuft ol setae at base of spur. “Major

form, moderate development; clypeus with anterior median vertical

horn (Pio. 290) rounded ai summit, apex neither bifurcate nor

bent posteriorly. sides: Of clypeus om either’ side evenly arcuate to

gena, net laterally flared, margin ©, géna in. line with clypeal

margin; ‘horns om vertex and fore tibia only very slightly reduced

in length; pronotum more distinctly bituberculate at summit. Minor

form: ¢lypeus feebly retlexed and acuminate anteriorly, horns on

vertex reduced to conical tubercles but with no carina between;

pronotal prominence represented by two distinct, “slightly longi-

tudinglly elomogte, rounded tubercles, one On either side of

narrow median longitudinal concavity; fore tibia similar to female,

lacking apreal Tuit of setae.

PEMALE, Dittering trom male inthe tollowing characters:

sides of clypeus nearly straight, converging to narrowly reflexed

bidentate apex, the apex moderately emarginate between rounded

teeth; clypeal surface transversely rugose; clypeal-frontal carina

represented by low ridge terminating on each side at junction

with Gena; ’cavina of vertex represented On cach side by a short,

Fairly stout) Vupright ‘horn ‘(in small females: horn reduced to

conical tubercle), vertex not carinate between horns; pronotum

Howden and Young: Panamanian Scarabaeinae Zh

with two longitudinal tubercles as described for male minor.

Fore tibia lacking apical tuft. of setae.

MATERIAL EXAMINED. PANAMA, 48 specimens from: CHIRIQUI

PROV. - Baldwin Forest; Begquetes)Cerro,Vandop Cerro Vunta;

Hartmann's Finca. PANAMA PROV. - San Isidro, 1600 ft. COLLEC-

TION DATES: January, «May, June), October, )November, December.

OTHER COUNTRIES, 19 specimens from: MEXICO, EL SALVADOR,

COSTA, RICA, GOLOMBIA, : ECUADOR,

| REMARKS. This species has been found at light, on rat

carrion and on unknown carrion, and at horse, human, and

opossum feces.

Ontherus Erichson

Ontherus’ Erichson, 1647, p. 1077) Janssens. 1949) ps 15.

Type-species: Ontherus didymus Erichson, listed by Lucas,

eos

Usually black, size moderate, 10°to20 mmr: secutellum: not

visible; elytron with eight striae (nine counting inflexed margin);

mesosternum moderate in length, somewhat triangular, median area

closely punctate; metasternum between middle coxae feebly to

moderately narrowed anteriorly; abdomen with six sternites visible

laterally, medially only three to five evident; abdomen along

ventral midline shortened, thus compressing sternites; hind, tibia

gradually expanded to apex, not transversely carinate on outer

margin; all tarsi with claws:

Key to species of Ontherus

Ae Metasternum between middle coxae distinctly

narrowed anteriorly, distinctly narrower than

WLAOLN Of, One coxa: Laccia midline not im-

DESSSER 4a aie 2

Metasternum beeen ale coxae eA ee et

narrowed anteriorly, approximately equal in

width :to, that of one coxa: pronotal imidiine

narrowly impressed in posterior half me

eH Da iSee LS The SD.

2 Vertex adjacent to inner edge ol eye concave;

elytra A ts twice as long as pronotum

, 2, Caaymus) Er.

Vertex dane ent to inner N ddge eh eve. al Least

slightly convex: ne ne uae longer

than! PPONOCU AD y)\) ay. 1. brevipennis | lar,

122 Contrib. Amere Ent... Insts, vol. 16. mo. 15+ .1981

1. Ontherus. brevipennis Harold

Pic. 200

Ontherus brevipennis Harold, 1867b, p. 97.

Type, Paris. Not seen,

Lenoith’ tlto 14 mim... Aiterior Mareain OL ‘clypeus arcuate,

not emarginate medially. Frons (Fig. 211) with low transverse

carina, termination tuberculate on each side; carina as wide as

three-fourths of distance between eyes. Pronotum (Fig. 211) with

inverted V-shaped swelling medially, adjacent to. anterior margin.

Metasternum with midline impressed for entire. length. © Posterior

margin Of hind femur with a.small tooth at. apical end... Parameres

of genitalia’ each with a small acute tooth just below apex.

| MATERIAL EXAMINED. PANAMA, 31 specimens from: CANAL

ZONE. — Black Tank Road... COLON.PROV: — Portebelo, 1911,.1912.

PANAMA. REP. (—) mo. other data... COLLECTION, DATES: ..Prebruary,

July.

OTHER COUNIRLES,: None.

REMARKS, All of the specimens. seen lacked host data and

only twe,of. these. were. collected after 1912.

2. Ontherus didymus Erichson

Face. -212,,.213

Ontherus didymus Erichson, 1847, p. 108.

Type, Berlin (?). Not seen.

Lenoth lb t0,20 mm. .Anterier margin, of clypeus shallowly

emarginate medially... —Frons tumid, varying from feebly bitubercu-—

late to distinctly horned, the upright horn (Fig. 213) bitubercu-

late at transverse apex. Pronotum (Figs. 212, 213) varying from

evenly convex (small specimens) to feebly bituberculate (females)

to. distinctly quadrituberculate, the lateral tubercles larger and

slightly more anterior 1m position. Metasternum lacking impressed

midline or midline feebly impressed Only in posterior half. Hind

femur of males lacking tooth or expanded flange near, apex of

hind, margin... Pavamerés of genitalia with acute apices.

MATERIAL BAAMINED, PANAMA, 219 specimens from: CHIRIQUL

PROV = Baldwin. forest: (Cerco Punta: 2 km W Cerro Punta, 1300 m,

1700. mi 2km EE Cerro Pinta, 2200 m:.Hartmann's Finca, 1300 .m,

1500, ms 1550; ms hae Laciumas, 1360 m. COLLECTION DATES:

January, May. june, July, Oetober.

OTHER COUNTRIES, 35 specimens from: COSTA RICA.

REMARKS. [his species has been captured at light: on un-

known, Carrion; amd at: coat, Norse, human, Opossum, and tapir

feces.

Oe OCWENErUS STFIUS f.. Sp.

Figs. 214-216

HOLOTYPE. Male, length 15 mm, greatest width 7.5 mm.

Dorsally shining black. Clypeus (Fig. 214) anteriorly very feebly

Howden and Young: Panamanian Scarabaeinae 123

emaroginate. Frons medially. with small conical tubercle... Prono

tum. moderately punctate Jdaterally, faintly tumid medially at

anterior. fifth, midline tmndentéed: medially. posterior to tumid area.

Elytron: with, six striae between suture and humeral umbone, striac

distinctly impressed and punctatée,. punctures separated by one. to

two diameters. °«Pygidium with’ complete marginal -bead, “surface

punctate... ‘Fore’ tibia: with four ‘distinet teeth oon over maroin;

apical tooth extended forward, nearly paralleling lone “axis .of

tibia. = Hind femur unusually broad, dorsally flatiened-with hind

margin ‘expanded, particularly apically,:the dorsal apical termin—

ation (Fig. 216). acutely angulate. Hind tibia’ with six or seven

lateral serrations, inner. margin withobtuse’ tooth at basal third;

longest tibial spur. longer tham first’two tarsal seaments — Hind

tarsus with basal segment. nearly rectangular. Genivalia’as in

fig. 215.

ALLOTYPE. Female, lengti 157mm, Greatest widii 7/7 am,

Differing from holotype in the followine’ major characters: brown

(teneral), tubercle on frons slightly smaller; fore tibia with

apical tooth with curvature similar to penultimate tooth; hind

femur lacking posterior expansion, a distinctive broad groove

present ventrally,:. paralleling hind, maroin;, hind tibia lacking

obtuse tooth at basal third, tibia more slender; abdominal sternites

less narrowed medially.

TYPE MATERIAL: Holotype, male,-Panama, Canal Zone, Barro

Colorado: Island, 1/7, 1X.19/5,. O.: Youne «(Washingion)..: Allotype,

female, same locality as holotype, 16-20.11.1976, A. Newton, human

dung trap (Cambridge).

PARATYPE: 1° male, same data as allotype (Howden) <

REMARKS. The male paratype, except for the sexual differ-

ences, is. the same size. and color (also teneral) as the-allotype.

We have seen one (or two) closely related species from Napo

Province, Ecuador, and from Distrito Federal, Brazil. The species

name, strius, is based’ on the-initials for We Smithsonian Tropical

Research Institute.

Dichotomius Hope

Dichotomius Hope, 1030, D. G2 sMartines. 1251. 9. 140) (complete

synonymy).

Type-species: Copris boreus Olivier, by original designation.

Pinotus Frichson;’ t647) po 0s.

Dark brown to black species (in Panama); size moderate

(10 mm) to large (35 mm); clypeus usually at least slightly rugose

laterally; scutellum not visible; elytron with eight striae; middle

and hind tibiae gradually expanded to apices, lacking distinct

transverse Carina” at’ euler Third “or falfs middie’ tibia’ with two

Spurs; tarsi with claws present on all legs; mesosternum wide,

middle coxae separated by distance greater than width of femur:

abdomen with six sternites, at least first five not greatly narrowed

medially.

124 Contrib. hmern. sate. imst.s wok. 06, no. 4), 1961

Key to species of Dichotomius

a First and second elytral striae not noticeably

widened in apical third, not noticeably

tomentose; size usually less than 24 mm. . . : 4

At least first and second elytral striae caca.,

widened in apical third, widened area often

brown, tomentose; size usually over 25 mm ‘

1. carolinus colonicus (Say)

v4 Metasternum with numerous setae; clypeus

POUNCE (OF CMa rGLNa lee iar eying ence Saber ge dere voenyse 3

Metasternum with few setae; clypeus medially

POT UGEIY MOLE EG mi ii aunie Gaberyiiier Wie soe be ie aes poe) alive 6

a. Pronotum with median posterior horn(s) or with

median ian hiatle tumid.,or carinate thansverse

UCI Wes hi ee 4

Pronotum lacking a. or ees dise evenly

convex, or slightly’ declivous, anteriorly .... ..« 5

4. First tarsal segment on middle and hind legs

strongly triangular, as wide as long; carina

or horn on head ee et medially . aut

soe satanas (Har.)

Piet co Somers on iadls and hind legs

elongately triangular, approximately twice

as. lone as. wide; carina: on head truncate or

POU Gk Nod ees mevermanni «(hLued.’)

a. Metasternum laterally closely, setose-punctate;

eize moderate ((lO+2k mm). cee i. 4. agenor ° (Har,.)

Metasternum laterally with widely scattered

punctures, setae sparse; size small (12-15 mm)

a Seatnacmis (Erich. )

6. Hind femur with 4:to 10 scattered” punctures on

ventral surface; pronotum with coarse punc-—

tures separated by more than one diameter,

surface between with minute secondary punc-

rures 7. 4 eo remoratie. | Ms Sp.

Hind femur munis in a) half; pronotum

moderately punctate, most punctures nearly

COV eUe a am be@aenets ml. Sp.

1. Dichotomius carolinus colonicus (Say)

Pig. 27

Copris colonica ‘Say; 1635; <p. 174.

Type, Cambridge: or destroyed... Not seen.

Dichotomius carolinus colonicus (Say); Martinez, 1951, p. 140.

MALE. Length 19 to 34 mm, greatest width 14 to 18 mm.

Black, often with the posterior third of discal striae brown,

Howden and Young: Panamanian Scarabaeinae E25

tomentose. Shape very convex, oval. Male majors (Fig. 217)

with short upright clypeal horn central (at middle of clypeus),

clypeus not or feebly emarginate. Pronotum pumetate,.. with

rounded, medially indented transverse swelling at posterior third;

a short, abrupt carina present just anterior to lateral pronotal

fovea; posterior marginal bead complete. Elytral striae fine and

with small punctures anteriorly, inner,two or more striae widened

in apical third, tomentose. Metasternum laterally and along

margins between middle coxae with numerous’ lone setae,

FEMALE. Shape and variation in size similar to male,

differing in the following major characters: low, transverse,

apically truncate horn or tubercle present on iweriex. Transverse

pronotal swelling more distinctly trilobed and positioned in

anterior third of pronotum; lateral: pronotal. carinae obsoicte.

Usually Tirst five striae on each elytron posteriorly temeniose.

MATERIAL EXAMINED. PANAMA, 59 specimens from: CANAL

ZONE = -Frijoles..: CHIRIQUL) PROVe+) Hartmann’ s Tanca,: 12700 m,

1300 m, 4500: ms Hato’ del. Volcan: Lags-Lagumascy. Los Vorcs: Talo

Santo; Santa “Claras) COCLE \PROVale Ubli Valle. CORON: PROV eos

Santa Rita Ridge... PANAMA: PROV .=) Cérypo. ‘Campana, (| COLLECTION

DATES: May, june. July, August, Novemuer.

OTHER COUNTRIES, 236 specimens from: UNITED STATES,

MEXICO; BELIZE, «GUATEMALA, El. SAUVADOR. COSa A, HCAs

REMARKS. In small specimens the horn on the head (in

either sex) may be ereatly reduced and the’same as true forthe

transverse: pronotal..swelling. Generally the large: size..and: the

two to five brown tomentose strial areas in the posterior third of

each elytron will readily distinguish colonicus from other Panaman-

lan species in the genus.

Dichotomius) c.: colonicus has: been. captured in vlight traps,

and at horse dung and human feces. It is most abundant in the

early wet season and is found mostly at moderate elevations in

forested areas.

2. Dichotomius nevermanni (Luederwaldt) New Combination

Fiey 218

Pinotus nevermanni Luederwaldt, 1935, p. 334.

Type, London (?).. Not seen.

FEMALE (only specimen, seen). Length 20 mm, greatest width

12.5 mm. Black, similar in shape and pronotal configuration to

D..c, colonicus, 9 Differing trom, colonmicus in the following major

characters. Head (Fig. 218) with low transverse carina extending

across anterior edge of frons, carina nearly straiehty of uniform

height, ‘vaguely tuberculate at lateral termination. Pronotum with

four swellings, one on either side of midline at median half, one

on each’ side above lateral fovea. Elvival stria: simple, feebly

punctate, first to fifth. striae, not widened: apically, lacking

tomentose areas,

MATERIAL EXAMINED. PANAMA, 1 specimen from: PANAMA

PROV. - Bl Malle, (134%). 2068. lonla« Bottimer, at: Meht, (Ottawa).

126 Contrib. Amér. Pnt.Imst:, vol 16,0ne.-?,° 1981

REMARKS. The single female specimen is similar to small

colonicus. In his description of nevermanni, Luederwaldt (1935)

based his description on a single female from Costa Rica. In his

five line description he related the species to carolinus (colonicus).

He stated that the metasternum lacks setae while in our female

the metasternum has setae. Since setae can be lost (by abrasion,

etc.) we are utilizing Luederwaldt's name. Only when more

material becomes available and the type is examined can the name

be. applied with certainty.

2. Dichotomius satanas. (Harold)

Fro. 219

Pinotus’satanas Harold, 186/b, p. 98.

ype or cotype, Paris: Nol’ seen.

Dichotomius satanas (Harold): Halffter and Matthews, 1966, p. 83.

MALE. Léenoth 17 16°23 mm, greatest width 10 to 13. mm.

Brownish black to biack; ‘shape elongate oval, very convex.

Clypeus rounded or very feebly emarginate; clypeal horn (Fig.

219) transverse, anteriorly—posteriorly flattened, apically rounded,

positioned “medially im center or clypeus. Pronotal disc, except

for lateral third, flattened; posterior edge of flattened area feebly

swollen (male minor) or with a median, slightly anteriorly bent

horw (Fie. 219) and a small tubercle on cach side anterior ‘to

fifth stria; pronotal surface granular. Elytral striae shallow,

moderately fine, punctures small; elytral intervals feebly convex,

finely rugose or granulate. Metasternum with numerous long

setae except near smooth, vaguely impressed median area.

FEMALE. Variation in size similar to male. Differing in the

following major characters: vertex with small conical horn;

pronotum with distinct bilobed median swelling, also on each side

near base a small tubercle or swelling usually present (in larger

females).

MATERIAL EXAMINED. PANAMA, 89 specimens from: BOCAS DEL

TORO PROV. - Rie Changuinola. CANAL ZONE — Barro Colorado

Island. -CHIRIOQUL PROV. — Hartmann s Binca, 1200 m, 1300 m,

1500 m:; bas Lagunas. COLON PROV. — Santa Kita Ridge. DARIEN

PROV. ~ Cerro Jacarcuna, G600 ft. PANAMA PROV. — Cerro’ Campana,

Aitos (isla) de.Maje. COLLECTION DATES: Every month of the year.

OTHER COUNTRIES, 157 Specimens from: MEXICO, BELIZE,

GUATEMALA, COSTA RICA, “COLOMBIA, ECUADOR, PERU.

REMARKS. In both sexes the horn on the head and the pro-

notal modifications may be greatly reduced in small specimens.

The granular pronotal surface should suffice to distinguish the

species from other members of the genus occurring in Panama.

This species has been recorded at rat and unknown carrion,

and on ¢oéati, human, spider monkey, and tapir feces. “On Barro

Colorado Island, satanas is nocturnally active, particularly during

the periods of 1800 fo 2000 hours and from 2400 to 0600 hours. <It

has been recorded for all months except March, July, and October,

with relative densities similar throughout the year. Teneral indi-

viduals have been recorded in April and November which suggests

Howden and Young: Panamanian Scarabaeinae tay

continuous reproductive, activity. « lt occurs, ino meist forest al all

elevations throughout Panama.

4. Dichotomius agenor (Harold) New Combination

Pigs. 220, 221

-Pinotus agenor: Harold, :1069c, pp. 441;

Type or colype, Paris.) Not seen.

MALE. “Length: 15 to 19 mm, greatest 10) 10 16 127i.

Black, shining; shape (Fig. 220) oblong-oval, moderately convex.

Clypeus broadly, shallowly emarginate; horn on vertex (Fig. 221)

extending transversely between anterior margins or eye, Dacal

third of horn broad, rising vertically, then abruptly angled (90°)

inward on each side to narrow, nearly cylindrical continuation: oF

horn; cylindrical two-thirds of horn slightly tapered to vaguely

binodose or. narrowly: truncate. apex; posterior basal portion of

horn shining, at most finely punctate. Pronotum moderately cde—

cliveus in anterior third, summit. rounded; pronotum: otherwise

evenly convex; Surface: minutely, punctate, appearing nearly

smooth, shining: posterior’ marginal bead: complete, the Indenia—

tion containing a4 row of punctures. Elytral’ striae shallowly

impressed, fine, seen as a double line only at meh magnification;

strial punctures vague, searcely wider than: Stria; elyitral imtervals

feebly convex, ‘surface nearly Smooth, shining. — Melacternum

laterally and adjacent to middle coxae with lone selae: medially

smooth, midline vaguely impressed.

FEMALE. Size> and shape similar’ to mate. Dilferins in ithe

following ‘major. characters: horn: om vertex small,, low, ‘Comical,

feebly ‘bidentate; pronotum slightly tumed im” anterior fourth behind

head, midline posteriorly Teebly tnpresced.

MATERIAL EXAMINED, PANAMA, G4 specimens’ trom: «CANAL

ZONE =, Ancon?) Barre Colorado: lsland: Fort Kopbe:, COCLE cPROV. =

El Valle; Rio: Hatow PANAMA PROV. = Bic La xlagua.). COLLECTION

DATES::: May, June; baly., August. eptember, October.

OTHER: COUNTRIES. 4 specimens trom: COsl A RICA, «COLOMBIA.

REMARKS. The only major variation noted is the reduction of

the: size and. development of sthe Dorn onthe Vertex im small, mates.

The minute’ punctures on tne pronoium and fine elytral striae are

quite constant and should serve to identify Panamanian specimens.

There 1s one very: closely: related. species that eceurs:. from

Mexico to Costa Rica, D. centralis (Harold), and there is a strong

likelihood that it may be taken in western Panama. It differs

from agenor im the tollowine ¢heracters: posterior’ base of horn

rugose in males, pronctum moderately and closely punctate, elytra!

striae wide, strial punctures contiguous’) and making ‘sides of striae

appear crenulate.

Both species are attracted to carrion and dung.” "The usual

habitat of agenor is lowland pastures and clearings.

128 Contvibs (Amer. Butictmstsy vols, no. Ol, d9SN1

5. Dichotomius inachus (Erichson) New Combination

Mio.) 222

Pinotus inachus Erichson, 1847, p. 108.

Type; Berlin (7)... Not seen.

MALE. Length 11 to 13 mm, greatest .width 7 to 8 mm. Black,

largely shining, shape oblong-oval, not strongly convex. Clypeus

feebly to moderately rugose-punctate. Frontal tubercle (Fig. 222)

conical, feebly indented apically... Vertex finely punctate.

Pronotum evenly convex, minutely. punctate taterally, smooth, other—

wise. Elytral striae moderately impressed, each stria delimited

by narrowly separated double line, lines broken by very shallow,

poorly delimited punctures. Pygidium smooth, marginal bead not

broken. Metasternum setose between middle coxae, sparsely so

laterally, lateral portions with scattered punctures separated by

two Or mone diameters, Middle ieémur with three to.five. setigerous

punctures on ventral apical surface.

PEMALE: Similan. in Gize and torm. to male, clypeus. medially

motched, (roniel tubercle. obcolete. Otner dilferences, minor.

MATERIAL EXAMINED. PANAMA, 1 male, 2 females from: COCLE

PROV = Rio Hato, 27 7.1950, Viol. tipton,..at liaht.

OCBER COUNTRIES, 25 specimens, trom: PERU. TRINIDAD,

REMARKS. The tew differences, in. the, Panamanian.,.specimens

have been included in the description. Specimens from. South

America vary more in size and the punctures on the metasternum

are Otten larger and more mumerous.. [here 18.4. complex of species

involyed and the imachiis croup meeds 4, thorough revision. betore

Hames canbe applied with certainty.

The relatively small size and smooth pronotum should dis-

tinguish inachus from other Dichotomius known to occur in Panama.

Titig species ‘has been. collected in. Panama, at light,.and at

human: feces in Trinidad,

6. Dichotomius femoratus n. sp.

Figs. 223-226

HOLOTYPE. Male, length 14 mm, greatest width 8.4 mm. Head

brownish black, pronotum and elytra black. Shape oblong-oval,

moderately convex. Clypeus bidentate with V-shaped notch between

teeth, clypeal surface punctate, less so medially. Vertex (Fig.

224) wiih, Tow conical shorn... Gena, and. posterior. third. of . vertex

with scattered coarse punctures. Pronotum with anterior third of

pronotal convexity behind head feebly concave, summit rounded,

fubercdies Lacking. midline, Of. Dronotum, posterior,.to .concavity,.im—

pressed; entire surface of pronotum with scattered coarse punc-

tures, very minute secondary punctures interspersed; posterior

marginal bead represented: Dy an,irregular double row of ocellate

punctures. Elytral striae tmpressed, each represented. by (a single

line or nearly contiguous double line enlarged by small punctures

beparated by two to three diameters; on. either side of each .stria

(in fresh, unworn, clean specimens) surface longitudinally finely

shagreened, opaque, the opaque area occupying approximately one-

Howden and Young: Panamanian Scarabaeinae 129

fifth: of »width cofinterval; velytral c<intervalsjeconvex, centrally

smooth and shining, occasionally with scattered minute punctures.

Pygidium with marginal bead obsolete near apex, surface with

scattered coarse punctures. Metasternum lacking setae, coarsely

punctate laterally and along margins between middle coxae.

Middle and hind’ femora nearly impunctate, 2 to 10 punctures

present in vapical thirds; femora each with posterior eroove

‘paralleling (hind margin, groove becoming obsolete near trochanter.

Abdominal sternites each with basal row of punctures extending

across midline except obsoletev'on apical: segment. "Genitalia ‘as in

fies.) 22550 226.

ALLOTYPh. Female, .leneth :i4.5)umm,) ereatest) widen; 9 mim,

Differing from male in following respects: clypeal surface (Fig.

223) moderately rugose, feebly so medially; vertex with low,

poorly defined obtusely angulate swelling, median anterior angle

of swelling anteriorly dissected, slightly more elevated and feebly

binodose; :pronotum jevenly convex except for ieebly impressed: mid—

line, more distinctly impressed posteriorly, pronotal surface

coarsely punctate, vaguely ocellate punctures’ clustered: along \mid-—

line; abdomen with sixth (apical) sternite with feeble median

swelling.

TYPE . MATERIAL: Holotype, mate; Renama yw Canal Zone, Barro

Colorado Island, 8.V1I.1978, N.E. Woodley, in trap baited with

Lepidoptera frass (Cambridge). Allotype, female, Panama (Panama

Prov.), Cerro Campana, 3000):ff.. GOV A970 nie eo wi Plowden

(Howden).

PARATYPES,: 3: females:):li; samevdata (as iellotype:) 2, .’same

locality as holotype, 13.V111.1978, H. Hespenheide, 20.V1.1979,

TCs POUrson,.

Paratypes ‘are inéiHowden, Poulsen,

REMARKS. The female paratypes measure 13.0 to 13.5 mm in

length and 8.5 mm in greatest width. In one paratype all

sternites exceptuthe basalione havewan extremely teeble: swelling or

irregularity atthe midline. Otherwise» the paratypes closely

resemble. the’ allotype,

The species: should. probably «be placed: near the: elobulus

complex, but the largely impunctate middle and hind femora with

the distinctive posterior groove separates) it trom this’ group. | The

coarsely punctate pronotum and very fine elytral striae and small

strial punctures are also quite distinctive. The unusual opaque

area on either: sidevol) the elytral’ striae whichis only ‘evident! in

fresh, clean specimens, is seemingly a unique character, and

distinguishes femoratus from related species in Costa Rica and

Colombia.

The two female: specimens from Cerro Campana were collected

on human. feces,

7. Dichotomius gamboaensis n. sp.

Figs. 227-229

HOLOTYPE. Male, length 11.5 mm, greatest width 6 mm.

Black, relatively dull; shape oblong-oval, not strongly convex.

Clypeus bidentate, V-shaped notch between moderately reflexed

130 Contrib. Amer. Entice Insts; volicld, ne. sho .'96!

teeth; clypeus Teebly -rugose laterally, coarsely punctate elsewhere

except near midline. (Prong (Pio. 227) with low, rounded ‘tubercle:

Vertex “moderately; closely punctate a. anterior halt, nearly im-

punctate posteriorly. FPronotum evenly. convex, lacking carina’ or

swellings, surface closely punctate, punctures of two types,

moderate’ sized and ocellate wilh interspersed, minute punctures;

posterior marginal bead represented by irrecular double: row of

coarse punctures. Elytral striae moderately impressed, each

Stria with feeble double line, broken. by single.row of small punc-

tures; elytval intervals moderately convex, dull, with “scattered,

minute shiny spots. Pygidium with complete marginal bead (line),

moderately punctate, more closely so Dasailly.. Metasternum lacking

setae, moderarely, Getisely puncvaie except along midline. Middle

and hind?femora, with distinc! Groove paralleling posterior edge;

middie: femur punctate in apical third, hind, femur*punctate: in

apical two fifths. Basal five abdominal sternites with basal row

of punctures extending across midline. Genitalia as in figs. 228,

229; dorsal lateral; edoes of parameres finely serrate.

ALEOUYPR. Hemale,: tength 12.5, mm, ereatest width 7 > mm.

Ditfering: from Holotype in the following major characters: clypeus

more abruptly notched between teeth; elytral striae feebly im-

pressed, intervals: fecbly convex; find temur closely punctate in

apical. Nall, scattered punctures: elsewhere.

TYEE GMATERIALS Teloiyoe, malic, Panama, .Canal. Zone, Gamboa,

Limbo Hunts Club, 237248 .1975,, O,P.. Young, tapir. feces: trap

(Washington). Allotype, female, Panama, Canal Zone, Gamboa,

23-24.X%.1975, O.P. Young, human feces trap (Washington).

PARATYPES: 3 males, 5 females; 4, same data as holotype;

3, same data as holotype except collected 28.1V.1975; 1, Panama,

Canal Zone, 5 mi. NW Gamboa, 9°09'N,; 79°249'W., 29.1K.1969, HA.

Hespenheide.

Paratypes' are in: Ottawa, Washington, Howden.

REMAKKS. The smatlest specimen measures li.mm, the largest

13 Mm. ~ Severavoclemales: have the tubercle\on the ‘vertex feebly

indented. Olherwise the series is relatively uniform:

Dichotomius’ gambogensis is: seemingly closely related to D.

globulus (Feleche). tt differs: from this species: in having the

punctures iIm-ihe celyival striae and of. the pygidium. less! pronounced,

the punctures on the middle and hind femora smaller and more

apically concentrated, and the genitalia wider: apically and with

the apices of the parameres bent downward.

Copris Muller

Copris’ Muller, 1764, ©. [1;. Matthews, 1961; p..-35.

Type-species: Scarabaeus lunaris Linnaeus, by subsequent

designation, Curtis, 1832.

Usually black, size moderate (12 to 24 mm); scutellum not

visible; elytron with nine striae (10 counting inflexed margin);

mesosternum short, transverse, with median area iMpunCtate; meso-

metasternal suture obtusely angulate; mesosternum between middle

coxae parallel-sided, lacking setae; abdomen with six distinct

sternites, at least basal five not greatly narrowed. medially,

distinct

Howden and Young: Panamanian Scarabaeinae

at midline; hind tibia gradually expanded to apex, with

transverse carina near middle on outer surface; all tarsi

with claws.

Key to species of Copris (after Matthews, 1961)

MAJOR MALES

gee

FEMALES

Ase

Median pronotal prominences, when present,

transversely truncated; lateral pronotal carina

absent; lateral pronotal margin evenly curved

Median pronotal prominences rounded or acute;

lateral pronotal carina present; lateral pro-

notal margin sinuate or ne! Papeee

near anterior third : : lee

Head horn arising approximately from middle of

head: surface: tubercle behind if erect or some-

times bent (never inclined) forward; dorsal

edges of median pronotal prominences, when

seen from the front, sloping down laterally at

an angle of about 45° from horizontal; © size

TASUG nits Wie eon ‘ ig Pes Le abris

Head Horn arising fon Shelore middle of head;

tubercle behind horn inclined forward; dorsal

edges of median pronotal prominences, when

seen from the front,“almest: horizontal; size

5 2 es cc eran: MM Ra Nig oC rol WE I M/E lo ag Hee

Anterolateral angle of pronotum with point made

salient by an inward curve or emargination

of the margin immediately behind; eighth

elytral stria nearly always complete; inner

border of grabs hie lh incomplete: ‘size

12-16 MO oe . 3. costaricensis

Anterolateral aioe! of protiotan subquadrate,

obtusely angled, or broadly rounded, occasion-—

ally acute in strongly developed specimens,

never with a sharp inward curve of the margin

immediately behind; eighth elytral stria incom-

plete, disintegrating posteriorly or largely

effaced; pygidial margin complete or obsolete

apically, sizevl7-24 mm oo7.05 "A.>subpunctatus

Lateral pronotal carina absent; lateral pronotal

margin evenly curved; pygidial margin incom-

plete, the inner border effaced ventrally .

Lateral pronotal carina present; lateral pronotal

margin sinuate or slightly angulate; pygidial

margin complete: or not iba ota an eae ee ea

Boh.

Say

Gahan

Gill.

132 Contrib. Amer. sEitis met.) wok lo. mo. hs uo PO8t

Ds Anterior portion of pronotum with surface before

and behind transverse swelling distinctly punc-

tate; punctate midline usually absent in anter-

hor thirds sizeyt5e1o mm a OT, lugtbric' Boh.

Anterior portion of pronotum on and anterior to

transverse swelling at most finely punctate;

punctate midline usually extending almost to

transverse swelling; vsize 146 mms. 2 incertus «Say

or Anterolateral angle of pronotum with point made

salient by an inward curve or indentation

of lateral margin immediately behind (some-

times indistinct), margin often sharply sinuate

anteriorly; eighth elytral stria nearly always

complete; inner border of pygidial margin

incomplete; size 13-15.5 mm... 3. costaricensis Gahan

Anterolateral angle of pronotum subquadrate,

obtusely angled, or broadly rounded, never

with sharp inward curve or margin immediately

behind; eighth elytral stria incomplete, disin-

tecrating «posteriorly: pyeidial margin complete

or obsolete: apically::size 22-25 mm . ene

pe A. subpunctatus Gill.

1. COpris lugubris Boheman

Pig 230

Copris lugubris Boheman, 1856, p. 42; Matthews, 1961, p. 49.

Type, stockholm, (Not seen.

Length 14 to 18 mm. Well-developed males (Fig. 230) with

slightly posteriorly directed arcuate horn arising from middle of

head; immediately behind an erect or anteriorly bent tubercle

(Fig. 230), the tubercle never inclined forward (see incertus).

Male median pronotal prominences approximate, broadly, trans-—

versely truncate. Anterior pronotal angles obtuse in both sexes.

Female head with short transverse carina or with narrow trans-

verse horn posteriorly excavated. Pronotum with two low rounded

prominences. For complete description see Matthews, 1961.

MATERIAL EXAMINED. PANAMA, 134 specimens from: CANAL

ZONE = Barro Colorado Island: Cocoe..Solo: .Fort Kobbe::Madden Dam:

Paraiso; Pipeline: Road. .CHIRIQUL PROV. = (David; Hartmann's

Finca, 1300 mr Woimeribios; Volcan, de (\Chiriquix.. COCLE RROV. =

Anton; El) Valle, PANAMA FROV.:— Cerro Campana; La Chorrera;

Maje: Ric La jagua; Rio Las Lajas. VERAGUAS PROV. ~ San Pedro.

COLLECTION DATES: Every month of the year.

OTRER COUNTIES, 202 Specimens from: MEATCO, BELIZE,

GUATEMALA, EL SALVADOR, HONDURAS, NICARAGUA, COSTA RICA.

REMARKS. This species\has been collected at light, and on

cow, horse, human, and tapir feces. On Barro Colorado Island

lugubris 16 active ‘at, night, particularly during the period 2100

to 0400 hours. It is present throughout the year with no signifi-

cant peaks of abundance. The species occurs in forested areas at

low and middle elevations.

Howden and Young: Panamanian Scarabaeinae 133

2. ‘COpPrisiiinecertws Say

Hee 2

Copris incertus) Say, !4835,) Dp.) 175s0 Matthews, 196) s/c an.

Type, Nambridge. “txamined,

Copris ‘procidua’ Say, 1835.9). jos) Matthewew 196n) and:

hype, Cambridge.) Examined:

Length 14 to 19 mm. Well-developed males (Fig. 231) with

slightly curved, tapering’ clypeal: horn: situated) anterior: to middle

of head; immediately behind an acute tubercle (Fig. 231) strongly

inclined anteriorly, almost recumbent. Pronotum with median

pronotal prominences low, transversely truncate anteriorly, approx-

imate (never as well developed as in male major lugubris).

Anterior pronotal angles obtuse in both sexes. Head of female

usually with narrow, transverse, truncate horn apically excavated.

For complete description see Matthews, 1961.

MATERIAL EXAMINED. PANAMA, 23 specimens from: BOCAS DEL

TORO PROV. -— Rio Changuinola. CANAL ZONE - Barro Colorado

Island. CHIRIQUI PROV. - Alanje; Hartmann's Finca,. 1300 m.

COCLE PROV. - EL Valle. COLON) PROV i= Sante (Rita, Ridge:

PANAMA PROV. - Cerro Campana; Maje. COLLECTION DATES:

January, February, March, May, June, July, August, September,

December.

OTHER COUNTRIES, 17. specimens, Troma. Mex PCO. BE Lig ee ue |

SALVADOR, COSTA URTCA, (COLOMBIA,

REMARKS, This. species has, .been, captured) at light, and. on

cow, horse, human, and tapir feces.

3. Copris ‘costaricensis 'Ganan

Pigs 2a

Copris costaricernsis, Gahan,) 1094) "po (1G: Matthews.) 1901. 5.) On.

Type, London. Not seen.

Copris’ furcillatus Pelsehe (1910, ou Sa echt tr hewe i 1 OGT plie

Type, Dresden. HOihlor seen:

Length 12: to) 16mm.) Male majors (Fie 232))) with ‘long,

slightly bent, tapering head horn; median pronotal prominences

merging into single bifurcate process. Both sexes with anterior

pronotal angles acute, adjacent margin often sharply indented;

pygidium with margin (bead) incomplete, inner border effaced

ventrally. For complete description see Matthews, 1961.

MATERIAL EXAMINED. PANAMA, (records from Matthews, 1961):

CHIRIQUL PROV.’ — Potrerilles;Volean de Chirigui, 2500-4000 ft.,

6-8 June, 1882.

OTHER? COUNTRIES, 2 (specimens irom: COST A\ RICA.

REMARKS. The species had been taken on carrion, cow dung,

and atilient.

oa 6. Contpip. Amer. Eni. [nst., vol. d6,. no.) ,. 1981

4. Copris subpunctatus Gillet

Fio., 233

COpris aSpericollis yar. euppunctatus Gillet, 1910,;. p.< 3

Type, Brussels. Not seen.

Copris subpunctatus Gillet; Matthews, 1961, p. 82.

Length 1/7 to 25 mm. Male majors (Fig. 233) with head. horn

massive and sharply bent at middle; pronotum humped behind

median pronotal prominence, prominence divergent in dorsal view.

Anterior pronotal angles acute in large specimens, rounded in

small ones. Pygidium of males sometimes with margin broken near

apex; margin always complete, unbroken, in females. For complete

description see Matthews, 1961.

MATERIAL EXAMINED. PANAMA, (records from Matthews):

CHDRIOUL, PROV... Chiraqui.

OTHER COUNTRIES, 5 specimens trom: COSTA RICA.

REMARKS. According to Matthews (1961) all specimens with

Gata were taken from. October to June at light.

Phanaeus Macleay

Phanacus Macleay, 1019, p. 124; Edmonds, 1972, p. 826 (synondomy

and species groups).

Type-species: Phanaeus vindex Macleay, by subsequent desig-

nation, Edmonds, 1972.

Panamanian members of the genus may be recognized by the

following combination of characters: clypeus usually with two

obtuse teeth separated by single median indentation; head of males

with posteriorly curved, nearly cylindrical horn; pronotum with

anterior marginal bead interrupted behind eye; pronotum of males

flattened, posterior angles often produced, flattened area of prono-

tum triangular in shape; both sexes frequently brightly colored

(green to reddish purple), dorsum rarely entirely black.

Key to species of Phanaeus

ie Elytral striae with large, distinctly separated

shallow punctures; color often cupreous to

reddish purple.) «>. fee ee hermeca tar.

Elytral striae lacking ee cee punctures;

color greenish or with cupreous pronotum and

Digee Clee a igi ee Clee ewe 2

2 Pronotum except in median posterior half or less

distinctly granulate or en dorsally uniform

e@recn or Diuc. «a .% pose Qe Deliianus Bates

Pronotum mostly a eee finely granu-

late, punctate or feebliy rugose laterally;

head, pronotum and pygidium often cupreous

Or. regaich ereen, Clyira Diack . . «3: pywois Bates

Howden and Young: Panamanian Scarabaeinae 135

1. Phanaeus hermes Harold

Fig. 254

Phanaeus hermes Harold, 1868, p. 83.

Coiypes, Paris. Not seen.

MALE. Lenoth 13°16 18 mm, ereates: witen 9 70 EF min.

Dorsally green to cupreous, sometimes reddish to purple. Well-

developed males with long, posteriorly curved horn on head.

Pronotum with flattened area anteriorly with small tubercle on

each side behind eye, posterior angles produced upwards and with

outer margins incurved; pronotal suriace distinctly granulate;

posteriorly, near .marein. with distinct fovea on ei{fer side of. mid-

line. Elytral stria (Fig. 234) with Gistinciive large punctures,

giving elytra slightly wrinkled appearance, intervals lfeebly convex,

are ale

FEMALE. Length 12 to 19 mm, greatest width 8 to 12 mm.

Differing from male, in the totlowing major Characters: mead with

low, anteriorly arcuate carina extending across frontal area,

carina .terminating on Gach side just anterior (Oo eye, pronoium

anteriorly with small, arcuate, tumid carina, (the Carina thickened

at midline and often with small concavity, benind, suriace irregu-

larly granulatée-rugose, becoming punctate. on disc in posterior

fourth.

MATERIAL EXAMINED. PANAMA, Zl specimens from: CANAL

ZONE’ — Ancon; Chiva Chiva; Empire. «Tn RiOUL PROV. — Bugana.

800-1500 ft.; David. PANAMA PROV. - Cerro Campana; Las Cumbres.

COLLECTION DATES: May,. June, July, Auguct, September, December.

OTHER COUNTRIES, 3 Specimen¢e Irom: (Ole RICA. COLOMEIA.

REMARKS. .Size is unusually Variable, particularly wher one

considers the short: series ‘seen, Small males may have the head

horn reduced to an arcudte carina and the pronctum almost evenly

convex. Small females may have the pronctal carina and concavity

obsolete. Size also .influences the size of the pronotal oranula-—

tions, but the. characteristic larvae punctures in tne elyiral striae

dre quite consistent and Serve to distinguish hermes. from related

Species Occurring in Panama.

The few specimens bearing accurate dates have been collected

at human feces.” Phangéus hermes.is diurnally active and occurs

only in the wet Season. The species is found in mid-elevation

moist forests.

2. Phanaeus beltianus Bates

Pigs s 2354 , 240

Phanaeus beltianus Bates, 1887, p. 63.

Cotypes, London and Paris. London specimens examined.

MALE. Length 15°46 10 > mm, areatest width 10mm.” ‘Color

green with cupreous reflections on pronotum. Head with long,

cylindrical, posteriorly curved horn. -Fronotum. (Fic. 235) with

flattened central area delimited anteriorly by conical tubercle on

either side about | mm behind margin posterior to inner edge. of

eye; POSlterior portion Of Tlattened area delimited laterally by

136 Contribs Amer, rate inet... wollte now deisel

inwardly arcuate ridge terminating abruptly on each side in an

acute point anterior to fourth ebviral siria, suriace of Tlattened

acea laterally and in anterior hal! transversely rugose, posterior

central half with nearly smooth triangular area bounded posteriorly

oneach: side by inner termination of arcuate ridge, narrowing

anteriorly to small tubercle at midline méar transverse median

half of pronctum; lateral declivous sides of pronotum finely punc-—

tate; fovea on each side of posterior midline absent. Elytral

striae. larcely ,impunctate, anteriorly terminating in distinct

indentation; intervals feebly convex, smooth, moderately shining.

Pygidium finely punctate. Genitalia as in fig. 236.

FEMALE. Length 14 to 18 mm, greatest width 10 to 11 mm.

Differing from male in the following major Characters:. color

dorsally. ereenish bine,.to Dlue; vertex with straight. transverse

Carina, the carina .trituberculate;s pronotum convex except concave

immediately posterior to short transverse arcuate or obtusely

angled carina behind head, posterior margin of concavity sinu—

ately tumid; surface of pronotum finely rugose, becoming finely

puUnCtaITe in median posterior, fiith.

MATERIAL EXAMINED. PANAMA, 18 specimens from: CANAL

ZONE — Barro Colorado Island» Chivos; Gamboa; Madden Forest;

Pipeline Road, PANAMA PROV. + 6.km Sk Bayano Bridge, 9°10'N,

FOC4O WW. (COLDECTION DATES: May, june, July, September,

October, November, December.

OTHER: COUNTRIES, specimens from: BELIZE, NICARAGUA,

COSTA. RECA. |

REMARKS. The above description was based only on Panaman-

ian specimens, which differ slightly from those from Belize and

Nicaragua. Typical beltianus (from Nicaragua) are often more

olive green, the male pronotum more strongly rugose, the small

anterior conical tubercles slightly more widely spaced and the

smooth pronotal area smaller and rhomboid in shape. Generally

the Panamanian specimens differ from typical beltianus in approx-

imately the same degree as beltianus differs from sallei. As

Edmonds (1972) has stated, the entire complex needs further study.

This species has been taken in bat mist nets, and at

howler monkey, Numan, spider monkey,.and tapir. feces.

On Barro Colorado Island, beltianus is diurnally active,

restricted as adults to the wet season, and more common in level

areas than in ravines. The species is limited in central Panama

to lowland moist forests.

3. Phanaecus pyrois Bates

Hig. 237

Phanaeus pyrois Bates, 165/, 0. 503 Olsoufiell, 1924,°p.° 92.

Cotypes, London and Paris. London specimens examined.

MALE. Length 4 to 16.mm, ereatest width 9 to 11 mm.

Head, pronotum and pygidium largely cupreous or greenish

cupreous; elytra dull black. Head of well-developed males with

long. eVlindrical, posteriorly curved hora. | Pronotum (Fig. 237)

with flattened triangular portion smooth, often finely punctate or

Howden and Young: Panamanian Scarabaeinae 137

oranulate laterally, posterior ‘angles: produced Taterally) not

noticeably elevated; pronotal) fovea on either side vor midline

adjacent’ to’ posterior maroin’ smallto obsolete. ( Elytralyintervals

feeblyconvex,;;dull’ Pysidium’ finely, ‘closely punctate, less: So

apically.

FEMALE. ‘Length 16 to 17 mm, greatest width 10) mm.” Differ—

ing from male in the following major characters: cupreous reflec-

tions present largely ‘on posterior ‘portion ‘or (head and laterat

third of pronotum; frontal area of head with low, slightly arcuate,

feebly trituberculate ‘carina: pronotum larcely convex, three’ small

tubercles present’ in transverse Tine vat anterior fourthor pronotum

behind: head; pronotal surface smooth to; finely punctate iarerally.

MATERIAL EXAMINED. PANAMA, 90 specimens from: CANAL

ZONE - Barro: Colorado’ ‘Island, Frijoles,: (Madden Forest, Pipeline

Road. COLON PROV.= Santa Rita Ridges’ (PANAMA PROV.) =) Cerro

Campana; 3000 m; Maje. COLLECTION DATES: January, February,

March, May, June,) July,’ August, September,’ October, November,

December «

OTHER COUNTRIES, (20 specimens! from: (NICARAGUA Wi COSTA

RICA, COLOMBIA.

REMARKS.:Small” males))ias’ eam bevex pected) may have tne

horn..on the head greatly reducedin) length and ithe triangular,

flattened area reduced in’ size... The cupreous’ color of the head

and pronotum may also be mostly lacking. A totally black form,

P. funereus Balth., is a very closely allied)\species or: wariety

occurring in Ecuador.

This’ species: has’ ‘been obtained: from: rotting: "truit\or Gustavia

sp.3\from;agoutl, lizard, >) mouse, vat, and tamandus carrion: and

from coati, howler monkey, human, and spider monkey feces. On

Barro Colorado Island pyrois is active diurnally and is most

abundant in the late wet season.

Sulcophanaeus Olsoufieff

sulcophandeus Olsoutieti TO im oor ROMO Ole ie oa oe.

Type-species: Scarabaeus sulcatus Drury (= carnifex L.), by

original designation.

The genus as defined by Edmonds (1972) is composed of three

species, groups, Only one emiwhicn. (the auricollis oroun, occurs in

Panama. The three Panamanian species in the group may be

recognized by the following combination of characters: clypeus

shallowly emarginate medially, at most feebly bidentate; anterior

lateral pronotal margin between eye and lateral angle sinuate,

marginal bead not broken behind eye; elytra dull black, intervals

very slightly convex; male majors with long, recurved horn on

head; females with distinctive sinuate V-shaped carina on pronotum

behind head.

Key to species of Sulcophanaeus

Ls Pygidium and legs largely black; pronotum of

major males with a low, broad, obtuse projec-—

tion pesterioriy on. ‘either side of midline. 4) 4) 2

138 Contrib. Amer. “Entecinst., wvweleclb 0.71, .1961

Pygidium and legs largely cupreous to greenish

red; pronotum (Of major> mates with a lone;

cylindrical anteriorly direcred horn on. either

side of midline: 2 oti G7. 20.0 do velutimus. (Murr: )

2 Prenctdin Uliek (ls ee a Bo nectis (Bates)

Pronotum greenish to cupreous® : 112. cupricollis (Nev.

1. Sulcophanaeus velutinus (Murray)

Fig. 236

Phanaeus velutinus Murray, 1856, p. 213.

iype;, London. - Examined:

Sulcophanaeus velutinus (Murray); Edmonds, 1972, p. 821.

MALE. Length 16 to 20 mm, greatest width 8 to 11 mm.

Black with margin of pronotum,) femora, and pygidium often

Cupreous 0 Orcem. Erontal area of nead of well-developed imales

with lone, nearly cylindrical Norn, arcuately bent posteriorly

near apex. FPronotum (Fig. 233) concave in anterior. half: behind

Mead; a lone, anteriorly directed cylindrical horm:on each side of

midline at summit of concavity; pronotal surface smooth to obsoletely

punctate laterally; pronotal fovea near. posterior midline vague to

moderately developed. Elytral intervals nearly flat, frequently

with transverse wrinkles. |

PEMALEE. Leneth 17 to 19 mm, ereatest. width. lO-to 11>mm:

Differing from males in the following major characters: frontal

ridge vaguely carinate, ‘feebly trituberculate; pronotal carina as

described for Genus; pronotal surface largely: punctate, many

punctures coalescing.

MATERIAL EXAMINED. PANAMA, 20 specimens from: CHIRIQUI

PROV) 2.2 bre N Santa Clara (]=:15%km W Hato del Volcan}; 1300: m,

Hartmann’ s Pinca: COLLECTION DATES» May.

OTHER COUNTRIES, 2 specimens from: COSTA RICA (COLOMBIA

and ECUADOR cited in literature).

REMARKS. Males vary allometrically, small specimens having

both the, head..and pronotal Norns greatly reduced. [n other

respects Panamanian specimens show little variation, the pygidium

at least always showing some metallic (green or cupreous) reflec-

tions.

Tie epecies Mas been taken in nearly equal numbers at feces

and Carrion trays anc ceems to be primarily nocturnal,

2. Sulcophanaeus cupricollis (Nevinson) New Combination

Fig. 239

Phanaeus cupricollis Nevinson, 1891, p. 209.

Cotypes, London. Not seen.

MALE. Length 16 -1to 20) mm, ereatest: width 10 to 11 mm.

Color Jarcely black; clypeus and most of pronotum ceupreous green,

py¢idium often faintly purple or violet. Frontal-area cof: head with

long, posteriorly curved horn. Pronotum (Fig. 239) with disc

Howden and Young: Panamanian Scarabaeinae 139

greatly flattened, the quadrate aréa bordered anteriorly on each

side with slightly compressed acute tooth and posteriorly by an

incurved broad ridge abruptly elevated on either side, of concave

median area; concave area posteriorly with small, distinct fovea

on each side of midline; lateral areas of pronotum moderately

rugose—punctate,’ Elytral intervals: nearly flat, sparsely: trans—

versely wrinkled, dull. - Pygidium distinctly punctate;

| PEMALES-Leneth 19> mm,> oreatest width.) ) mmss Dilteringe

from male in following major characters: frontal area tumid,

vaguely cariniform, tubercles obsolete; pronotal carina as described

for genus, abruptly rounded anteriorly; surface of pronotum

rugose, feebly.-so pesterior|y.

MATERIAL EXAMINED. PANAMA, 25 specimens from: BOCAS DEL

TORO PROV. - Rio Changuinola. CANAL ZONE —- Barro Colorado

Island; Madden Forest. PANAMA PROV. —- Cerro Azul; Cerro

Campanas COLLECTION, DATES: January,; February, March, April,

May, July, August, September, October, November, December.

OTHER COUNTRIES, 2 specimens from: NICARAGUA.

REMARKS. There are slight. differences in the configuration: .of

the pronotal ridge and undoubtedly small males (none seen) will

have the horn and pronotal protuberances reduced. All specimens

seen of both sexes have the pronotum largely cupreous green.

The species, S. cupricollis, has incorrectly been. synonymized

with chryseicollis Harold and by inference with auricollis Harold.

Males of both of these latter species have the posterior pronotal

concavity separating the posterior 'horns’’ wider and the “horns’’

more acute and elevated than in cupricollis, in omrvepinion

noctis is the species most closely related to cupricollis; see dis-—

cussion under noctis for comparison.

This species has been found on agcutt carrion, and at

howler monkey, human, and tapir feces.

On Barro Colorado: Island, 3. %cupricollis is active. diurnally,

particularly during the period of 0900 to 1200 hours. It occurs

on Barro Colorado Island only duving the late wet season and

early dry season.

3. Sulcophanaeus noctis (Bates)

Fre. 240

Phanaeus noctis Bates, 1887, p. 56.

Cotypes, London and Paris. London specimens examined.

Sulcophanaeus noctis: (Bates );, Edmonds ,)'1972, p.. 826.

MALE. Length 15 to 19 mm, greatest width 9 to 12 mm.

Frequently entirely black, occasionally with traces of cupreous

or green near posterior lateral pronetal- margin. Horn on head,

quadrately flattened pronotal disc (Fig. 240) and surrounding

protrusions as: described for S, cupricollis: pronotal surface

laterally smooth to feebly wrinkled (obsoletely punctate). Elytral

intervals nearly flat, dull. Pygidium not punctate or feebly so

basally.

FEMALE. Length: 152:to 20 mm, greatest width 6.5 to 11.5 mm.

Very similar to other females in genus, particularly cupricollis

140 Contrib Vier. “BntoTasi.) voloode.) mo. 198!

from which i diulfers int lackine (the: metallic coloration of the

pronotum and by having the pronotum only slightly rugose, more

obviously punctate.

MATERIAL EXAMINED. -PANAMA, 2° specimens from: CHIRIQUI

PROV. - Bugaba; Volcan de Chiriqui (cotypes).

OTHER COUNTRIES, 50 specimens from: COLOMBIA, ECUADOR.

REMARKS. A series of 29 (males collected in Ecuador: shows

extensive allometric growth. Small (15-16 mm) males have the

horn on: the head reduced ‘to alow, straight, transverse (carina;

the pronotum in the same specimens is almost evenly convex,

pronotal “protrusions Deing reduced to three small tubercles: forming

a transverse ©ow ‘about | mm behind the anterior margin.” One

moderately developed male from Ecuador has the pronotum largely

cCupreous,) approaching the coloration of ‘cupricollis.

ihere is little doubt) that. cupricollis. and noctis ‘are: éxtremely

close and possibly icupricollis: should be: placed’as a synonym of

noctis. However in the series we have seen, the sides of the

pronotum are always more distinctly rugose in cupricollis, the

pyetdium’ is) more ‘distinctly punctate, there is <a'‘slight difference

in the position and shape of the pronotal horns or protrusions and

usually cupricollis has a distinctly cupreous to green pronotum

While NOCtIS #s Mostly blacks (hic: color difference is: particularly

evident in the Téemates::« Because of (these differences we have

continued to treal) the twoforms as distinct species.

The Ecuadorian specimens were taken in almost equal numbers

in duns and carrion) traps ‘set. in: lowland wet forest ‘on the west

side of the Andes,

Coprophanaeus Olsoufieff

Copropnanaeus Olsouliei,, 1924, p.°22; Bamonds, 19/2, p. 343.

Type-species: Scarabaeus jasius Olivier, by original desig-

nation.

All Panamanian species are in the subgenus Coprophanaeus

as established ‘by Edmonds, 19/72... Both the jasius and dardanus

Species GrOUDs are Fepresented and may be recognized by the

following combination of characters: mostly black species, occasion-

athy with Slight metatiic oréeen, purple, or ‘cCuUpreous reflections on

the lateral pronotal margins; clypeal margin with two distinct

acute feet, “Sharply tneised’ not only medially but laterally.

Key to species of Coprophanaeus

1. Biviral intervals, (except at. “base, mearly flat

to very feebly ‘convex, uniformly dull; pro-

nmotumnDlackh: or with ereenish cach sy) 2s ee ag

Elytral intervals slightly to moderately sone,

dull“primarily near) striae; pronotum’ often

Wit DlUiSh purple Casi Oa) al vee Ohipiquensis: .Ols.

fan Male with low, néarly vertical, transverse tri-

tuberculate carina extending across anterior

Howden and Young: Panamanian Scarabaeinae 141

edge of frons; female with similar carina,

but tubercles close, often separated by

approximately their own diameter or less;

pronotal margins lacking metallic reflections. . . S

Male with posteriorly inclined flattened carina,

a small tubercle on each side medially with

central tubercle usually developed into a short

cylindrical horn; female with low transverse

trituberculate carina, the tubercles separated

by more than own diameter, the carina only

slightly anterior to eyes; pronotum of both

sexes frequently with metallic reflections ... .

. 1. telamon corythus (Har.)

ce Well-developed males with shallow longitudinal

depression on each side adjacent and anterior

to transverse pronotal carina, the depression

distinctly separated from lateral pronotal

fovea; both sexes with excavate posterior

edge of hind trochanter with scattered setae

basally Bh a LUD Maik AOC MeN 3. ohausi (Fels. )

Well-developed males lacking distinct longitudinal

depression on pronotum or with depression

contiguous with lateral pronotal fovea; both

sexes with excavate posterior edge of hind

trochanter! lacking, setae pie tii no Cis PORE i Ie. SDs

1. Coprophanaeus telamon corythus (Har.)

Figs. 241-243

Phanaeus corythus' Harold, 1963; py 103.

Type. or ‘cotype; Paris. Not seen,

Phanaeus (Coprophanaeus) corythus Har.; Olsoufieff, 1924, p. 68.

Coprophanaeus telamon corythus Har.; Pereira and Martinez, 1956,

Dy) Anes

MALE. Length 20 to 26 mm, greatest width 12.5 to 15.5 mm.

Black, frequently with trace of greenish or cupreous reflections

along anterior and lateral margins of pronotum. Clypeus (Fig.

241) with lobe on each side of acute teeth angulately truncate to

obtuse angle. Horn on vertex anteriorly broadly flattened,

sloping posteriorly, sides narrowing to small lateral denticle then

narrowed to short, median, cylindrical, slightly anteriorly curved

horn. Pronotum shallowly concave and shining below binodose

swelling; area between nodes evenly, posteriorly arcuate when

viewed from above; distance between nodes approximately equals

distance between middle of each eye. Pronotal surface laterally

rugose, finely punctate to smooth behind swelling, surface just

anterior to posterior pronotal margin on each side of midline with

small fovea. Elytral intervals uniformly dull, very feebly convex.

Male genitalia as in figs. 242, 243.

FEMALE. Length 18 to 24 mm, greatest width 11 to 15 mm.

Differing from male in following major characters: head with low

142 Conthib. Amer. nth dnsivs velo woe ose. 196 |

trituberculate transverse carina at anterior edge of frons, carina

laterally curving posteriorly on each side mesad of eye, termin-—

ating near posterior margin. ol evye;.tubercles on carina. separated

by two to three times own width;. pronotum with anteriorly arcuate

carina approximately L.mm,behind. anterior margin,. the carina

extending no more than width of eyes, usually slightly less;

pronotum briefly concave behind carina, concavity and lateral

surfaces uniformly rugose, pronotum becoming punctate on basal

cu gaale

MATERIAL EXAMINED. PANAMA, 34 specimens from: CANAL

ZONE —- Barro Colorado Island; Corozal; Frijoles; Gamboa; Madden

Porest.. “COLON PROV... 2/0 m, 0 Ki Sb Golon,..Santa..Rita Ridge.

DAR URN 2 RRO. (c)-RiG-hacanii,; }90U it, , COLLECTION. DATES: .February.,

merit, May, June, july, Seplembe:, October, November, December.

OTHER COUNTRIES, 32, specimens .from: -MEX1ICO,. BELIZE,

GUATEMALA, NICARAGUA, COSTA RIGA (1. corythus); BRAZLL,

COLOMBIA, PERU, VENEZUELA (t. telamon Er.).

REMARKS. Small males may have the head horn greatly

reduced in height, forming a trituberculate carina, the middle

tubercle being only :moderately produced. . There is also a similar

reduction in the degree of pronotal swelling, but the shape and

position is not drastically altered. Some specimens of both sexes

may lack metallic reflections on the pronotum, this occurring most

frequently in Mexican specimens.

The key characters and particularly the shape of the male

head horn should identify telamon corythus. Specimens of the

South American t. telamon Er. have the median lobe of the male

horn abruptly posteriorly offset basally, then curved upward; in

females the basal third of the pronotum is more heavily punctate.

In Panama specimens were attracted primarily to carrion

with a few coming to dung. The species is most abundant in

the late rainy season. Flight activity seems to be mostly noctur-

nal. Most specimens were taken in lowland forests below 1000 m.

2. Coprophanaeus chiriquensis Olsoufieff

Figs. 244-247

Phanaeus (Coprophanaeus) chiriquensis Olsoufieff, 1924, p. 73.

type. -Paniss. Not seen,

MALEs Leneth 14.,to<ie mm, greatest width 9. to 12: mm.

Dorsally black, often with violet reflections on sides of pronotum

and elytra. .Clypeus (Fig.. 244). with lobe.on .each side of acute

teeth low, broadly arcuate to feebly angled, frontal carina tri-

tuberculate, dateral tubercle on each..side: often slightly «larger

than median one. Major males with broad median pronotal pro-

tuberance (Fig. 245) delimited laterally by longitudinal depression

extending toward lateral edge of eye; anterior face of protuber-

ance smooth, dorsal edge irregularly cariniform, edge thickened

on either side of slight depression at midline. Pronotum poster-

iorly with small fovea on either side of midline just anterior to

posterior margin: «Elytral intervals; slightly, but.noticeably convex;

each interval dull near striae, feebly shining medially. Genitalia

asin Mes. oe, 247.

Howden and Young: Panamanian Scarabaeinae 143

FEMALE. Length 16 to 20 mm, greatest width 11 to 13 mm.

Differing from male in the following major characters: frontal

carina with three: small tubercles; median one slightly elevated,

tubercles separated’ by a distance approximately equal to taree

times width of tubercle; prondotum >with ‘transverse ‘carina about 1

mm, behind anterior margin, léenoth-of carina equal/to’ or slichtly

exceeding outside width of eyes: carina with small, usually dis-—

tinct tubercle or-angle on either Side’ of shallow, medranm imdenta—

tion? area immediately, behind carina shallowly concave, rfugose,

sometimes less so near carina; posterior third of pronotal disc

nearly smooth, dull.

MATERIAL EXAMINED. PANAMA, 28 specimens from: CHIRIQUI

PROV. =° 15> km NW -Hato del: Volcan; 1500 mm, “Hartmann’s Finca:

10 kmeW, Cerro: Pando?’ * ‘COLLECTION DATE: May. |

OTHER COUNTRIES, One ‘closely related species oceurs in

Mexico, another in Ecuador: We have not Deen able to -assien

names to these; the one in, Ecuador being possibiy ©, foubali

Balthi. |

REMARKS. In small males the pronotal protuberance is repre-

sented by a low carina, shallowly depressed medially. In small

females the area immediately behind the transverse pronotal carina

may:be only feebly concave. ~The. faint vielel. color may. be timited

to the sides of the pronotum in either Sex.

This species has seémingly gone unrecognized since Olsoufieft's

description based on three females from ’'Chiriqui”. | Morphologically,

females of chiriquensis are very close to females of telamon corythus

and have been incorrectly synonomized under that species. The

faint violet color of chiriquensis should serve to separate it from

t. corythus which has greenish or cupreous reflections (if any).

Also the form of the male pronotum is very different from males of

t. ‘corythus, and the pronotal carina’ of female oniviquensis is

more distinctly bidentate.

At present chiriquensis is known only from the vicinity of

the Volcan de. Chiriqui at elévations-of*approximately 1500) m.%'\1t

has been taken in traps baited with human feces and with carrion,

apparently favoring the datier.

3. Coprophanaeus ohausi (Felsche)

Figs. 248-250

Phanaeus ohausi: Felsche, 1911, p. 138.

Type, Dresden, (2) 4 Nok seen,

MALE. Length 22: te, 26°mm, sereatest swidth 12. to: 14 mm.

Black, lacking metallic reflections. Clypeal lobe on either side

of acute teeth broadly, obtusely angulate. Transverse frontal

carina as wide as distance between eyes, at least one eye dia-

meter anterior to eyes; carina’ low, /trituberculate,. tubercles

separated by approximately two times their diameter. Frons and

most of vertex finely punctate. Crest of transverse pronotal

carina in well-developed males (Fig. 248) with four swollen or

thickened areas, one on each side of midline and one at each

side at termination of carina; in smaller males only lateral

144 Contribs Amer, sbi nei) Meda UO snes hoo!

thickening of carina evident, the carina slightly bowed medially.

Lateral \termination of carina, delimited) anteriorly..by elongate

depression, 6éxtending, anteriorly. (toward outer margin of eye, . the

elongate depression distinctly separate from lateral fovea; pronotal

surface moderately rugose laterally, smooth to obsoletely punctate

elsewhere; distinct posterior depressions near pronotal margin

absent) or very Teebly indicated. Elytral striae, nearly flat, dull.

Excavated posterior edge of hind trochanter with setae.

Genitalia. asin) fios, 249,:°250..

FEMALE: «Length 20: to 23 (mm, .areatest. width 11.5. to 13..mm.

Differing trom mateo in the following major ‘characters: frontal

Carina.jlow,: feeble, tubercles darce, well-developed, separated: by

approximately one diameter; \transverse ‘pronotal carina broadly

obtuse, ‘blunt at anterior median edge.) vacuely indented ..in some

specimens; surface behind carina: (‘shallowly concave, surface rugose

to rounded summit; laterally posterior third of pronotal disc

mostly smooth.

MATERIAL EXAMINED: PANAMA; 13: specimens from: BOCAS DEL

TORO PROV... =) Rio, Chancenas 2400) 10. o.CANAL. ZONE -—- Madden

Forest... COLON, PROV. —:16 km SE Celon, 270 m,:Santa Rita Ridge.

PANAMA PROV. - Cerro Campana, 800 m. COLLECTION DATES: May,

Sepreniber.

OTHER: COUNTRIES, Ovspecamens:. from: ECUADOR.

REMARKS.» Fhe transverse:pronotal carina ,in the.males varies

considerably both in local populations and geographically. Males

from Ecuador have the carina laterally produced, sometimes

extending slightly over. the. oblong lateral concavity. . In, other

males, particularly Panamanian specimens, the carina may be

nearly straight medially and lack median swellings; in some the

thickening of the caring on either, side may be anterior, in position

to the. lateral thickenings or angles, giving the carina an. irregu—

lan arcuate ‘shape. ‘Other chatacters,’ including. the ‘genitalia,

Show little variation. Males from Ecuador can often be distin-

guished from those from Panama, but the differences do not seem

Sufficient tO\warrant nomenclatorial, recognition of »the: geographic

variation.

| We ‘characters.@iven im the key.) the’ lack of metallic

retlections and the shape of ithe pronotal carina should serve to

distinguish the species from related forms occurring in Panama.

The species has been taken in traps baited with human

feces and carrion, Seemingly preferring the latter.

A) COULOphaMaeus Decki., nN. Sp.

Figs. 251-253

HOLOTYPE. Male, length 26 mm, greatest width 14.5 mm.

Dorsally dull black, lacking any metallic reflections. © Clypeus

(Fig. 251) with lobe on either side of the two acute teeth obtusely

angulate; clypeal surface anterior to frontal carina transversely

rugose. lransverse’ frontal’ carina’ abruptly vertical, slightly

over two. eye diameters anterior to: anterior margins of eyes,

transverse width of carina less than distance between eyes; carina

Howden and Young: Panamanian Scarabaeinae 145

trituberculate, median tubercle slightly elevated above lateral

ones; surface of frons and vertex posterior to carina smooth.

Pronotum at anterior third with bilobed transverse carina extending

slightly more than distance between outside of eyes; (the carina

most prominent on\either Side“ol midline, “curved ‘anrvertorly Jater-—

ally. ‘Pronotal surfaces laterally moderately rugose) smooth beiore

and’ behinds carina; “surface ‘om either side ori midline “ust anterior

to posterior margin feebly depressed. Elytral intervals nearly

flat; smooth,. dull. “Pygidium feebly,: timely punctate. (Pore tibia

quadridentate on outer margin. Excavated posterior edge of

hind trochanter lacking setae. Metasternum nearly flat, smooth,

very finely punctate, midline vaguely impressed. Genitalia as

bn figs. 25a. 23,

ALLOTYPE. Female, length: 28° mm, greatest width 1/7 mm.

Differing .from male in the following..major, characters: iubercies

of frontal carina.,distinctly closer, median: tupercle slightly more

pronounced than lateral ones, width of carina distinctly reduced;

surface behind carina finely punctate except in »osterior third of

vertex: transverse pronotal |/caring obiusely ancled. the anterior

median jangle ‘abruptly emaroinate at miciiie, bidentate; carina

medially about 0.8 mm behind posterior edge of marginal bead;

surface. immediately posterior. to’ carina concave, .moderately rucgose;

surface of pronotum behind concavity rugose, punctate to finely

so laterally and medially, nearly impunctate centrally in’ posterior

fourth; indentation’ ‘on either Side ef midlinecnear posterior marcin

Obsolete.

TYPE MATERIAL: Holotype, male, Panama, Chiriqui Prov., 15

km NW Hato del Volcan, 1200 m, Hartmann's Finca, 20-31.V.197/7,

S.: Peck, in’ dung’ trap: (Howden) Allotype, female,’ same data as

holotype (Howden).

PARATYPES: 8 males, 11 females. PANAMA: 2 females, same

data’ as. holotypes'3° males, "4 fenales ;(Chiriqur row. (lO kim) W

Cerro Pande, 2Z4AsV.197) ,VS.sPecky) carrion trap.) Ceol RICA t

female, Puntarenas Prov., 15 km NE Potrero Grande, Finca

Helechales, 2.1X.1967, RiWe MeDiarmid?y tumatle, 2 Temates:,

Puntarenas Prov.; 6 km S$‘ Sane Vi1toy 08042 Ne) S3000' WW. 25-2801.

1967, Dik. Veirs: 4 males, 2 females, Puntarenas Prov... 6 km 5

San Vito, 1000°m,\(27.1V-7.V.19607, (DoF. Vers; in forest in’ traps ‘-

meat and human) feces:

Paratypes are in: Ottawa,’ Washington, Edmonds; Howden.

REMARKS. Body’ length in:the specimens seen’)ranges) Irom 22

to 28 mm. The three male paratypes from Panama have the carina

on ‘the ‘frons ‘slightly narrower than inthe holotype and) in ore

male with the most reduced carina the median tubercle is distinctly

elevated} \° The greatest variation “in ‘the males ‘occurs inthe

pronotal carina which is binodose in large males, nearly truncate

and lacking) any: lateral’ concavity in smaller: males.’ The same

type: Or variation ‘may’ be! found an Temales,: Large specimens have

three ‘distinct tubercles on the frontal carina and the pronotum is

distinctly ‘tumid ‘on either side of the midline; small females may

have the frontal carina arched with tubercles obsolete, and the

pronotum may ‘be only ‘very ‘Teebly 'tumid) on either. side behind “the

concave area.

146 Contrib. Amer. Ent. lasts, Vol. 16, fio. 4,.1981

Small specimens of either sex may be confused with the

related C. ohausi, but ohausi has at least a few setae on the

posterior excavated edge of the hind -(rochanters: (nese setae are

lacking In pecki:

The species is mamed for bat stewart Peck who collected

all of the Panamanian series.

Oxysternon Laporte (= Castelnau)

Oxysternon Laporte, 1040, 0. 62. Edmonds, 1972. ). 838.

Type-species: Scarabaeus festivus Linnaeus, by subsequent

designation, Martinez and Pereira, 1967.

Edmonds, 1972, divided Oxysternon into two subgenera.

Only two species of the nominate subgenus are known to occur in

Fanama, Both of these are largely oreénish, broadly oval species

that can be distinguished from the related genus Phanaeus by the

following characters: metasternum with anterior median projection

forming a long, sharp spine that terminates anteriorly between

the anterior coxae; posterior margin of pronotum medially extended

posteriorly, forming am acute angle.

Key lo species of. Oxysternon

t Length 18 to 25 mm; male with conical or cylin-

drical horn on head: both “sexes: with, anterior

pronotal ces thickened adjacent to anterior

ameles i. ~ 1. conspicillatum~ (Web..)

Eength 12°-te 15 mm, Te ae low, transverse

tuberculate carina on head, both sexes with

anterior pronotal margin not noticeably thick-

ened adjacent t©. anterior angles... .». «2. zikani. Per.

1. Oxysternon conspicillatum (Weber)

Figs. 254-256

Copris conspicillatum Weber, 1801, p. 36.

Type, Kiel (?). Not seen.

Oxysternom conspicillatum: (Weber); Olsoufieff, 1924; p. 113.

MALE.; Leneth 19 to 25 mm, greatest width=12: to: 17. mm.

Dorsally green, usually with black pronotal markings. Well-

developed males (Fig. 254) with long, cylindrical posteriorly

curved horn on head. Pronotum on each side in lateral third

with inwardly curved, longitudinally flattened horn, a transverse

depression present between horns; pronotal surface smooth. Elytral

intervals distinctly convex. Genitalia as in fies! 255, 256:

FEMALE. Length 19 16 23. mm, ereatest width 12to 16 mm.

Differing from male in the following major characters: head with

low transverse carina on frons; pronotum inregularly convex; a

shallow longitudinal depression in anterior third on either side of

midline; a feeble, arcuate or obtusely angled carina extending

between pronotal depressions.

Howden and Young: Panamanian Scarabaeinae 147

MATERIAL EXAMINED. PANAMA, 2 specimens from: DARIEN

PROVe.—" Rio Tacartiy 1900. ft. « COLLECTION Dates july.

OTHER COUNTRIES, 50 specimens from? COLOMBIA,” VENEZUELA,

ECUADORG BRAZIL, BOLIVIA; PERU.

REMARKS. There is considerable variation in size. Very

small males resemble females. Also the configuration and extent

of the black markings on the pronotum is extremely variable.

The species is easily recognized by its large size, basic

green color, convex elytral striae, and the characters mentioned

for the genus. In Colombia the species is commonly found flying

during the day in open or second growth areas where it is readily

attracted to human feces. It can also be found at cow and horse

dung, and occasionally at carrion.

Z. Oxysternon zikani Pereira

Figs. 257-259

Oxysternom zikani Pereira,;: 1943, p.: 108.

Type, Pereira collection (7). . Not seen.

MALE (Panama.and.Ecuador). <Lenmeth 15.1fo0 1/ mm, greatest

width 10 mm... Dorsally @reen .with black markings in cenizal

area.of. pronotal .dise and on -elyiral vmbone.. /Ciypeus.ieebly

emarginate medially, beneath emargination lacking spiniform pro-

cess. Frontal area of head (Fig. 257) with transverse, binodose

ridge. ..Pronotum,in anterior half, with: low longitudinal, rounded

ridge on. either side of midline, ,concave between and shallowly. so

on outside, of ridges; posterior hall of prenotum convex, smooth.

Elytral-striae moderately, indented, not distincily. punctate, intervals

moderately. convex. . Pygidium: feebliy punctate, Pore (Lo1a with

four: teeth on outer margin... ,Gériitalia as in digs. 256. 259;

FEMALE (Panama specimens). Length 14 to 16 mm, greatest

width 9 to 10 mm. Differing from male in the following major

characters: transverse frontal ridge trituberculate; longitudinal

pronotal ridges lower, more broadly rounded, concavity between

narrow.

MATERIAL EXAMINED. PANAMA, 4 specimens from: CANAL

ZONE - Barro Colorado Island, Madden Forest. PANAMA PROV. -

Maje. COLLECPION DATES: January. Fepruary, September.

OTHER COUNTRIES, (11. specimens. from: -COLOMBIA,, ECUADOR,

BRAZ LL PERU.

REMARKS. As pointed out by Edmonds (1972) there is some

confusion in the silenus group of species. Olsoufieff (1924) implies

that. the,.males of silenus and related species have horns on the

head. However. there is\.at. least ome -species:where, the male .has..a

binodose transverse carina on the head and the pronotum is

similar to that of the female. Since we have not seen the type

of silenus that is supposedly a horned male, and since females of

all species are similar, we do not believe the name can be applied

to Panamanian specimens. In 1943 Pereira (p. 109) described O.

zikani from Brazil and Peru and this seems to be the correct name

to apply.to the species. lacking horns in the. males... Unfortunately

confusion concerning the application of the. various names will

148 Contrib, (Amer. Ent.) last.) vol. 28,% mo. ok) 1981

remain until the silenus complex is revised, The smaller size is

usually sufficient to separate zikani from the larger conspicillatum

which also occurs in Panama.

: Specimens of zikani have been taken on rodent fecal pellets,

monkey dung and human feces.

TOPOGRAPHIC FEATURES AND FAUNAL PATTERNS

Panama, a country of approximately 47,600 sq km, has a

rich and varied biota. Topography, geographic position, and at

least its Tertiary history are factors that have contributed to its

biotic “diversity.

Panama, lies between /° and DOYN latitude and 77° and 83°

30'W longitude, the long axis of the country running approximately

east-west (Fig. 260)... The north onm,Garibbean side’ has, in

general, a high rainfall and contains extensive areas of lowland,

evergreen, (opical, tain forest. Extending most of the leneth of

the Country lies an irregular spine of volcanic peaks and hills.

The highest elevation, 3427 m.\is in’ the west on the Volcan. de

Chiriqui tor Hl Baru), the lowest point in the divide, 65 min

the Canal Zone. In the eastern province of Darien near the north

shore a series of basaltic ridges attain elevations ranging between

1000:;and TGOO"m with’ 4° low Dom Of TiO vm inthe divide” *(Haffer,

19/0. Aboot’ 65% of ihe land “grea is lower than 1000 m, ©

The north facing slopes of most of the ridges above about

800 m interrupt the cloud-trade wind pattern of the Caribbean.

As a result of this many north facing slopes are moist throughout

the year and have a rainfall often in excess of 3000 mm per year.

In areas where the elevation tanges between 000° m and 2500 m, a

cloud forest type of formation may develop. Above 3000 m (mainly

on Volcan de Chiriqui) an elfin type of forest mixed with open

treeless areas exists. The south-facing slopes of the various

Sierras are weually well forested but with a fairly pronounced dry

season extending, on the average, trom January to April. Below

800 m and on the south coast the dry season is pronounced and

large lowland areas are covered with a deciduous lowland forest

formation (Fig. 261). ~The various forest zones and some of the

more conspicuous species of these zones are discussed and listed

by Porter (1973);

While the location, topography, and climate undoubtedly allow

and contribute to the present diversity of the fauna, the historical

relationship of the Panamanian fauna is a particularly interesting

one. The distribution patterns of the 113 Panamanian species of

Scarabaeinae that we recognize are placed into five categories in

Table 2." Of these species, 42 are widespread, 13 occur in Panama

and Souti America, 22 are found in “Panama and Central America,

18 are restricted to Panama and Costa Rica, and 18 are limited

to Panama. If one assumes that both the widespread and Panama-

south American groups have a probable Pliocene-Pleistocene South

mmerican Origin, “then fom (55 Of 113) of the Species: show ‘this

affinity. On the other hand 58 species or 52% represent a Central

American element with nearly 16% of the 113 species being restricted

to Panama. The strong endemic nature of the Central American

Howden and Young: Panamanian Scarabaeinae 149

fauna has been discussed by Duellman (1966) for the herpetofauna,

and Howden (1969) has discussed the variation in generic endemicity

found in American Scarabaeidae as it might relate to Pleistocene

climatic disturbances. If one considers the species endemicity

eccurring in Panama, (10) plus that of western Vanage i) eastern

Costa Rica (18), the 36 endemic species seem to concentrate in one

of several of the areas of Panama that have been emergent for at

least several million years; i.e., some of the oldest portions of

Panama.

Because of this pattern, a brief mention of the geological

history of Panama seems pertinent. [he early history. of ine

Caribbean area and the plate movements associated with continental

drift have perhaps best been summarized by Malfait and Dinkleman

(1972). Im this paper, they depict a Caribbean plate moving

generally from west to east, ‘with the present Panamanian area

occurring on.the western edge of the plate. ~ However... lateral

plate movement over large distances may be more relevant to genera

and higher ‘taxa patterns.” Thée.fauna.of-interestto.us is seem-—

ingly much more closely tied to: Teriary events." iiese events

have been discussed dn.some detail’ for eastern Panama and ine

neighboring portion of Colombia by Haffer (1970). According to

Haffer, in the upper -Lecene, initial orogeny extended. at leacp

into eastern Panama but the actual exposure of any (and mass

may have been relatively temporary... Prom the upper. Eocene:.to

middle Miocene, Panama and neighboring Colombia were largely

submerged with small and largely scattered emergent areas or

islands. During the upper Miocene there’ was a further orogeny

with narrow water gaps persisting primarily in western Panama.

Again’ there was subsidence ‘during the lower Vliocene, with the

formation of three major water gaps, consisting of the Nicaraguan,

Panamanian, and Colombian portals, These were closed either in

the: upper Pliocene or early Pleistocene approximately two million

years ago. During some of the Glacial perieds of the Pleistocene

the coastal lowland areas of Panama and adjacent countries were

greatly expanded so that the land area was perhaps Jarcer. by

20% to 25% than at present. These various events have been dia-

grammed and discussed by Haffer (1970) along with the problems

that exist: in the’ geaiogical interpretation of the sediments ‘in the

area. More recent events; tneludinge sea. level changes (and climatic

fluctuations of the last 35,000 years, have been discussed as they

relate’ to Panama by Bartlett and Barehoorn: (1973).

The geological evidence thus indicates that Panama has been

an area Of instability, 7 For, much. ol: the early Tertiary only small

isolated areas of exposed land ‘were present so that any faunal

interchange must have ‘been by overwater ‘dispersal. This, changing

topography undoubtedly strongly influenced speciation and also

contributed to numerous extinctions. The fauna; at least in the

Scarabaeinae, reflects this. The ranges of the 36 endemic species

occurring in’ Panama and adiacent Costa’ Rica ‘seem ‘to coincide

with one of the main’ Pliocene — early Pleistocene emergent areas,

with the Nicaraguan trench to the north west ‘and the Bolivar

(Darien) trench to the east. The sizes of the emergent areas and

the length of time that’ they have remained emergent is not certain.

bo) Wi. bor 901

, Mids

Inst.

Ents

Contrib. Amer.

150

*(qs0} UNS YURI popIs OM4 B) 480} N AoUzTYM-UUBI oY} BUISN [9A9] %G OY} Je JUBOTFIUSIS ST I]

*[OA9] %), OU} JNoge 3 JUVOTFIUSIS ST 4S94 J V SUISN WW E°g PUS SG°OT USOMIOG DOUSTOJJIP 9YUL

(WW §6°0T)

(wi ¢°g) (WU FT TT) (WU gc * OT)

WU (2°), wUUL 19 °6 ULUL 6S °OT ULUL 80 °CT wu ZT° Or

OZIS OSVIOAB

*dds gT ‘dds 91 ‘dds 7Z ‘dds ¢] ‘dds ZF

VINVNVd "Ud O-VINVNVd VOIWAINV °O-VINVNVd VOIMAINV °S-VINVNVd GVaudSddIM

“AZIS SHIOddS OL SNYALLIVd

GHL FO dIHSNOLLVIGY AHL ONV AVNIAVEVEVOS NVINVAVNVd

AHL TO SNYALLVd NOLLAETULSIG TVYANAD HHL FO AUVWINNS °¢ WIAVL

Howden and Young: Panamanian Scarabaeinae 15i

If Haffer's generalized diagrams are reasonably accurate, then

at least some of the Chiriqui or adjacent areas have been exposed

since Upper Miocene; i.e., for about the last 5,000,000 years.

this: length of tine certainly has been Sufficient te valtow a dis—

tinctly endemic biota to develop. The length of isolation (or

period of emergence) apparently has not been sufficient to be

reflected’ in any degree of endemism at the generic or tribal

level within Panama, at least in the Scarabaeinae. However,

particularly’ in the genus Canthidium, groups: of related species

occurring in the Chiriqui area apparently radiated in tie western

Panama—Costa’ Rita upli{fty® lt2is) interésting te note tat the

degree of endemism is higher among genera composed of small-sized

species (under 10 mm) than among genera containing only larger

species (Table 2). This in turn may reflect. differences in flight

behavior as it relates to foraging, Which in turn may relate 10

dispersal.

If, until 2,000,000: years cago) tne exposed portions of

Panama resembled an archipelago or even, an isclated 1stand

situation the present fauna, particularly the endemic, elements,

should reflect ‘this. There are a number of different ciaracter—

istics’ exhibited Dy. island, faunas, but if: one: examines the species

listed in Matthews’ (1966) work on the Scarabaeinae of the West

Indies, .it»is--obvious: that a’ large peéréentace, of Iie taunea ais

composed of relatively small species (10 mm or less). Genera

such ‘as Dichotomius, Phanaeus, »Eurysternus, and: Deéltochilum are

common: in-surrounding mainland: localities and: generally are of

moderate to large size (15 mm or more). In the West Indies only

Jamaica has one large endemic species, Phanaeus carnifex (L.).

This ‘speciés,: measuring from 17 10 23 mav4in. tenoth; (lacks, any

really close’ relative, the most closely -rélated species oceurring in

nerthern South Americas: Ty general the smal size of istand

Scarabaeinae.is not a unique feature as Darlington (19/0) nas also

noted the same characteristic for Carabidae occurring in the West

Indies. ‘If “species size does have any relationship with island or

habitat isolation, then,» based on Haffer’s (19/70) geological history

of Panama, endemic forms in Panama should have a smaller

average size than more wide ranging forms if Panama were an

archipeligo in the recent past. Also, genera occurring in Panama

that contain a number of endemic species, in theory, should be

composed of mainly small species with their closest relatives in

nearby localities. ; This 4s true-atvleast for Uroxys and Canthidium.

Also the species endemic to Panama and the adjacent area of

Costa Rica average 8.3 mm in length while the widespread (1),

Panama—South America (2), cand Panama-—Central America § (3)

groups average 10.12, 12.08, and 10.59 mm respectively. Thus

the size as well as the location of the endemics seems to coincide

with the past geological events of the mid to late Tertiary as

they relate’ to Panama,

Because of the preliminary nature of the present study, the

numerous unsolved taxonomic problems, and spotty collecting

coverage, further comments on faunal relations are not justified.

152 Contrib. Amer. nti sihmsee) woluods, cme. 4s..1081

HABITS: COMMENTS ON THE BEHAVIOR OF A FEW SPECIES

: Since this work is primarily concerned with the identifica-

tion of the Panamanian Scarabaeinae, details of behavior that

have been observed on a number of species will be omitted as they

have been or will be published elsewhere. It does seem appro-

priate, however, to briefly summarize the major feeding habits of

the Panamanian species and to mention briefly a few of the more

interesting aspects of some of the: current behavioral work that

has been completed or that is still under investigation.

The known habits of the Scarabaeinae have been well sum-

marized by Halffter and Matthews (1966). Their work outlines the

variety of food utilized by «some members of the subfamily... In

our study of the Panamanian species, we have found that the

food preferences largely seem to follow the general patterns as

described by Halffter and Matthews (1966). In general the

species are attracted to mammal dung, while some show a prefer-

ence for carrion. Frequently species may be attracted to both

dung and carrion. Other foods utilized by fewer species are decay-

ing Vegetation, fungi, and rotting fruit. Again, in some ‘cases,

species going to rotting fruit may also be attracted to dung and/or

carrion. Examples of. this, typeof behavior can be. found within

the genus Onthophagus. As noted by Halffter and Matthews (1966),

it seems that most species.of Scarabaeinae have the capability of

utilizing morée.than one type of dung or vetting material... Many

species undoubtedly have a food preference but this aspect has

not been thoroughly investigated for the majority of species.

Once the dung or other food material has been located it

appears 10 be “expedient [or dhe beetles to. move it. as. rapidly .as

possible. This need for rapid removal seems to be a response, at

least in part, to the number of species competing for the resource.

In Panama, as many as 22 species have been taken in one locality

at traps baited with human feces. The number of species competing

for the resource is probably only part of the story since individu-

als of the same species may also compete (see Matthews, 1963).

The response to this competition has been the development of

several methods of rapid food handling that essentially remove

the food from the competitive arena. One method is to immediately

dig a burrow under the food mass and bury a quantity of the

dung or carrion. Once the food is in the burrow it may be eaten

or formed: into. a brood cell.

seme, species may actually push (butt) pieces; of .the dung

for a short distance and then bury it, while other species

(members of the genus Canthon, for example), may form the food

material into a ball and roll it for a considerable distance before

burying it. In all cases, after burial, the food may be consumed

by @ither the adults or .utelized in) a ~breod cell for.larval, food.

Details of the types of brood cells and mention of brood

care may be found in papers by Bornemissza (1969), Halffter

(1977), and Lengerken (1954). Normally, development of the

brood is fairly rapid with the period from egg to adult frequently

occupying between 30 and 50 days. In areas with a prolonged

dry season, either the pupa or teneral adult may remain in the

Howden and Young: Panamanian Scarabaeinae L539

brood cell antil there issufficient moisture/in the round to. allow

the: adult "to: ‘emerge, thus: limiting the: reproductive eyvere to tone

generation per year. However) in Panama there Vane: frequently -a

number) of generations’ per year’ bul (there: may pe saiiierernces iim

the size of species populations between the wet and dry seasons.

Seasonal population fluctuations of dung beetles have been studied

carefully in“only afew) areas.) Oni ibarro Volerddon island." tie

scarabaeine biomass in the mid-dry season (Feb.) is estimated to

be six times larger than in the mid-wet season (Sept.). The

biomass’ of all insect'species' captured at liehiy traps, however,’ is

approximately the same for February and September (Smythe,

1974). The increase in biomass that occurs in most insect groups

with the start of the wet season (Wolda, 1978) does not occur in

most: species of ‘Scarabaeinae,’ at) least von Barre Colorado | island,

Several factors’ canbe ‘correlated with ‘the higher biomass of

scarabaeines in ‘the: dry season: 'on’ Barro Colorade island. the

population density of vertebrate dung-producers is probably

lowest ‘during the mid-late wet season, due'at) least) in part: to

high.mortality ‘because of the (scarcityic! tril, new leaves sy flowers,

insects, and other foods typically consumed by vertebrates (Young,

1978). Thus the production of vertebrate dung is also corres-

pondinely ‘lowest in’ the wet season.) The ‘eavailanlity or thar, aune

to scarabaeines is probably lowest during the late wet season,

due to the action ‘of heavy rains on exposed dung with the sub

sequent ‘rapid loss of aromatic properties and dissolution. With

both the: production ‘andthe availability of ivertebrate cung: lowest

in) the mid-late wet ‘season, the Aite-hastory strategy. olieach

dung-consuming scarabaeine must include some mechanism for

surviving ‘food scarcity, at’ that) time. The dominant) diurnal and

nocturnal species, Canthon angustatus and C. aequinoctiale

respectively, utilize a greater proportion of the available dung

during the late wet season and are reproductively active through-

out) the entire years) Some "ether species! are Mon ipresent as adults

during the late" wetsencon. /(istead they larey helow) aru (im

the larval or pupal (stage and emerge arthe stant on thendary

season. ‘Still other species change their: Teeding Habits.) ouring

the late’ wet season* they ‘teed as adults ony vertebrate "carrion,

which’ is relatively more’ avarlaple to scarabacines Guring) this

period /(VYoung, ta 7o:

The availabilty of) food ‘and the ‘numper ot indiyidua ts

competing for’ the’ food” may have net only led to the development’ of

methods of rapid food removal but have apparently also influ-

enced the evolution ion different foraging techniques.) il yoteally,

beetles ‘locate duns, ‘carrion,| or rotting vegetation’ by the detec—

tion. of odors. The euecies generally “pursue a laterally sinuous

flight pattern moving upwind, the pattern becoming narrower as

they approach’ the’ source'lor ‘the ‘odor. Normally, ‘the maiority of

species fly only a few feet above the ground and are able to

land ‘either onvor close’ to: the ‘odor source, "While this type of

foraging behavior: ile. ‘flying upwind, is typical’ of most species,

there are a number of variants that have developed on the basic

pattern; an example of this 1s’ Tound in the’ species’ Canthon

angustatus. °On Barro Colorada Island it 1s perhaps the most

154 Contrib, Amer. Ents. dnstis. vol. 43, ome, «),. 1931

common species during all or most of the year. Normally, when

food odors are first detected, individuals leave their burrows,

take flight, and. follow the airborne oder.trail upwind. to: the

odor source... Individuals that arrive: in: the vicinity of the odor

source .within the first. few. minutes after defecation. by a verte-

brate are usually able to locate and land on the food. Individuals

that arrive later are usually. unable: to determine, the exact

location. of the.food and. after several minutes, of flying in the

vicinity dand on leat surfaces...tailuve to detect.ihe. odor. source

is probably due to decreasing production of odor and confusing

local convection currents: (Individuals’that are: thus, unsuccessful

im locating. food are the ones.that- perch, on leaves... Jhey are in

the best position, however, to locate food subsequently produced

in the same, area:

While foraging near the ground is common, not all species -

utilizine this method, can locate, rotting iruit.or dune lodged

above the surface of the ground. .Howler monkey dung, for

example, may lodge om branches or leaves in the upper levels of

the canopy. A few species apparently have developed capabil-—

ities ef locating and ulilizing arboreal food sources....On. Barro

Colorado Island, observations made from a 65 foot high weather

tower in the forest indicate that at least two species, Canthon

angustatus and C. subhyalinus, have a three dimensional capacity

to locate food odors. This was apparent when howler monkey

dung was placed at the top of one of the towers. . All responding

species could be seen flying towards the tower at a height above

the ground of 15.10.60, cmiin Near, the base.ol the tower, €.

angustatus and C. subhyalinus changed their elevation, flying

upwards to the top of the tower. The other species maintained

their original elevation, flying, avound.in circles in. the area

where the odor trail from the tower feces apparently reached the

ground... Thus-only in situations where arboreally-produced feces

fell ail the way to the, ground: would. those: species, with: two-

dimensional dor detection: capabilities. compete for..its, possession.

Another type of. possible foraging behavior is leaf. perching

as suggested by Howden and Nealis (1978). Actual reasons for

Perching on Jeaves are seeming!y complex, and varied but, un-

published data from Panama and Ecuador has shown that small

Species im certain oenera, particularly species in Canthidium,

tend to be found perchine, more: frequently amd closer to, the.ground

than do larger species in other genera. Howden and Nealis (1978)

postulated that flying above the-low level vegetation: or .along

trails, at an. average height of about |..meter allows: a species to

detect relatively strong oders.that.would, be given off by a large

quantity .of food. Small iood sources.micght be better detected: by

perching mear the ground in favorable areas... -[his: postulate was

later tested by selling large dune, traps baited with 100-200 cc of

human feces, and small traps.with 2 ec of othe «same: bait. . Both

average size and species composition varied between the 100-200 cc

and: 2 .cc. baits; with some species: béine taken only in.traps of. one

size (unpublished data of H. Howden and S. Peck). Exactly what

types .of .behavior cause the species and size: differences in the

different traps remains an interesting question and one that obvi-

ously meeds further investigation.

Howden and Young: Panamanian Scarabaeinae 155

Another type of behavior that has only been partly investi—

gated might be categorized as a form of olfactory communication.

Young, working in Panama, noted that when Canthon angustatus

individuals were active at monkey droppings, individuals of its

principal competitor, Canthon lamprimus, surrounded the fecal

material at a distance of approximately 15 cm. They then remained

at this’ distance until most individuals of CC. “angustatus teft the

site. Laboratory experiments by Young indicated that in a one-

to-one- confrontation at ‘a limited tcod site, ©. ancugtaius was

competitively superior to C. lamprimus, depriving the latter of

food when it was in short supply. Field observations supported

this data by indicating an avoidance of C. angustatus by C.

lamprimus. Further investigation led to the observation that a

white viscous fluid, secreted from the anal area of C. angustatus

and coated over most of the body, appeared to be an aromatic

substance.” This substance functioned as an allelochemic since

in laboratory olfactometer tests C. lamprimus attempted to avoid

filter paper saturated with the substance. The ecological conse-

quence of this is that when monkey fecal material becomes less

available in the wet season, ©. lamprimus adults may either die

or shift to carrion as a food source whereas C. angustatus continues

to utilize the monkey feces. fa

There are many other aspects of Scarabaeinae behavior that

need investigation. Mammals often tend to utilize trails and move

in groups. Do dung beetles {follow these potential tcod sources or

locate them visually or by sound? Jmere 13 some very tenuous

evidence that at least a few species may do so by either method.

To date, information concerning the behavior and ecology of

neotropical scarabaeines is generally of a fragmented and anecdotal

nature, with very few studies on single species and a complete

absence of published research on community dynamics. This

situation has been due in part to an unstable taxonomy of the

group. Material presented in this review will hopefully provide

that stable taxonomic base and also give some indication of the

fascinating and diverse biologies found in the Scarabaeinae.

Hopefully this work will encourage and facilitate further research.

ACKNOWLEDGMENTS

The following individuals have made a particular effort to

accumulate material that greatly augmented our studies: Dr. Henry

Hespenheide, Department of Biology, University of California, Los

Angeles; Drs. Stewart and: Jarmila Peck, and Mrs. Anne Howden,

Carleton. University, Ottawa: Dr, -1,C. Poulson,:. Biological :Sciences,

University of illimers, Chicaoo Civcie: 0n., Breit OC. Ratelitie.

Division of Entomology, University of Nebraska, Lincoln; Mr.

Edward Riley, Department of Entomology, University of Missouri,

Columbia;. and» Dr. Henry Stockwell, Gorges: Hospital, Balboa

Heights, Canal Zone.

Dr. W.De Edmonds; Biological Selences, California, State

Polytechnic College, Pomona, assisted us with some of the taxonomic

problems encountered in the phanaeine genera and also loaned us

material. Some of the photographs used herein were taken by Mr.

156 Contrib. “Amert “Rate ciastiiy wel. i. ve. chs. 1061

Anthony Davies, Biosystematics Research Institute, Ottawa, and

all of the scanning electron microscope photographs were taken by

“Mr. Lewis Ling, Carleton University. Mrs. Betty Perry and Miss

Jennifer Read assisted with the preparation of the manuscript and

plates.

The following curators and their institutions kindly loaned

material: Drs; .E.C., Becker sand J.M. Campbell, Canadian. National

Collection, Biosystematics Research Institute, Ottawa, Ontario; Dr.

Henry Dybas, Field Museum of Natural History, Chicago, Illinois;

Dr. A. Newton, Museum of Comparative Zoology, Cambridge,

Massachusetts; and Dr. R. Gordon, United States National Museum

(Natural Mistery )) \Wasnineten,): D.C... Dr. J. jelinek,. Narodni

Muzeum, Prague, Czechoslovakia, kindly sent on loan a number of

Balthasar types. To those named above and others inadvertently

omitted, our sincere thanks.

Support for the work has been derived from a number of

sources, both personal and institutional. The laboratory portion

of the work, largely done in Ottawa, has been supported by the

Natural Sciences and Engineering Research Council of Canada.

This included the funding for a Postdoctorate Fellowship:.at

Carleton University for ©. Youne): Field work’ done in Panama. by

O. Young was supported by funding from the Smithsonian I[nstitu-

tion (STRI). The above assistance and support, both individual

and institutional, is gratefully acknowledged.

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1951..Notas ecoleopterologicas V. La invalidez. del nombre

generico Pinotus Erichson y dos nuevas sinonomias (Col.

Scarab.) Amal, Soc. Cient. Argentina <152:. 138-142.

1952. Scarabaeidae nuevos o poco conocidos III. Publ.

Mission Estud. Pat. Reg. Argentina 23: 53-118.

1967. Notas para una monografia del genero Trichillum

Harold, 1868 (Col., Scarab., Scarabaeinae—Coprini).

Rev. Soc. Mex. Hist. Nat. 28: 119-147.

Martinez, A., G. Halifter, and V. Haltiter.

1964. Notas sobre el genero Glaphyrocanthon (Coleopt.,

Goarabs, Canthonina):. Agta. Zool. .Méx... 7s 1-42.

Martinez, A.,°G: Halffter, and. F.S..° Pereira.

1964. Notes on the genus Canthidium Er. and allied genera.

Pativl. (Col, Searabacidae). Studia Ent.. 7: 161-178.

Martinez, A., ¢ and F.S. Pereira.

1967. Notas Escarabeidologicas I11. (Col. Scarabaeidae -

Scarabaeinae). Rev. Soc. Ent. Argentina 29: 53-69.

Matthews, E.G. :

1961. A revision of the genus Copris Muller of the western

hemisphere. Ent. Americana 41: 1-139.

1963. Observations on the ball-rolling behavior of Canthon

pilularime (.), (Coleeptera,. Scarabaeidae)... Psyche /0:

75-93.

1966. A taxonomic and zoogeographic survey of the Scarabaeinae

of the Antilles (Coleoptera: Scarabaeidae). Mem. Amer.

hit. Sees Zit talS4.

Muller, O.F.

1764. Fauna: insectorum Fridrichsdalina: Frid. Gleditschii,

Hafniae et Lipsiae. 96 pp.

Howden and Young: Panamanian Scarabaeinae 161

Murray; As

1856. Descriptions of new Coleoptera from the Western-Andes

and’ the’ neighbourhood of Quito. Proc.. Roy. Phys. Soc.

Edinburgh U: 20/-222,

Nevinson, B.G.

1891. On two undescribed species of the genus Phanaeus

Macleay. Ent. Monthly Mag. 27: 208-209.

Olsoufieff, Gd.

1924. Les Phanaeides (Coleoptera: Lamellicornia). Insecta,

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Paulian. oh.

1936. Une nouvelle espéces de Trichillum (Col. Scarabaeidae).

Bull; Soc. Ent. France 41: 205 2c.

1938. Contribution a l'étude des canthonides americains

(Coléopt. Lamellics). Anns Soe, Ent. Prance 107: 213-296.

1939. Three new neotropical coprophagous Coleoptera. Psyche

46: 141-146.

Pereira, F.S.

1943. Duas novas espécies e uma variedade de Oxysternon

Cast. (Coleopt. Scarab.). Arqu. Mus, Paranaense 3:

109-118.

Pereira, F.S., and A. Martinez.

1956. Algunas notas sinonimicas en Phanaeini. Rev. Brasil

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Pereira, F.S., M.A. Nulcano, end A. Martinez.

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1? 155-164,

Porter, . 1.

1973. The vegetation of Panama: a review. In Vegetation

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Alan Graham, ed. Elsevier Scientific Publ. Co.,

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feu Camille van Volxem pendant son voyage au Brésil et

a La Plata en 1872 suivie de la description de dix huit

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Belgique 30: 103-120.

Robinson, M.

1948. Two new species of neotropical Scarabaeidae (Coleoptera).

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Say, F

1835. Descriptions of new North American coleopterous insects,

and observations on some already described. Boston

Jour. NateoMist, lr 1515203;

Schamidt. hy

1920 (1921). Beitrag zur Kenntnis der Gattungen Canthon

Hifse., Sypax Boh.,’ Aphodius 11,7 Simoegenius Har:,

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Z. Verbretungsgebiete der Canthon-Arten. 3. Neubeschrei-

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162 Contoib Amer. int: nei. Meals s V8 y, me. 745.1981

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1962. A Gazetteer to accompany the ''Insecta'' volumes of the

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1826. In Eneyelopedie: methodique 4! i.e), @mtemolegie, Paris.

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1877. Description of some new species of beetles (Scarabaeidae)

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1974. Terrestrial, studies -— Barro Colorado Island. In 1973

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1964. Catalogue of the Canthonini (Col. Scarab.) inhabiting

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1801... Observationes entomologicae, continentes novorum quae

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1967. The effect of cattle dung patches on pasture growth,

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1842. Descriptions of some new exotic genera belonging to

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1847. Characters of various new groups and species amongst

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Wille, A.

1973. Coleoptera: Scarabaeidae. Observations on the behav-

L0Or’ Ol. a Tropical. rain forest, dune beetle,..Megathoposoma

candezi (Harold). Rev. Biol. Trop. 21: 41-5/;

Wille, Aw) Gos Guentesy. “BE. Oroezos.and). E.. Selis,

1974. Additional: observations om the; behavior iof a tropical

forest dung beetle, Megathoposoma candezi (Coleoptera:

Scarabaeidae). Rev. Biol. Trop.) 22: 129-133:

Wolda, H.

1978: Fluctuations in abundance of tropical insects. Amer.

Nat. i822: 1017-1045.

Young, .0: Fi.

1978. Resource partitioning in a neotropical necrophagous

scarab guild.: Ph.D. .thesis,«University of. Maryland,

College: Park.

Howden and Young: Panamanian Scarabaeinae 163

Fig. (20 Lunyeternus /plepejus ihearcyrdersum.. Fig 8.

burysternus: Caribacus Most., (thoracic sternma.’ Fie?) 4.>Burysternus

mexicanus (Germ.), thoracic sterna. Fig. 5. Megathoposoma candezei

Har.), dorsum. °Fig.'6. Canthom ‘moniliatus “Bates, dorsum, /Fig.

7s Canthon Vituratus (Germ,), dorsum.

164 Contrib. Amer. Ent. Inst., vol. 16, no. :1, 1981

Figs. 8-13. Eurysternus spp., left hind femur, ventral view.

8. claudicans Kirs., female. 9. plebejus Har., male and female.

10. velutinus Bates;,> female. 11: mexicanus: Har., .male. 2:

caribaecus Hbst., female... 13. magnus Lap., female.

165

Howden and Young: Panamanian Scarabaeinae

Fics 14. Dorsum,

Fig. 15. Male genitalia. Figs. 16, 17. Canthon juvencus Har.

Fies: 14, 45. Canthon morsel group.

17. Head and pronotum. Figs. 18, 19. Pseudocanthon

16. Dorsum.

18. Dorsum. 19. Head and pronotum.

perplexus (LeC.).

166 Contribs Amer: Ente sinsted wolkudS, bne. a4h981

° eo .

eo? oeeee,

28 29

Pigs. «20-29.) (Canthon .eppes, head@vandymate fore tibia: respec—

tively wo 20,«:21es montliatus. Bates... (22; 23. septemmaculatus (Latr.).

24.25. Geyrollei. Hare. 20~)27. te vehevrolatic Hara! 28). 29./literatus

(Germ. ). nh

Howden and Young: Panamanian Scarabaeinae 167

EF”

DOO

Figs. 30-37. Canthon spp., head and fore tibia respectively.

SOY CRT.” MUta DL ee Smears aber RD 1g ss Coe bed ; Mhars siete’.

34-37. aequinoctialis Har.; 34, 35. Male. 36, 37. Female.

168 Contrib. “Amer .o ENte inst. , volo 10, “nor bj. 7951

42> |

5 Pas,

J \,

44 45

Figs. 38-45. Canthon spp., head and male fore tibia respec-

tively. . 30,098 juwencus Har... 40.412 morsel group. 42; 43:

angustatus Har. 44, 45. subhyalinus Har.

Howden and Young: Panamanian Scarabaeinae 169

M Sy

46 47

be ne 53

Figs. 46-53. Canthon spp., head and male fore tibia respec-

tively. 46, 47. euryscelis Bates. 48, 49. femoralis (Chev.).

50, Sia vbeamprimus Bates. . 92, os. -v. mevidtonalis Ma, i. Seal.

170 Contribs ©Amer. Bate inet. Vole 18) onol 4981

Figs. 54-59. Deltochilum spp. 54. ¢. panamensis Howd.,

dorsum. 55. mexicanum Burm., head and pronotum. 56. parile

Bates, head and pronotum. 57. pseudoparile Paul., head and

pronotum. 58-59. v. acropyge Bates. 58. Head. 59. Abdomen

and pygidium, lateral view.

Howden and Young: Panamanian Scarabaeinae BL

Fig. 60. Cryptocanthon newtoni Howd., dorsum. Figs. 61-63.

Agamopus lampros Bates. 61. Dorsum. 62. Head and pronotum.

63. Pygidium. Figs. 64, 65. Bdelyrus seminudus (Bates). 64.

Dorsum. 65. Head and pronotum.

1/2 Comtribi: Amer. Ent. inst. ,2vol. 118, mo75), 1981

Figs. 66-68. Pedaridium pilosum (Rob.). 66. Dorsum. 67.

Head. 68. Male genitalia, apical dorsal view. Figs. 69-71.

Pedaridium brevisetosum ns sp. 69. Dorsum, .70. Head. 71. Male

genitalia, apical dorsal view.

Howden and Young: Panamanian Scarabaeinae LPS

Pies. (2-7/4. Pederidium bettimeri ninsp.. (2. Doreume 623.

Head. 74. Male genitalia, apical dorsal view. Figs. 75, 76.

Onthocharis panamensis Paul. 75. Dorsum. 76. Head and pronotum.

Fie. 77. Scalimus, quadridentatus Balth., head and pronotum.

174 Contrib, sAmer. Ent: “iist., ‘vol. 10, mo. 1, 1961

Figs. 78, 79. Scatimus quadridentatus Balth. 78. Head.

79. Male genitalia, lateral view. Figs. 80, 81. Scatimus ovatus

Har. 90...cead..- Ol... Male genitalia, lateral view, “Figs. 82, -83.

Uroxys gorgon Arrow. 82. Head and pronotum. 83. Male genitalia,

lateral. wiew..

Howden and Young: Panamanian Scarabaeinae 175

88 89

Figs. 84-93. Uroxys spp., head and pygidium respectively.

04, 05.. DiatyDVea. Ms ep: Sh, 0/.macrocwlaria in. . sda, Ob, Oo.

bidenltigums spy 690, Uli mibves, Bates. 92, 92. microeularis' n. sp.

176 Contrib. “Amer. Ent) Inst., wole 18,001) )19601

101

103 é

Figs. 94-103. Uroxys spp., head and pygidium respectively.

G4, 05. Gatumensis n. Sp. OG. OP dyvbast im. Sp. 08, 99,

depressifrons n. Sp. 100, 101. metagorgon nm, sp. 102, /103.

gorgon Arrow.

6=9

Howden and Young: Panamanian Scarabaeinae 17}

Figs. 104-109. Uroxys spp. 104. depressifrons n. sp., head.

105. depressifrons n. Sp., Male eenitalia...lateral view: °106.

melagorgon mn, 6p... male genitalia, lateral wiew,. 7107: platypyga

n. Sp., male genitalia, lateral view. 108, 109. micros Bates.

108. Dorsum. 109. Head and pronotum.

178 Contrib. Amer. ont. Insts, vole 76) no. 1, 4981

Figs. 110-114. Uroxys spp., mate genitalia. 110. micrcs

Bates, lateral views. ill:. bidentis. mn. sp., lateral.view. dl2.

microctilaris nn. 6p.; lateral view.« bli, microoularis n. sp»,

dorsal view. «114; macrocularis mn. sp., lateral view. 115. dybasi

NH. SDu, Head and pronotum.

Howden and Young: Panamanian Scarabaeinae Lg

Pigs. 116, 117. Uroxys dybasi ni: sp., male genitalia,. 116.

Lateral wiew.) ~ 117. Dorsal view. « Pigs...110-121. ‘Ateuchus. spp.

118. candezei (Har.); dorsum: 119) candezei(Har:), head and

pronotum. 120. aeneomicans (Har.), head and pronotum. 121.

guatemalensis (Bates), head and pronotum.

180 Contriows Amer ss “Ente Aneel. ols) lemon dy 2901

Piper 122-125 sCanthidium centrale Bouc. \( 122. Head...’ 123.

Apex, male fore tibia. 124. Male genitalia, lateral view. 125.

Male genitalia, dorsal view... Figs. 126i 127... Canbiidium

tuberifrons nm. spin 126. Head and: pronetummeoi 12). Posterior margin

of pronotum.

Howden and Young: Panamanian Scarabaeinae 181

Figs. 128, 129. Canthidium tuberifrons n. sp., male genitalia.

128. Lateral view. 129. Dorsal view. Figs. 130-133. Canthidium

harold: Preud,: 11306. Heads): 18lu- Male; fore ti bila! °¥32. Male

genitalia, lateral view. 133. Male genitalia, dorsal view.

182 Contribs: Amer) (Ent. inst., “vol. 18) no. 1, 1981

Figs. 134-136.. Canthidium angusticeps Bates. 134. Head.

135. Male genitalia, lateral view. 136. Male genitalia, dorsal

View. Figs. 13/-199. Canthidium elegantulum Balth. 237. Head.

138. Posterior margin .éf pronotums: 139." Male genitalia, lateral

view.

Howden and Young: Panamanian Scarabaeinae 183

Fig. 140. Canthidium elegantulum Balth., male genitalia,

dorsal view. Figs. 141-143. Canthidium bituberifrons n. sp.

141. Head. 142. Male genitalia, lateral view. 143. Male genitalia,

dorsal view.. Figs; 1445. 145. Canthidium disecoycidiale in. sp.

144. Head. 145. Male genitalia, lateral view.

184 Contrib. Amer sbnte Inst.) owelviclGs nol ial 961

Fig. 146. Canthidium discopygidiale n. sp., male genitalia,

dorsal view. Figs. 147-149. Canthidium subdopuncticolle n. sp.

147. Head. 148. Male genitalia, lateral view. 149. Male genitalia,

dorsal View. cFigsi (50; 0151. ‘Canthidium téenebrosum n. ‘sp.

150) ‘Head. 151... Male \wenitalia;: lateral view.

Howden and Young: Panamanian Scarabaeinae 185

Fig. 152. Canthidium tenebrosum n. sp., male genitalia,

dorsal view. Figs. 153-155. Canthidium planovultum n. sp.

153. Head. 154. Male genitalia, lateral view. 155. Male genitalia,

dorsal view. Figs. 156, 157. Canthidium perceptibile n. Sp.

156. Head. 157. Male genitalia, lateral view.

186 Contrib. Amer. Ent. fast... vole 18, no. 4) 1981

Fig. 158. Canthidium perceptibile n. sp., male genitalia,

dorsal view. Figs. 159-161. Canthidium vespertinum n. sp.

159. Head. 160. Male genitalia, lateral view. 161. Male genitalia,

dersal view. Figs.1672,-163..Canthidium pallidoalatum n.. sp.

162. Head. 163. Male genitalia, lateral view.

Howden and Young: Panamanian Scarabaeinae 187

Fig. 164. Canthidium pallidoglatum n. So... male Genitalia,

dorsal view. Figs. 165-167. Canthidium variolosum n. Sp. 165,

Head. 166. Male genitalia, lateral view. 167. Male genitalia,

dorsal view. Figs. 168, 169. Canthidium leucopterum n. sp.

168. Head. 169. Male genitalia, lateral view.

188 Contribi Ameri int. lst. owel U8) anos ls 1981

Fig. (170. Canthidium leucopterum ms) spi, imale. genitalia,

dorsal view. © Fies. 17i-173.> Canthidium ardens Bates. 1/1. Head.

172. Male genitalia, lateral view. 173. Male genitalia, dorsal

View. (bigs) 2/4, 175. Canthidium eurtiex Bates, 1/4. Dorsum.

17>. Head.

Howden and Young: Panamanian Scarabaeinae 189

Figs. 176-178. Canthidium aurifex Bates. 176. Apex male

fore tibia. 177. Male genitalia, lateral view. 178. Male genitalia,

dorsal view. Figs. 179-181. Canthidium hespenheidei n. sp.

179. Head. i 160, "Posterior margin of orenetum. 161.) Male

genitalia, lateral view.

190 Contrib. Amer. “bnte dimsts, vols 123; no. 1, 21981

Fico. 102. Canthidium heepenheidern..sp., male genitalia,

dorsal view. Figs. 183-186. Onthophagus spp., male head and

pronotum.. 163, ineensus Say... 184. Datesi Hy .&C. 1) 185.0c:

panamensis Bates. 186. stockwelli n. sp. Fig. 187. Onthophagus

stockwelli n. sp., male fore tibia.

Howden and Young: Panamanian Scarabaeinae Ol

Figs. 188-193. Onthophagus spp., head and pronotum. 188,

189. stockwelli n. sp., female.” 190. nyctopus Bates; male. 191.

acuminatugs Har. male... 192.) transisthmius m.-sp.', male.’ 193.

transisthmius n. sp., female.

192 Contribs Aner. Ent. tmet. ss vobo 18) ones, 1981

Figs. 194-198. Onthophagus spp., head and pronotum.

(94. maroinicollis Har., male. 195. imereinicoblis, Har. female.

196. lebasi Bouc., male. -197. lebasi Bouc., female. 198. chryses

Bates, male. Fig. 199. Onthophagus coscineus Bates, dorsum.

Howden and Young: Panamanian Scarabaeinae 0

Figs. 200-205. Onthophagus spp., male head and pronotum.

200... ‘eoseineus” Bates. (201, idicranius Bates...) 202.) landolti Har.

203. atrosericeus Bowe, 204. °cyanellus Bates. ° 205. (‘mirabilis

Bates, male, major form,

194 Contrib. Amer. Ents inst... vol! 16.0 nes 1, 196!

Figs. 206-210. Onthophagus spp., head and pronotum. 206.

mirabilis’ Bates, male, minor form. 20/7. mirabilis Bates, demale.

208. sharp Har., male, 209. praecellens Bates, male. ..210.

belorhinus Bates, male. Fig. 211. Ontherus: brevipennis Har., head.

Howden and Young: Panamanian Scarabaeinae 195

21 17

Figs. 212,.4135 (Ontherus, didymus Ers, male. . 212. Dorsum.

213. Head and pronotum. Figs. 214-216. Ontherus strius n. sp.,

male. 214. Head and pronotum. 215. Male genitalia, lateral

view. 216. Hind femur and tibia. Fig. 217. Dichotomius c.

colonicus (Say), dorsum. ee

196 Contrib.) Amere Bints slat. jy vol? bOy, mo. 1, 1981

Figs. 218-223. Dichotomius spp. 218. nevermanni (Lued.),

female. ZITO satsnas (dar. ) male. o.220, agenor (Har.), dorsum

ane

of male. “221. acenom (|Har.), male head: 222. “inachus:(Er.),

head and pronotum. 223. femoratus n. sp., head of female.

Howden and Young: Panamanian Scarabaeinae Vo7

229

Figs. 224-226. Dichotomius femoratus n. SO. Maley, 224,

Head and pronotum. 225. Male genitalia, lateral view. 226. Male

genitalia, dorsal view. Figs. 227-229. Dichotomius gamboaensis

n. sp., male. (227. Head and pronotum. 228, Male genitalia,

lateral view. 229. Male genitalia, dorsal view.

198 Contribe “Amer; Ent. Inst.2voly 16, no.c 1) >.) 1981

235

Figs. 230-233. Copris spp., male head and pronotum. 230.

lugubris Bohs v.23]. incertus Say. . 232. ¢costaricensis Gahan.

230.. subpunctatus: Gillis. figs. 234, 255. FPhanaeus.spp.,. male

dorsum. 234. hermes Har., minor form. 235. beltianus Bates.

Howden and Young: Panamanian Scarabaeinae 199

Higs. 236,207, BPhanaeus sop.. male. . 236... beltianus. Bates,

genitalia, dorsal view. 237. pyrois Bates, head and pronotum.

Figs. 230-240. Suleophanacus sop., male. 238. velutinus (Murr.),

dorsum, 439. .cupricolliis, (Nev.), head and pronotum, 240. noctis

(Bates], dorsum. Fie. 221. Coprophnandeus t. coryvthus (Har.). male

head and pronotum. ie

200 Gontrib. Amero vEnt. bnst., wolo (16,09 nord) 1981

Figs. 242, 243. Coprophanaeus t. corythus (Har.), male

genitalia. 242. Lateral view. 243. Dorsal view. Figs.) 244-247.

Coprophanaeus chiriquensis Ols., male. 244. Dorsum. 245. Head

and pronotum. 246. Genitalia, lateral view. 247. Genitalia,

dorsal view.

Howden and Young: Panamanian Scarabaeinae 201

Figs. 248-250. Coprophanaeus ohausi (Felsche), male. 248.

Head and pronotum. 249. Genitalia, lateral view. 250. Genitalia,

dorsal view, . Kilgs.. 251-253. Coprophanaeus, pecki n. sp... male.

2ol, Doveum. Bae Genitalia. lateral View, 25a. Genitalia.

dorsal view.

202 Contribs Amer. Eni... nist, vol. 162. 90, 1.196)

Figs. 254-256. Oxysternon conspicillatum (Web.), male.

254° Dorsum. 255. Genitalia, latera. view. 256. Genitalia; dorsal

Vipwe Pies. 25/-259; Oxysternon Zikani Per. , «male. 25/7. /Dorsum.

250. Genitalia; lateral view. “259. Genitalia, dorsal’ view.

203

Howden and Young: Panamanian Scarabaeinae

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A Revision of the Genus Aphanistes of America North

of Mexico (Hymenoptera: Ichneumonidae)

H. Pearson Hopper

3713 35th Street, NW, Washington, DC 20016

ABSTRACT

A review of the species of the genus Aphanistes of America north of

Mexico has been made on the basis of over 800 specimens from eleven collec-

tions. Nineteen species are described, of which fourteen are new.

INTRODUC TION

This study was originally intended to be the first step in a revision of the

tribe Anomalini (now elevated to subfamily rank as Anomaloninae; the change

in form of the base of the name is justified by a different interpretation of the

genus name--see Hopper (1959), p. 159, Anomalon, Note--and required be-

cause the other form is preoccupied). However, owing to personal problems,

the analysis of the present genus has taken so long that the larger project has

had to be abandoned, and I am therefore glad to learn that Dr. Clement E.,

Dasch of Muskingum College is undertaking a revision of the subfamily.

Since for the most part the characters distinguishing the species of this

genus are habitus characters, I have made a great many measurements in an

attempt to represent the various proportions in numerical form. Unfortunately,

in many cases the measurements either were incorrectly designed or for other

reasons do not show what was wanted, or merely show a trend in the direction

of what was expected and are not discriminative. In some cases, a measure-

ment that is discriminative for the female is not so for the male (especially

those concerning the hindbasitarsus), Consequently, several sets of measure-

ments on which I had spent considerable time have not been included in this

paper. In addition, I have been compelled to some extent to use in the keys

the relative kind of terminology I had hoped the measurements would eliminate.

Possibly other measurements that were not made might have been productive,

but either I have not thought of them or they have come to mind too late, after

pressure to finish had become too strong. This latter factor has also resulted

in less consistency in the specjes descriptions than I consider desirable, since

further studies not strictly necessary would have been needed to make up for

the differences in point of view due to the rather large time spread between

my study of different species. Some additional studies that I did find necessary

were, of course, made,

The catalog data are limited to original descriptions and have been taken

from Krombein, Hurd, Smith, Burks, et al. (1979), although all the original

descriptions have been consulted (and the types examined).

2 Contrib. Amer. Ent. Inst., vol. 18, no. 2, 1981

The description of the genus is a synthesis of published descriptions con-

sulted (especially Gauld, 1976, pp. 41-42 and Townes, 1971, pp. 131-132) plus

my observation of the material used for this paper. The species descriptions

are based on all specimens at hand.

Collection data of the new species are arranged alphabetically by state or

province, then alphabetically by locality, and under this in date order. Cer-

tain labels on Forest Insect Survey (F.I.S.) specimens from the Canadian

National Collection bearing only a number have not been listed,

Host records have been taken from pin labels and have not been verified.

It should be noted that almost all specimens of the genus that I have seen

with identification labels have been misidentified as to species.

ACKNOWLEDGEMENTS

I am particularly obliged to Dr. Henry K. Townes of the American

Entomological Institute, Ann Arbor, Michigan and Dr. Robert W. Carlson,

Systematic Entomology Laboratory, United States Department of Agriculture,

Washington, D. C. for extensive advice and other assistance, toG. S. Walley

and W. R. M. Mason, Biosystematics Research Institute, Agriculture Canada,

Ottawa, Canada for assistance with loans from the Canadian National Collec-

tion and other help, to Luella M. Walkley and Robert W. Carlson, Systematic

Entomology Laboratory, United States Department of Agriculture, for assis-

tance in obtaining material from the United States National Museum, to

Dr. Townes for the loan of material from the Townes collection, R. R.

Dreisbach collection, and the collections of Michigan State University (East

Lansing, Michigan), The University of Michigan (Ann Arbor, Michigan),

University of California (Berkeley and Davis, California), and the Carnegie

Museum of Natural History (Pittsburgh, Pennsylvania), to Dr. Charles

Remington (Peabody Museum, Yale University, New Haven, Connecticut) for

access to the Norton types, to Dr. Curtis Dunn (Academy of Natural Sciences

of Philadelphia, Pennsylvania) for access to the Davis types and for the loan

of one of them, to the respective institutions concerned, and to Robert T.

Mitchell, University Park, Maryland, for the loan of a number of specimens.

The material examined numbered over 800 specimens, the majority from

the Townes collection and the Canadian National Collection, and included the

types of all published species.

The following abbreviations have been used to identify the institutions and

collections in which the types and paratypes of the new species described will

be found.

CHT = collection of Henry and Marjorie Townes, American Entomological

Institute, Ann Arbor, Michigan }

CNC = Canadian National Collection, Ottawa, Ontario, Canada

Dreisbach = collection of R. R. Dreisbach, Midland, Michigan (now

incorporated in the Michigan State University collection).

HPH = collection of the author, Washington, D. C.

MSEL = Michigan State University, East Lansing, Michigan.

Pittsburgh = Carnegie Museum of Natural History, Pittsburgh, Pennsylvania

RTM = collection of Robert T, Mitchell, University Park, Maryland

UCD = University of California, Davis

UMI = University of Michigan, Ann Arbor, Michigan

USNM = U. S. National Museum of Natural History, Washington, D. C.

Hopper: Nearctic Aphanistes 3

TERMINOLOGY

The terminology used in this paper agrees in general with that given in

Townes (1969, pp. 36-48). Below I give some definitions of terms not found

there or for which my usage differs from that of Townes.

Area dentipara: I use this term (used by European authors, especially for

the Ichneumoninae, in which it sometimes gives rise to a tooth), for the sake

of brevity, for the second lateral area of the propodeum, which in most speci-

mens is not separated from the first lateral area, If separated, the first

lateral area is more than twice as wide as long,

Cheek: The lower part of the gena (as defined in general works, e.¢.,

Pratt, 1940--‘‘temple’’ of Townes, see below) from the base of the mandible

to the point where the orbit begins to turn toward the horizontal, .

Crista: The crest or horn (called ‘‘lamella’’ by some authors) on the

frons between and above the antennal sockets, between the antennal scrobes,

and continuing as a mesal carina to the anterior margin of the stemmaticum.

Gena: The part called by this name in morphological works (‘‘ temple’’ of

Townes).

Inner orbit: Either one of the two lateral longitudinal regions of the face

adjacent to the orbits, the parafrontal of Pratt (1940, pp. 157- -158, plate V,

fig. 1), usually set off from the median field (which see) by a more or less

distinct longitudinal depression (this is visible also in Townes, 1969, p. 48,

fig. F). Called << facial orbit’? by Townes.

Interocellar line: The line between the inner margins of the lateral ocelli

(or its length).

Median field of face: The mesal eAntiol of the face, the antefrons of Pratt

(1940). I have taken the term from Heinrich (1961, 1967).

Metakatepimerum: The upper part of the lateral portion of the propodeum

labeled ‘ ‘lower division of metapleurum”’’ in Townes (1969, pp. 44-45, fig. D,

15) and ‘‘kemg’’ in Pratt (1940, plate VI, fig. 9), abutting on the hindcoxa,

Metakatepisternum: The lower part of the metapleurum, adjacent to the

metasternum (Pratt, 1940, plate VI, fig. 9, kes3). The juxtacoxal area of

Townes (1969, pp. 44-45, figure D, 16), but note that when the so-called juxta-

coxal carina is complete, it reaches the metasternum before the insertion of

the coxa.

Metanepimerum: The more or less pillow- or lozenge-shaped structure

between the lateral portion of the metanotum (also called ‘ ‘ postscutellum’’)

and the metapleurum (Pratt, 1940, plate VI, fig. 9, aemg3). The upper divi-

Sion of the metapleurum of Townes (1969, pp. 44-45, fig. D, 14).

Metanepisternum: The structure resembling an expanded carina caudad

from the suture between mesopleurum and metapleurum, extending from the

base of the hindwing to behind the midcoxa (Pratt, 1940, plate VI, fig. 9, aesg).

Metapleurum: The metakatepimerum and metakatepisternum taken

together. Although the anepimerum and anepisternum of the metathorax are

morphologically part of the metapleurum, in Ichneumonidae they are structural-

ly associated with the mesothorax and are therefore not included in my use of

the word metapleurum.,

Obsolete: Barely distinguishable, but not completely absent,

Ocellocular line: (The length of) an imaginary line from the lateral margin

of a lateral ocellus to the nearest point of the eye margin.

Ocellocular space: An area centering on the ocellocular line, usually

differing in sculpture from frons, stemmaticum, and vertex and containing the

4 Contrib, Amer. Ent. Inst., vol. 18, no. 2, 1981

orbital spot.

Orbital spot: A spot of color differing from adjacent areas, adjacent to

the eye margin and occupying part or rarely all of the ocellocular space.

Paracostula: The portion of the apical transverse carina of the propodeum

separating the area dentipara from the petiolar area and third lateral area

(the portion bounding on the petiolar area is part of both the apical transverse

carina and median longitudinal carina),

Parapsides: The two lateral portions of the mesoscutum (in the strict

morphological sense) separated from the mesoprescutum by the notauli.

Called lateral lobes of mesoscutum in Townes (1969).

Postocciput: The reflexed flange around the foramen magnum, more or

less perpendicular to the occiput. It can be seen in Townes (1969) in figure F,

p. 48 but is not labeled, It is the flange that can be seen around part 15 (it is

not part 14, ‘‘ postocciput’’ in the legend as originally printed but changed to

occiput in the Corrigenda).

Postpetiole: The portion of the apparent first abdominal segment from the

anterior margin of the spiracle (including the rim usually present) to the apex

of the tergite. Note the inclusion of the spiracle (in contrast to Townes,

1949), which makes a difference in measuring the proportions involving this

part in the descriptions.

Prescutum: (mesoprescutum): The portion of the mesonotum between the

notauli (median lobe of mesoscutum in Townes, 1969).

Propodeal fossa: I am proposing this term for the longitudinal troughlike

depression occurring mainly in the pleural area of the propodeum in most

species of Aphanistes. The fossa often appears to involve an adjacent portion

of the metapleurum. It actually can be identified in all our species, although

in some it is very weakly developed (a flat surface between convex ones,

rather than a depression. )

Propodeal pedicel: The pair of processes forming the socket for the

petiole.

Scutellar depression: The transverse depression between mesoscutum

and scutellum,

Stemmaticum: A more or less distinct portion of the top of the head,

encompassing the ocelli and usually a narrow strip outside them, It is often

more or less elevated and distinguished from the frons and vertex by sculp-

ture (the ocellar triangle of Townes, 1969).

Subantennal carina: The carina surmounting the ridge found below and in

front of the antennae.

Temple: The upper part of the gena (in the morphological sense, see

above) from about the summit of the eye to where the line of the gena begins

to become less strongly curved. (Note difference from definition of Townes,

1969).

MEASUREMENTS

The following criteria apply to the measurements given: The length of

the forewing is measured from the distal margin of the tegula to the tip of

the wing. The bodylength has been determined progressively (no doubt giving

a result somewhat exceeding the length of the living insect in a natural attitude,

but one which is replicable), i.e., from the edge of the subantennal carina (note

that the head is included here in the term ‘‘body’’) to the apex of the propodeal

pedicel, from that point to the caudal margin of the first tergite, and thence to

Hopper: Nearctic Aphanistes 5

the apex of the abdomen (excluding the terminalia) in as many instalments as

required by it positioning. The antenna length has in some cases been measured

by relaxing the appendage and straightening it, thus providing a precise mea-

surement, or in many instances the length has been approximated by succes-

sive measurements along the curves or, less often, by estimating--the total

includes the scape and pedicel. The number of antennal segments includes

only those of the flagellum. The length of the first flagellar segment in rela-

tion to its width is measured viewed in the narrowest aspect, taking the width

at the distal end and the length along the centerline including the proximal

bulb of the segment that usually appears to be part of it and is considered to

be so by Pratt (1940), but I have known it to separate as if it were a distinct

segment, which it is considered to be by Gauld (1976, p. 5, ‘‘ anellus’’--

doubtless this is the correct interpretation). The combined length of the

second and third flagellar segments times the first is also taken from mea-

surements made along the midline (not to the he farthest point of the oblique

margin of the segments),

The various proportional measurements are made as indicated below.

The head thickness-width ratio is the maximum span across the eyes

divided by the distance between a transverse line tangent to the central prom-

inence of the subantennal carina and a line parallel to it drawn across the

outer edges of the occipital carina with the head oriented to bring the lateral

extremes of the eyes, the central prominence of the subantennal carina, and

the postocciput into focus simultaneously (differing orientation of the plane of

the occiput gives very different results in the measurements),

The head length-width ratio is the eye span determined as in the fore-

going divided by the length of the head from the tip of the clypeal tooth to

the summit of the lateral ocellus. The eye span may be measured either from

in front or from above, and the length I have measured with the head in pro-

file oriented to bring the tip of the clypeal tooth and the tip of the frontal crista

into focus simultaneously.

The head thickness-camber ratio is the quotient of the thickness of the head

determined as for the head thickness-width ratio divided by the length of the

perpendicular from the midpoint of the occipital carina to a line drawn across

the outer edges of the occipital carina, with the head oriented as above.

The occipital camber is the quotient of the length of a line connecting

the outer edges of the occipital carina divided by the length of the perpen-

dicular from that line to the midpoint of the occipital carina, with the head

oriented as above.

The head-occiput width ratio is the eye span as measured above divided

by the distance across the outer edges of the occipital carina, with the head

oriented as indicated in the foregoing.

The eye-temple ratio is the quotient of the length of a line parallel

to the midline of the head measured between perpendiculars to the midline at

the anterior and posterior points of the eye, divided by the distance between

a line across the outer edges of the occipital carina and a line parallel to it

through the posterior margin of the eye, with the head oriented as above.

The mesoscutum index is the quotient of the length of the mesoscutum as

defined above by its width, the length measured from the cephalic margin of

the prescutum to the bottom of the scutellar depression, with these two points

in simultaneous focus, and the width across the widest point (usually formed

by two slight eminences in the upturned marginal carina of the mesoscutum

and always just mesad of the forward edge of the tegula) with the two sides in

simultaneous focus (usually all four points can be brought into focus simultane-

6 Contrib. Amer, Ent. Inst., vol. 18, no. 2, 1981

ously).

by the latter. The measurements are made as already described,

The scutellum proportions are given in the form 1:x:y, where 1 represents

the width of the disc of the scutellum measured across the anterior edge (not

including the apophyses joining it to the scutum), x is the length from the

cephalic margin (usually marked by a more or less distinct carina) to the cau-

dal margin (including the more or less flangelike apical carina), and y is the

apical (caudal) width of the disc between the lateral carinae (if present) in front

of the apical carina,

Depth:width:length of propodeum is given in the same form as the foregoing,

taking the depth as unity. The depth is measured from the highest point of the

median longitudinal carina at approximately the anterior end of the areola to

the tip of the small, flat lobe at the point where the lower anterior margin of

the metapleurum meets the metasternum near the midcoxa, the width is

measured at the widest point as viewed from above or below, and the length is

measured in profile from the depression that appears to be the anterior bound -

ary to the tip of the propodeal pedicel.

‘« Hindbasitarsus n times as long as thick’’ is the quotient of the visible

length divided by the maximum thickness of the segment in side view (perpen-

dicular to the plane of motion of the tibia relative to the femur), which may be

either at or near the distal end.

‘¢ First tergite n times as long as hindbasitarsus’’ is determined from the

visible length of the former measured from the tip of the propodeal pedicel to

the caudal margin of the postpetiole, divided by the length of the latter mea-

sured as in the preceding paragraph.

‘« Width of postpetiole n times length’’ is the quotient of the maximum width

of the postpetiole (whether across the spiracles or behind them) divided by its

length according to the definition. :

‘«First tergite n times as long as postpetiole’’ is determined according to

the specifications in the two previous paragraphs.

Genus Aphanistes Foerster

Aphanistes Foerster, 1868. Naturh. Ver. preuss. Rheinl., Verh. 25: 145.

Type: Anomalon bellicosum Wesmael. Desig. by Viereck, 1914.

Anochilacrum Enderlein, 1921. Stettin. ent. Ztg, 82: 12,

Type: Anochilacrum flavigena Enderlein. Monotypic and orig. desig..

Apex of clypeus prolonged into a moderately large tooth, clypeus separated

from the face in our species by a shallow, rounded indentation, sometimes

reduced to merely a slight change of direction, never a sharp groove or suture,

usually with considerably fewer and larger punctures than face, with difference

between species in this usually not readily perceptible. Frons with a short,

high median longitudinal carina or lamella between and just above antennal soc-

kets and extending to the median ocellus and forming a crista (flattened horn or

tooth--absent in some exotic species). Occipital carina sometimes not as near

outer margin of head as in most other genera of the subfamily, the width across

the eyes in ours being from 1,14--1.55 times the width across the occipital

carina (the former value typical of the subfamily, the latter not so--Some spe-

cies with intermediate head-occiput width ratio have the temples bulging, so

that the head appears wider behind than indicated by the ratio). Ocelli usually

Hopper: Nearctic Aphanistes 7

of medium size (their diameter approximately equal to ocellocular and inter-

ocellar lines), in some species (only one of ours) very much larger, Eye

sometimes completely hairless, more often with sparse hairs of about the

length of the diameter of an ommatidium (visible only at about 30x). Upper

tooth of mandible slightly longer. Occipital carina complete, joining oral

carina at base of mandible. Epomia moderately to strongly divergent from

the carina along collar, then curved dorsad; enclosure from slightly larger

than tegula to about one-fourth that size. Mesoscutum descending abruptly

cephalad with a narrow flat shelf or concavity just behind the anterior margin

and before the abrupt portion; mesoscutum more or less punctate, Notauli

rarely sharp, more often narrow and distinct to broad and shallow to indistinct

or obsolete (occasionally indicated only by elevation of the prescutum above the

parapsides); they rarely meet before the prescutellar patch (usually black

and ruguloso-reticulate, with transverse rugulae predominant), more often

disappearing into it. Disc of scutellum from slightly concave to fairly strongly

convex, lateral carina from strong to obsolete (I believe it is never wholly

absent in our species, but may be very difficult to distinguish); disc some-

times with a median longitudinal depression (Sometimes relatively narrow,

sometimes extending the full width, in which case it is cylindroid). Prepectus

sometimes barely differentiated on mesopleurum, carina always present on

sternum, extending either just barely above the sternaulus or farther accord-

ing to species, but never clearly reaching margin of pronotum above; ster-

naulus rather distinct to obsolete, one-third to one-half the length of meso-

pleurum, Propodeum rugose-reticulate, usually without distinct carinae and

areolation, although portions of most can be distinguished, especially the

pleural carina, the portion of the lateral and median longitudinal carinae along

the areola and area dentipara, these areas (the areola never distinctly closed

behind and the area dentipara rarely separated from the first lateral area)

and the basal area (very short and descending abruptly into the groove between

propodeum and postscutellum); irregularity of the bounding carinae and the

presence of internal reticulations may considerably obscure the area dentipara,

but never completely in our species, and actually most of the other portions

of the carinae can be made out as zigzag lines stronger than the intersecting

rugulosity, which, for instance (in combination with the conformation of the

propodeum) permit the petiolar area to be made out. The caudal extremity of

the propodeum is produced as a pedicel to which the petiole of the abdomen is

articulated. Forecoxa without carina, occasionally barely convex below, some-

times moderately convex below, in one North American species with an oblique

ridge (not sharp) below. Midtibia with two spurs, hindtibia and -tarsus unspe-

cialized, the latter more slender and parallel-sided or rarely tapering distally

in female, stouter and usually diverging distally in male (in female sometimes

slightly compressed in plane of tibia-femur, in male often broader at right

angles to this plane). First abscissa of discoideus not nearly in line with and

distinctly shorter than discocubitus; postnervulus broken slightly below middle

(subdiscoideus below middle of brachial cell); second recurrent distinctly post-

furcal (distad of second intercubitus); first abscissa of radiella longer than

intercubitella; nervellus distinctly broken well below middle; areolet (second

intercubitus) absent. Ovipositor 0, '75-1.3 times the length of the first tergite

in our species,

Insects of moderate size (in our species, forewing 4. 86-14.55mm, body

including head 8, 3-24,3 mm), varying in color in North American species

from yellowish fulvous to ferruginous, with tip of mandible, top of head more

or less (at least part of stemmaticum), apical one-third to one-half of hind-

8 Contrib. Amer, Ent, Inst., vol. 18, no. 2, 1981

tibia, and second tergite except lateral margins, black; usually also black

are upper part of occiput, patch on mesoscutum before scutellum, and hind-

coxa above; often also black are a spot below wings, metanepimerum, spot at

base of propodeum, sterna and hindtrochanters more or less, and spot or spots

on apical tergites; rarely thorax wholly black; clypeal tooth (mucro) fulvous to

piceous, Tegula always luteous to fulvous., In one species, abdomen wholly or

almost wholly piceous.

Hosts of specimens examined (according to pin labels) are Geometridae,

Noctuidae, and Notodontidae.

Consideration of the various characters shows that the interrelations of

the species are by no means linear, so that the sequence adopted is a mere

approximation to the actual relationships. The key that follows is artificial

and does not express the relationships of the species.

Key to the species of Aphanistes of North America north of Mexico

1. Ocelli very large, ocellocular line not more than 0.5 times diameter

of lateral ocellus; eye very large.

1, crepuscularis, new species (p. 13)

Ocelli not so, ocellocular line 1,0-2,0 times diameter of lateral ocellus;

Oya Veale TOA IVely SIN Aer i. ae es es Geek ese ene we

2, Second tergite dark to lateral margins. Abdomen usually wholly dark

brown; at least apical tergites partly or wholly brown to piceous.

Midcoxa nearly always infuscate below. (Hindfemur stout, usually

dark brown, at least above. First tergite usually distinctly deeper

than wide before spiracles or both deepened and widened. Face and

clypeus in profile forming nearly straight line; top of head, mesoscutum,

and mesopleurum rather strongly punctate, opaque; interocellar line

1,14 times ocellocular line; temple usually distinctly bulging; head

width 9 1.27-1.4, of 1. 24-1. 38 times mesoscutum width. )

4, edemae (Davis) (p. 19)

second tergite not dark to lateral margins. Third and fourth tergites

never dark brown. Midcoxa always yellow or ferruginous below. .. 3

3, Petiolar area and third lateral area of propodeum usually with only

transverse rugae or rugulae. Scutellum usually rather rounded at

apex, Propodeum usually with transverse black band at base,

metakatepisternum usually yellow; thorax and propodeum more or

less tinged with yellow. (Head moderately convergent behind, temple

nearly flat; median field of face with few or no longitudinal wrinkles;

head width 9? 1. 4-1. 6, o1. 40-1. 43 times mesoscutum width; frons

rugulose; stemmaticum and vertex behind ocelli closely deeply punctate,

more or less rugulosopunctate; interocellar line 1.125 times ocellocular

line; hindfemur slightly stout; hindbasitarsus ? 9-11, o° 8-11 times as

long as thick; first tergite usually apically black or piceous, lighter

anteriorly--entire tergite Oe TeS fulvous in small specimens. )

3, hyalinis (Norton) (p. 16)

Petiolar area rarely with only transverse rugae. Scutellum truncate at

apex. Propodeum rarely with transverse black band, metakatepister-

num not yellow (Metapleurum yellowish in one species, in which

temple rather convex and sculpture rather fine). (First tergite rarely

BG Se a a eG Gea ed a Gods elles Rant ke eas

Hopper: Nearctic Aphanistes 9

Forecoxa with strong, oblique ventral ridge (narrowly rounded, not a

carina), surface on either side of ridge not smoothly rounded,

distally rather flat, proximally slightly concave, (Mesothorax rather

strongly punctate; face rather narrow and markedly convergent below).

6. coxatus, new species (p. 23)

Forecoxa usually uniformly rounded (rarely with an indistinct ridge barely

differentiated from the rounded surface). ..... Piyiauied uia o,

Head strongly convergent behind, relatively thin front to back, temple

nearly flat; top of head, mesoscutum, and mesopleurum moderately

strongly punctate, subpolished. (Notauli broad, distinct; side of

propodeum only weakly convex; hindfemur slender; interocellar line

shorter than ocellocular line; hindbasitarsus 9 12.25-15. 6, o 8, 8-9. 6

times as.long.as thick. sjsu vw daira 2. gracilis, new species (p. 14)

Head not so or head, mesoscutum, and mesopleurum either weakly

punctate and rather polished or very strongly punctate and opaque.

(Notauli either quite sharply impressed or indistinct, or if broad

but distinct, scutellum appears at least as wide as long; propodeum

usually more rounded; hindfemur usually medium to stout), ..... 6

Hindbasitarsus brown at base; prepectus and adjacent area of meso-

pleurum yellow; first tergite at least slightly wider than deep through-

out, (Small insects.)..... .. 15, fuscipes, new species (p. 45)

Hindbasitarsus not brown (though sometimes deeper yellow) at base;

prepectus not yellow; first tergite usually not wider than deep through-

uibs fal? gcse eed mermairnep PL Ge Tet ee ny rire Tne oe "2 aR:

Scutellum appears square or wider than long, little narrowed behind;

epomia short, enclosure small; propodeum with median longitudinal

carina distinct and other traces of areolation; mesoscutum index

0. 93-1.14 (rarely 1.22); thorax, hindfemur, and hindbasitarsus very

stout; pronotum quite narrow front to back; postpetiole rather long in

proportion to petiole. (Head subbuccate to buccate.)......... 8

Scutellum appears longer than wide or distinctly narrowed behind, or

epomia and propodeum not so; mesoscutum index usually over 1.15;

thorax, hindfemur, and hindbasitarsus variable; pronotum not markedly

narrow; postpetiole usually relatively Shorter i. os oe: eye dawn in 10

Scutellum distinctly elevated anteriorly, with rounded transverse angle

between steep anterior surface and sloping disc with distinct median

longitudinal depression; space enclosed by epomia about one-half as

wide as long; face much wrinkled and punctate, inner orbit moderately

punctate; upper third of clypeus with more than one-half as many punc-

tures per unit area as face except in a few of smallest specimens;

petiole distinctly elevated before spiracles.

17. crassulus, new species (p. 50)

Scutellum flat, sometimes very slightly depressed; space enclosed by

epomia less than one-half as wide as long; face more weakly rugulose,

inner orbit broadly nearly impunctate; clypeus not so; entire first tergite

distinctly flattened throughout, petiole rarely very faintly elevated

ADCO Ae iis ei iil iba ig volt at Nae og Nh 9 cal edger ein ih A PAG ak 9

10 Contrib. Amer. Ent. Inst., vol. 18, no, 2, 1981

9, Rather extensively black; notauli obsolete but prescutum distinctly raised

above parapsides; stemmaticum rather strongly elevated at lateral

ocelli in female (hardly in male); top of head rugulose-reticulate-

punctate, most strongly on vertex behind ocelli, grading from sub-

polished on frons to subopaque on vertex; head rounded-rectangular

and moderately transverse, lower part of face and clypeus moderately

short in profile; cheek narrow, especially below; mesoscutum and

mesopleurum moderately finely densely deeply punctate (more

coarsely and shallowly on pleurum), subopaque; ovipositor longer

than first tergite, distinctly decurved in apical two-fifths.

19. decurvihastatus, new species (p, 54)

Usually with little more black than typical of genus; notauli fairly distinct

though not sharp and prescutum little or not at all raised; stemmaticum

little elevated; top of head, mesoscutum, and mesopleurum polished,

punctation sparse and weak; head buccate, transverse, lower part of

face and clypeus quite short in profile; ovipositor variable in length,

not decurved, rather deep in proportion to length, nodus rather long

and large, andtip long... 18. flaviscutellaris, new species (p. 53)

10. Eye projecting little from head outline in plan view, head moderately to

strongly buccate; gena and vertex weakly punctate and polished; face

and clypeus in profile forming nearly straight line; curve of occiput

relatively flat (occipital camber 4. 6-6. 1); temple usually forming

rather sharp, sometimes very sharp angle with occiput; notauli

weakly indicated, represented chiefly by change of direction of surface

from parapsides to prescutum; scutellum sometimes slightly wider than

long but convergent behind, moderately elevated, lateral carina obso-

lete; propodeum usually rather polished with weak rugulosity; hindbasi-

tarsus 6,5-8.3 times as long as thick. (Known so far only from

California excepting one specimen from Texas. )

16. puparum (Ashmead) (p. 48)

Eye projecting from outline of head in plan, head not buccate, or notauli

distinct, or scutellum carinate and relatively narrow, or otherwise

GUO RONE eh Gee ee ce Pe re See ee eee 11

11. Scutellum rather gibbous, relatively wide but convergent behind, lateral

carina more or less reduced; propodeum rather distinctly bulging

laterally, usually with reduced rugulosity and only fine

POU CUEIO ee eee a ee ee 12

Scutellum usually flat or medially depressed (or if slightly elevated,

highly polished and lateral carina distinct) appearing distinctly or

moderately narrow, lateral carina usually distinct; propodeum with

side weakly convex, usually moderately extensively reticulate with

RADU SUC ORE Ee ee Gee be ee 8s 13

12. Hindbasitarsus stout (9 7-10, o 5, 6-8.3 times as long as thick); 32-41

flagellar segments, first segment 9? 3, 7-4.6, o3,8-5, 2 times as long

as wide; thorax somewhat coarsely punctate, often extensively black.

9, enargiae, new species (p. 29)

Hindbasitarsus slender (? 11.04-12.8, o°9.0-10.5 times as long as thick);

42-43 flagellar segments, first segment 9 4,8-6.0, %4,2-6.1 times

as long as wide; thorax otherwise. . 10. masoni, new species (p. 33)

13.

14,

15,

16.

Hopper: Nearctic Aphanistes 11

Head rather strongly narrowed behind, temple flattish; temple, frons,

vertex and stemmaticum with rather small, shallow punctures

separated by more than their diameter, polished, crista small, frons

and vertex rather distinctly convex, especially stemmaticum; meso-

scutum and mesopleurum polished, punctures shallow and separated by

about one-half their diameter; outer surface of hindtibia at most yellow-

ish, never yellow. (Small insects.). . 5. politus, new species (p. 21)

Head not narrowed behind, temple not flattish; top of head more strongly

punctate, subopaque, not or weakly convex, crista fairly prominent,

usually not small; hindtibia yellow on outer surface......... 14

Head and thorax wholly coarsely punctate, opaque; notauli indistinct;

scutellum moderately elevated, highest at one-fourth to one-third

its length, usually with strong (sometimes very strong) lateral

carina and always somewhat longitudinally concave; thorax wholly

black except tegula; first tergite strongly infuscate or piceous above,

postpetiole elliptical in section with axis vertical.

7. nigrorufus (Norton) (p. 25)

Head and thorax less strongly punctate, subopaque, subpolished, or

polished; notauli distinct; scutellum little or not at all elevated,

highest very close to cephalic margin; thorax rarely wholly black; first

tergite usually not infuscate or piceous, form of postpetiole vari-

OOO css ae ee ee Aa ar a dak WMI shia ian Sp ms a ie Nahe a eto die a),

Mesoscutum without black markings except usual patch before scutellum

and occasionally a minute medial line cephalad; side of propodeum very

WEOKLY CONVENE Ce nee ge ec eet rs ; See) aaa 16

Mesoscutum with at least a black medial band or distinct line or anterior

spot, usually more extensively black; side of propodeum distinctly

convex though not more than slightly bulging. ........... 17

Head in plan view more or less elliptical and when tilted to bring anterior

edge of median ocellus to forward edge of head, edge of head appears

nearly straight across frons from forward edge of one eye to the

other; posterior margin of eye not transverse as in next alternative;

notauli rather sharply impressed. (Head not thick in profile; epomia

forming smooth curve, not parallel to collar; petiole rather slender,

not raised before spiracles.)....... 12. basilicon (Davis) (p. 38)

Head more or less buccate (except some small males), frons not straight

across when viewed as above; posterior margin of eye more or less

transverse when occiput in line of sight; notauli indistinct or obsolete,

(Head in profile rather thick, often more or less nearly circular;

epomia with abrupt curve below, remainder nearly parallel to collar;

petiole usually distinctly raised before Spiracles, not particularly

slender, ) 11. walleyi, new species (p. 36)

Head and thorax highly polished; frons, stemmaticum, and vertex rather

weakly punctate, with a few weak rugulae around stemmaticum, never

reticulate, all three weakly distinctly convex except for smallish

antennal scrobes; notauli indistinct, shallow, marked by an irregular

line of punctures; scutellum polished, not elevated, slightly longitu-

dinally depressed, with usually relatively sparse but rather large

punctures that sometimes partially run together to form longitudinal

12 Contrib. Amer, Ent. Inst., vol. 18, no. 2, 1981

aciculations, lateral carina almost always very strong, disc sometimes

wholly black; petiole elevated before spiracles in most specimens; hind-

femur stout; nodus of ovipositor usually decidedly abrupt distad.

14, heinrichi, new species (p. 43)

Head subpolished to opaque, frons often rugulosoreticulate and fairly

strongly rugulose around stemmaticum, slightly concave or only very

slightly convex before median ocellus; notauli distinct, usually sharply

impressed; scutellum subopaque, slightly elevated, usually not longitu-

dinally depressed, with relatively smaller, considerably denser punc-

tures, never aciculate, lateral carina moderate or occasionally weak,

rarely wholly black or piceous; petiole not elevated; hindfemur variable;

OVEPOSICOF MOL SO. bei wo bie ib Cane Laid ten ieee? Lie ceoits 18

18, Head usually subopaque with rather dense moderately deep punctures,

often rugulosopunctate on frons, which is slightly concave behind

antennae (in addition to the antennal scrobes present in all species);

temple progressively less punctate and more polished than vertex;

epomia forming smooth curve and enclosing rather narrow space with

maximum width usually less than one-half length, without noticeable

rugulae; notauli usually not sharply impressed, but distinct; maximum

width of prescutum greater than that of one of parapsides, prescutum

distinctly raised above parapsides; scutellum slightly wider (length

0.95-1.15 times width), usually with black mark at base; propodeal

fossa rather distinctly impressed, pleural carina usually stronger than

others; outer carina of area dentipara moderate; metapleurum slightly

bulging; petiolar area rather distinctly longitudinally depressed; hind-

basitarsus 9 7. 4-10. 8 times as long as thick; first tergite usually

partly or wholly piceous or black, in o’0. 9-1. 1328 times as long as

DA DAB IAP SUB soa. ues we eaves 8. townesi, new species (p. 27)

Head more subpolished, frons slightly convex with little rugulosity; temple

abruptly less punctate and more polished than vertex; lower end of

€pomia approximately at right angle to remainder, with short ramus

at angle, enclosure moderately large, usually one-half or more as

wide as long, with ladderlike crosswise rugulae; notauli usually

sharply impressed; maximum width of prescutum usually approximately

equal to that of one of the parapsides, prescutum not noticeably raised

above parapsides; scutellum usually narrower (9 1.1-1.4, o0.93-1.14

times as long as wide--but appears narrow in both sexes), usually with-

out black mark at base; propodeal fossa slightly to distinctly impressed,

pleural carina indistinct or obsolete; outer carina of area dentipara

usually quite strong; metapleurum only moderately convex, not bulging;

petiolar area nearly flat; hindbasitarsus 9 10. 8-14 times as long as

thick; first tergite very rarely black or piceous, in o' 1. 326-1. 65 times

as long as hindbasitarsus, .... 138. dreisbachi, new species (p. 40)

The Crepuscularis Group

The group is characterized by the very large ocelli, short temples conver-

gent caudad, fulvous color with reduced black markings, extensive pale-yellow

markings on thorax in male, and slender form. It contains only the species

crepuscularis,

Hopper: Nearctic Aphanistes 13

1. Aphanistes crepuscularis, new species

Female: Forewing 10.3-14.5 mm, body [see ‘Measurements‘ above]

16, 5-24 mm; antenna 11-16 mm, 43- 48 segments, first flagellar segment 4, 9-

Do. 1 times as long as wide, combined length of second and third segments

0.87-1.0 times first; head thickness-width ratio 1, 80-1.87, head length-

width ratio 1,2-1,.26, head thickness-camber ratio 2, 65-3,6, occipital camber

4,3-4,5, head-occiput width ratio 1.49-1.55, eye-temple ratio 3.0-3.9; meso-

scutum index 1,.25--1.34, head width 1, 64- 1. 66 times mesoscutum width, scu-

tellum 1:1.05--1, 34:0. A7--0, 77; depth: width:length of propodeum 1:0. 98--

1,06:1. 06--1.17; hindbasitarsus 9,0--12,3 times as long as thick, first tergite

1,1-1.5 times as long as hindbasitarsus; width of postpetiole 0. 50--0, o7 times

length, first tergite 3, 72--8. 84 times as long as a

Habitus rather slender. Eye and ocelli very large, ocellocular line not

more than 0.5 times diameter of lateral ocellus; face rather flat; head rather

strongly convergent behind, temple nearly flat, cheek narrow, head rather

thin. Notauli moderately strongly, rather sharply impressed: scutellum very

slightly convex, Sometimes with a slight longitudinal depression anteriorly;

propodeum with anterior portions of pleural, lateral longitudinal, and median

longitudinal carinae present, not very closely rugose, rugulose between rugae;

areola distinct but open behind, second lateral area recognizable but indistinctly

bounded; mesoscutum and mesopleurum subpolished, rather densely punctate,

punctures separated by approximately one-half their diameter; epomia indistinct

at lower (outer) end, often wholly indistinct, inner end weakly curved forward.

Fulvoferruginous, Antenna fulvous; vertex, temple, and mesoscutum

ferruginous; face, clypeus except tooth, mandible except tip, cheek, fore- and

midlegs (sometimes stramineous), lower surface of hind distal trochanter, and

proximal two-thirds of hindtibia yellow; tip of mandible, distal one-third approxi-

mately of hindtibia, dorsum of second tergite except lateral margins, sometimes

stemmaticum at posterior edge of median ocellus, and sometimes spot on meso-

scutum before scutellum and spot on base of propodeum, black; hindcoxa dor-

sally or wholly, proximal hindtrochanter wholly, and distal hindtrochanter dor-

sally black or piceous, Side of scutellum wholly fulvous except anterior black

spot in most Canadian and Transition Zone specimens and the male from Great

Smoky Mt, Nat. Pk. (black in most specimens of most other species). Wings

subhyaline.

Male (only two specimens seen): Forewing 10. 6--10.7 mm, body 19. 8--

20.5 mm; antenna 14,2 mm (tip lacking in one specimen), 48 segments, first

flagellar segment 4, 56--4, 84 times as long as wide, combined length of second

and third segments 0, 98--1.0 times first; head thickness-width ratio 1. 69--

1.75, head length-width ratio 1,16--1.18, head thickness-camber ratio 3, 4--

4,2, occipital camber 4,0--5.0, head-occiput width ratio 1. 4--1.45, eye-

temple ratio 2, 85--2,9; mesoscutum index 1.12--1.27, head width 1.575--

1.6 times mesoscutum width, scutellum 1:1. 13--1.3:0. 63--0. 73; depth:width:

length of propodeum 1:1,05--1,0625:1, 125--1, 1875; hindbasitarsus 9, 02--

10, 26 times as long as thick, first tergite 1.257--1,.259 times as long as hind-

basitarsus; width of postpetiole 0.5--0. 6 times length, first tergite 3. 72--3. 88

times as long as postpetiole.

Form and sculpture as in female except usual difference in tarsi.

Color as in female except marks on pronotum, margins of mesoscutum,

scutellum more or less, lower portion of mesopleurum, especially cephalad

(extensively), and metapleurum more or less, very pale yellow (nearly white),

Material examined: 2899, 200%

14 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

Type: 9, Wake Co., NC, VII-1 1951, H&M Townes (CHT); allotype: g,

Cleveland, SC, VI-5 1961, G. F. Townes (CHT); paratypes: 99, Tuscaloosa,

Ala., V-13 1948, R. L. Chernock (CNC); Atlanta, Ga., 5-4-36, P. W. Fattig

(CHT); Atlanta, Ga., 5-13-36 (USNM); Atlanta, Ga., 5-11-37, P. W. Fattig

(CHT); Atlanta, Ga., 5-18-39, P. W. Fattig (USNM); Atlanta, Ga., 5-24-39,

P. W. Fattig (USNM); Atlanta, Ga., 5-30-40, P. W. Fattig (USNM); Atlanta,

Ga., 5-16-41, P. W. Fattig (USNM); Atlanta, Ga., 5-21-41, P. W. Fattig

(USNM); Atlanta, Ga., 5-22-41, P. W. Fattig (29, USNM); Lakemont, Ga.,

6-5-40, P. W. Fattig (29, USNM); Neel Gap, Ga., 6-4-51, P. W. Fattig (29,

CHT); Ranger, Ga., 5-25-50, P. W. Fattig (CHT); ILL.: DuPage Co.,

Argonne Natl. Lab., at light, 1-VII-1967, leg. J. Wagner & W. Suter, 1967

AMU-ANL Ecological Survey (USNM); Principia College, Elsah, Jersey Co.,

Ill., V-26-41, C. L. Remington (USNM); PlummersI., Md., 6-27-38, A. B.

Gurney (USNM); Oxford, Miss., 5-15-1949, H. V. Weems, Jr. (USNM); Dean

For., O., 5--’41, C. R. Neiswander, Collr. (CHT); Highlands, N. C.,

31-V-1957, W. R. M. Mason (CNC); Highlands, N. C., 19-VI-1957, W. R. M.

Mason (CNC); Sylva, N. C., VI-7-47, H. Townes (CHT); Springfield, Del. Co.,

Pa., 6-VI-1934, H. P. Hopper, woodland (HPH); Cleveland, S. C., VI-1-1961,

G. F. Townes (CHT); paratype: of, Alt. 2500’, Gr. Smoky Mt. Nat. Pk., Tenn.,

V-8-1955, H&A Howden (CHT).

The Hyalinis Group

In this group, the diameter of the lateral ocelli is equal to or less than

the ocellocular line, the mesoscutum and mesopleurum have moderately to

rather deep punctures of medium size separated by more than one-half their

diameter but appearing moderately dense, epomia with a smooth curve that

becomes almost perpendicular to the collar below, enclosing a space of medium

size, notauli broad and shallow but distinct, especially broad where they meet

margin of mesoscutum, petiolar area with transverse rugulae predominating

(there are usually only transverse rugulae in hyalinis, sometimes also in

gracilis, sometimes last few in edemae). The species included are gracilis

hyalinis, edemae, and politus.

2, Aphanistes gracilis, new species

Female: Forewing 6. 4--11.4 mm, body 10. 8--19.5 mm; antenna 7, 5--

15mm, 37--47 segments, first flagellar segment 5, 81--6, 5 times as long as

wide, combined length of second and third segments 0, 9--1.01 times first;

head thickness-width ratio 1, 63--1.66, head length-width ratio 1,176--1. 213,

head thickness-camber ratio 3, 6--3, 7, occipital camber 3,9--4, 4, head-

occiput width ratio 1.397--1,52, eye- temple ratio 2,1--2,5; mesoscutum

index 1. 2--1.35, head width 1, 6375--1, 64 times mesoscutum width, scutellum

1:1.08--1.2:0. 59--0, 6; depth:width:length of propodeum 1:1. 025--1. 08:1. 175--

1.2; hindbasitarsus 12. 25--15, 6 times as long as thick, first tergite 1,07--

1; 12 times as long as hindbasitarsus; width of postpetiole 0, 48--0, 565 times

length, first tergite 3, 6--3. 74 times as long as postpetiole.

Habitus moderately slender. Head above and thorax including scutellum

rather closely, moderately deeply punctate, subopaque; head moderately

strongly convergent behind, relatively thin from front to back, temple rather

flat, cheek moderately narrow, median field of face with longitudinal wrinkles.

Hopper: Nearctic Aphanistes 15

Notauli broad and shallow but distinct, especially broad where they meet

anterior margin of mesoscutum; scutellum slightly elevated at anterior margin,

elevation rounded, one-fourth length of scutellum, but disc rather flat to very

slightly depressed otherwise, Sides of propodeum very weakly convex; areola-

tion mostly indistinct because of extensive rugulosoreticulation, lateral and

median longitudinal carinae present at area dentipara and areola, latter usually

open behind, Postpetiole little wider than petiole, its depth at spiracles equal

to its maximum width. Hindfemur slender.

Antenna fulvous; face and clypeus except apical tooth, mandible except tip,

scape, orbital spot, all tarsi, fore- and midtibiae entirely, base of hindtibia,

forecoxa, mid- and hindcoxae below, fore- and midtrochanters, propleurum,

collar, usually mesopleurum in part (especially around prepectus), often

scutellum in part, and tinge on metapleurum, yellow; black are: tip of man-

dible, temple, frons and vertex largely, occiput, pedicel, and base of first

flagellar segment above, usually broad longitudinal stripe along midline of

mesoscutum, often parapsides (but not anterior ends of notauli), patch before

and side of scutellum, usually marks on mesopleurum and mesosternum (these

sometimes almost entirely black), usually spot at base of propodeum (some-

times entire dorsum), usually metakatepisternum, usually upper surface of

hindcoxa, and always about apical third of hindtibia; sometimes spot at base of

disc of scutellum, first tergite extensively, and hindtrochanters above piceous

(last-named often black); tooth of clypeus piceous.

Male: Forewing 7.55--11.12 mm, body 14. 3--22 mm; antenna 12, 9--

14.9mm, 40--45 segments (the smallest specimens lack complete antennae),

first flagellar segment 4, 6--6.0 times as long as wide, combined length of

second and third segments 0, 98--1.19 times first; head thickness-width ratio

1, 62--1. 73, head length-width ratio 1.14--1.17, head thickness-camber ratio

3,8--5,.25, occipital camber 4, 8--6,375, head-occiput width ratio 1, 35--1. 46,

eye-temple ratio 2. 44-2, 95; mesoscutum index 1, 26--1.325, head width 1, 5--

1.53 times mesoscutum width, scutellum 1:1.18--1.3:0.526--0, 69; depth:

width:length of propodeum 1:1, 0--1.1:1,08--1. 25; hindbasitarsus 8, 8--9. 6

times as long as thick, first tergite 1,14--1.21 times as long as hindbasitarsus;

width of postpetiole 0, 46--0.55 times length, first tergite 3, 4--3,8 times as

long as postpetiole.

Sculpture essentially as in female. Color similar to that of female, but

with mesopleurum more extensively yellow; tergites 6--8 and clasper often black

or piceous (even in one specimen with otherwise greatly reduced black mark-

ings).

Material examined: 6799, 3006%

Type: 9, Takoma Pk., Md., VII-1, 1943, H. & M. Townes (CHT);

allotype: o, Takoma Pk., VII-2, 1944, H. & M. Townes (CHT); paratypes:

09, Hartford, Conn., 29-VI-1966, G. H. Heinrich (HPH); Voluntown, Conn.,

Aug. 19, 1951, H. & M. Townes (CHT); Washington, D. C., 12-IK-1942,

H. P. Hopper (29, HPH); Wash., D. C., 7-11-43, M. Ng [remainder illegible]

(USNM); Washington, D. C., VI-19-44 (UCD); Washington, D. C., 8-IX-1957,

H. P. Hopper (HPH); Iowa City, Iowa, IX-4-44, RTMitchell, Collector (29,

RTM); Patuxent Ref., Bowie, Md., 11-IX-1946, RTMitchell, Collector (RTM);

Plummers Isl., Md., 27-IX-1949, G. E. Shewell (CNC); Takoma Pk., Md.,

VII-30, 1942, H. & M. Townes (CHT); Takoma Pk., Md., VI-20, 1943, H. &

M. Townes (39, CHT); Takoma Pk., Md., VI-27, 1943, H. & M. Townes

(CHT); Takoma Pk., Md., VII-1, 1943, H. & M. Townes (39 in addition to

type, CHT); Takoma Pk., Md., I[X-11, 1943, H. & M. Townes (39, CHT);

Takoma Pk., Md., VII-15-1945, H. & M. Townes (CHT); Takoma Pk., Md.,

16 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

VIII-27-1945, HK Townes (CHT); Takoma Pk., Md., VI-5, 1947, H. & M.

Townes (29, CHT); Colrain, Mass., VIII-17-35, H. D. Pratt (USNM);

Fitchburg, Mass., VII-5-37, M. Chapman (CHT); Winchendon, Mass., July 2,

1892 (USNM); Alger Co., Mich., VIII-7-59, R. & K. Dreisbach (Dreisbach);

Pelham, N. H., IX-5-1905, Bridwell, Collector (USNM); Pinkham Notch,

N. H., Aug. 24°51, HM&D Townes (CHT); Moorestown, N. J., 13-VIII-1939,

H. P. Hopper (HPH); Greene Co., N. Y., Aug. (Pittsburgh); Hancock, N. Y.,

Aug. 10 1935, H. K. Townes (29, CHT); Hancock, N. Y., Aug. 12 1935, H.

K. Townes (CHT); Ithaca, N. Y., 30 Aug. 35 (CHT); Oliverea, N. Y., Catskills,

IX, 3--8 ’18 (USNM); Saranac Lake, N. Y., 30-VI ’46, RTMitchell, Collector

(RTM); Saranac Lake, N. Y., 9-VII ’46 RTMitchell, Collector (RTM); Crab-

tree MdsNC, Yancey Co. 3600’, Aug 21, 1950, HMD&J Townes (CHT); Hamrick

NC, Aug 19 °50, HM&D Townes (CHT); Hamrick NC, Aug 28 ’50, HM&D Townes

(CHT); Philadelphia, Pa., VI-24, 1941, H. & M. Townes (CHT); Wissahickon

Val., Philadelphia, Pa., 12 Oct., 1938, H. P. Hopper (HPH); Spring Brook,

Pa., Aug. 13 °44, H. K. Townes (CHT); Spring Br Pa, VIII-27-1945, HK

Townes (CHT); Ex Hadena turbulenta, Westerly, R. I., VIII-19-27, GipMothLab

12435 M. 1 (USNM); Westerly, R. I., Aug. 20, 1951, H. & M. Townes (CHT);

Bolivar, W. Va., Sept. 20, 1942, H. K. Townes (29, CHT); Bolivar, W.Va.,

Sept. 22, 1942, H. K. Townes (29, CHT); Bolivar, W. Va., Sept. 23, 1942,

H. K. Townes (59, CHT); Bolivar, W.Va., Sept. 24, 1942, H. K. Townes

(29, CHT); Bolivar, W.Va., Sept. 25, 1942, H. K. Townes (29, CHT); Bolivar,

W.Va., Sept. 26, 1942, H. K. Townes (29, CHT); Loc. Restigouche Co.,

N. B., Host. Semiothisa granitata, Rec. (K) F. I. Survey 1949, Em Mar 13,

1950 «‘ Aphanistes sp. Det. R. S. Forbes,’’ 61P3, F. B. Lab, F.Con.N.B.,

F.1.S. (CNC); paratypes: oo, lowa City, Iowa, IX-4-44, RTMitchell, Collector

(2c, RTM); Patuxent Ref., Bowie, Md., 9-VIII-1948, RTMitchell, Collector

(RTM); Patuxent Ref., Bowie, Md., 13-IX-1948, RTMitchell, Collector (RTM);

Takoma Pk., Md., VI-17, 1942, H. & M. Townes (CHT); Takoma Pk., Md.,

VI-20, 1943, H. & M. Townes (CHT); Takoma Pk., Md., VII-18, 1943, H. &

M. Townes (CHT); Takoma Pk., Md., VII-25, 1943, H. & M. Townes (CHT);

Takoma Pk., Md., IX-7, 1943, H. & M. Townes (CHT); Takoma Pk., Md.,

IX-9, 1943, H. & M. Townes (CHT); Takoma Pk., Md., VII-5, 1947, H. & M.

Townes (CHT); Alger Co. Mich., 7-31-46, R. R. Dreisbach (Dreisbach);

Aurelius, Mich,, 20 Sept 1938 (MSEL); Huron Mts., Mich., July 30, 1959,

Henry Townes (CHT); Pinkham Notch, NH, Aug. 25 751, HM&D Townes (CHT);

Greene Co., N. Y., Aug. (Pittsburgh); Flat Rock, N. C., VIIM-28-39, H&M

Townes (CHT); Ex Hadena turbulenta, Westerly, R. I., VIM-22-27, GipMothLab

12435 M 1 (USNM); Westerly, R. I., Aug. 13, 1951, H.&M. Townes (CHT);

Westerly, R. I., Aug. 20, 1951, H.&M. Townes (CHT); Arlington, Virginia,

14 October 1950, J. G. Franclemont (CHT); Bolivar, W.Va., Sept. 22, 1942,

H.K. Townes (2c, CHT); Bolivar, W.Va., Sept. 23, 1942, H.K. Townes (4¢,

CHT); Knowlton, Que., 20-VII-1936, G. S. Walley (CNC); Knowlton, Que.,

25-VII-1936, G. S. Walley (CNC).

Hosts: Noctuidae: Hadena turbulenta; Geometridae: Semiothisa granitata,

3. Aphanistes hyalinis (Norton)

Anomalon hyaline Norton, 1863, Ent. Soc, Phila., Proc. 1: 361.

Type: 9, Farmington, Conn, (in Peabody Museum, Yale University,

New Haven, Connecticut).

Hopper: Nearctic Aphanistes 17

Female: Forewing 6,9--12.2 mm, body 11. 85--21.1 mm; antenna 11, 5--

14mm, 43--47 segments (antennae lacking in smallest specimens); first

flagellar segment 4, 2--5,5 times as long as wide, combined length of second

and third segments 0. 89 (large specimens)--1.07 (smallest specimen) times

first; head thickness-width ratio 1.575--1.8, head length-width ratio 1.1--1. 2,

head thickness-camber ratio 3.5--4,2, occipital camber 4,1--5.2, head-occiput

width ratio 1,325--1.5, eye-temple ratio 1. 6--2,0; mesoscutum index 1,15--

1,275, head width 1, 425--1,58 times mesoscutum width, scutellum 1:0, 8--1.15:

0.55--0. 7; depth:width:length of propodeum 1:1,05--1, 2:1. 15--1, 45; hindbasi-

tarsus 9,0--11.0 times as long as thick, first tergite 1,05--1, 25 times as long

as hindbasitarsus; width of postpetiole 0, 4--0. 75 times length, first tergite

3,4--4, 7 times as long as postpetiole.

Habitus moderately slender (usually slightly less so than in gracilis). Face

rather finely densely punctate (punctures separated by one-half to one time

diameter), very shallowly laterally, more deeply (moderately shallowly) toward

midline, usually becoming vertically rugulose or wrinkled (about four wrinkles)

in mesal one-fourth (if wrinkles are weak or absent, there is a low rounded

vertical (longitudinal) prominence extending from subantennal carina to base of

clypeus); gena with rather close, shallow punctures below (one-half or less

their diameter apart), becoming sparser toward middle (separated by one to

two times their diameter), and becoming denser again on the rather flat

temple and much deeper without a change of size on stemmaticum; vertex

behind ocelli closely deeply punctate with a slight tendency for punctures to run

together to form rugulae between; cheek in profile at least twice as wide as

temple, lower posterior corner usually distinctly below inner/upper angle of

mandible when long axis of head vertical; frons distinctly rugulose except more

or less laterally (sometimes only orbits, sometimes wider area); ocellocular

Space with only a few shallow punctures, interocellar line 1.125 times ocellocu-

lar line, head moderately convergent behind. Pronotum rather densely punctate,

with punctures running together in the trough to form rugulae between; meso-

pleurum and mesoscutum rather closely moderately deeply punctate, subopaque;

notauli distinct, rather broad and shallow, with an irregular line of punctures

rather than a groove, widening into a broad cove without a distinct line at about

one-fourth length of mesoscutum from anterior margin; epomia distinct, curv-

ing toward collar rather sharply at the outer (lower) end and curving slightly

in the opposite direction at upper (inner) end (not directed toward anterior

margin of pronotum), length from outer end to midline 0, 7--0.8 times distance

from outer end to margin of pronotum, enclosed space with indistinct sculpture,

width about 0. 46 times width of adjacent collar; mesoscutum and mesopleurum

rather densely punctate, punctures rather small and moderately deep, separated

by less than one-half their diameter; black area of mesoscutum in front of

scutellum less distinctly transversely rugulose than usual in the genus, in

some specimens rather rugulosopunctate; scutellum more or less elevated, usu-

ally with distinct lateral carina, disc less closely punctate than mesoscutum,

punctures slightly larger, center of disc often subpolished to polished, usually

with more or less distinct median longitudinal indentation and rather rounded

at apex; speculum represented only by a slight elevation with only a narrow

anterior edge polished; sternaulus distinct, short, about two-fifths length of

mesopleurum, but usually not sharply impressed. Propodeum subpolished with

only very minute indistinct punctures except in areola and area dentipara (lat-

ter usually not separated from first lateral area), where they are shallow and

very closely placed and tend to appear as fine reticulations; areola between

square and pentagonal, sometimes separated from the petiolar area only by a

18 Cont. Amer, Ent. Inst., vol. 18, no. 2, 1981

difference in sculpture, sometimes more or less distinctly closed behind;

area dentipara (including first lateral area) occupying two-fifths length of

propodeum, when partially separated from first lateral area, area dentipara

tetragonal, almost an irregular triangle, adjacent portion of paracostula more

or less straight and at about 45° to midline, lateral longitudinal carina particu-

larly strong at this point; sides of propodeum rather bulging, pleural carina

usually indistinct or obsolete, weaker than any longitudinal ruga on meta-

pleurum and near bottom of propodeal fossa; petiolar area and third lateral

area usually with (rather straight) transverse rugae only. Hindfemur slightly

stout. Petiole and postpetiole not or little deeper than wide, never widened,

postpetiole subelliptical to subcompressed.

Antenna dark fulvous to fuscous; head yellow oe clypeal tooth fulvous

and tip of mandible, frons, stemmaticum, vertex (with usual yellow orbital

spots), and temple (ferruginous behind orbital spot in type), and upper part of

occiput, which are black. Thorax, propodeum, and abdomen luteous or dull

fulvous (thorax and propodeum often with yellow-tinged areas; in one specimen

thorax and propodeum wholly yellowish except narrow piceous to black median

line on mesoscutum) with black longitudinal stripe on mesoscutum, sometimes

confluent with the black patch before scutellum, and with more or less piceous

on parapsides, extending forward from the black patch (rarely mesoscutum

almost entirely black except anterior ends of notauli); spot below forewing,

mesosternum (extending above sternaulus), areola, and area dentipara nor-

mally black, metanepimerum usually more or less piceous or black, meta-

sternum rarely black (only one such specimen seen, a few have limited piceous

area directly above coxa); side of scutellum (black in most other species)

usually yellow or with only a black spot, at least upper anterior corner yellow;

rarely apex of disc of scutellum black, still more rarely also blotch at base

and median longitudinal streak piceous to black. Wings hyaline to subhyaline.

Fore- and midlegs yellow except sometimes tips of tarsi and often femora and

coxae more or less fulvous, midcoxa never infuscate below; hindtarsus and

approximately proximal two-fifths of hindtibia yellow, remainder of tibia black,

hindfemur fulvous or sometimes ferruginous or rarely brown above, hindtro-

chanters and hindcoxa largely black. Postpetiole or sometimes entire first

tergite usually piceous to black, commonly lighter anteriorly (in small speci-

mens only infuscate or entire segment fulvous); venter of first two to four

abdominal segments usually black, also portions of the last two or three ter-

gites sometimes piceous to black; dorsum of second tergite largely black with

luteous lateral margins.

Male: Forewing 7.6--11.4 mm, body 13.3--21.7 mm; antenna 8. 7--14.1

mm, 35--45 segments, first flagellar segment 4, 9--5.5 times as long as wide,

combined length of second and third segments 0, 93--1.15 times first; head

thickness-width ratio 1.6--1.7, head length-width ratio 1.14--1.2, head

thickness-camber ratio 3, 65--3, 75, occipital camber 4, 3--4, 45, head-

occiput width ratio 1,35--1.4, eye-temple ratio 1, 67--1, 78; mesoscutum index

1.16--1.275, head width 1. 40--1, 43 times mesoscutum width, scutellum

1:0. 9--0, 975: 0. 6--0. 7; depth:width:length of propodeum 1:1. Disco. 175:

0.975--1.16; hindbasitarsus 8,0--11.0 times as long as thick, first tergite

0.97--1.15 times as long as hindbasitarsus; width of postpetiole 0, 45--0. 625

times length, first tergite 3,3--3,9 times as long as postpetiole.

Habitus and sculpture generally as in female, with less tendency (in the

six specimens examined) to vertical wrinkling in the middle of the face,

rather deeper punctures on temple and vertex, and stronger sr Me ar on

frons. Space enclosed by epomia with weak rugulosities; punctures on meso-

Hopper: Nearctic Aphanistes i

pleurum with rather linear, more or less longitudinal separations (more

elevated than separations transverse to the former); rugulosity of space before

scutellum more as in other species of the genus; scutellum less polished, but

otherwise as in female. Propodeum almost wholly devoid of punctures in the

median, lateral, and pleural regions, including the area dentipara and areola--

these last instead with some rather distinct rugulae forming some reticulations.

Color as in female except prepectus and portion of mesopleurum immedi-

ately behind yellow, and one specimen has entire lower portion of mesopleurum

and upper posterior margin of pronotum yellow, remainder of mesopleurum

except subalar black spot and entire mesoscutum except black patch before

scutellum ferruginous and hindcoxa without black or ferruginous,

Material examined: 2699, 606%

Distribution: Alaska, Connecticut, Illinois, Maryland, Michigan, New

York, Pennsylvania, Virginia, ‘‘Collection Belfrage’’ [| Texas?|, New Bruns-

wick, Nova Scotia, Ontario, Quebec.

Hosts: Noctuidae: Baileya ophthalmica, Catocala sp.; Notodontidae: Sym-

merista albifrons, Symmerista canicosta.

4, Aphanistes edemae (Davis)

Anomalon edemae Davis, 1898 (1897). Amer. Ent. Soc., Trans. 24: 360.

o, 9[o'not examined]. Lectotype: @ (designated by Cresson, 1928),

Michigan, reared from Edema albifrons (in the Academy of Natural

Sciences of Philadelphia, Philadelphia, Pa.).

Female: Forewing 11--13.3 mm, body 18, 7--23, 7 mm; antenna 12--15,2

mm (only three specimens with complete antennae), 43--48 segments, first

flagellar segment 4.0--5.0 times as long as wide, combined length of second

and third segments 0, 85--1.0 times first; head thickness-width ratio 1, 625--

1.675, head length-width ratio 1.13--1,.275, head thickness-camber ratio 3, 4--

4,6, occipital camber 3,8--5,5, head-occiput width ratio 1,29--1.4, eye-

temple ratio 1.5--1.9; mesoscutum index 1,186--1.275, head width 1,27--1. 4

times mesoscutum width, scutellum 1:0, 8--0,. 9:0. 5--0. 6; depth:width:length of

propodeum 1:1.037--1.175:1, 06--1.125; hindbasitarsus 8,15--10. 86 times as

long as thick, first tergite 1,02--1.2 times as long as hindbasitarsus; width of

postpetiole 0. 4--0.6 times length, first tergite 2.575--3, 477 times as long as

postpetiole.

Habitus slightly stout, especially thorax. Clypeus and face in profile form

rather straight, nearly unbroken line with inconspicuous dividing furrow; face

with rather dense, somewhat shallow punctures, usually forming a few low

longitudinal wrinkles on the median field just below subantennal carina; frons

rugulose, stemmaticum rugulosopunctate, vertex rather densely deeply punc-

tate, ocellocular space with very few small shallow punctures, interocellar line

1.14 times ocellocular line; gena with rather small dense punctures, rather

shallow below, but becoming rather deep on temple, temple rather bulging (ex-

cept in type, in which temple only moderately convex). Pronotum, mesoscutum,

and mesopleurum moderately finely closely deeply punctate, punctures separated

by one-half their diameter or less, those on pronotum and mesopleurum often

partially running together to become rugulosopunctate, but without really dis-

tinct rugulae; epomia enclosing moderately large space, length from mesal line

0.65 times distance from outer end to lateral margin of pronotum; notauli

rather broad and deep, without a furrow (which is replaced by very closely

20 Cont, Amer, Ent, Inst,, vol, 16, no. 2, 1981

Spaced punctures not in a smooth line); sternaulus broad and shallow without a

distinct furrow; speculum obscure, mesopleural impression indistinct; scutellum

somewhat elevated, usually with a weak median longitudinal impression, lateral

carina from moderately distinct to obsolete, punctation rather uniform and

pretty much like mesoscutum; transverse rugulae in front of scutellum very fine

but distinct. Propodeum subopaque with very fine shallow indistinct punctures

throughout--slightly deeper and more distinct in areola (which is not closed

behind) and area dentipara and usually more or less distinctly separated first

lateral area, area dentipara frequently containing some supplementary rugulae;

areola more or less hexagonal; third lateral area with only transverse rugulae,

separated from petiolar area, which has chiefly only transverse ridges caudad,

but some irregular rugulae with other orientations anteriorly; area dentipara

extending from two-fifths to slightly more than half length of propodeum ;

pleural carina distinct, usually stronger than any ruga on metapleurum,

situated ventrolaterad from bottom of propodeal fossa. Postpetiole deepened,

subcompressed, entire first tergite usually distinctly deeper than wide except

in type and one other specimen, in which the portion of postpetiole adjacent to

spiracles and the apical half of petiole are abnormally enlarged, both deepened

and widened, upper surface of postpetiole strongly arched in all specimens,

Hindfemur stout,

Antenna fuscous to piceous; head yellow except tip of mandible, temple,

frons and vertex (except orbital spots--sometimes very small), upper side of

scape and pedicel, and upper half of occiput black. Mesosternum and medial

portion of mesoprescutum and parapsides black, latter interconnected by usual

black area before scutellum, black on mesoscutum often more extensive, only

lateral margins (extending slightly into anterior ends of notauli) always dull

fulvous or luteous; mesopleurum from wholly dull fulvous to almost wholly

black with only sutures and mesopleural impression dull fulvous; scutellum

with disc from wholly black to dull fulvous with only small basal and apical

black marks, side of scutellum usually black, but a few specimens have side

fulvous with only small black spot in upper anterior corner; collar dull fulvous

as is also mesal part of pronotum and often entire pronotum, or latter may be

black. Metanotum dull fulvous except usually postscutellum and metanepi-

merum black. Propodeum with first lateral area, area dentipara, basal area,

areola, and most of petiolar area black; metapleurum (except sometimes part

of metakatepisternum next to coxa) fulvous, Fore- and midlegs yellow except

femora more or less fulvous, forecoxa with black spot and midcoxa largely

black (usually infuscated to black beneath); hindcoxa usually wholly black,

hindtrochanters usually almost wholly black; hindfemur dark brown or piceous

at least above; hindtibia yellow at base, black at apex; hindtarsus yellow.

Wings usually moderately infuscate, occasionally subhyaline. First tergite

wholly black or piceous, as is second including lateral margins, remainder of

abdomen wholly piceous except hypopygium and ovipositor sheath fulvous

(except that in type tergites 3--5 dull fulvous, 6--7 black above).

Male: Forewing 10. 8--12.4 mm, body 19. 8--24.3 mm; antenna 13.1 mm

(only one specimen--not largest or smallest--with complete antennae), 42 seg-

ments, first flagellar segment 4,1--4.8 times as long as wide, combined

length of second and third segments 0. 95--1.1 times first; head thickness-

width ratio 1, 64--1. 75, head length-width ratio 1.15--1. 25, head thickness-

camber ratio 3,0--3, 45, occipital camber 3, T--4, 6, head- -occiput width ratio

1.275--1. 425, eye- temple ratio 1. 45--1. 9; mesoscutum index 1,14--1.25, head

width 1. 24--1. 38 times mesoscutum width, scutellum 1:0, 8--1, 25:0. 45--0, 8;

depth:width:length of propodeum 1:1.0--1. 325: 1.0--1. 325; hindbasitarsus 17, qi.

Hopper: Nearctic Aphanistes 21

8.9 times as long as thick, first tergite 1,06--1, 22 times as long as hindbasi-

tarsus; width of postpetiole 0, 45--0.6 times length, first tergite 2, 7--3,5 times

as long as postpetiole.

Habitus and sculpture as in female. Color as in female with slightly higher

incidence of fulvous on thorax, often including wholly fulvous scutellum without

darker marks; one specimen also has fore- and midcoxae wholly yellow.

Material examined: 1099, 1306

Distribution: Maine, Massachusetts, Michigan, Missouri, New Jersey,

New York, North Carolina, Ontario, Pennsylvania, Quebec, South Carolina,

Virginia,

Hosts: Notodontidae: Datana contracta, Datana integerrima, Datana

ministra, [ ?]Edema albifrons [ =Symmerista a. ?].

0. Aphanistes politus, new species

Female: Forewing 5. 6--7.15 mm, body 9, 4--11. 7 mm; antenna 5, 5--8,0

mm, 31--36 segments, first flagellar segment 5, 08--5, 89 times as long as

wide, combined length of second and third segments 1, 0--1.014 times first;

head thickness-width ratio 1.55--1.596, head length-width ratio 1,085--1.112,

head thickness-camber ratio 3,3--4,3, occipital camber 3, 68--5,16, head-

occiput width ratio 1.36--1. 42, eye-temple ratio 2, 25--2, 83; mesoscutum index

1.097--1.21, head width 1. 46--1.57 times mesoscutum width, scutellum 1:1, 0--

1.1:0. 5--0. 8; depth:width:length of propodeum 1:1,08--1. 12:1, 16--1. 36; hind-

basitarsus 6, 2--10. 75 times as long as thick, first tergite 1.25--1.5 times as

long as hindbasitarsus; width of postpetiole 0.5--0.56 times length, first

tergite 3, 43--3, 56 times as long as postpetiole.

Habitus medium slender, Head above in general highly polished with scat-

tered small weak punctures, rather strongly narrowed behind, temple flattish;

frons and vertex rather distinctly convex, especially stemmaticum, which is

weakly rugulosopunctate, ocellocular space rather large, punctures inconspicu-

ous except at about 30x. Face projecting between eyes (in profile, the far eye

disappears considerably before it comes in line with the near one); midpoint of

occipital carina (viewed in plane of occiput) in the line between upper rear inner

corners of eyes. Pronotum, mesoscutum, mesopleurum, and scutellum mod-

erately densely shallowly punctate, polished, punctures separated by one-half

their diameter or less; notauli rather strongly sharply impressed, prescutum

slightly raised above parapsides, angle between notauli 15°--25°: scutellum

with fully developed lateral carina, anterior margin rather angular but without

evident carina; space enclosed by epomia shortish, narrow, tapered below,

approximately twice thickness of the carina. Propodeum rugose with lateral

longitudinal and median longitudinal carinae distinct; area dentipara (coalesced

with first lateral area) distinctly bounded; petiolar area broad, moderately

excavated, with tendency to transverse rugulae; pleural carina indistinct, pro-

podeal fossa very shallow. Hindfemur moderately stout. First tergite slightly

to distinctly wider than deep, flattish above, more or less distinctly elevated

before spiracles, widening gradually in posterior half or third before spiracles,

postpetiole distinctly wider than petiole; abdomen rather long and slender;

length of ovipositor about 1.25 times depth of apex of abdomen, distinctly

slightly decurved from anterior end of nodus.

Fulvoferruginous; antenna fulvous (sometimes slightly darker); head (except

tip of mandible, stemmaticum, and upper part of occiput black), forecoxa and

-trochanters, posterior face of foretibia, all tarsi, scutellum (except side and

22 Cont. Amer, Ent. Inst., vol. 18, no. 2, 1981

apical edge black), lower surfaces of hindtrochanters, and sometimes inner

surface of hindtibia yellow; collar yellow or fulvous; in addition to areas

already mentioned, the following black: spot on mesoscutum before scutellum,

usually spot under wings, upper surfaces of hindcoxa and hindbasitrochanter

(the last two sometimes piceous), dorsum of second tergite (except lateral

margins), sometimes longitudinal line on mesoscutum (joining prescutellar

patch), often mark at base of propodeum, often upper surface of hinddistitro-

chanter, metanotum and metanepimerum more or less, often splotches near

apex of abdomen, rarely mesosternum, and in one specimen upper surface of

propodeum entirely and first tergite (perhaps shellac); first tergite otherwise

sometimes slightly infuscate to piceous. Coloring of hindtibia exceptional:

black at apex spreading toward base on outer side, becoming progressively

piceous and ferruginous, never yellow on outer surface, rarely yellowish at

base, about one-third of circumference on inner surface at base usually yellow.

Wings hyaline to subhyaline.

Male: Forewing 5.26--6.4 mm, body 9.0--12.0 mm; antenna 6,5--7.5 mm,

32--35 segments, first flagellar segment 5. 96--6.09 times as long as wide,

combined length of second and third segments 1.0--1.07 times first; head

thickness-width ratio 1.56--1.6, head length-width ratio 1,112--1,125, head

thickness-camber ratio 4, 3--4, 4, occipital camber 3. 68--5.17, head-occiput

width ratio 1,37--1.396, eye-temple ratio 2. 85--3, 28; mesoscutum index

1. 06--1.26, head width 1.5--1. 66 times mesoscutum width, scutellum 1:1.1--

1. 2:0. 6--0. 7; depth:width:length of propodeum 1:1. 08--1,125:1, 25--1,. 26;

hindbasitarsus 4, 6--6. 25 times as long as thick, first tergite 1.5--1.83 times

as long as hindbasitarsus; width of postpetiole 0.53--0. 65 times length, first

tergite 3, 6--4, 2 times as long as postpetiole.

Sculpture generally as female. First tergite as in female except petiole

flattish dorsally, but not wider than deep. Color generally as in female with

additional yellow spots at lower edge of prepectus and on mesopleurum at base

of midleg; midcoxa, all trochanters except upper surface of hindtrochanters,

and lower surface of hindcoxa, yellow. Black at apex of scutellum more exten-

sive and apex of abdomen usually black including clasper.

Material examined: 1899, 1000.

Type: 9, Ann Arbor, Mich., Trap, VI-6 ’63, H. & M. Townes (CHT);

allotype: o, Westmoreland St. Pk., Westmoreland Co., Va., 6 July 1951,

Karl V. Krombein (CHT); paratypes: 99, Takoma Pk., Md., VI-20, 1943,

H. & M. Townes (CHT); Moorestown, N. J., July 16, 1939, H. & M. Townes

(CHT); Bemus Pt., N. Y., Jul. 17, 1937, H..K. Townes (CHT); Farmingdale,

N. Y., Jul. 2°38, H. & M. Townes (29, CHT); Farmingdale, N. Y., Jul. 3

38, H. & M. Townes (29, one CHT, one HPH); Ithaca, N. Y., 17 VIII-47

(CNC); Poughkeepsie, N. Y., Jl. 6 1936, H. K. Townes (CHT); Poughkeepsie,

N. Y., Jl. 181996, H. K. Townes (CHT); Baddeck Forks, N. S., 9-V1I-1926,

T. N. Freeman (CNC); Ottawa, Ont., 27 VIL 1947, W. R. M. Mason (CNC);

Crisp, Pa., C. M. Acc. 321, July (29, Pittsburgh); Westm’nd Co., Pa.,

July (29, Pittsburgh); Youngwood, Pa., V-29-41, H. K. Townes (CHT); para-

types: oo, Takoma Pk., Md., VI-15, 1943, H. & M. Townes (CHT); Takoma

Pk., Md., VI-20, 1943, H. & M. Townes (CHT); Takoma Pk., Md., VI-24,

1943, H. & M. Townes (CHT); Ionia Co., Mich., R. & K. Dreisbach (Dreis-

bach); Randolph, N. H., July 2, 1946, J. Peck & M. Townes (CHT); New

Brunswick, N. J., VI- 29 (USNM); Buffalo, N. Y., 4 July ’33, Krombein

Coll. (USNM); Farmingdale, N. Y., Jul. 3°38, H. & M. Townes (CHT);

Ithaca, N. ¥.; dum, 20, 1937, BH. K. Townes (CHT).

Hopper: Nearctic Aphanistes 23

The Nigrorufus Group

This group has ocelli of normal size for the genus, diameter of lateral

ocellus equal to or less than ocellocular line; top of head moderately densely

punctate (punctures separated by about their diameter) all over except in the

rather small orbital spot, punctures moderately large and deep; frons often

slightly depressed, stemmaticum not or scarcely raised above the surrounding

parts, temple usually moderately to strongly bulging, so that head more or

less buccate; mesoscutum and mesopleurum densely punctate, punctures from

moderate in size and depth to strong and deep; epomia enclosing space of mod-

erate size, sometimes somewhat narrow, usually quite distinct, sometimes

bounded by a number of rather weak rugulae instead of a carina, lower end of

epomia not nearly perpendicular to collar; notauli from somewhat sharply

impressed (but not deep, and consisting of a line of punctures rather than a

groove) to diffuse and barely distinct or wide and rounded (approaching the

preceding group) to flat and distinguished only by a line of punctures of differ-

ent size and depth from the adjacent ones; propodeum from neither broad nor

long to slightly broad, rugosity and carinae rather strong, sides slightly to

moderately bulging, petiolar area rarely with transverse rugae predominant;

hindfemur sometimes somewhat incrassate, hindtarsus rather long and slender,

Postpetiole usually elliptical in cross section, slightly deeper than wide, first

tergite usually infuscate to piceous. Species included are coxatus, nigrorufus,

and townesi.

6. Aphanistes coxatus, new species

Female: Forewing 7. 7--10.8 mm, body 13, 6--21.2 mm; antenna 8, 4--11.5

mm (the two specimens with longest forewings and the one with longest body lack

complete antennae), 39--46 segments, first flagellar segment 4.1--4,9 times

as long as wide, combined length of second and third segments 0, 83--0, 878

times first; head thickness-width ratio 1, 09--1.78, head length-width ratio

-1.12--1.27, head thickness-camber ratio 3,0--3,95, occipital camber 3, 9--

5,35, head-occiput width ratio 1,213--1.25, eye-temple ratio 1. 3--1. 7;

mesoscutum index 1.15--1.24, head width 1, 3--1.36 times mesoscutum width,

scutellum 1:0, 9--1.15:0. 55--0. 7; depth:width:length of propodeum 1:1. 04--

1.2:1.04--1.18; hindbasitarsus 9,1--10. 88 times as long as thick, first ter-

gite 1,13--1.32 times as long as hindbasitarsus; width of postpetiole 0, 49--

0.8 times length, first tergite 3,04--4, 2 times as long as postpetiole,

Habitus slightly stout. Face in profile slightly prominent, rather smoothly

curved, in front view somewhat narrow and a little more than usually conver-

gent below, closely finely rather deeply punctate, punctures tending to line up

to form longitudinal wrinkles; clypeus, especially in Canadian specimens,

with somewhat smaller and more numerous punctures in the central area

than usual in our species; temple rather strongly and deeply punctate, cheek

more sparsely and shallowly (but more closely and deeply than in other species

of the genus, especially those of other species groups), the occipital carina

strongly convergent toward base of mandible from the middle of the eye, its

line straightish to slightly recurved and with rather abrupt change of direction

at midlength of the eye; frons and vertex rather strongly to very strongly

deeply punctate or rugulosopunctate. Thorax moderately (New Jersey speci-

mens) to very (Canadian specimens) strongly deeply moderately closely punc-

24 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

tate, in Canadian specimens with considerable development of rugulosity,

especially beneath forewing and at anterior margin of mesopleurum; epomia

distinct, enclosure about one-third length of margin of pronotum, less than

one-half as wide as long; prepectus in Canadian specimens rather highly

polished with little interior sculpture and with distinct carina, in New Jersey

specimens depressed area opaque and only slightly Less closely punctate than

mesopleurum and carina more or less indistinct; speculum with only very

small polished area; scutellum moderately elevated, anterior margin abrupt

or somewhat rounded, highest at one-fifth to one-fourth length, upper surface

flat to slightly depressed, closely and strongly punctate with very strong later-

al carina (often a double line with a narrow space between on each side, the

inner stronger) and anterior transverse carina in Canadian specimens, obso-

lete (most distinct in location of outer row of Canadian specimens) in New

Jersey specimens; notauli distinct but broad and shallow, prescutum Little

elevated above parapsides; sternaulus strongly impressed, extending beyond

midlength of mesopleurum. Propodeum coarsely rugose with only faint sculp-

ture in the spaces; area dentipara rather distinct, usually at least partially

separated from first lateral area; all longitudinal carinae more or less distinct.

Forecoxa rather strongly flattened in apical ventral two-thirds, flattened area

bounded proximally by a distinct narrow rounded oblique ridge (not a carina),

higher at its inner (posterior when legs at right angle to midline) end (at the

point where some other species have highest point of a rounded conical forma-

tion sometimes with a faint suggestion of a ridge, but with the entire ventral

surface convexly rounded) with a flattish or cylindrically concave surface

proximad. First tergite of Canadian specimens with a median longitudinal

(rounded) furrow, usually from near base to near apex, occasionally from

just before to just behind spiracles, lacking altogether in New Jersey speci-

mens.

Antenna from dark ferruginous to piceous, with first flagellar segment

above, usually also pedicel, sometimes scape above (latter otherwise yellow)

black. Head yellow except tip of mandible black, upper one-fourth to two-

fifths of gena usually black (brown or with large brown spot in New Jersey

specimens); frons, vertex, stemmaticum, and upper part of occiput black

except yellow orbital spot (in New Jersey specimens, yellow extends from

face along sides of frons and vertex nearly to occipital carina). Mesoscutum

usually black except anterior ends of notauli and often lateral edges of parap-

sides ferruginous; sterna usually wholly black (only black just in front of coxae

in New Jersey specimens); spot under forewing black in all specimens, in

Canadian specimens mesopleurum more extensively black, sometimes wholly

black including prepectus; pronotum from wholly fulvous (strongly tinged with

yellow in New Jersey specimens) to almost wholly black; scutellum ferruginous,

often with small amount of yellow around edges, rarely with some black

toward base, extending caudad medially; thorax otherwise fulvoferruginous.

Propodeum with transverse black marks at base of area dentipara (correspond-

ing to first lateral area), small in New Jersey specimens, usually larger in

Canadian specimens and extending from side to side, sometimes expanded over

almost entire dorsal surface, in which case metakatepisternum also black.

First tergite medially infuscate in all specimens, rarely with some black;

lateral edges of second tergite and remainder of abdomen fulvous. Fore- and

midlegs yellow except femora ferruginous, hindtarsus and base of hindtibia

yellow, hindtibia apically black, extending well beyond halfway to base on at

least one side, hindfemur ferruginous, dorsally infuscate. Wings subhyaline,

rarely hyaline or slightly infuscate.

Hopper: Nearctic Aphanistes 25

Male: Forewing 7.8--9.3 mm, body 15.0--17.4 mm; antenna 9.4--10.8

mm, 39--43 segments, first flagellar segment 4.06--4.75 times as long as

wide, combined length of second and third segments 1.0--1.05 times first;

head thickness-width ratio 1.58--1.74, head length-width ratio 1.12--1.39,

head thickness-camber ratio 2. 86--3.2, occipital camber 3.97--4.5, head-

occiput width ratio 1.16--1.25, eye-temple ratio 1.6--1.8; mesoscutum index

1.1--1.2, head width 1.27--1.41 times mesoscutum width, scutellum 1:0. 89--

1.15:0. 58--0.6; depth: width:length of propodeum 1:1.1--1.19:1.025--1.1;

hindbasitarsus 6.625--7.55 times as long as thick, first tergite 1. 176--

1.396 times as long as hindbasitarsus; width of postpetiole 0.5--0.6 times

length, first tergite 3.2--3.58 times as long as postpetiole.

Sculpture and color substantially as in female.

Material examined: 1822, 10¢°°.

Type: *, Ex E. crepuscularia, Moffette, Que., Rec. 2208A, F. I. Survey

1940, Em. (Incubator) 19- III 1941 (CNC); allotype: Say <>. ay 2a porcelaria,

Treesbank, Man., Rec.: 6298A, F. I. Survey 1939, Em. (Incubator) 20-V

1939 (CNC); paratypes: 99, No. 5-1626A, Date 3- ne P.i1. 8.1951, - Zale sp..,

Sinclair, B. C. (CNC); Spruce Woods, Man. , May 26, 1939, H. A. Richmond

(F. I. S., CNC); Ex P. porcelaria, Treesbank, Man., Rec.: 6298A, F. I.

Survey sh Em. (Incubator) 20-V 1939 (CNC); Ex P. porcelaria, Treesbank,

Man., Rec.: 6298A, F. I. Survey 1939, Em. (Incubator) 22-V 1939 (48,

CNC): Ex E. crepuscularia, Treesbank, Man., Rec.: 6298A, F. I. Survey

1939, Em. (Incubator) 22-V 1939 (CNC); Wildwood, N. Js) DT-VIH 1985, Lae d.

Bottimer (5°, four HPH, one CHT); Prince Albert S. [= Saskatchewan], Em.

10-3-53, F. I. S: W-2320A, Ex Eufidonia notataria (CNC); Ex F. jocosa,

Ct. Labrecque [P.Q.], R.I.F. '40, 4337-A, Em. 26-2-41 (CNC);

[no label, but associated in collection with specimen of A. townesi labelled

'W.Was."' without date, but evidently of similar age] (USNM); [also without

collection data, with one label ''Anomalon nigro-rufum Nort. ¢"' in writing

suggesting a date approximately between 1880 and 1910, and a second label

"Anomalon nigro-rufum Nort., C. W. T. Gahan 1920" | (USNM); paratypes:

Oo, Rec.: 44-662A, F. I. Survey 1946, Em. (incubator), 11-II-1947,

Hazelton, B. C., Phalaenidae | =Noctuidae] (CNC); Spruce Woods, Man.,

May 16, 1939, H. A. Richmond, F. I. S. Man. 1942-1943 (CNC); Spruce

Woods, Man., May 26, 1939, H. A. Richmond, (F.I.S., CNC); Ex P.

porcelaria, Treesbank, Man., Rec: 6298A, F. I. Survey 1939, Em. (Incuba-

tor) 17-V 1939 (CNC); Ex Anacamptodes sp., Treesbank, Man., Rec: 6298A,

F. I. Survey 1939, Em. (Incubator) 20-V 1939 (CNC); Ex P. porcelaria,

Treesbank, Man., Rec: 6298A, F. I. Survey 1939, Em. (Incubator) 22-V

1939 (CNC); Ex Zale helata, Angus, Ont., No. 045-521, Date 21-I1-46, F. I.

S. 1945 (CNC); Ex Zale sp., Pointe au Baril, Ont., 550, No. 3873-01-1,

Date 16-III-51, F. I. $.1950, 5437 (CNC); Rec; 6298, F. I. Survey 1939, Em

(incubator), 18-V-1939 (CNC).

Hosts: Geometridae and Noctuidae: Anacamptodes sp., Ectropis crepus-

cularia, Eufidonia nee Feralia jocosa, Protoboarmia porcelaria, Zale

helata, Zale sp.

7. Aphanistes nigrorufus (Norton)

Anomalon nigro-rufus Norton, 1863. Ent. Soc. Phila., Proc. 1: 361.

Type: 2, N. Y. (in Peabody Museum, Yale University, New Haven,

Connecticut).

26 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

Female: Forewing 11.03--12.8 mm, body 18.9--22.3 mm; antenna

11.4--12.9 mm (only two specimens with complete antennae), 45--47 segments,

first flagellar segment 3.9--5.7 times as long as wide, combined length of

second and third segments 0. 83--1.04 times first; head thickness-width ratio

1.6--1.7, head length-width ratio 1.14--1.21, head thickness-camber ratio

3.475--4.3, occipital camber 3.65--5.55, head-occiput width ratio 1.23--

1.38, eye-temple ratio 1.4--1.75; mesoscutum index 1.2--1.3, head width

1.25--1.5 times mesoscutum width, scutellum 1:1.0--1.15:0.55-0.66; depth:

width:length of propodeum 1:1.012--1.125:0.94--1.1; hindbasitarsus 8.0--

12.375 times as long as thick, first tergite 0.95--1.125 times as long as hind-

basitarsus; width of postpetiole 0.54--0.85 times length, first tergite 3.2--

4.2 times as long as postpetiole.

Habitus somewhat stout to moderately slender. Antenna very slightly com-

pressed. Face with a number of vertical wrinkles, moderately closely finely

punctate; gena somewhat convergent toward base of mandible, relatively finely

sparsely punctate, subpolished; frons and vertex rugulosopunctate to ruguloso-

reticulate, rather opaque, sculpture rather coarse, frons slightly concave;

eye rather large and prominent. Thorax rather densely deeply punctate, sub-

polished to rather opaque; notauli indistinct or obsolete, little or not at all

impressed, represented chiefly by a closely spaced line of punctures differing

from the general punctation, prescutum little elevated above parapsides, often

imperceptibly; scutellum usually with very strong lateral carina, disc narrowly

squarish, convergent caudad, usually broadly shallowly longitudinally concave

(these features present even when lateral carina weak) with large deep moder-

ately close punctures, moderately elevated, highest at one-third to one-half its

length from anterior margin; epomia usually quite distinct and a number of

rather weak rugulae rather than a carina, enclosure moderately large, rather

narrow; sternaulus not longer than one-half length of mesopleurum, little or

no rugulosity under forewing, speculum with small subpolished area not parti-

cularly less densely punctate than remainder of pleurum. Propodeum without

any really distinct areolation, strongly rugose-reticulate with fair amount of

faint punctation in spaces, opaque, without distinct propodeal fossa; one speci-

men has about posterior one-third of what would be petiolar area with curved

transverse rugae. Hindfemur at least seven times as long as thick. Postpetiole

usually rather elliptical in cross section with long axis vertical.

Antenna fuscous to piceous; head yellow except tip of mandible, temple,

frons, stemmaticum and vertex (except orbital spot), and upper part of occiput

black. Thorax black except collar, anterior edge of pronotum (sometimes

also upper edge), and metanepisternum piceous or rarely dull fulvous (in type,

spot on pronotum and spot on collar rufofulvous); apex of disc of scutellum often

piceous. Propodeum black except metapleurum from above hindcoxa and pedi-

cel fulvoferruginous (in type, this area and almost entire metapleurum yellow-

ish rufous). Wings subhyaline. Fore- and midlegs yellow except femora ful-

vous; hindleg with tarsus and upper two-thirds of tibia yellow, apex of tibia,

coxa, and most of upper surface of trochanters black, hindfemur dark brown

above. Abdomen fulvoferruginous, first tergite more or less strongly infus-

cate to piceous above and more or less of upper surface of apical tergites

infuscate or black.

Male: Forewing 7.75--10.35 mm, body 13.8--20.5 mm; antenna 10+ mm

(only three specimens, none with complete antennae), 43+ segments (first seg-

ment missing in all); head thickness-width ratio 1.59--1.69, head length-

width ratio 1.177--1.2, head thickness-camber ratio 2.875--3.95, occipital

camber 3.38--4.33, head-occiput width ratio 1.405--1.415, eye-temple ratio

Hopper: Nearctic Aphanistes 27

2.0--2.97; mesoscutum index 1.15--1.3, head width 1.36--1.5 times meso-

scutum width, scutellum 1:1.0--1.1:0.52--0.55; depth: width:length of pro-

podeum 1:1.15--1.26:1.15--1.23; hindbasitarsus 6.25--10.0 times as long

as thick, first tergite 1.05--1.32 times as long as hindbasitarsus; width of

postpetiole 0.492--0.646 times length, first tergite 3.48--4.03 times as long

as postpetiole.

Habitus, sculpture, and color as in female on the whole. Eye extremely

large; scutellum with some lighter markings in the two larger specimens,

somewhat polished in the smallest.

Material examined: 11992, 30°.

Distribution: British Columbia, New York, North Carolina, Nova Scotia,

Ontario, Quebec, Tennessee.

Hosts: C. [Caripeta] divisata? (Geometridae).

8. Aphanistes townesi, new species

Female: Forewing 5.3--11.88 mm, body (estimated 9.7)--20.55 mm;

antenna 5. 2--11.5+ (estimated 12.5) mm, 30--39 segments (note that largest

specimen has incomplete antennae), first flagellar segment 4.16--5.65 times

as long as wide, combined length of second and third segments 0.91--1.09

times first; head thickness-width ratio 1.6--1.7, head length-width ratio

1.14--1.21, head thickness-camber ratio 3.68--5.4, occipital camber 4. 56--

6.675, head-occiput width ratio 1.3--1.4, eye-temple ratio 1.73--2.7; meso-

scutum index 1.05--1.18, head width 1.37--1.56 times mesoscutum width,

scutellum 1:0. 95--1.15:0.5--0.7; depth: width:length of propodeum 1:1.02--

1.2:1.1--1.23; hindbasitarsus 7.4--10.8 times as long as thick, first tergite

0.989--1.21 times as long as hindbasitarsus; width of postpetiole 0.588--0. 72

times length, first tergite 3.5--4.0 times as long as postpetiole.

Habitus moderately slender. Antenna rather distinctly compressed, about

1.25 times as deep as wide. Face rather densely finely punctate with few and

weak vertical wrinkles; gena rather sparsely finely punctate, more or less

polished, temple progressively less punctate and more polished than vertex;

frons and vertex rugulosoreticulate or rugulose, sometimes with some areas

merely punctate, sculpture rather dense and moderately deep, subopaque,

frons slightly concave behind antennae (in addition to the usual antennal

scrobes). Mesoscutum and mesopleurum rather finely weakly densely punc-

tate; notauli fairly distinct, varying from shallow, diffuse, and barely distinct

(lateral spread less than in hyalinis group) to narrow, shallow, somewhat

sharply impressed lines of punctures; maximum width of prescutum greater

than that of one of parapsides, prescutum distinctly raised above parapsides;

epomia usually single, forming a smooth curve, enclosure usually rather

narrow, maximum width usually less than one-half length, without noticeable

rugulae within; scutellum flattish, weakly elevated, with a slight rather

straight anterior declivity, sometimes with an indistinct median longitudinal

depression never extending from side to side, lateral carina weak to indistinct

and below upper surface, punctation of disc somewhat sparser, deeper, and

larger than on mesoscutum; sternaulus usually distinctly impressed, one-half

or more the length of mesopleurum, latter usually with a number of horizontal

rugulae beneath wings. Propodeum rather coarsely rugose, rather polished,

only area dentipara (not separated from first lateral area) defined, portion of

median longitudinal carina bounding these areas and lateral longitudinal carina

recognizable, pleural area moderately distinctly impressed (i.e., propodeal

28 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

fossa developed), pleural carina usually stronger than others; metapleurum

slightly bulging; petiolar area rather distinctly longitudinally depressed.

Hindfemur stoutish, about 6.4 times as long as deep. Postpetiole usually

rather elliptical in cross section, slightly deeper than wide.

Antenna fuscoferruginous to piceous, scape either wholly yellow or black

above. Head yellow except clypeal tooth fuscous, tip of mandible, temple,

and upper part of occiput black. Thorax black, with collar and propleurum

extensively, parapsides and mesopleurum more or less extensively, disc of

scutellum largely (usually with black mark at base), and propodeum largely

(except more or less of dorsal surface and sometimes metakatepisternum )

fulvous or ferruginous. First tergite usually partly to wholly piceous or black.

Legs yellow except fore- and midfemora more or less fulvous, hindcoxa, more

or less of hindtrochanters, and tip of hindtibia black, hindfemur fulvoferru-

ginous to ferruginous, often darker above and infuscate toward apex. Wings

hyaline to lightly infuscate, usually subhyaline.

Male: Forewing 5.9--11.3 mm, body 11.35--21.2 mm; antenna 7. 4--

12 mm, 33--40 segments (largest specimens have incomplete antennae), first

flagellar segment 3.88--5.12 times as long as wide, combined length of second

and third segments 0.95--1.15 times first; head thickness-width ratio 1. 64--

1.77, head length-width ratio 1.133--1.358, head thickness-camber ratio 3.05--

4.86, occipital camber 3.6--5.86, head-occiput width ratio 1.3--1.4, eye-

temple ratio 2.0--2.5; mesoscutum index 1.13--1.26, head width 1. 436--

1.486 times mesoscutum width, scutellum 1:0.95--1.1:0.5--0.6; depth:width:

length of propodeum 1:1.05--1.116:1.06--1.176; hindbasitarsus 5.33--9.5 |

times as long as thick, first tergite 0.9--1.3 times as long as hindbasitarsus;

width of postpetiole 0. 548--0.634 times length, first tergite 3.48--3.94 times

as long as postpetiole.

Sculpture as in female except punctation of scutellum little different from

mesoscutum. Color much as in female, but scutellum always with at least

posterior edge black, more usually black with two lateral fulvous or piceous

spots or rarely wholly black; sometimes thorax wholly black except tegula,

upper anterior and posterior margins of mesopleurum and usually some portion

of upper edge of pronotum and a fairly large blotch at base of coxae; propodeum

more extensively black: most of dorsum and metakatepisternum always black.

Apical tergites of abdomen sometimes black.

Material examined: 822, 200°.

Type: 2°, Ashford, Wash., Aug. 18, 1940, H. & M. Townes (CHT);

allotype: “, Ashford, Wash., Aug. 18, 1940, H. & M. Townes (CHT);

paratypes: °°, Ex Nyctobia limitaria, Athabasca, Alta., No. A 471 C, Date:

13-11-50, F.I.S. 1949 (CNC); No. P49-3634, Date: 13-III-50, F.1.S. 1949,

Hydriomena sp., Kitwanga R., B.C. (CNC); Wissahickon Val., Philadelphia,

Pa., 12 Oct. 1938, H. P. Hopper (HPH); Stoneham, P.Q., June 21, 1938,

H. & M. Townes (CHT); Ashford, Wash., Aug. 18, 1940, H. & M. Townes

(29 in addition to type, CHT); W. Was. (no other data, USNM); paratypes: od’

Blood Mt., Ga., 7-6-51, P.W. Fattig (CHT); Flin Flon, Man., Em. 17-I-47,

F.1.S. :W-934, Ex Carapeta [sic!] divisata (CNC); Waskesiu, Man., Em.

5-3-51, F.1.S:W-4389, Ex Semiothisa sexmaculata (F.1.S., CNC); Huron

Mts., Mich., July 20, 1959, Henry Townes (CHT); Allarduilie, N.B., Rec:

40-L691A. F.I.Survey 1940, Em. Apr. 13 1941 (CNC); Loc. Sunbury, N.B.,

Host [blank], Rec: 45 L1084A, F.I.Survey, Em. Apr 71946, Ex Semiothisa

sexmaculata (F.I.S., CNC); Franconia, N.H., VII-1-46, M. Townes (CHT);

Mt. Madison, N.H., June 23 1938, H. & M. Townes (CHT): Mt. Madison, N.H.,

June 26 1938, H. & M. Townes (CHT); Pinkham Notch, N.H., June 25, 1938,

H. & M. Townes (CHT); Ottawa, Ont., 18-VII-1946, A. Brooks (CNC); Spring

Hopper: Nearctic Aphanistes 29

Br., Pa., VII[-1-1945, H.K. Townes (CHT); Portage, P.E.I., 18-VII-1940,

G.S. Walley (CNC); Green Lake, Sask., Em. 9-III-50, F.1.S:W-2408A, Ex

Semiothisa sexmaculata (CNC); Gr.Sm.Mt.Nat. Park, Tenn., J.G. Chillcoit,

Indian Gap to Clingman’s Dome 5200-6600’, Tenn., 6-VIII-1957 (CNC); Lake

Willoughby, Vt., Alt. 3200 ft., June 17-29, 1945, C.P. Alexander (CHT);

Ashford, Wash., Aug. 18, 1940, H. & M. Townes (30 in addition to allotype,

CHT).

ae Geometridae: Caripeta divisata, Hydriomena sp., Nyctobia

limitaria, Semiothisa sexmaculata.

The Enargiae Group

Ocelli of normal size, eye fairly large but only moderately projecting from

head outline in plan view, head oblong elliptic or buccate, temple rather

bulging; gena widening from temple to near base of mandible and without con-

spicuous change of direction or angle (usually none), rather large area

between stemmaticum and orbit almost impunctate; epomia usually weak and

multiple, not reaching collar at upper end before midline, enclosure usually

very narrow; punctation of mesocutum and mesopleurum fairly strong and

dense, usually stronger and denser just above change of direction from pleurum to

sternum; prescutum somewhat wider anteriorly than parapsides, little raised above

them, but set off by rather broad indentations at anterior ends of notauli; scutellum

moderately strongly convexly elevated, lateral carina obsolete in larger specimens,

somewhat distinct in smaller ones; propodeum quite convex in all aspects,

rather large in proportion to thorax, usually with few carinae or strong rugae,

and rather finely weakly reticulate; petiole usually distinctly wider than deep,

rarely slightly deeper than wide; postpetiole usually slightly convergent

caudad, occasionally deeper than wide, often wider than deep, in the latter

case often with one or more shallow rounded longitudinal grooves; hindleg

somewhat incrassate. Only species included is enargiae.

9. Aphanistes enargiae, new species

Female: Forewing 5.65--12.95 mm, body 9.2--18.75 mm; antenna 5. 6--

10.1 mm, 32--41 segments, first flagellar segment 3.68--4.6 times as long as

wide, combined length of second and third segments 0. 86--1.12 times first;

head thickness-width ratio 1.62--1.77, head length-width ratio 1.14--1.23,

head thickness-camber ratio 2.95--4.25, occipital camber 3.6--5.7, head-

occiput width ratio 1.18--1.39, eye-temple ratio 1.23--2.5; mesoscutum

index 1.05--1.23, head width 1.2--1.6 times mesoscutum width, scutellum

1:0. 86--1. 1:0. 5--0.6; depth: width:length of propodeum 1:1.0--1.25:0. 95--

1.2; hindbasitarsus 7.045--10.0 times as long as thick (in one specimen

compressed with ratio to the two thicknesses 6.88 and 8.44), first tergite

1.014--1.258 times as long as hindbasitarsus; width of postpetiole 0.6--0. 88

times length, first tergite 3.4--4.2 times as long as postpetiole.

Habitus somewhat stout. Inner orbit relatively finely fairly closely punc-

tate, sculpture of median field of face from similar to that of inner orbit

with a few vertical wrinkles (chiefly smaller specimens) to wholly somewhat

closely finely wrinkled; frons rugulosopunctate to rugulosoreticulate; gena

moderately closely finely punctate, basically widening from temple toward

base of mandible (but widest a little above lower end of eye, though not dis-

30 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

tinctly narrowing below this point) and without conspicuous change of direction

or angle near midpoint of genal carina (usually none); rather large area be-

tween apex of eye and stemmaticum almost impunctate, stemmaticum and ver-

tex strongly punctate. Pronotum rugulose around edges, rather closely moder-

ately finely punctate otherwise, with polished impunctate spot near center of

lateral portion, especially in small specimens, or rather distinctly rugulose

throughout; epomia usually weak and multiple (most often triple), not reaching

carina of collar at upper end before midline, enclosure usually very narrow,

straight-line length from outer end to midline 9--10 times maximum interior

width, sometimes indistinctly closed; mesoscutum fairly closely strongly

punctate, notauli rather distinctly defined, moderately sharply impressed, but

consisting of a line of very closely spaced punctures rather than a groove and

rather shallow; prescutum not much raised above parapsides, but set off by

depressions at anterior ends of notauli, prescutum rather wide anteriorly

with notauli fairly strongly divergent; mesopleurum rather densely slightly

coarsely punctate, usually more strongly and densely just above change of

direction from pleurum to sternum and more or less rugulose under forewing;

scutellum moderately strongly convexly elevated, distinctly rising from anteri-

or carina for about one-third its length, sometimes with a slight conicity or an

obscure transverse ridge, sometimes more or less quadrangular, especially

in small specimens, in which also lateral carina (usually indistinct or obso-

lete) may be somewhat distinct, rather densely distinctly punctate. Propodeum

quite convex in all aspects, rather large in proportion to thorax, from rather

closely finely weakly rugulosoreticulate with only area dentipara fairly distinct

(rather short and wide) and a trace of pleural carina to more openly and less

finely rugulosoreticulate with a portion of the lateral longitudinal carina

behind the area dentipara also more or less distinct; lower portion of pleural

area and upper portion of metapleurum rather strongly bulging, metakate-

pisternum more or less distinctly inflexed. Forecoxa subglobose, undersur-

face from flattish convex to slightly elevated with an indistinct conicity, which

when present may form the inner end of an indistinct rounded ridge without

distinct change of direction of the surface on either side. Hindleg somewhat

incrassate. Petiole rarely slightly deeper than wide, most frequently dis-

tinctly to strongly wider than deep; postpetiole occasionally rather deep, usu-

ally wider than deep, in the latter case often with one or more shallow

rounded longitudinal grooves, postpetiole usually slightly convergent caudad

behind spiracles,

Antenna from fulvoferruginous to piceous, with part of upper surface of

first flagellar segment, usually pedicel, and sometimes upper surface of scape

black, lower surface or whole of scape yellow. Head yellow except tip of man-

dible black, margin of clypeus including mucro fulvous, temple fulvoferrugin-

ous or black, frons, stemmaticum, and vertex black except yellow orbital spot

and sometimes ferruginous lateral margin; upper part of occiput black.

Thorax may be from wholly fulvous except spot before scutellum, spot beneath

forewing, mesosternum, side and apical margin of scutellum, to wholly black

except for reddish tinge on disc of scutellum, small spot of reddish tinge on

mesopleurum before speculum, and the portion of the pleural and metapleural

region of propodeum immediately in front of hindcoxa. First tergite more or

less infuscate with postpetiole sometimes strongly so but not black; second

tergite broadly black along midline, lateral margin dull fulvous as is remainder

of abdomen, last two or three tergites with rather heavy infuscation. Legs

yellow except inner surface of fore- and midtibiae and -femora, sometimes

fore- and midcoxae, and hindfemur fulvoferruginous; hindfemur rather infus-

Hopper: Nearctic Aphanistes 31

cate; apex of hindtibia and at least upper surface of hindcoxa and hindtrochan-

ters black; base of hindbasitarsus sometimes orange-tinged. Wings sub-

hyaline to occasionally slightly infuscate.

Male: Forewing 5.8--10 (?--incomplete, estimated 10.3) mm, body

12.4--19.2 mm (specimen with forewing 5.8 mm lacks abdomen); antenna

(more than 6. 4)--9.6 mm (missing or incomplete in largest specimens),

(more than 29) 35--37 (probably more: see foregoing) segments, first flagel-

lar segment 3.8--5.2 times as long as wide, combined length of second and

third segments 0.91--1.15 times first; head thickness-width ratio 1. 63--

1.785, head length-width ratio 1.17--1.28, head thickness-camber ratio 2.47--

4.0, occipital camber 3.3--5.3, head-occiput width ratio 1.2--1.4, eye-

temple ratio 1.6--2.76; mesoscutum index 1.06--1.24, head width 1.26--

1.49 times mesoscutum width, scutellum 1:0.9--1.1:0.45--0.65; depth:width:

length of propodeum 1:1.05--1.25:0.94--1.26; hindbasitarsus 5.6--8.3 times

as long as thick, first tergite 1.0--1.36 times as long as hindbasitarsus;

width of postpetiole 0.6--0.78 times length, first tergite 3.22-3.86 times as

long as postpetiole.

Sculpture and color generally as in female, but with general wrinkling of

face only ina minority of specimens, and in a number of specimens the meso-

pleurum rather rugulosoreticulate below the subalar rugulosity.

Material examined: 5822, 400,

Type: ¢, Ex E. decolor, Cedar Lake Field Station, Ont., Em. Mar. 8/60,

Inc. (CNC); allotype: °, Ex Enargia decolor, Cedar Lake Field Station, Ont.,

Em. Feb. 22/60 (CNC); paratypes: 9%, Ex Nyctobia limitaria, 59A 809-01,

Em. 15-II-60, Inc., F. I. S., Hinton [Alta.] (CNC); McMurray, Alta., 22 VI

03, G. E. Ball (CNC); No. 48-2652A, Date 4-III-48, F. I. S. 1949, Melano-

lophia imitata, Alice Arm, B. C. (CNC); No. 8051-1786, Date 5-IIl, F. I. S.

1952, Melanolophia imitata Mi31, Big Bend, B. C. (CNC); No. 50-694, Date

{9-I,, F. 1. 8. 1951, Geometrid,; Cambie, B. C- (CNC);Clinton, B. C., 18-

V1 1938, J. K. Jacob (CNC); No. P49-548G, Date 11-IV-50, F. I. S. 1949,

Geometrid, Comox, B. C. (CNC); No. P49-490C, Date 18-III-50, F. I. S.

1949, Geometridae, Denman Island, B. C. (CNC); No. BC49-1376, Date 23-

fit-50, F. 1. S. 90, N. limitaria, Fernie, B. C. (CNC); No, B. C. 49-2017F,

Date 5-IV-50, F. I. S. 1950, Itame hulstaria, Golden, B. C. (CNC); No.

90-633A, Date 31-I-, F. I. S. 1951, M. imitata, Grand Beach, B. C. (CNC);

No. BC48-2170, Date 19-II-, F. I. S. 1949, Enypia moilleti?, Grenville

Channel, B. C. (CNC); 56-846-01A, R'rd, Kakweiken R, BC, Ex Aseptis

binotata (CNC); No. P49-1860A, Date 13-IV-50, F. I. S. 1949, Geometrid,

Kildonan, B. C. (CNC); No. BC48-2314, Date 24-II-, F. I. S. 1949, Enypia

moilleti?, Lowe Inlet, B. C. (CNC); No. 51-1685A, Date 4-II-, F. I. S. 1952,

N. limitaria, Miocene, B. C. (CNC); No. 50-211C, Date 2-II-, F. I. S. 1951,

N. limitaria, Pritchard, B. C. (?9, one CNC, one HPH); No. B. C. 48-1005,

Date 3-2-49, F. I. S. 1949, Phalaenid, Quesnel, B. C. (CNC); No. P49-739D,

Date 23-I-50, F. I. S. 1949, Geometrid, Sproat River, B. C. (CNC); No.

BC51-2320A, Date 29-II, F. I. S. 1952, Trout Lake, B. C. (CNC); No.

P49-1854A, Date 13-IV-50, F. I. S. 1949, Geometrid, Ucluelet, B. C. (CNC);

No. P49-1285C, Date 22-III-50, F. I. S. 1949, Geometrid, Wellington, B. C.

(CNC); No. BC49-866E, Date 27-III-50, F. I. S. 1950, N. limitaria, White-

mans Cr., B. C. (CNC); No. 50-886A, Date 5-II-, F. I. S. 1951, N. limi-

taria, Wynndel, B. C. (CNC); nr Estes Pk, Colo, June 15, 1948, HMGD&J

Townes (CHT); Aweme, Man., R. M. White, 13.VI.1922, 38 (CNC); 5 mi.

SW Shilo, Man., 5-VI-1958, C. D. F. Miller (22, CNC); Sprague, Man., Em.

18-II-52, F. I. S.: W-1709A, Ex Orthosia hibisci (CNC); ex P. porcelaria,

oa Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

Spruce Woods, Man., Rec; 6301, F. I. Survey 1939, Em. (Incubator), 16-V

1939 (CNC); ex P. porcelaria, Spruce Woods, Man., Rec; 8301, F. I. Survey

1939, Em. (Incubator), 17-V 1939 (CNC); ex P. porcelaria, Treesbank, Man.,

Rec; 6298A, F. I. Survey 1939, Em. (Incubator), 19-V 1939 (CNC); Cadillac

Mt. Me, VI- 14-35, C. P. Alex [sic!] (USNM); MICHIGAN, Cheboygan Co.,

VI-27-28-1949, G. W. Byers (UMI); Loc. Madawaska Co., N. B., Host:

Nyctobia limitaria, Rec.(B, F. I. Survey 1949, Em. Feb. 22, 1950 (F. 1. S.,

CNC); Loc. St. Georges, NFLD., Host. Nyctobia limitaria, Rec. @) ?, F. I.

Survey 1950, Em. Mar 19 1951 (F. I. S., CNC); Loc. mi. 56 Cabot T., Inv.

Co., N. S., Ex. undet. Host. 52-L618, Rec. 52-L618, F. I. Survey, Em.

Mar. 20 1953 (F. I. S., CNC); Ex Geometridae, Barry's Bay, Ont. 82, No.

Sso 3459-03, Date 13- Ix- 505° Fo I. SY-1900 (CNC); Ex. Enargia decolor,

Cedar Lake Field Station, Ont., Em. Feb. 15/60, Inc. (CNC); Ex. Enargia

decolor, Cedar Lake Field Station, Ont., Em. Feb. 22/60, Inc. (32, two CNC,

one HPH); Ex. Enargia decolor, Cedar Lake Field Station, Ont., Em. Feb.

23/60, Inc. (39, CNC); Ex. Enargia decolor, Cedar Lake Field Station, Ont.,

Em. Feb. 26/60, fe. (22, CNC); Ex - Geometrid sp., Cloyne, Ont., No. 050-

1303C, Date 2-IV-51, F. I. S. 1950 (CNC); ex Ectropis crepuscularia, Halls

Lake, Ont., No. 052- 1607B, Date 3-ITI-53, F. I. S. 1952 (CNC); Ex Geomet-

ridae, North Bay, Ont. 71, No. S49- 2323F, hat ge Ix-1949, F. I. S. 1949

(CNC); Ex Phalaenidae, Thessalon, Ont., . S47--785E, Date 1-III-1948,

F.d. S. 1947 (F. I. S., CNC); Ex Ectropis a ONS Henny Tory Hill, Ont.,

No. 050-1256C, Date 14- I-51, F. I. S. 1950 (CNC); Ex Phalaenidae, Twp.

Prince, Sec. 32, [Ont. , ] No. S46- 903, Date 24-III-1947, F. I. S. 1946 tei,

Ss. , CNC); Linglestown, Pa., V-30, Champlain (USNM); Ex S. granitata, Lac

Pentecote emg. how. L. FB, 140, 4677 Em. 20-II-41 (CNC); Pierceland, Sask.,

Em. 13-I[-52, F. I. S: W-1488, Ex Iphimorpha pleonectusa (F. I. S., CNC);

paratypes: Jo, Waterton Lakes, Alta., 30-VI-1923, J. McDunnough (CNC):

No. 60-1004, Date 29-I-, F. I. S. 1951, N. limitaria, Aleza Lake, B. C.

(CNC); No. 50- 1736E, Date 5- Ill-, F. I. S. 1951, M. imitata, Anstey Arm,

B. C. (CNC); No. BC49- 1260C. Date 20-11-50, FL J. S. 1950, N. limitaria,

Ft. Fraser (CNC); No. P50-1804, Date 14-TIII- 51, ae ook 1950, , M. imitata,

Grouse Mt., B. C. (CNC); No. 50- 656A, Date 31- FSS. 1951,

N. limitaria, Inonoaklin, B. C. (CNC); No. P49-550A, Date 1-IV-50, Pt SS:

1949, Nyctobia Sp., Merville, B. C. (CNC); No. 50-1018, Date 28-I-, F. I.

S. 1951, N. limitaria, Monashee, B. C. (CNC); No. P49-1737A. Date 31-III-

50, F. I. S. 1949, Geometrid, Nuchalat Arm, B. C. (CNC);No. P49-2894A,

Date 19-IV-50, F. I. S. 1949, Geometridae, Prince Rupert, B. C. (CNC); No.

P49-1318, Date 20-III-50, F. I. S. 1949, Geometrid, Royston, B. C. (CNC);

No. 50-692A, Date 29-I-, F. I. S. 1951, N. limitaria, Solsqua, B. C. (CNC);

No. P49-1788, Date 4-IV-50, F. I. S. 1949, Geometrid, Tofino Inlet, B. C.

(2c, one CNC, one HPH); No. P49-1858A, Date 6-IV-50, F. I. S. 1949,

Lepidoptera, Ucluelet, B. C. (CNC); No. 50-1414A, Date 28-I-, F. I. S.

1951, N. limitaria, Wingdam, B. C. (CNC); No. BC51-1138C, Date 10-IIL,

F. I. S. 1952, Phalaenidae, Wynndel, B. C. (CNC); No. 50-628A, Date 29-I-,

F. I. S. 1951, N. limitaria, Yard Creek, B. C. (CNC); Antrim Co., Mich.,

Vll-4-60, R., K. Dreisbach (Dreisbach); W. Fish Pt., Chippewa Co., Mich.,

1 Aug. 14, F. Combs (Dreisbach); Mt. Washington, N. H., a-4000, VI-4,

1914 (USNM); Ex Geometridae, Angus, Ont., Rec: 4224B, F. I. Survey 1938,

Em. 11-X 1938 (CNC); Ex Enargia decolor, Cedar Lake Field Station, Ont.,

Em. Feb 15/60, Inc. (20°, CNC); Ex Enargia decolor, Cedar Lake Field Sta-

tion, Ont., Em. Feb 22/60, Inc. (50, including allotype, CNC); Ex E. decolor,

Cedar Lake Field Station, Ont., Em. Mar. 18/60, Inc. (CNC); Ex Phalaenid,

Hopper: Nearctic Aphanistes 3 33

Hawk Lake, Ont., No. S49-984, Date 8-II[-1946, F. I. S. 1945 (CNC); Ex

Phalaenid sp., Kaladar, Ont., No. 049-494A, Date 7-II-50, F. I. S. 1949

(CNC); Ex Ectropis crepuscularia, Lassarade, Ont., No. 050-1285B, Date

16-IV-51, F. I. S. 1950 (CNC); Marmora, Ont., 29-V-1952, J. C. Mitchell

(CNC); Ex Geometridae, North Bay 71, Ont., S51-3969-01-1, 4-Ix-1951,

Eu 8.8148 «1. 82, °CNC)y.ex EB. hastata, Sand I.,-Ont., Ree, 1 2950C, F.

I. Survey 1940, Em. (Incubator), 27-II-1941 (CNC); S58-5013-01-1, Ex

Enargia decolor, Vermilion Bay, #18 Ont., Em. 20-II-59, Inc., F. I. S.

(CNC); Ex Geometridae, Crow Bridge, Que., R. I. F. '40, 8961B, Em. 18/8,

F. I. S. Que '42 (CNC); Rec.: 1741 B, F. I. Survey 1940, Em. (Incubator),

19-Il] 1941, ex E. crepuscularia, St. Ambroise, Que. (CNC); Claslyn, Sask.,

Em. 22-2-54, F. I. S: W-X 1352 (02), Host Phalaenid sp. (CNC).

Hosts: Geometridae and Noctuidae (''Phalaenidae''): Aseptis binotata,

Ectropis crepuscularia, Enargia decolor, Enypia moilleti? [moillieti], Eulype

hastata, Iphimorpha pleonectusa, Itame hulstaria, Melanolophia imitata,

Nyctobia limitaria, Orthosia hibisci, Protoboarmia porcelaria.

The Basilicon Group

In this group, the ocelli are of normal size for the genus and the eye of

moderate size, distinctly extending beyond outline of head in plan view but not

prominent; temple rather convex, head elliptical (weakly convergent behind) or

buccate; gena from nearly parallel to orbit to somewhat convergent toward

mandible, occipital carina weakly to indistinctly angled by cheek; sculpture of

frons, stemmaticum, and vertex fine, moderately dense, the whole subpolished;

mesoscutum, scutellum, and mesopleurum rather finely, shallowly, moder-

ately densely punctate; notauli moderately sharp to obsolete, prescutum not or

little wider than parapsides; space enclosed by epomia short or of moderate

length (one-fifth to one-third the length from midline to outer edge of pronotum),

of moderate width or narrow; sternaulus rather short, usually not sharply

impressed; hindbasitarsus rather long and slender; propodeal fossa shallow:

uniformly in one species, in the other two almost flat cephalad, more distinct

caudad; petiolar area nearly flat, weakly concave, often with a very shallow

narrow medial ''gutter"’ bounded by weak straight parallel longitudinal rugulae;

postpetiole without sculpture. Species included are masoni, walleyi, and

basilicon.

10. Aphanistes masoni, new species

Female: Forewing 8.2--12.5 mm, body 14.4--20.2 mm; antenna 11.2--

12.25 mm (may not cover the true range of the material examined, since few

specimens have complete antennae), 42--43 segments, first flagellar segment

4,8--6.0 times as long as wide, combined length of second and third segments

0. 80--0.99 times first; head thickness-width ratio 1.625--1.797, head length-

width ratio 1.09--1.17, head thickness-camber ratio 2.6--3.9, occipital

camber 3.4--4.8, head-occiput width ratio 1.13--1.39, eye-temple ratio

1.43--2.05; mesoscutum index 1.18--1.275, head width 1.34--1.5 times

mesoscutum width, scutellum 1:0. 95--1.2:0.6--0.7; depth: width:length of

propodeum 1:0.97--1.09:1.02--1.125; hindbasitarsus 11.04--12.8 times as

long as thick, first tergite 1.02--1.18 times as long as hindbasitarsus; width

of postpetiole 0. 55--0.66 times length, first tergite 3. 41--4. 36 times as long

34 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

as postpetiole.

Habitus intermediate. Face rather finely and sparsely punctate, with

punctures in lines, giving effect of being rugulosopunctate, without any rugu-

lae being present except ina small number of specimens, in which longitudinal

wrinkles found in several other species occur; frons finely closely punctate,

forming rugulae toward and on stemmaticum; cheek rather finely weakly punc-

tate, impunctate near orbits. Lower part of epomia abruptly turned toward

collar, sometimes angled, enclosure short, less than 0.4 times length of

cephalic margin of pronotum from outer end to midline, narrow; length from

outer/lower end to midline of pronotum at least four times interior width; a

more or less distinct polished impunctate spot on sides of pronotum; mesoscu-

tum rather finely closely and deeply punctate; notauli distinct but not sharply

impressed; scutellum usually rather distinctly elevated, either convex or with

weak median longitudinal depression, appearing wider in proportion to its

length than indicated by measurements above, usually rather steeply descending

to the lateral carina; mesopleurum rather polished, with distinct small punc-

tures separated by about their diameter; sternaulus usually rather broad and

shallow, very rarely sharply impressed, less than two-fifths length of pleurum;

prepectal carina present only at extreme lower end. Propodeum rather

polished, rugae usually rather reduced; area dentipara clearly defined, nearly

triangular, little longer than wide, rather distinctly less than one-half length

of propodeum, usually with few, weak rugulae in interior and rather broad

shallow punctures, usually rather polished; areola not clearly defined, posteri-

or carina wholly lacking; propodeal fossa nearly wanting cephalad, rather

shallow caudad, pleural carina strong, lateral longitudinal carina indistinct;

petiolar area nearly flat, very weakly concave, sometimes with (weak) trans-

verse ridges continuous across its surface and often with narrow very shallow

longitudinal parallel-sided "'gutter"' from areola to insertion of petiole; metaka-

tepisternum rather strongly inflexed.

Fulvoferruginous or sometimes ferruginous. Antenna ferruginous to fus-

cous with some black and yellow on scape and pedicel. Head with the usual

distribution of yellow and black. Mesoscutum ferruginous, prescutum some-

times infuscate, in one specimen extensively black; scutellum usually yellow

or yellowish, occasionally fulvoferruginous; mesopleurum usually slightly

yellow-tinged, sometimes with yellow spots toward anterior and posterior ends

of line between pleurum and sternum; mesosternum usually more or less

black, as are usually spot beneath wings and always patch before scutellum.

Propodeum fulvoferruginous, with metapleurum more or less tinged with

yellow and sometimes with metakatepisternum or parts of the upper surface

black. Abdomen fulvous, more or less yellow-tinged. Wings hyaline to

subhyaline. Legs with usual coloration: all tarsi, fore- and midlegs almost

wholly, and base of hindtibia yellow; hindcoxa above and apex of hindtibia

black.

Male: Forewing 7.3--10.6 mm, body 14.5--18.8 mm; antennae incomplete

in all specimens, first flagellar segment 4.2--6.1 times as long as wide, com-

bined length of second and third segments 0.96--1.0 times first; head thickness-

width ratio 1.6--1.7, head length-width ratio 1.17--1.2, head thickness-

camber ratio 2.95--3.7, occipital camber 3.9--4.5, head-occiput width ratio

1.27--1.39, eye-temple ratio 1.36--2.075; mesoscutum index 1.2--1.35,

head width 1.46--1.51 times mesoscutum width, scutellum 1:1. 06--1. 26:0. 53--

0.66; depth: width:length of propodeum 1:1.0--1.15:1.05--1.15; hindbasitarsus

9.0--10. 53 times as long as thick, first tergite 1.0--1.19 times as long as

hindbasitarsus; width of postpetiole 0.52--0.604 times length, first tergite

Hopper: Nearctic Aphanistes 30

3.2--4. 35 times as long as postpetiole.

Sculpture as in female, but often with slightly coarser punctation of meso-

pleurum and rugulosity of propodeum. Color as in female, but often with

slightly more black; however two specimens have black restricted to stemmati-

cum and a very small spot before scutellum, hindtibia and second abdominal

tergite. These two also have more extensive yellow on sternum and pleurum;

in some other specimens yellowish coloration of metapleurum weak, but with

a small yellow spot always present above base of hindcoxa, and in a few speci-

mens there is a fair-sized yellow spot at about center of mesopleurum.

Material examined: 3722, 150°".

Type: 2, Highlands, N. C., 3-VI-1957, W. R. M. Mason, Horse Cove,

3000' (CNC); allotype: ©, Mt. Madison, N. H., June 23, 1938, H. &M.

Townes (CHT); paratypes: 2°, Waterton, Alta., Jul. 3, 1924, H. L. Seamans

(CNC); Hartford, Conn., 29-VI-1966, G. H. Heinrich (HPH); Flying, VI-15-18,

Lyme, Conn., WmMiddleton, Collector (USNM); Tray Mt., Ga., 6-19-36,

P. W. Fattig (USNM); Algonquin, Ill., Col. CF Baker (USNM); Dryden, Me.,

VII-1956, G. H. Heinrich (HPH); Magnolia, Mass. (Holland), Aug. 2, 1902,

Holland Collection (Pittsburgh); Huron Mts., Mich., June 27, 1961, H. &M.

Townes (22, CHT); Huron Mts., Mich., June 30, 1961, H. & M. Townes

(CHT); Midland Co., Mich., VI-21-38, R. R. Dreisbach (CNC); Ontonogan Co.,

Mich., 6-18-60, R. & K. Dreisbach (Dreisbach); Ithaca, N. Y., 23 May '33

(USNM); Ithaca, N. Y., 5 June '33 (USNM); Ludlowville, N. Y. (Pittsburgh);

Whiteface Mt., Adirondacks, N. Y., JM Aldrich Coll., VII-4, 3800 ft. (USNM);

Highlands, N. C., 21-V-1957, W. R. M. Mason (CNC); Highlands, N. C.,

26-V-1957, W. R. M. Mason (CNC); Highlands, N. C., 3800', 27-V-1957,

J. R. Vockeroth (CNC); Highlands, N. C., 3800', 8-VI-1957, J. R. Vockeroth

(CNC); Highlands, N. C., 1-VII-1957, J. R. Vockeroth (CNC); Highlands,

N. C., 14-VII-1957, C. J. Durden (CNC); Barrys Bay, Ont., 5-VII 1950,

T. N. Freeman (CNC); Ottawa, Ont., 12-VI-1946, G. S. Walley, Dow's

Swamp (CNC); Coraopolis, Pa., July 13, 1907 (Pittsburgh); Fairmount Pk.,

Philadelphia, Pa., 9-VII-1937, H. P. Hopper (HPH); Gleasonton, Pa.,

VII-20-38, G. E. Wallace (Pittsburgh); Spring Br. [=Brook], Pa., VI-8-1945,

H. K. Townes (CHT); Spring Br., Pa., VI-11-1945, H. K. Townes (2°, CHT);

Spring Br., Pa., VI-12-1945, H. K. Townes (CHT); Spring Br., Pa.,

VII-3-1945, H. K. Townes (CHT); Georgeville, Que. , 23-VI-1936, G. S.

Walley (CNC); Kazubazua, Que., 28-VI 1933, G. S. Walley (CNC); Cleveland,

S. C., VI-19-1961, G. F. Townes (CHT); Wattacoo, S. C., Pickens Co.,

VI-28-59, G. F. Townes (CHT); paratypes: “o, North Berwick, Southern

Maine, USA, June 1964, G. H. Heinrich (HPH); Aweme, Man., N. Criddle,

16-VI-1921, Ex no. 826-3 (CNC); Ann Arbor, Mich., May 23, 1962, H. &M.

Townes (CHT); Wayne Co., Mich., Detroit, VI-9-1936, A. W. Andrews

(UMI); Durham, N. H., Weed & Fiske (USNM); Ithaca, N. Y., VI-23-1935,

H. K. Townes (CHT); Medina, N. Y., 13 June 1952, LL Pechuman (CHT);

Great Smoky Mt. N. P., N. C., 28-V-1957, W. R. M. Mason, Indian Gap,

5200' (CNC); Crisp, Pa., C. M. Acc. 321., July 190 [sic!] (Pittsburgh);

Spring Br., Pa., VI-12-1945, H. K. Townes (CHT); Ingersoll, Ont., 25-VI-

1939, G. S. Walley (CNC); Macdiarmid, Ont., L. Nipigon, VII-10-21, N. K.

Bigelow (CNC); Beechgrove, P. Q., 7-VI-1955, R. Lambert (CNC); Stoneham,

P. Q., June 21, 1938, H. & M. Townes (CHT).

36 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

11. Aphanistes walleyi, new species

Female: Forewing 6.95--10.1 mm, body 12.1--19.2 mm; antenna 7. 6--

10.5 mm (lacking in largest specimen), 36--39 segments, first flagellar seg-

ment 4.77--5.87 times as long as wide, combined length of second and third

segments 0.918--0. 98 times first; head thickness-width ratio 1.6--1.7, head

length-width ratio 1.11--1.17, head thickness-camber ratio 3.45--4.1, occi-

pital camber 4.38--5.2, head-occiput width ratio 1.24--1.33, eye-temple

ratio 1.5--2.13; mesoscutum index 1.2--1.266, head width 1.35--1.53 times

mesoscutum width, scutellum 1:1. 06--1.25:0. 46--0.69; depth: width:length of

propodeum 1:1.03--1.125:1.11--1.2; hindbasitarsus 8.33--12.44 times as

long as thick, first tergite 1.23--1.42 times as long as hindbasitarsus; width

of postpetiole 0. 56--0. 69 times length, first tergite 3. 659--4.0625 times as

long as postpetiole.

Habitus somewhat slender. Clypeus and face moderately convex, head

profile very broadly elliptical; face moderately finely distinctly punctate with

some longitudinal wrinkles and with rather short, dense, conspicous perpen-

dicular pubescence (facial hairs of most other species are sparser, longer,

and inclined forward); frons and vertex punctate and rugulosoreticulate;

cheek moderately finely distinctly punctate, temple more deeply and coarsely.

Mesoscutum and mesopleurum rather finely fairly closely punctate (punctures

separated by one-half to one times their diameter); notauli rather indistinct,

especially posteriorly, represented by quite shallow longitudinal depressions

with slightly closer punctures than adjacent areas, becoming shallower and

with less contrast in punctation caudad, sometimes obsolete with slight change

of elevation between prescutum and parapsides anteriorly; scutellum with

rather distinct anterior declivity, slightly elevated, lateral carina evident but

somewhat indistinct, sometimes a shallow longitudinal depression present,

usually extending across disc; lower end of epomia abruptly turned toward

collar, remainder nearly parallel to collar, enclosure very short, about one-

fifth length of cephalic margin of pronotum, and less than one-half as wide as

long; sternaulus broad, shallow, not sharply impressed, not more than two-

fifths length of pleurum. Propodeum usually rather coarsely reticulate, area

dentipara (not separated from first lateral area) and areola (open behind) de-

fined, former irregularly trapezoidal, truncate caudad, about one and one-half

times as long as wide and about one-half length of propodeum, rugose; propodeal

fossa broad and shallow but distinct, more or less uniform throughout length of

propodeum, pleural carina usually indistinct or obsolete, lateral longitudinal

carina indistinct behind area dentipara; petiolar area nearly flat, very weakly

concave, often with narrow very shallow longitudinal parallel-sided "gutter"’

from areola to insertion of petiole. Hindfemur rather incrassate, hindtarsus

slender. Petiole often rather elevated before spiracles.

Color fulvoferruginous. Antenna brown with black on first flagellar seg-

ment above, scape and pedicel more or less yellow; face, clypeus, mandible

except tip, cheek, and orbital spot yellow; temple fulvoferruginous, sometimes

black above; tip of mandible, frons, vertex, and upper part of occiput black.

Thorax fulvoferruginous, with patch before scutellum, side of scutellum, spot

on mesopleurum adjacent to base of hindwing, mesosternum, side of metanotum,

metanepimerum, metakatepisternum, metasternum, and often extreme base

of propodeum black; area along boundary between pronotum and mesoscutum

usually yellow-tinged, disc of scutellum yellow-tinged or yellow, propodeal

fossa yellow-tinged to yellow. Legs yellow with fore- and midfemora tinged

Hopper: Nearctic Aphanistes 37

with ferruginous, hindfemur fulvoferruginous, hindcoxa more or less black as

are hindtrochanters, apical portion of hindtibia black. Abdomen fulvoferrugin-

ous, sometimes with some black at apex. Wings subhyaline to moderately in-

fuscate or rarely hyaline.

Male: Forewing 7.3--9.9 mm, body 14.6--20.5 mm; antenna 7.6--(more

than) 11.5 mm (smallest and largest specimens have antennae incomplete),

37--39 segments, first flagellar segment 4.4--5.5 times as long as wide,

combined length of second and third segments 1.024--1.088 times first; head

thickness-width ratio 1.57--1.72, head length-width ratio 1.08--1.17, head

thickness-camber ratio 3.9--4.3, occipital camber 5.16--5.5, head-occiput

width ratio 1.25--1.35, eye-temple ratio 1.4--2.1; mesoscutum index 1.19--

1.31, head width 1.295--1.475 times mesoscutum width, scutellum 1:0. 93--

1.27:0.45--0. 666; depth: width:length of propodeum 1:1.01--1.18:1.03--1. 33;

hindbasitarsus 5.48--7.94 times as long as thick, first tergite 1.297--1.55

times as long as hindbasitarsus; width of postpetiole 0. 56--0.75 times length,

first tergite 3.76--4.25 times as long as postpetiole.

Sculpture as in female. Color differing only in that a number of specimens

have no yellow on thorax and propodeum except disc of scutellum (in one speci-

men this is fulvoferruginous) and on the other hand several specimens have

rather extensive yellow-tinged blotches and/or yellow spots on thorax.

Material examined: 24922, 16°C.

Type: ¢ and allotype, “(on same pin): Kazubazua, Que., 21-VII-1927,

G. S. Walley (CNC); paratypes: 2°, FIS W Ex 2127(07), S. F. R. [= Sandilands

Forest Reserve], Man., Em. 9-4-57, Host Anavitrinella pampinaria (CNC);

Ann Arbor, Mich., May 28, 1962, H. & M. Townes (CHT); Ann Arbor, Mich.,

VI-11-61, Trap, H. & M. Townes (CHT); Grand Traverse Co., Mich. VII-9-

60, R., K. Dreisbach (Dreisbach); Ontonogan Co., Mich., 6-18-60, R. & K.

Dreisbach (Dreisbach); Yellow Dog Plains, Marquette Co., Mich., July 21,

1959, Henry Townes (CHT); Yellow Dog Plains, Marquette Co., Mich., July

28, 1959, Henry Townes (CHT); Yellow Dog Plains, Marquette Co., Mich.,

July 31, 1959, Henry Townes (3°, two CHT, 1 HPH); Aug. 3, 1902, on foliage

of Veratrum, Beulah, 8000 ft., New Mexico, Cockerell (USNM); Saranac Lake,

N. Y., 4-VIlII-'46, RTMitchell, Collector (RTM); Deerlock, Ont., 18-VII-1960,

S. M. Clark (CNC); Ft. Frances, Ont., 2-VIII-1960, S. M. Clark (CNC);

Galetta, Ont., 22-VII-1942, G. S. Walley (CNC); Ottawa, Ont., 17-VII-1946,

G. S. Walley, Dow's Swamp (CHC); Torrance, Ont., 16 June 1959, L. L.

Pechuman (CHT); Portage, P. E. I., 18-VII-1940, G. S. Walley (CNC);

Aylmer | probably Que.], 11-8-26, R. P. (CNC); Kazubazua, Que., 26-VI-

1933, G. S. Walley (CNC); Kazubazua, Que., 18-VII-1935, F. A. Urquhart

(CNC); Knowlton, Que., 16-VI-1936, G. S. Walley (CNC); Laurel Lake, nr.

Jacksonville, Vt., VIL or VIII-1936, Harry D. Pratt (USNM); paratypes: °C,

Ninette, Manitoba, 15-VII-1958, R. L. Hurley, Ex Betula glandulosa (CNC);

Crawford Co., Mich., VII-10-59, R. and K. Dreisbach (Dreisbach); Huron

Mts., Mich., July 25, 1959, Henry Townes, In heath bog (CHT); Ionia Co.,

Mich., 7-14-57, R.&K.Dreisbach (Dreisbach); Midland Co., Mich., 7-18-22-

09, R. R. Dreisbach (Dreisbach); Olmsted Co., Minn., CNAinslie, Collector

(USNM); Osceola Co., Mich., 7-7-47, R. R. Dreisbach (Dreisbach); Wayne

Co., Mich., Grosse Ile, July 24, 1961, Geo. Steyskal (CHT); Yellow Dog

Plains, Marquette Co., Mich., July 31, 1959, Henry Townes (CHT); Yellow

Dog Plains, Marquette Co., Mich., June 28, 1961, H. & M. Townes (CHT);

St John N B, July 21, 1902, A G Leavitt (USNM); Saranac Lake, N. Y., 12-VI-

'46, RTMitchell, Collector (2°, RTM); S. Milford, N. S., 25-VI-1934, J.

McDunnough (CNC); Ft. Frances, Ont., 2-VII-1960, S. M. Clark (CNC).

38 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

Hosts: Anavitrinella pampinaria (Geometridae).

12. Aphanistes basilicon (Davis)

Anomalon basilicon Davis, 1898 (1897). Amer. Ent. Soc., Trans. 24: 359.

Type: %, Moscow, Id. (Aldrich). Type in Academy of Natural Sciences

of Philadelphia, Philadelphia, Pa.

Female: Forewing 7.45--11.7 mm, body 12.0--19.4 mm; antenna (less

than) 9--(more than) 12.5 (smallest and largest specimens without or with

incomplete antennae), 36--44 segments, first flagellar segment 4. 59--5. 629

times as long as wide, combined length of second and third segments 0. 873--

1.047 times first; head thickness-width ratio 1.58--1.73, head length-width

ratio 1.106--1.163, head thickness-camber ratio 3.0--3.55, occipital camber

3.55--4.4, head-occiput width ratio 1.3--1.45, eye-temple ratio 1. 7--2. 4;

mesoscutum index 1.166--1.275, head width 1.38--1.54 times mesoscutum

width, scutellum 1:1.0--1.2:0.53--0.73; depth:width:length of propodeum

1:0.94--1.1:1.1--1.23; hindbasitarsus 10.5--15.0 times as long as thick,

first tergite 1.0--1.24 times as long as hindbasitarsus; width of postpetiole

0.59--0.73 times length, first tergite 3.8--4.3 times as long as postpetiole.

Habitus moderately slender. Head in plan view more or less elliptical

and when tilted to bring anterior edge of median ocellus to forward edge of

head, line across frons from forward edge of one eye to other appears nearly

straight. Punctures on clypeus rather few as usual (about 50 aside from those

at margins), but not more than twice diameter of punctures on face (in other

species three or more times), and those of this size few and near clypeal

tooth, others little larger than those on face; face moderately finely and closely

punctate, density decreasing only slightly toward orbits, those toward middle

tending to run together into longitudinal wrinkles; occipital carina by gena

nearly parallel to orbit except at temple or moderately convergent toward man-

dible, cheek with moderately sparse punctures (but somewhat denser than in

most species of the genus), increasing in density and depth onto temple; frons

and vertex mostly punctate-reticulate, becoming rugose-reticulate immediately

in front of lateral ocelli, occasionally more extensively rugulose. Mesoster-

num polished, largely impunctate; mesopleurum moderately densely punctate,

from subpolished to moderately polished, punctures fairly deep, separated by

less than their diameter; prepectus usually similar in sculpture to pleurum,

rarely with partial carina above sternum; epomia not parallel to collar, lower

part forming a smooth curve, enclosure fairly large, extending one-fourth to

one-third distance from midline to lateral margin of pronotum, width one-third

to one-half length; mesoscutum subopaque, Somewhat more finely and weakly

punctate than pleurum, rugulose in front of scutellum; notauli rather sharply

impressed; disc of scutellum usually rather flat, scarcely raised above lateral

carina, with a small anterior declivity, rarely slightly to moderately elevated,

sometimes with shallow longitudinal depression, subopaque, slightly more

deeply densely punctate than mesoscutum. Propodeum strongly rugose-

reticulate, not distinctly punctate; pleural carina strong, lateral longitudinal

carina distinct, and those portions of median longitudinal carina bounding area

dentipara (usually not separated from first lateral area) distinguishable; area

dentipara about one-half length of propodeum, one and one-half to two times as

long as wide, truncate caudad, reticulated, opaque; propodeal fossa shallow,

barely or not depressed anteriorly, moderately shallow posteriorly; petiolar

Hopper: Nearctic Aphanistes 39

area nearly flat, very weakly concave, often with very shallow narrow parallel-

sided longitudinal ''gutter'' medially from areola to insertion of petiole.

Petiole not elevated before spiracles.

Antenna fulvoferruginous to dark fuscous, usually fuscous; head yellow,

with tip of mandible and usually frons, stemmaticum, and vertex (except

orbital spot) and upper part of occiput black; clypeal tooth ferruginous to

piceous, temple usually ferruginous, rarely black (in one specimen, only tip

of mandible, small areas between lateral and median ocelli, and transverse

stripe on occiput black). Thorax in one specimen wholly fulvoferruginous,

otherwise usually with patch before scutellum and at least spot on side of

scutellum, sometimes also spot beneath forewing and more or less of base of

propodeum black; sterna and metakatepisternum at least with black markings;

disc of scutellum and collar usually yellowish. Fore- and midlegs yellow

except coxae and at least outer surface of femora and sometimes one side of

tibiae fulvous; hindleg with upper surface of coxa and trochanters black or

piceous, femur fulvoferruginous, base of tibia, and tarsus wholly, yellow;

apex of tibia black. Wings usually subhyaline, often hyaline, rarely slightly

infuscate. Abdomen with mesal band on second tergite black, remainder

fulvous or sometimes fulvoferruginous, sometimes with black blotches (that

may be artifacts). Occasionally metakatepisternum black or marked with

black, metanepimerum often black and rarely a black mark on sides of prono-

tum and on midline of cephalic margin of mesoscutum; in one case black mark

in petiolar area. Sometimes metakatepisternum extensively yellow and some-

times yellow spot on metakatepimerum at base of hindcoxa and on propodeal

pedicel.

Male: Forewing 7.5--10.25 mm, body 14.0--19.5 mm; antenna (lacking

in one, incomplete in remainder: more than 9 mm), (more than 28 segments),

first flagellar segment 4.3--5.5 times as long as wide, combined length of

second and third segments 0.95--1.125 times first; head thickness-width

ratio 1.59--1. 74, head length-width ratio 1.11--1.19, head thickness-camber

ratio 3.13--3.86, occipital camber 3.9--4.4, head-occiput width ratio 1.35--

1.47, eye-temple ratio 1. 77--2.33; mesoscutum index 1.175--1.25, head

width 1.4--1.55 times mesoscutum width, scutellum 1:1.0--1.06:0. 46--0. 6;

depth: width:length of propodeum 1:1.025--1.15:1.05--1.175; hindbasitarsus

9.26--10.4 times as long as thick, first tergite 1.065--1.277 times as long

as hindbasitarsus; width of postpetiole 0.52--0. 83 times length, first tergite

3.6--4. 45 times as long as postpetiole.

Sculpture and color generally as in female. Head usually thicker and eyes

distinctly larger (Some tendency in this direction throughout family). In most

(of only six) specimens, rather marked indentation between clypeus and face.

In one specimen, scutellum rather strongly elevated. Thorax in three speci-

mens with less black than usual in female, but one specimen has more (includ-

ing propodeum); in all, metakatepisternum either wholly black or with black

spot or at least lower margin black. Three specimens have yellow marks

behind prepectus and at base of mid- and hindcoxae; only two have hindcoxa

black above, three fuscous, and one ferruginous; all six have hindcoxa yellow

below (ferruginous in female).

Material examined: 6392, 60°",

Distribution: Maryland, Michigan, Newfoundland, New Hampshire, New

York, North Carolina, Ontario, Pennsylvania, Quebec, Saskatchewan, South

Carolina, Vermont, Virginia, West Virginia.

(No host records. )

40 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

The Fuscipes Group

In this group the ocelli are of normal size for the genus and the eye of

moderate size, distinctly extending beyond the outline of the head in plan view

but not prominent; temple rather convex, head elliptical to buccate, weakly

convergent behind; gena indistinctly to distinctly convergent toward mandible,

genal portion of occipital carina with slight distinct or indistinct change of

direction but not angled. Top of head rather smoothly moderately convex,

rather polished; face from rather flat to slightly to moderately prominent.

Notauli usually sharp, occasionally indistinct to obsolete; scutellum flat to

slightly elevated, distinctly longer than anterior width, disc flattish, usually

with distinct shallow longitudinal depression, lateral carina distinct; meso-

pleurum with rather fine, sharp, moderately dense to rather sparse punctures;

epomia enclosing small space, one-fourth or less length of pronotum, one-half

or less as wide as long, triangular with ramus or smoothly curved. Propodeum

rather polished, only weakly convex laterally; propodeal fossa sometimes mod-

erately impressed, usually shallow, sometimes obsolete, especially cephalad;

petiolar area essentially flat. Hindfemur slightly to distinctly incrassate.

Species included are dreisbachi, heinrichi, and fuscipes.

13. Aphanistes dreisbachi, new species

Female: Forewing 7. 74--11.13 mm, body 12.5--18.7 mm; antenna (less

than) 9.3--(more than) 11.4 mm (smallest and largest specimens have incom-

plete antennae), 36--43 segments (see foregoing), first flagellar segment

4.76--5.75 times as long as wide, combined length of second and third seg-

ments 1.0--1.1 times first; head thickness-width ratio 1.658--1.75, head

length-width ratio 1.1125--1.198, head thickness-camber ratio 3.2--5.0,

occipital camber 4.2--6.3, head-occiput width ratio 1.27--1.37, eye-temple

ratio 1. 7--2.3; mesoscutum index 1.18--1.27, head width 1. 43--1.54 times

mesoscutum width, scutellum 1:1. 1--1.42:0.53--0. 83; depth:width:length of

propodeum 1:0. 98--1.11:1.15--1.2; hindbasitarsus 10. 8--14.0 times as long

as thick, first tergite 1.057--1.235 times as long as hindbasitarsus; width of

postpetiole 0.57--0.63 times length, first tergite 3.5--4.1 times as long as

postpetiole.

Habitus moderately slender. Face slightly prominent, forming nearly

smooth line with clypeus in profile, with very shallow indistinct punctures

separated by more than their diameter and with a weak ridge from each end of

subantennal carina to upper edge of clypeus and a few longitudinal wrinkles

in the middle; frons, stemmaticum, and vertex rather flat or (chiefly small

specimens) very slightly convex, stemmaticum forming smooth line with rest,

not or very slightly raised above frons (in latter case only within ocelli); frons,

stemmaticum, and vertex distinctly rather closely rugulosopunctate, spaces

between punctures less than one-half diameter of punctures, rather polished;

ocellocular space and temple (especially former) more weakly and sparsely

punctate, temple abruptly less punctate and more polished than vertex; ocellocu-

lar line slightly longer than diameter of lateral ocellus, latter separated from

occipital carina by less than one-half its diameter. Mesoscutum and meso-

pleurum rather densely shallowly punctate, punctures separated by slightly

less than their diameter, appearing rugulosopunctate at certain angles of

illumination; pronotum rather extensively rugulose; epomia distinct, with more

Hopper: Nearctic Aphanistes 41

or less of an angle at outer posterior point, often with ramus, enclosure about

one third length of cephalic margin of pronotum, and with transverse (ladder-

like) rugulae within; notauli rather sharply impressed, width of prescutum at

widest point about equal to greatest width of one of parapsides, angle between

notauli ca. 159; scutellum with punctation similar to mesoscutum, punctures

in center of disc sometimes slightly larger than the rest, disc flat or with very

‘slight longitudinal depression, scutellum very slightly elevated anteriorly,

anterior carina obsolete, low, lateral carina distinct. Mesopleurum rather

polished to subpolished (in smaller specimens), punctation similar to meso-

scutum, area below wings with vertical rugulae anteriorly and horizontal rugu-

lae posteriorly; speculum with slightly finer punctures, but no more polished

than remainder except on the slight angle; prepectus rugulose; sternaulus

shortish, about one-third length of pleurum, not sharply incised. Propodeum

usually rather polished, sometimes subopaque, rather strongly rugose and

rugulose, usually with stronger (but sometimes irregular) rugae corresponding

to longitudinal carinae of areolation, usually anterior and lateral carinae of

areola well-defined, areola and petiolar area forming a continuous rather flat

surface; area dentipara fully bounded, usually not separated from first lateral

area, outer carina usually especially strong, in particular at posterior angle,

but not forming a tooth; propodeal fossa slightly to moderately impressed, often

with very weak sculpture, pleural carina indistinct or obsolete; metapleurum

only slightly convex, not bulging, metakatepisternum inflexed toward sternum.

Hindfemur usually somewhat incrassate. Petiole narrow throughout, widening

rather abruptly but only moderately to postpetiole, which is little or not at all

wider than deep.

Fulvoferruginous. Antenna fulvous. Temple and most of thorax ferrugi-

nous (rather red), propodeum more or less and abdomen entirely fulvoferru-

ginous. Face, mandible except tip, orbital spot, fore- and midlegs extensively,

tegula, hindtarsus, and base of hindtibia yellow; tip of mandible, uppermost

part of temple, vertex except orbital spot, stemmaticum, and upper part of

occiput black; also black are: prepectus, spot under wings, mesosternum,

patch before and side of scutellum, upper surface of hindcoxa, apical part of

hindtibia, usually side of metanotum, metanepimerum, metakatepisternum,

usually spots at base and apex of propodeum, usually most of prescutum, and

sometimes mesoscutum, mesopleurum, and propodeum more extensively.

Male: Forewing 7.3--9.8 mm, body 12. 8--17.0 mm; antenna 8.1--11.7

mm, 37--42 segments, first flagellar segment 4.0--5.4 times as long as

wide, combined length of second and third segments 1.02--1.07 times first;

head thickness-width ratio 1.59--1.7, head length-width ratio 1.13--1.2, head

thickness-camber ratio 3.55--4.79, occipital camber 4.5--6.1, head-occiput

width ratio 1.25--1.36, eye-temple ratio 1. 8--2.5; mesoscutum index 0. 9625--

1.2375, head width 1.225--1.5 times mesoscutum width, scutellum 1:0. 93--

1. 14:0. 56--0.64; depth: width:length of propodeum 1:1.01--1. 125:1.03--1. 2;

hindbasitarsus 5.0--8.57 times as long as thick, first tergite 1.326--1.65

times as long as hindbasitarsus; width of postpetiole 0. 58--0.66 times length,

first tergite 3.685--4. 58 times as long as postpetiole.

Structure and sculpture generally as in female. Ocellocular line distinctly

longer than diameter of lateral ocellus, latter about two-thirds its diameter

from occipital carina; head in profile sometimes showing a slight indentation

between face and clypeus. Color as in female, except one specimen (which

also has scutellum rising from anterior margin to one-third length) with thorax

and propodeum wholly black except ferruginous-edged yellow spot at center of

mesopleurum and sutural margins of mesopleurum fulvous, side and apex of

42 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

propodeum yellow edged with fulvous, first tergite fuscous; clasper piceous to

black in all specimens.

Material examined: 4899, 37¢°°.

Type: 2, Crabtree MdsNC, Yancey Co 3600', Aug 25 1950, HMD&J

Townes (CHT); allotype: °, Yellow Dog Plains, Marquette Co., Mich., June

28 1961, H. & M. Townes (CHT); paratypes: ¢2, Edmonton, Alta., 24 VII

1945, E. H. Strickland (CHT): Agassiz, B. C., 9-VI-1922, E. Glendenning

(CNC); Kleanza Creek, nr. Terrace, B. C., 12-VII-1955, G. P. Holland

(CNC); Chippewa Co., Mich., 6-25-60, R. and K. Dreisbach (Dreisbach);

Gogebic Co., Mich., 6-29-55, R. R. Dreisbach (3°, two Dreisbach, one

HPH); Houghton Co., Mich., 6-20-60, R. and K. Dreisbach (2°, Dreisbach);

Huron Mts., Mich., July 22 1959, Henry Townes (CHT); Manistee Co., Mich.,

7-5-57, R.&K. Dreisbach (Dreisbach); Ogemaw Co., Mich., 7-28-57, R. &K.

Dreisbach (Dreisbach); Roscommon Co., Mich., 7-7-48, R. R. Dreisbach

(UMI); Schoolcraft Co., Mich., 6-24-60, R. and K. Dreisbach (2¢, Dreisbach);

Mt. Madison, N. H., June 23 1938, H. & M. Townes (22, CHT); Mt. Madison,

N. H., June 26 1938, H. & M. Townes (22, CHT); Pinkham Notch, N. H.,

June 25, '38, H. & M. Townes (22, CHT); Randolph, N. H., July 2 1946,

J. Peck &M. Townes (CHT); Ithaca, N. Y., 6 VII-47 (CNC); Saranac Lake,

N. Y., 28-VI-'46, RTMitchell, Collector (RTM); Saranac Lake, N. Y.,

30-VI-'46, RTMitchell, Collector (RTM); Saranac Lake, N. Y., 2-VII-'46,

RTMitchell, Collector (RTM); NEW YORK, Ulster Co., Slide Mt., VII-11 1970,

Fred Hough (USNM); Hamrick NC, Aug 17 '50, HM&DTownes (CHT); Wayah

Bald, N. C., 4700', 29-VII-1957, W.R. Richards (CNC); Black Sturgeon Bay,

L. Nipigon, Ont., VII-21-22, N.K.B. (CNC); Deerlock, Ont., 18-VII-1960,

S. M. Clark (CNC); Macdiarmid, Ont., L. Nipigon, VII-10-21, NKBigelow

(CNC); Spring Brook, Pa., Aug 13, '44, H. K. Townes (2°, CHT); Spring Br

Pa, VI-27-1945, H K Townes (CHT); Spring Br Pa, VI-29-1945, H K Townes

(CHT); Spring Br Pa, VI-30-1945, H K Townes (CHT); Spring Br., Pa.,

VII-3-1945, H. K. Townes (CHT); Spring Br Pa, VIII-27-1945, H K Townes

(CHT); Alberton, P.E.I., 13-VII-1940, G.S.Walley (CNC); Georgeville, Que.,

27-VI-1936, G.S.Walley (CNC); Knowlton, Que., 2-VII-1936, G.S.Walley

(CNC); Nr. Cookshire, P.Q., VI-23-38, H. &M. Townes (2%, CHT); Ste.

Agathe des M., Que., 7-viii-1937, G. S. Walley (CNC); Stoneham, P.Q.,

June 18 1938, H. & M. Townes (CHT); Sutton, Que., 26-VII-1936, G.S.Walley

(CNC); paratypes: %°, Alger Co., Mich., VII-4-46, R.R. Dreisbach

(Dreisbach); Gladwin Co., Mich., 6-15-52, R.R.Dreisbach (Dreisbach);

Gogebic Co., Mich., 6-15-60, R. and K. Dreisbach (10°, nine Dreisbach,

one HPH); Houghton, Co., Mich., 6-20-60, R. and K. Dreisbach (3°,

Dreisbach); Midland Co., Mich., VI /12-21/61, R.R. Dreisbach (Dreisbach);

Ontonogan Co., Mich., 6-18-60, R. and K. Dreisbach (Dreisbach); School-

craft Co., Mich., 6-24-60, R. and K. Dreisbach (Dreisbach); Nerepis, NB,

11 Jul, AG Levitt, Collector (USNM); Mt. Madison, N.H., June 24 1938,

H. &M. Townes (CHT); Mt. Madison, N.H., June 26 1938, H. &M. Townes

(CHT); Montvale, N.J., 27-VI-70, RTMitchell, Collector (RTM); Connecticut

Hill, Tompkins Co., N.Y., 2000', 6-21-47 (CNC); Ithaca, N.Y., 11 VI 1947,

W. Mason (CNC); Saranac Lake, N. Y. 17-VI-'46, RTMitchell, Collector

(RTM); Saranac Lake, N. Y., 20-VI-'46. RTMitchell, Collector (RTM);

Saranac Lake, N. Y., 23-VI-’46, RTMitchell, Collector (RTM); Saranac Lake,

N. Y., 26-VI-'46, RTMitchell, Collector (RTM); Saranac Lake, N. Y.,

6-VII-'46, RTMitchell, Collector (RTM); Saranac Lake, N. Y., 2-VIII-'46,

RTMitchell, Collector (RTM); NEW YORK, Ulster Co., Overlook Mt., 7-16

1969, Fred Hough (USNM); Ex Geometridae, Gordon Lake, 36, Ont., No.

Hopper: Nearctic Aphanistes 43

S50-0092-03, Date: 2-VI-1950, F. I. S. 1950 (CNC); Spring Br., Pa.,

Vil-4-1945, H. K. Townes (CHT); Pa 2067, Collector, CFBaker (USNM);

Stoneham, P. Q., June 18 1938, H. & M. Townes (2%, CHT).

Hosts: Geometridae (only one record in the material).

14. Aphanistes heinrichi, new species

Female: Forewing 6.04--8.3 mm, body 10.2--14.6 mm; antenna 6.14--

8.5mm, 32--35 segments, first flagellar segment 4.48--5.9 times as long

as wide, combined length of second and third segments 0. 86--1.03 times first;

head thickness-width ratio 1.63--1.67, head length-width ratio 1.11-1.15,

head thickness-camber ratio 3.6--4.95, occipital camber 4.3--6.4, head-

occiput width ratio 1.215--1.36, eye-temple ratio 1.3--1.8; mesoscutum index

0.98 (? badly bent, perhaps should be 1.1)--1.2, head width 1.37--1.57 times

mesoscutum width, scutellum 1:1.0--1.1:0.6--0.9; depth: width:length of pro-

podeum 1:1.1--1.28:1.16--1.35; hindbasitarsus 8.5--9.7 times as long as

thick, first tergite 1.23--1.43 times as long as hindbasitarsus; width of

postpetiole 0.57--0. 606 times length, first tergite 3.5--3.96 times as long as

postpetiole.

Habitus moderately slender. Antenna slightly compressed, width about

0.8 depth. Face from rather flat to slightly prominent, median field not

closely punctate, punctures separated by one to one and one-half times their

diameter, inner orbit more sparsely punctate; cheek and temple sparsely

shallowly punctate; frons moderately deeply punctate, punctures separated by

about one-half to one times their diameter, usually becoming weakly to strongly

rugulose along forward edge of stemmaticum: cheek, temple, frons, stemma-

ticum, and vertex rather highly polished. Thorax usually rather highly

polished; pronotum rugulose along anterior margin only or sometimes all over;

epomia occasionally difficult to discern (although carina not obsolete), usually

with posteroinferior angle, sometimes with ramus; enclosure rather small,

especially in small specimens; prepectus indistinct with only an inferior carina;

sternaulus indistinct; mesoscutum and mesopleurum distinctly but not deeply

(neither densely nor sparsely) punctate, punctures sharp and separated by one

to one and one-half times their diameter, pleurum usually more or less rugu-

lose under wings--usually only a small area, but sometimes more extensive;

notauli indistinct, shallow, marked by an irregular line of punctures, some-

times almost wholly wanting (in which case anterior end marked by elevation

of prescutum above parapsides), or sometimes sharp but consisting only of a

line of punctures so close as to form furrows; scutellum slightly elevated with

distinct anterior declivity with its summit at about one-third length of scutellum,

sometimes disc has broad shallow longitudinal depression, lateral carina usu-

ally very strong, punctures on disc rather coarser and deeper than on mesoscu-

tum, quite dense, often lying in longitudinal furrows separated by rugulae.

Propodeum rather highly polished, almost impunctate, rather strongly

rugose, first lateral area smooth, not separated from area dentipara by

carina, second lateral area with a few rugulae; propodeal fossa distinct but

shallow and rather narrow; petiolar area not distinctly separated from areola

and third lateral area, broadly and very shallowly longitudinally depressed

(nearly flat) usually with an indistinct narrow longitudinal furrow in middle.

Hindfemur rather stout, hindtarsus usually distinctly compressed, especially

in small specimens (two or three of the largest specimens examined had hind-

tarsi not perceptibly compressed). Postpetiole noticeably extended before

44 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

spiracles, petiole usually slightly widened and elevated in posterior two-fifths

or more, no marked change of direction of outline between petiole and post-

petiole in plan view. Ovipositor rather deep in proportion to its length and

almost always with abrupt declivity in upper valve at distal end of nodus, the

tip relatively long. 3

Flagellum of antenna fulvoferruginous (dark brown in one specimen), first

segment tinged with piceous or black above, pedicel yellow and black, scape

yellow. Head yellow except tip of mandible, frons except orbital spot, stem-

maticum, vertex, and occiput above foramen black. Prescutum more or less

black or piceous, mesoscutum otherwise ferruginous or in a few cases wholly

black except extreme margins of parapsides; scutellum from wholly black to

only side of scutellum and extreme apex of disc (apical flange) black, remainder

ferruginous; mesopleurum black with fulvoferruginous area next to prepectus

and another by base of midcoxa. Dorsum of propodeum and metakatepisternum

black, remainder of metapleurum and pleural area fulvoferruginous. Fore-

and midlegs yellow with femora fulvous; hindcoxa dorsally black or ferruginous,

ventrally fulvous, hindfemur fulvoferruginous, hindtibia yellow at base, ex-

tensively black at apex, hindtarsus yellow. Wings usually subhyaline, some-

times hyaline or lightly infuscate. First tergite ferruginous (sometimes

infuscate), second tergite medially black as usual for genus, laterally ferru-

ginous (in one specimen entirely black); apex of abdomen more or less black.

One specimen has thorax and propodeum wholly black except along sutures and

spot behind base of hindcoxa.

Male: Forewing 5.9--8.2 mm, body (11.6?)--14.1 mm; antenna 6.57--

8.4mm, 34--37 segments, first flagellar segment 3.78--4.57 times as long

as wide, combined length of second and third segments 1. 02--1.14 times first;

head thickness-width ratio 1.54--1.63, head length-width ratio 1.06--1.2,

head thickness-camber ratio 4.0--5.2, occipital camber 4. 9--6.56, head-

occiput width ratio 1.2--1.35, eye-temple ratio 1.45--1.85; mesoscutum index

1.02--1.12, head width 1.29--1.49 times mesoscutum width, scutellum

1:0.95--1. 2:0. 5--0. 75; depth: width:length of propodeum 1:1.12--1.2:1.1--1. 22;

hindbasitarsus 5.6 (not compressed)--7/7.7 (compressed) times as long as

thick (of the five specimens measured--selected to give a size distribution--

all had compressed tarsi except the one with 5.6 ratio and one with ratio

7.33), first tergite 1.36--1.56 times as long as hindbasitarsus; width of post-

petiole 0.57--0.7 times length, first tergite 3.46--3.98 times as long as post-

petiole.

Structure and sculpture generally as in female, but frons, stemmaticum,

and vertex more finely shallowly punctate and polished; antenna barely com-

pressed except first flagellar segment; scutellum with longitudinal ridges and

furrows only in slightly less than one-half of specimens and then usually less

marked; the compression of the hindtarsus found in most specimens of this

species is unusual for males of this genus (in which the typical enlargement

in males relative to females is normally more prominent laterally).

Color generally as in female, but usually with greater prevalence of black,

especially on mesoscutum and propodeum, the latter usually wholly black

except third pleural area and tips of propodeal pedicel. First tergite usually

wholly fulvous (leaning toward fulvoferruginous), clasper black or piceous.

One specimen is much more extensively fulvoverruginous, with only a restricted

black stripe on prescutum, an upper anterior black patch on mesopleurum

running into a longitudinal posterior one, and with color of propodeum as in

female; another male has thorax wholly black except sutural margins and

propodeum also except at base of hindcoxa (notauli in this specimen almost

Hopper: Nearctic Aphanistes 45

imperceptible).

Material examined: 2699, 11¢°.

Type: ¢, Randolph, N. H., July 6 1946, J. Peck & M. Townes (CHT);

allotype: ©“, Ithaca, N. Y., V-31-1936, H. K. Townes (CHT); paratypes: $9,

Vancouver, B. C., 30-6-1914, R.H.Chrystal (CNC); Woolwich, Me., 1-VIII-

1956, G. H. Heinrich (HPH); Pine Ridge, Man., Em. 13-IV-48, F. I. S:

W-815, Ex Bapta vestaliata (CNC); Alcona Co., Mich., 7-3-48, R.R. Dreis-

bach (Dreisbach); Ann Arbor, Mich., May 18 1962, H. & M. Townes (CHT);

Benzie Co., Mich., 7-4-57, R.&K. Dreisbach (Dreisbach); Clare Co., Mich.,

7-23-28-59, R. R. Dreisbach (Dreisbach); Gogebic Co., M., 6-29-55,

R.R. Dreisbach (Dreisbach); Missaukee Co., Mich., 6-30-57, R.&K. Dreis-

bach (Dreisbach); Oscoda Co., Mich., 2-4-48, R.R.Dreisbach (Dreisbach);

Pinkham Notch, N. H., June 25 '38, H. & M. Townes (CHT); Cranberry Lake,

N. Y., 6-30-24, E.A. Hartley, coll. (USNM); Ithaca, N. Y., 17-VIII-47

(CNC); Saranac Lake, N. Y., 12-VI-'46, RTMitchell, Collector (RTM);

Saranac Lake, N. Y., 26-VI-'46, RTMitchell, Collector (RTM); Saranac Lake,

N. Y., 2-VII-'46, RTMitchell, Collector (RTM); Ottawa, Ont., Hog's Back,

17-VI-52, R.S. Bigelow (CNC); Simcoe, Ont., 26-VI-1939, T.N. Freeman

(CNC); Mary's Peak, Ore., Jul. 4 1904, JCBridwell, Collector (USNM);

MtHollySpgs, Pa, VII-7-1918, RMFouts, Collector (32, USNM); Great Smoky

Mt. N. P., N. C. Tenn., 18-VI-1957, W. R. M. Mason, Indian Gap, 5200’

(CNC); Gr.Smoky Mt. Nat.Pk., Tenn., 18-VI-1957, J.R.Vockeroth, Indian

Gap, 5200' (“ on same pin, CNC); Mason Co., Wash., Lake Cushman, VII-2

1919, F.M.Gaige, 34 (UMI); paratypes: °°, Birds Hill, Man., Em. 18-II-47,

F. I. S: W-617, Ex Melanolaphia [sic!] canadaria (CNC); Clare Co., M.,

V-23-59, R.Dreisbach (Dreisbach); Cranberry Lake, N. Y., 6-20-24,

E.A.Hartley, Coll. (USNM); Cranberry Lake, N.Y., VII.1.24, E.A.Hartley,

Coll. (USNM); Ithaca, N. Y., 6-VI-1947, W. Mason (CNC); Ottawa, Ont.,

11-VI-1947, G.S. Walley, Hog's Back (CNC); Berks Co., Pa., 5-VI-38,

S. C. Schell, Coll. (HPH); Knowlton, Que., 11-VI-1930, L.J.Milne (CNC);

Nominingue, Que., 5-VI-1941, G.S.Walley (CNC); Gr.Smoky Mt. Nat.Pk.,

Tenn., 18-VI-1957, J.R. Vockeroth, Indian Gap, 5200' (on pin with ¢, CNC).

Hosts: Bapta vestaliata, Melanolophia canadaria (Geometridae).

15. Aphanistes fuscipes, new species

Female: Forewing 4. 86--6.57 mm, body 8.57--11.9 mm; antenna (wanting

in smallest specimen) 6.4--7.7 mm, 32--35 segments, first flagellar segment

4.65--5.5 times as long as wide, combined length of second and third segments

1.02--1.17 times first; head thickness-width ratio 1.56--2.16, head lLength-

width ratio 1.18--1.2, head thickness-camber ratio 2.8--3.65, occipital

camber 3.5--4.4, head-occiput width ratio 1.27--1.4, eye-temple ratio 1. 88--

2.16; mesoscutum index 1.05--1.12, head width 1.48--1.6 times mesoscutum

width, scutellum 1:0.89--1.1:0.6--0.7; depth: width:length of propodeum 1:

1.04--1.14:1. 16--1.26; hindbasitarsus 8.25/9.43--9.22/9.76 times as long

as thick (all compressed: most extreme ratio of thickness to length 8.9/10. 4),

first tergite 1.35--1.53 times as long as hindbasitarsus; width of postpetiole

0.52--0.59 times length, first tergite 3.4--4.2 times as long as postpetiole.

Habitus moderately slender. Head subbuccate to slightly convergent behind

(broadly elliptical); face rather polished, but with numerous fine punctures;

cheek and temple very finely sparsely punctate; frons and vertex finely shal-

lowly punctate, punctures separated by own diameter or more, wholly without

46 Cont. Amer. Ent. Inst. , vol. 18, no. 2, 1981

rugulae or with punctures between rugulae or relatively closely rugulose and

subpolished, most frequently polished, top of head always slightly convex;

head profile very convex, usually with a rather distinct change of direction

between face and clypeus. Epomia distinct, form rounded; enclosure from

about size of tegula to about one-half that size; pronotum more or less

rugose; mesoscutum and mesopleurum with rather fine sharp rather sparse

punctures; notauli fairly sharply impressed, meeting before prescutellar

black patch (usually finely transversely rugulose) in distinct depression in

front of which prescutum often longitudinally rugulose, prescutum rather

strongly raised above parapsides; mesopleurum usually with more or less

rugulosity beneath wings; scutellum very slightly elevated with slight anterior

declivity and broad shallow longitudinal depression the width of disc. Propode-

um impunctate with only indistinct areolation; petiolar area and third lateral

area with distinct transverse rugae, sometimes transverse rugae dominate

propodeum--rugosity other than in areola and third lateral area often very

weak. Hindfemur incrassate; hindtarsus usually compressed (at least one

specimen--not included in measurements above--has no perceptible difference

between transverse and vertical dimensions). First tergite at least slightly

wider than deep throughout.

Antenna brown. Head yellow except tip of mandible, more or less of frons,

stemmaticum, vertex (except orbital spot), and upper part of occiput black.

Body generally fulvous except for the usual black patch before scutellum,

tip of hindtibia, and midline of second tergite. Boundary between pronotum

and mesoscutum often yellowish; prepectus and adjacent area of mesopleurum

yellow or yellowish, disc of scutellum sometimes yellowish; midline of pre-

scutum sometimes infuscate anteriorly. First tergite, hindfemur, and abdo-

men usually darker. Spot beneath forewing, base of propodeum, and hindcoxa

above sometimes infuscate or black; abdomen usually blotched with black

apically. All tarsi yellow except hindbasitarsus brown at base.

Male: Forewing 5.03--6.54 mm, body 9.5--12.14 mm; antenna 6.2--

7.9mm, 33--35 segments, first flagellar segment 4.1--5.4 times as long as

wide, combined length of second and third segments 1.01--1.2 times first;

head thickness-width ratio 1.55--1.73, head length-width ratio 1.11-1.19,

head thickness-camber ratio 2.46--3.63, occipital camber 3.3--4.5, head-

occiput width ratio 1.24--1.34, eye-temple ratio 1.75--2.46; mesoscutum

index 1.08--1.16, head width 1.43--1.54 times mesoscutum width, scutellum

1:0.9--1.0:0. 5--0.65; depth: width:length of propodeum 1:1.08--1.175:1.06--

1.2; hindbasitarsus 6.6/7.2 (compressed)--8.5 (compressed in basal fifth,

depressed at apex) times as long as thick, first tergite 1.46--1.58 times as

long as hindbasitarsus; width of postpetiole 0.479--0.58 times length, first

tergite 3.23--3.935 times as long as postpetiole.

Structure and sculpture generally as in female except frons and vertex

usually rugulosopunctate or sometimes with only a few short rugulae. Notauli

especially deep, often with large punctures at bottom producing a ladderlike

effect. Propodeum more distinctly rugose, somewhat areolated.

Color generally as in female, but usually with somewhat more black and

yellow: head often more extensively black, mesoscutum almost always with

black on prescutum, sometimes wholly black, black area beneath wings, some-

times blotches on mesopleurum, at least base of propodeum black, sometimes

wholly black except pleural area; hindcoxa and first tergite more or less

infuscate or black; seventh and eighth tergites and clasper and usually also

sixth tergite black; disc of scutellum usually yellow or yellowish, rarely with

only slight yellow tinge; pleural area of propodeum yellow or sometimes only

Hopper: Nearctic Aphanistes 47

yellowish.

Material examined: 2199, 240°.

Type: 9, York Co. near Highspire, Pa., 7-18-’09, WSFisher, Colr

(USNM); allotype: “, MARYLAND, Glen Echo, R. M. Fouts (USNM); para-

types: °°, Cabin John, Md., June 26, 191(4?), RMFouts, Collector (USNM);

Glen Echo, Md, July 23 1921, J.R.Malloch, Collector (22, one USNM, one

HPH); MARYLAND, Glen Echo, R. M. Fouts (USNM); Takoma Pk., Md.,

VI-23, 1943, H. & M. Townes (CHT); Takoma Pk., Md., VI-27, 1943, H. &

M. Townes (CHT); Takoma Pk., Md., VII-1, 1943, H. & M. Townes (CHT);

Takoma Pk., Md., VIII-9, 1943, H. & M. Townes (CHT); Takoma Pk., Md.,

VIll-10, 1943, H. & M. Townes (22, CHT); Takoma Pk., Md., VIII-12, 1943,

H. & M. Townes (CHT); Chesilhurst, N. J., June 25, 1939, H. K. Townes

(CHT); Riverton, NJ, 7-12-08, GMGreene, Collector (USNM); Rossville,

Staten Isl NY, July 18, 1925, FM Schott (USNM); Wake Co.NC, VI-10 1951,

H&M Townes (% on same pin, CHT); Wake Co.NC, VI-20 1951, H&M Townes

(CHT); DunnLoring, Va., VIII-28 1948, Karl V. Krombein (CHT); DunnLoring

Va., VI-26 1949, K. V. Krombein (CHT); DunnLoringVa., VIII-21 1949, K. V.

Krombein (CHT); Dunn Loring Va., VII-15-51, K. V. Krombein (CHT); para-

types: °%, F66 (USNM); Beltsville, Md, 6-23-18, on Quercus alba, WLMcAtee,

Collector (USNM); MARYLAND, Glen Echo, R. M. Fouts (40° including allo-

type, three USNM, one HPH); Glen Echo, Md., June 11 1922, JRMalloch,

Collector (3%, USNM); Glen Echo, Md., June 18 1922, JRMalloch, Collector

(USNM); Glen Echo, Md., June 25 1922, JRMalloch, Collector (USNM); Glen

Echo, Md., July 2 1922, JRMalloch, Collector (USNM); Takoma Pk., Md.,

Vil-1, 1943, H. & M. Townes (CHT); Takoma Pk., Md., VIII-3, 1943, H. &

M. Townes (2%, CHT); Takoma Pk., Md., VIII-9, 1943, H. & M. Townes

(CHT); Takoma Pk., Md., VIII-12, 1943, H. & M. Townes (3°, CHT); Wake

Co. NC, VI-10 1951, H&M Townes (on same pin with , CHT); Greenville, SC,

VI-14 1952, G.&L. Townes (CHT); Wattacoo, Pickens Co., 8.C., VI-21-59,

G. F. Townes (CHT); Arlington, Va., VII-5 1948, KVKrombein (CHT); Dunn

Loring, Va., VIII-28 1948, Karl V. Krombein (CHT).

The Puparum Group

In this group, the ocelli are of normal size for the genus, eye of moderate

size but projecting very Little from the head outline in plan view; head moder-

ately to strongly buccate, curve of occiput relatively flat, temple usually form-

ing rather sharp, sometimes very sharp angle with occiput; face and clypeus

in profile form nearly straight line; frons, stemmaticum, vertex, and temple

form continuous nearly unbroken surface except for ocelli, sometimes indis-

tinct ridges (composed of rugulae) between lateral ocelli and median ocellus,

and unusually small antennal scrobes; epomia often somewhat indistinct,

usually consisting of multiple rugulae rather than a distinct single carina,

enclosure slightly larger than tegula, more than one-half as wide as long;

mesopleurum rather coarsely and densely punctate in contrast to mesoscutum

rather finely and sparsely punctate, except in Texas specimen, where rela-

tionship reversed (most other species have sculpture of mesopleurum and

mesoscutum similar); scutellum sometimes slightly wider than long, but con-

vergent caudad, moderately elevated, lateral carina obsolete; notauli very

indistinct; propodeum rather polished with weak rugulosity, median longitu-

dinal carina obsolete except at areola. Contains only the species puparum.

48 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

16. Aphanistes puparum (Ashmead)

Agrypon puparum Ashmead, 1890 (1889). U. S. Natl. Mus., Proc. 12: 424.

Type: 2, Alameda, California, reared from unknown noctuid (in U. S.

National Museum, Natural History, Washington, D. C.).

Female: Forewing 6.6--9.6 (10) mm (figures in parentheses refer to

Texas specimen), body 10.3--15.2 (18.4) mm; antenna 6.4--8.2 mm (only ©

three specimens with complete antennae), 33--36 segments, first flagellar

segment 4.0--4.42 times as long as wide, combined length of second and

third segments 0.92--0.98 times first; head thickness-width ratio 1.58--1.8,

head length-width ratio 1.12--1.16, head thickness-camber ratio 3.1--4. 5,

occipital camber (4.6) 5.25--6.14, head-occiput width ratio 1.14--1. 166

(1.28), eye-temple ratio (1.2) 1.3--1.85; mesoscutum index 1.1375--1.21

(1.26), width of head (1.304) 1.39--1.45 times mesoscutum width, scutellum

1:0.9--1.1:0. 6--0.7; depth: width:length of propodeum 1:1.0--1.2:1.02--1.15;

hindbasitarsus 6.5--8.19 (8.33) times as long as thick, first tergite 1.16--

1.33 times as long as hindbasitarsus; width of postpetiole 0.61--0.65 (0. 76)

times length, first tergite 3.4--4.12 (4.19) times as long as postpetiole.

Habitus somewhat stout. Head moderately to strongly buccate, curve of

occiput relatively flat, eye projecting little from head outline in plan view;

temple weakly to moderately convex, usually forming rather sharp, some-

times very sharp angle with occiput; frons, stemmaticum, vertex, and temple

forming continuous nearly unbroken surface except for ocelli and sometimes

indistinct ridges (composed of rugulae) between lateral ocelli and median

ocellus and unusually small antennal scrobes; face and clypeus in profile

forming nearly straight line; face with rather dense, rather shallow punctures,

rather polished, frons rugulosoreticulate (i.e., with rather large shallow

punctures separated by much less than their diameter, leaving only rugulae

between), usually forming rugulae or rugae along forward edge of stemmati-

cum; vertex and gena moderately closely rather shallowly and finely punctate,

punctures separated by more than their diameter, polished, gena with orbital

band one-fifth to one-half its width with considerably reduced punctation (fine,

shallow, sparse). Pronotum and mesoscutum rather finely distinctly sparsely

punctate (punctures separated by one to two times their diameter) and sub-

polished; mesopleurum and scutellum polished, rather more closely and deeply

punctate than mesoscutum, especially in larger specimens with predominantly

brown color (except Texas specimen, in which punctures of both mesopleurum

and mesoscutum are of about same size and depth, but very dense on the latter

and somewhat sparse on the former, mesoscutum subopaque, mesopleurum

rather polished--specimens from all localities agree in having a difference in

density of punctation between dorsum and pleurum in contrast to the relative

identity in most other species); epomia often somewhat indistinct and usually

consisting of multiple rugulae rather than a distinct single carina (in pre-

dominantly black specimens, epomia may be subobsolete and adjacent edge of

collar nearly absent), often lower end more or less distinct (epomia most

complete and most nearly a single carina in Texas specimen), enclosure

slightly larger than tegula, more than one-half as wide as long; notauli rather

broad and very shallow and weakly indicated, chiefly represented by change of

direction from parapsides to moderately to considerably elevated prescutum,

especially in black specimens: faintly impressed in brown specimens; scu-

tellum moderately elevated, without distinct lateral carina, with a distinct

Hopper: Nearctic Aphanistes 49

anterior slope, sometimes slightly wider than long but distinctly convergent

caudad; prepectus without pleural carina, not different in sculpture from

pleurum; some rugulae under wings. Propodeum usually with weak to obsolete,

rarely with moderate rugulosity (least in predominantly black, most in pre-

dominantly fulvoferruginous specimens), at most pleural and lateral longi-

tudinal carinae and the portion of median longitudinal carina at areola distinct,

first two sometimes strong, sometimes only the anterior portions of these

present, areola obsolete; in all specimens propodeum is subpolished (slightly

finely weakly reticulate in one or two fulvoferruginous specimens), dorsal

surface rather smooth and rounded and posterior portion of petiolar area with

transverse rugulae. Hindfemur somewhat stout. Postpetiole rather deep

(0. 81--0.87 times width), often widest at apex; petiole more or less elevated

for some distance cephalad of spiracles. .

Antenna brownish or fulvous. Clypeus, face, orbital spot, lower surface

of scape, mandible except tip, and cheek yellow; temple fulvous with black

margin adjacent to occiput or wholly black; frons, stemmaticum, and vertex

black (except Texas specimen, in which rather broad yellow band entirely

encircles eye, only middle of frons «nd stemmaticum black); upper part of

occiput and tip of mandible black. ‘fhorax fulvoferruginous, with more or less

of propleurum and pronotum, patch before and side of scutellum, spot beneath

wings, sterna, band across base of propodeum, and metakatepisternum black;

the black may be more extensive to the point of the entire thorax being black

except parapsides, spot at apex of scutellum, metakatepimerum, propodeal

pedicel, and some small spots on mesopleurum. Fore- and midlegs yellow

with more or less fulvous on tibiae and femora, hindleg fulvous except upper

surface of coxa usually black, upper surface of femur usually ferruginous,

apex of tibia usually black (in a few specimens merely fuscous), and tarsus

yellow (no yellow on tibia!). Wings sometimes hyaline, usually subhyaline,

sometimes slightly infuscate. Abdomen fulvoferruginous, mesal portion of

second tergite and upper surface of apical tergites black, first tergite usually

ferruginous, sometimes infuscate to black above (according to original descrip-

tion, postpetiole and second and following tergites above black in type--this

cannot be checked, since the type specimen now lacks abdomen).

Male: Forewing 7.0--9.6 mm, body 12.3--17.0 mm; antenna (only two:

one apparently broken after I measured it) 7.4--8.5 mm, 34--36 segments,

first flagellar segment 3. 94--4.375 times as long as wide, combined length

of second and third segments 0. 87--0.97 times first; head thickness-width

ratio 1.54--1.71, head length-width ratio 1.16--1.18, head thickness-camber

ratio 3.3--4.1, occipital camber 4.6--5.66, head-occiput width ratio 1.16--

1.22, eye-temple ratio 1.2--1.95; mesoscutum index 1.11--1.24, head width

1.4--1.49 times mesoscutum width, scutellum 1:1.0--1.1:0. 53--0. 625; depth:

width:length of propodeum 1:1.02--1.24:1.08--1.24; hindbasitarsus 4. 66--

7.69 times as long as thick, first tergite 1.136--1.4 times as long as hind-

basitarsus; width of postpetiole 0.55 (?--damaged)--0.68 times length, first

tergite 1.41--1.46 times as long as postpetiole.

Habitus generally as in female, but head in profile rather round, the line

of face and clypeus slightly convex but with a very inconspicuous separation.

Sculpture generally as in female, but rugulosity beneath wings usually more

distinct and more extensive. Color in general as in female but with greater

amount of black: thorax may be wholly black except for metanepisternum,

more or less of metakatepimerum immediately before hindcoxa, and pro-

podeal pedicel. First tergite tends to be ferruginous or fuscous and its dorsum

may be piceous to black, especially upper surface of postpetiole; apical ter-

20 Cont. Amer. Ent.Inst., vol. 18, now 2, 1981

gites more extensively black than in female; clasper black.

Material examined: 1522, 120°.

Distribution: middle California from north of Sacramento to north of Los

Angeles; northeastern Texas (near Fort Worth).

Hosts: "unknown noctuid" (type).

The Crassulus Group

In this group, the ocelli are of normal size for the genus, eye of moderate

size and distinctly projecting from outline of head in plan view, head subbuc-

cate to buccate, face in profile slightly to moderately prominent; space

enclosed by epomia moderately to very small, narrow; pronotum narrow

(width about one-third length), mesoscutum rather wide in proportion to its

length (except Florida specimen), slightly to moderately gibbous; notauli

shallowly narrowly distinct to obsolete; scutellum appears square or wider

than long, little narrowed behind; propodeum with median longitudinal carina

distinct and other traces of areolation, propodeal fossa very shallow to wanting;

thorax and hindfemur and hindtarsus stout; postpetiole rather long in propor-

tion to petiole, usually convergent caudad from spiracles. Species included

are crassulus, flaviscutellaris, and decurvihastatus.

17. Aphanistes crassulus, new species

Female: (Data in parentheses refer to specimen from Florida.) Forewing

5. 46-8. 4 (10.4) mm, body 8.5--14.65 (18.6) mm; antenna 5.7--8.8 (ca. 10)

mm, 29 (estimated)--40 (more than 40) segments, first flagellar segment

(4.0) 4.1--4.5 times as long as wide, combined length of second and third

segments (0.825) 0. 86--1.04 times first; head thickness-width ratio 1.6--

1.74, head length-width ratio (1.11) 1.12--1.15, head thickness-camber ratio

3.4--5.0, occipital camber (4.05) 4.45--6.7, head-occiput width ratio 1.2--

1.26 (1.38), eye-temple ratio (1.4) 1.5--1.75; mesoscutum index 1.05--1.06

(1.22), head width (1.31) 1.37--1.56 times mesoscutum width, scutellum

1:0. 7--0. 825 (0. 95):0.5--0. 7; depth: width:length of propodeum 1:(1.1) 1.12--

1.18:0.96--1.16, hindbasitarsus 5.6--7.9 times as long as thick, first tergite

(1.28) 1.42--1.5 times as long as hindbasitarsus; width of postpetiole 0. 54--

0.655 times length, first tergite 3.4--3.6 times as long as postpetiole.

Habitus rather stout. Head rounded rectangular, moderately transverse;

upper third of clypeus moderately closely punctate (more than one-half as

many punctures per unit area as on face--rare in our species of genus) except

in a very few specimens (chiefly the smallest); face much wrinkled and punctate,

slightly prominent in profile, inner orbit moderately punctate with only a nar-

row band of reduced punctation next to orbit; cheek rather densely punctate

with a decrease in density just below middle and an increase in density and

depth (but punctures small throughout) toward temple; frons, stemmaticum,

and vertex moderately densely and deeply punctate, usually with distinct but

not strong rugulae on frons, ocellocular space essentially impunctate (in

small specimens, punctation of cheek, temple, and stemmaticum sparser and

weaker); mandible rather strongly punctate. Thorax generally rather densely

and deeply punctate, including prepectus, punctures on mesopleurum larger

(separated by about one-third their diameter) than those on mesoscutum

(separated by about one-half their diameter); notauli moderately distinct,

Hopper: Nearctic Aphanistes ol

narrow but not sharply impressed, extending at least halfway to scutellum,

prescutum more or less distinctly elevated above parapsides; scutellum rather

strongly rather densely punctate, punctures separated by less than one-half

their diameter, scutellum distinctly elevated anteriorly, with rounded trans-

verse angle between steep anterior surface and sloping disc with distinct

shallow median longitudinal depression, usually extending from side to Side,

sometimes reduced to short longitudinal depression in midline near base,

lateral carina obsolete to wanting; epomia distinct, forming smooth curve,

usually completely closed, enclosure rather small, one-fourth or less length

of anterior margin of pronotum, roughly one-half as wide as long; mesopleurum

with some rugulae or reticulations beneath wings, usually extending back onto

upper part of speculum, lower half of speculum polished impunctate; prepectal

carina present on at least lower third of pleural margin; sternaulus short,

weak to obsolete (in small specimens more sharply--sometimes sharply--

impressed; polished area on speculum sometimes nearly absent). Propodeum

strongly rather coarsely rugose-reticulate, with rather distinct but small and

shallow punctures between the rugae (reticulations about one-eight length of

propodeum); area dentipara and areola very little more strongly punctate than

remainder, areola usually not closed behind; petiolar area moderately

depressed longitudinally, with narrow parallel-sided ''gutter,'' slightly further

depressed, from areola to base of pedicel; first and second lateral areas usually

not separated, combined area dentipara about as wide as long; anterior portions

of lateral longitudinal carina and pleural carina strong, median longitudinal

carina rather distinct. First tergite in plan view widening to spiracles, then

narrowing to apex; postpetiole long and elevated, elevation extending cephalad

of spiracles, total elevated portion more than one-half length of tergite, upper

surface of tergite more or less flattened, petiole usually wider than deep, with

a median longitudinal rounded groove that extends onto postpetiole, where

there usually are two short similar grooves between it and spiracles. Hindleg

very stout. Ovipositor much deeper than wide.

Antenna dark fulvous to light fuscous. Mandible except black tip, clypeus,

face including space between antenna and eye margin and extending short dis-

tance onto frons, orbital spot, and lower part of cheek yellow; frons, stemma-

ticum, and upper part of occiput black; upper part of cheek, temple, and vertex

fulvoferruginous (vertex and temple often piceous or black). Thorax fulvofer-

ruginous, with patch on mesoscutum before scutellum, side of scutellum, and

mesosternum black, scutellum very rarely slightly yellowish at base; black on

sternum may extend up onto lower part of mesopleurum; usually a black spot

beneath wings; metanepimerum, metasternum, metakatepisternum, basal area,

areola, area dentipara, and petiolar area may also be black. Tip of hind-

tibia black; hindcoxa and -trochanters may be more or less black above; fore-

and midlegs yellow with more or less fulvous on tibiae, femora, and coxae;

base of hindtibia more or less yellow or yellowish, hindtarsus yellow. Abdo-

men fulvoferruginous with broad midline of second tergite black; sometimes

first tergite infuscate or with some piceous, apical tergites sometimes black.

Specimen from Florida differs from the others in that only the ridges be-

tween lateral ocelli and median ocellus, spots on mesosternum before coxae,

and midline of second tergite are black; tip of mandible, small patch before

scutellum, and mesal line of mesosternum piceous; apex of hindtibia fuscous,

propodeal pedicel yellow, head wholly yellow except as just indicated and about

middle half of frons and upper part of occiput fulvoferruginous.

Male: Forewing 5.6--7.6 mm, body 9.5--13.4 mm; (no complete antennae),

first flagellar segment 4.35--4.66 times as long as wide, combined length of

o2 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

second and third segments 0. 87--0. 94 times first; head thickness-width ratio

1.62--1.75, head length-width ratio 1.14--1.24, head thickness-camber ratio

3.4--4.8, occipital camber 4.5--6.3, head-occiput width ratio 1.24--1. 34,

eye-temple ratio 1.575--1.9; mesoscutum index 1.10--1.12, head width 1.36--

1.44 times mesoscutum width, scutellum 1:0. 90--0. 93:0. 53--0.6; depth: width:

length of propodeum 1:1.14--1.2:1.04--1.06; hindbasitarsus 5.4--6.3 times as

long as thick, first tergite 1.44--1.64 times as long as hindbasitarsus; width

of postpetiole 0.55--0.66 times length, first tergite 3.45--4. 24 times as long

as postpetiole.

Habitus and sculpture in general as in female, but wrinkles on face and

rugulae or reticulations on frons slight or absent, gena including temple with

much weaker punctation, middle portion often almost impunctate. Epomia

closed, enclosure very narrow (less than one-third as wide as long); speculum

sometimes almost without polished area; mesopleurum often without rugulae or

reticulations under wings. Cheek and temple usually wholly yellow or yellowish-

fulvous; mesoscutum sometimes with a thin piceous-to-black median longitu-

dinal line; mesopleurum usually with narrow oblique black line tilting backward

toward top of speculum and sometimes with large roundish spot under fore-

wings. Metasternum black (rare in female).

Material examined: 7292, 70°.

Type: °, Kingston, R.I., VII-18 1937, M Chapman (CHT); allotype: ¢,

Millwood, N. Y., Jun. 20 1936, H. K. Townes (CHT); paratypes: 292, 2 mi.

W of Archer, Fla., Mar. 25 1953, H. F. Howden (CHT); Club Hill, Md.,

7/14/32 (CHT); Beltsville Md., VII-21 '47, R. Boettcher (USNM); Plummerlsl.

Md, 4-VII '21 (USNM); Takoma Pk., Md., VII-18, 1942, H. & M. Townes

(CHT); Takoma Pk., Md., VI-11, 1944, H. & M. Townes (CHT); Ex leaf-

roller, Lynnfield, Mass 2-6-22, GipMothLab, 12164G68 (USNM); Ontonogan

Co., Mich., 6-18-60, R. and K. Dreisbach (Dreisbach); Wayne Co., Mich.,

Grosse Ile, July 15, 1961, Geo. Steyskal (CHT); Moorestown, N. J., June 11,

1939, H. & M. Townes (CHT); Moorestown, N. J., June 14, 1939, H. &M.

Townes (CHT); Moorestown, N. J., June 22, 1939, H. & M. Townes (CHT);

Moorestown, N. J., June 25, 1939, H. & M. Townes (CHT); Moorestown,

N. J., June 27, 1939, H. & M. Townes (CHT); Moorestown, N. J., July 1,

1939, H. & M. Townes (2°, CHT); Moorestown, N. J., July 4, 1939, H. &

M. Townes (CHT); Moorestown, N. J., July 5, 1939, H. & M. Townes (CHT);

Moorestown, N. J., July 6, 1939, H. & M. Townes (CHT); Moorestown, N. J.,

July 9, 1939, H. & M. Townes (CHT); Farmingdale, N. Y., Jul 26 '38, H. &

M. Townes (CHT); Millwood, N. Y., Jun. 27 1936, H. K. Townes (CHT);

Yonkers, N. Y., 4 July 1935, L. L. Pechuman (CHT); Clinton, N. C., May

24 1951, H&M Townes (22, CHT); Poplar, N. C., 15-VII-1936, D.L. Wray

(USNM); Deerlock, Ont., 18-VII-1960, S. M. Clark (CNC); Coraopolis, Pa.,

July 13, 1907 (3°, Pittsburgh); Crisp, Pa., C.M.Acc.321, July 190 |[sic!]

(Pittsburgh); All'y Co., Penna., Kenneywood, July 9, 1907 (2°, Pittsburgh);

MtHollySpgs, Pa., VII-7-1918, R.M. Fouts, Collector (USNM); F31Pa, Mt.

Holly Spgs., Pa. 8-25-20, R.M. Fouts, Coll. (USNM); North East, Pa., 1914,

RACushman, Collector (USNM); Pittsburg [sic!], Pa, June (2¢, Pittsburgh);

Rockville, Pa., 14-VII-10, P. R. Myers (USNM); Westmoreland Co., Pa.,

July (19°, Pittsburgh); Westmor.Co., Pa., July 190 [sic!] (32, Pittsburgh);

Wissahickon Val., Philadelphia, Pa., 26-VI-1935, H. P. Hopper (HPH);

Westerly, R. I., VII-5 1936, M. Chapman (2°, CHT); Westerly, R. I., VI-

6-1936, M. Chapman (CHT); Cleveland, S. C., V-7-1961, G. F. Townes

(CHT); Va near PlummerslIsl, 3-VII '21 (2°, USNM); Lost River St. Pk.,

Hardy Co., W.Va., VII-5 1953, Karl V. Krombein (22, CHT); Polk Co., Wis.,

Hopper: Nearctic Aphanistes o3

July, Baker, Collector CFBaker (USNM); paratypes: °°, Plummersid, Md,

VI-24-17, GMGreene, Collector (USNM); Ontonogan Co., Mich., 6-18-60,

R. and K. Dreisbach (Dreisbach); All'y Co., Penna., Kenneywood, July 9,

1907 (3°, Pittsburgh). }

Hosts: "leaf-roller."

18. Aphanistes flaviscutellaris, new species

Female: Forewing 5.45--6.65 mm, body 8.26--11.06 mm; antenna 5. 75--

6.37 mm (only two complete), 31--32 segments, first flagellar segment 3. 8--

4.4 times as long as wide, combined length of second and third segments 1.0--

1.2 times first; head thickness-width ratio 1.54--1.84, head length-width ratio

1.15--1.24, head thickness-camber ratio 3.75--5.54, occipital camber 4. 86--

6.86, head-occiput width ratio 1.27--1.37, eye-temple ratio 1. 56--2.2875;

mesoscutum index 0.96--1.02, head width 1.406--1.537 times mesoscutum

width, scutellum 1:0.94--1.1:0.55-0.9; depth: width:length of propodeum

1:1.1--1.14:0.96--1.0; hindbasitarsus 5.1--7.1 times as long as thick, first

tergite 1.6--1.7 times as long as hindbasitarsus; width of postpetiole 0.56--

0.72 times length, first tergite 3.58--4.0 times as long as postpetiole.

Habitus somewhat stout. Head rather buccate, transverse, lower part of

head in profile quite short, profile suborbicular; face slightly prominent,

median field rather rugulosopunctate, inner orbit broadly nearly impunctate;

clypeus with few punctures, especially below; gena very sparsely finely shal-

lowly punctate; frons, stemmaticum, and vertex rather weakly and sparsely

punctate, especially frons and vertex and especially toward edges, with very

weak rugulae on frons, rather polished, stemmaticum little elevated. Epomia

enclosing short and narrow space (one-third to one-fifth as long as wide), pos-

terior margin rather straight and parallel to margin of collar; notauli usually

fairly distinct at least in anterior third, though not sharp, prescutum little or

not at all raised; mesoscutum fairly densely finely distinctly punctate, punc-

tures on mesopleurum larger, sparser, and shallower; speculum with only very

narrow polished anteroventral edge; scutellum flat, anterior margin not at all

elevated, shallowly longitudinally depressed from side to side, lateral carina

more or less distinct. Propodeum rather finely weakly rugulosoreticulate.

First tergite widened before spiracles but very Little or not at all elevated,

postpetiole quite flat with three distinct longitudinal furrows. Legs very

stout. Ovipositor much deeper than wide, rather deep in proportion to its

length, not decurved (or very slightly and beyond nodus), nodus rather long and

large, ovipositor tip proportionately long.

Color generally fulvous. Antenna fulvous. Head yellow with tip of man-

dible, stemmaticum, and upper part of occiput black, temple fulvoferruginous.

Mesopleurum and propodeum may have fairly extensive black markings; scu-

tellum more or less yellowish; first tergite often infuscate. Wings hyaline to

subhyaline with brown hairs.

Male: Forewing 5.35--6.0 mm, body 10.25--11.8 mm; antenna 6.35--7.0

mm, 31--34 segments,first flagellar segment 4. 5--4.66 times as long as wide,

combined length of second and third segments 1.036--1.185 times first; head

thickness-width ratio 1.67--1.796, head length-width ratio 1.128--1.217, head

thickness-camber ratio 3.46--4.166, occipital camber 4. 45--6.9, head-occiput

width ratio 1.208--1.326, eye-temple ratio 2.08--2.1; mesoscutum index 1.0--

1.05, head width 1.43--1.5 times mesoscutum width, scutellum 1:1.0--1.1:

0.75--0.85; depth:width:length of propodeum 1:1. 08--1.2:1.04--1.14; hind-

D4 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

basitarsus 5.3--5.7 times as long as thick, first tergite 1.6--1.7 times as

long as hindbasitarsus; width of postpetiole 0.52--0.63 times length, first

tergite 3.57--3.93 times as long as postpetiole.

Sculpture and color as in female, but with somewhat more black; meso-

pleurum and metapleurum often with some yellow markings.

Material examined: 892, 3¢°,

Type: ?, Sand Lake, Ont., 3-VII-1926, F. P. Ide (CNC); allotype: ~,

Pittsburg |sic!], Pa, June 3 (Pittsburgh); paratypes: 22, Gull Lake Biol. Sta.,

Kalamazoo Co., Mich., 11 July 1955, Collector, A. D. Dawson (MSEL);

Midland Co., Mich., 7-10-52, R.R.Dreisbach (Dreisbach); Blackburn, Ont.,

18-VI-1941, G.S.Walley (CNC); One Sided Lake, Ont., 2-VI-1960, S.M.Clark

(CNC); Simcoe, Ont., 22-VI-1939, T.N. Freeman (CNC); Brome, Que.,

31-V-1936, G.S.Walley (CNC); Lac Brule, P.Q., 8-VIII-1945. O. Peck (CNC);

paratypes: So, Simcoe, Ont., 22-VI-1939, G.S.Walley (2%, CNC).

19. Aphanistes decurvihastatus, new species

Female: Forewing 6.2--7.05 mm, body 10.3--12.2 mm; antenna 7.3--

7.5 mm (only two specimens with complete antennae), 36--37 segments, first

flagellar segment 3.5--4.5 times as long as wide, combined length of second

and third segments 1.0--1.07 times first; head thickness-width ratio 1.65--

1.74, head length-width ratio 1.16--1.19, head thickness-camber ratio 3.5--

4.66, occipital camber 4.5--6.4, head-occiput width ratio 1.23--1.32, eye-

temple ratio 1.7--2.375; mesoscutum index 1.0--1.14, head width 1.34--1. 4

times mesoscutum width, scutellum 1:0. 9--0.98:0.65--0. 7; depth: width:length

of propodeum 1:1.1--1.2:0.98--1.125; hindbasitarsus 5.3--6.6 times as long

as thick, first tergite 1.55--1.6 times as long as hindbasitarsus; width of post-

petiole 0.52--0.58 times length, first tergite 3.33--3.7 times as long as post-

petiole.

Habitus rather stout. Head rounded-rectangular and moderately trans-

verse; median field of face moderately closely but not deeply rugulosopunctate,

inner orbit broadly nearly impunctate; cheek narrow, increasingly so toward

base of mandible, rather finely sparsely punctate, becoming more closely and

deeply punctate above, temple becoming rather rapidly closely deeply less

finely punctate, genal portion of occipital carina with more or less distinct

angle; frons, stemmaticum, and vertex directly behind stemmaticum very

distinctly rugulose-reticulate-punctate, most deeply on vertex, with some

rugulae at anterior edge of stemmaticum, grading from subpolished on frons

to subopaque on vertex; stemmaticum rather strongly elevated at lateral ocelli,

ocellocular line about equal to diameter of lateral ocellus, slightly shorter

than interocellar line (ocellocular line:diameter of ocellus:interocellar line

6:6:7), lateral ocellus less than three times thickness of occipital carina from

it. Space enclosed by epomia somewhat small, narrow (one-fourth to one-

fifth as wide as long); mesoscutum rather closely moderately finely and deeply

punctate; prescutum rather strongly raised above parapsides, notauli obsolete;

scutellum moderately strongly and closely punctate with a longitudinal con-

cavity usually extending from base to apex and from side to side, disc rather

flat with short vertical anterior declivity, lateral carina indistinct, not at

apparent lateral margin of disc; prepectus rather polished, punctate or

rugulose, without carina on pleurum; mesopleurum moderately densely punc-

tate, somewhat less finely and deeply than mesoscutum; sternaulus rather

distinct and sharp but shallow, about one-third length of pleurum; center of

Hopper: Nearctic Aphanistes 55

mesopleurum just in front of speculum with very sparsely punctate and polished

area, speculum with a distinct polished anteroventral ridge. Propodeum rather

strongly rugose, first lateral area usually separated from second, area denti-

para not or only slightly longer than wide; areola open behind, petiolar area

depressed. Postpetiole approximately parallel-sided, rather narrow and only

slightly flattened, with three longitudinal furrows anteriorly, the middle one

short and weak; petiole rather flat. Ovipositor rather compressed, rather

strongly decurved in apical two-fifths.

Antenna fulvous, head yellow except tip of mandible, mucro of clypeus,

frons, stemmaticum, vertex except orbital spot, temple, and upper part of

occiput black. Mesoscutum ferruginous, spot on cephalic margin of prescutum

and patch before scutellum black; side and apical edge of scutellum black, disc

yellow along sides and fulvous medially (in one specimen blackish, but I be-

lieve this specimen has been discolored by shellac or the like--see comment

on propodeum and first tergite below); pronotum black with upper end ferrugi-

nous; mesopleurum largely black with an oblique ferruginous band from base

of midcoxa to a little below upper anterior corner, extending upward thence to

upper margin and a certain distance backward from there; sterna black. Wings

hyaline to subhyaline, with brown hairs. Fore- and midlegs fulvoferruginous

with tarsi, trochanters, and posterior surface of tibiae yellow; hindcoxa

largely black, hindfemur ferruginous, base of hindtibia yellow, merging

through fulvous and ferruginous to black at apex (outer half of tibia black),

hindtarsus yellow. Propodeum ferruginous with metakatepisternum, basal

area, areola, and spot in petiolar area black (one specimen appears to have

propodeum entirely black with pleural area piceous and first tergite black, but

latter especially is a reddish translucent kind of black suggesting the effect of

shellac or the like--see comment on scutellum of same specimen). Abdomen

fulvoferruginous with second tergite except lateral margins and with blotches

at apex black.

Male: Forewing 5.8--6.35 mm, body 10.3--11.5 mm; antenna 6.6 mm

(only two specimens with complete antennae), 32 segments, first flagellar

seement 4.3--5.2 times as long as wide, combined length of second and third

segments 1.0--1.12 times first; head thickness-width ratio 1. 635--1. 75,

head length-width ratio 1.154--1.189, head thickness-camber ratio 3. 64--

4.5, occipital camber 5.0--5.9, head-occiput width ratio 1.23--1.35, eye-

temple ratio 1.66-2.46; mesoscutum index 0.93--1.1, head width 1.25--1.5

times mesoscutum width, scutellum 1:0.96--1.02:0.69--0.9; depth: width:

length of propodeum 1:1.06--1.116:0. 95--1.03; hindbasitarsus 4.4--6.4 times

as long as thick, first tergite 1.436--1.627 times as long as hindbasitarsus;

width of postpetiole 0.57--0.76 times length, first tergite 3.4--3.9 times as

long as postpetiole.

Sculpture and color much as in female, although punctation slightly weaker,

especially top of head, which rather polished; interocellar line about equal to

diameter of ocellus and ocellocular line; stemmaticum barely elevated, lateral

ocellus one-half its diameter or more from occipital carina. Antenna fulvous

to light fuscous, black apex of hindtibia not half its length.

Material examined: 422, 50°C.

Type: 2, Ithaca, N. Y., 22 July 1940, J.G.Franclemont (CHT); allotype:

o, Robson, B.C., 3-VI-1947, H.R. Foxlee (CNC); paratypes: 2°, Ithaca,

N. Y., VI-7 1940, H.&M. Townes (CHT); Ithaca, N. Y., 9 Aug. 1940, J.G.

Franclemont (CHT); Ithaca, N. Y., 17 July 47 (CNC); paratypes: °°, Emmet

Co., Mich., 5-27-60, R. and K. Dreisbach (Dreisbach); Ontonogan Co., Mich.,

6-18-60, R. and K. Dreisbach (Dreisbach); North Branch, Ont., 23-VII-1960,

o6 Cont. Amer. Ent. Inst., vol. 18, no. 2, 1981

S.M. Clark (CNC); Sand Lake, Ont., 3-VII-1926, F.P.Ide (CNC).

LITERATURE CITED

Cresson, E. T. 1928. The types of Hymonoptera in the Academy of Natural

Sciences of Philadelphia other than those of Ezra T. Cresson. Amer.

Ent. Soc., Mem. 5, 90 pp.

Gauld, I. D. 1976. The classification of the Anomaloninae (Hymenoptera:

Ichneumonidae). Brit. Mus. (Nat. Hist.), Ent., Bul 33(1): 1-135.

Heinrich, G. H. 1961 (1960). Synopsis of Nearctic Ichneumoninae Stenopneus-

ticae with particular reference to the Northeastern Region (Hymenop-

tera). Can. Ent. Sup. 15: 1-87.

1967. Synopsis and reclassification of the Ichneumoninae

Stenopneusticae of Africa south of the Sahara (Hymenoptera). v. 1,

pp. 1-250.

Hopper, H. P. 1959. The pronunciation and derivation of the names of the

genera and subgenera of the family Ichneumonidae found in North

America north of Mexico (Hymenoptera). Ent. Soc. Wash., Proc.

61(4): 155-171.

Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks. 1979.

Catalog of Hymenoptera in America north of Mexico. Smithsonian

Institution Press. v. 1, xvitl198 pp.

Pratt, H. D. 1940. Studies on the Ichneumonidae of New England (Hymenop-

tera). Part 1. The external morphology of Arotes amoenus Cresson.

N. Y. Ent. Soc., Jour. 48: 155-193.

Townes, H. andM. 1959. Ichneumon-flies of America north of Mexico: 1.

Subfamily Metopiinae. (Hymenoptera). US. Nat. Mus. Bull. 216,

ix+318 pp.

Townes, H. 1969. The genera of Ichneumonidae. Part 1. Amer. Ent. Inst.,

Mem. 11, ii +300 pp.

Townes, H. 1971. The genera of Ichneumonidae. Part 4. Amer. Ent. Inst.,

Mem. 17, iii+372 pp.

INDEX

basilicon 38 gracilis 14

coxatus 23 heinrichi 43

crassulus 50 hyalinis 16

crepuscularis 13 masoni 33

decurvihastatus 54 nigrorufus 25

dreisbachi 40 politus 21

edemae 19 puparum 48

enargiae 29 townesi 27

flaviscutellaris 53 walleyi 36

fuscipes 45

Contributions

of the

American Entomological Institute

Volume 18, Number 3, 1981

CLASSIFICATION OF THE BEE FAMILY

MELITTIDAE

WITH A REVIEW OF SPECIES OF

~~ MEGANOMIINAE

Charles D. Michener

CLASSIFICATION OF THE BEE FAMILY MELITTIDAE,

WITH A REVIEW OF SPECIES OF MEGANOMIINAE

Charles D. Michener

Snow Entomological Museum and Department of Entomology

University of Kansas

(The Hesperapis group largely by Gerald |. Stage

Stafford Springs, Connecticut)

ABSTRACT

This is a revision, to the subgeneric level, of the bee family

Melittidae. The family is here divided into three subfamilies, the

Meganomiinae, Melittinae, and Dasypodinae, the last being divided into

three tribes, the Promelittini, Dasypodini, and Sambini. Several new

genera and subgenera are described. Appendices include a revision of

the species of Meganomiinae and descriptions of various new species of

the other subfamilies.

CONTENTS

Introductions 2 "r + ss # £ = = tye ae ee Bee ew oe 1

Authorship= - co 5-ton om "ost Serer Se Aldtcti res 6 oe oe te 2

Acknowledgements- - --------+--*+-++-+--+---+-+----- 3

Methods - ------------------+-+-+-+-+------- 3

Gladogramss. = =. -.+." - timer Ge cin ae toe et te S

Family Melittidae: -. +. -dagenisl wieder 8 = taal - ce 14

Key to the subfamilies of Melittidae - - - - - -------- fe

Meganomiinae new subfamily - - - - -------+-+------ 18

Key to the genera of Meganomiinae- - - - - - - - - - 19

Ceratomonia Michener new genus - - - - - ------ 20

Uromonia Michener new genus- - - - - -----+--- 23

Agemmonia Michener new genus - - ------+-+--- 26

Genus Meganomia Cockerell- - - - - - -------- 30

Subfamily Melittinae - - - - - ----+-+-++-+-+-+-+-+-+2#-+--+ 34

Key to the genera of Melittinae- - - - - - --- - - 36

Genus Melitta Kirby- - - - - - ----+--+-+---- 37

Genus Dolichochile Viereck - - - - - ----+---- - 41

Redivivoides Michener new genus- - - - - - ----- 42

Genus Rediviva.Pries@r 2 =) sommelier ee Ae

Genus Waecropis Kluge + + 6 5 + ee ee ee ee 47

Key. to the subgenera of..Maeropis- 5," --° - > > 49

ii Contrib. Amer. Ent. Inst., vol. 16, no. 3, 198)

Subgenus’ Macropis: Klug 8." Str. st roster t - - 50

Sinomacropis Michener new subgenus- - - - - - - 51

Subgenus Paramacropis Popov and Guiglia - - - - 951

Subranily DasVhOciiGe- - 8 FR Se eR eS oe ee ae ee oe

Key to the tribes or Usesyeodinde--- “<= =. - =--*- +--+ - he

PROMEITLINI New “TNGe= Ste aletcere Tao eee ee Bee oe 54

Genus Promertia Warncne vee = = = = = eee ee 94

SHU eA ie ee ee ee eee Oe ee 60

Key to the genera of the tribe Sambini - - - - - - - 60

Genus Maplomeitts Cockere! “oto -~ = - = = - = = 60

Key to the subgenera Haplomelitta - - - - - - - 63

Prosamba Michener new subgenus- - - - - - - - - 63

Atrosamba Michener new subgenus - - - - - - - - 65

Haplosamba Michener new subgenus- - - - - - - - 66

Subgenus Haplomelitta Cockerell s. str. - - - - 68

Metasamba Michener new subgenus - - - - - - - - 69

GENUS -Sambs FRESE ss She. SR SP me ee 70

DasyDouin THEW TRIBES Sos = Ste ea ate ee ae 73

Key to the genera of the tribe Dasypodini- - - - - - Px

CehUS Oss yOOCS Laren Vee erie iaree: Be = Fae @ 14

Genus: Eremaniance FOpey~ = = & + Rees ta ee Tee oe ar

Key to the subgenera of Eremaphanta - - - - - - 78

Subgenus Eremaprante’ Popov. ¢. Sir.« = = * = - - 79

Popovapis WMichener new subgenus - - - - - - - - 80

COs (eoicge hee A ee a Rr a le 80

PPC e Ue Capo eer Re Re ee 82

Capicoloides Michener new subgenus- - - - - - - 83

-SeeRWS Pesberapis Goccerell = =< = - oss = = + = = = > 85

Key to the subgenera of Hesperapis- - - - - - - 90

Key to the subgenera of certain male Hesperapis 92

Subgenus Afivivapis Cockerell> <"-"<" +0 ee Be)

Subgenus Hesperapis Cockerell s. str. - - - - - 94

Subgenus Zacesta Ashmead= “- -"- = =< oe oR Re} 95

Disbarapis Stace new Subcenus = - = -.-/- - - > 96

SuUGGeNUS FanUraumia. Viereck <" - -eOw rp a alte a7

Carimeos Stage few Sibogenus+ - - ------ > 98

Xeralictoides Stage new genus- - - - - ------ - 99

Appendix |. The species of Meganomiinae- - - - - --------- 102

Genus < Orato. iat oe eee CMe ee ee ee 102

Ceratomonia rozenorum Michener new species 102

Genus Uromonia - - - - - - - - - ss 2 ese 2 ee eo ee 104

Uromonia stagei Michener new species - - - 104

GENUS "Rocio Re Se E Ae Ee ee eH 107

Agemmonia tsavoensis (Strand)- - - - - - - 107

Agemmonie “Laveteneis ‘(Cockerell).- - - - - 109

GSNUS Megat a & SOR TH BE Tee BM ee mo 109

Key “TOCreigies = SVK OSTA A se ee 109

Michener: Classification of Melittidae

Key to males -.- “06 oe Sone = we ew eee eee

Meganomia binghami (Cockerell) - - - - - -

Meganomia gigas Michener new species - - -

Meganomia andersoni (Meade-Waldo)- - - - -

Meganomia (Meganomia a rossi NGHEDEE

new siete sig eee ee eae ee

Appendix Il. New and little-known species of Melittinae and

Dasypodingé << -"S) 4 <'8) 8 40808 Bo ae we Ree ee ee a

Melitta, callornica VBPEC i tk oN ee oy. aie

Redivivoides simulans Mine er

new species- -. =.+ .*..* -- - -- = = -

Rediviva peringuey| (Friese) - - -- == =

Rediviva longimanus Michener new species -

Rediviva colorata Michener new species - -

Haplomelitta (Prosamba) grissonigrs

Michener new species - - Med Lock Pe a

Haplomelitta (Atrosamba) etre Michener

new species- - - - - -- - ee ee ee

Haplomelitta (Haplosamba a) tridentata

Michener new species - - - 25 5 >

Haplomelitta (Metasamba) fasciata

NICMENG! SW SDECIBG( @.* pm tea ot Sitar

Literature Cited- - -- ----- - ef ett eee etter te ee

pee Gee

ens

Michener: Classification of Melittidae 1

INTRODUCTION

As demonstrated previously (Michener and Greenberg, 1980), the

Melittidae is a critical family in bee evolution. It not only exhibits

many primitive features, but is perhaps the sister group of the

Ctenoplectridae plus long-tongued bees. If McGinley (1980) is right

that the bifid tongue of colletids is a derived rather than a plesiomor-

phic wasp-like feature, then the Melittidae could be the ancestral

family of bees.

Because it is a rather small family, many elements of which are not

well known, and because its classification as proposed by me in 1944 now

seems quite inadequate, | here present a new classification. Two recent

changes in the content of the family require notice here. First, when

melittids were first assembled and separated from other bees,

Ctenoplectra was included. The Ctenoplectridae has now been separated

and shown to be more similar to the long-tongued bees (Fideliidae,

Megachilidae, Anthophoridae, Apidae) than to Melittidae (Michener and

Greenberg, 1980). Second, the African genus Meganomia has been removed

from the Halictidae and placed in the Melittidae by Stage (1971). He

was clearly right in so doing; the matter is of special importance

because Meganomia and its relatives (described below) turn out to have

a very distinctive combination of plesiomorphic and apomorphic features.

They may be the sister group to all the other Melittidae, although the

synapomorphic characters of the latter group (all other Melittidae) are

few and not very satisfying.

Melittidae mostly have the superficial aspect of andrenid or cer-

tain halictid bees. Before the familial attributes of the melittids

were known, the melittids with three submarginal cells were often placed

near Andrena (e.g., when Friese in 1911 described Rediviva as a subgenus

of Andrena). Meganomia, however, was described as a subgenus of Nomia

(Halictidae) by Cockerell, 1909, and its species were described in Nomia

not only by Cockerell, but also by Friese (1909), Meade-Waldo (1916),

and Strand (1920), a generic placement that continued for decades in

spite of the lack of nomiine characteristics in Meganomia, until

Cockerell (193la) raised it to generic rank. The yellow and black

coloration is more suggestive of an anthidiine bee than of any Nomia or

other melittid. Its placement in Nomia can only be attributed to the

yellow metasomal bands which, although preapical on each tergum, are

suggestive of the apical yellow, green, blue, or white bands of some

species of Nomia.

Forms with two submarginal cells were often placed among the panur-

gine bees (Andrenidae) or dufoureine bees (Halictidae). For example

Cockerell described various species of Hesperapis in Panurgus, Friese

described the species of Promelitta as a Dufourea, and Morawitz

described the species of Eremaphanta in Rhophites (Dufoureinae). Most

of the species with two submarginal cells (e.g., Dasypoda, Hesperapis,

Capicola) look like Andrena, or the smaller ones like Calliopsis

Panurginae, Andrenidae). Some, for example Haplomelitta, are superfi-

cially very like some species of the colletid genus Scrapter. Perhaps

most noteworthy of all are the tiny species of Eremaphanta which

resemble the panurgine genus Perdita or the halictine genera Nomioides

and Habralictus in size and coloration.

2 Contre: Amer. “Ent. nists? vole 16, no. 3, 1961

After studying a number of characters for a selected list of apoid

genera, Warncke (1977) placed the genus Pararhophites Friese (=

Ctenoapis Cameron) in the Melittidae. The particular characters on

which he based this action are not stated. However, there is nothing

in Pararhophites morphology specifically to support Warncke's action,

and there is much to indicate that it is wrong. The mouthparts are like

those of a typical long-tongued bee (Winston, 1979; Michener and

Greenberg, 1980). Thus the labial palpus is made up of two long sheath-

like basal segments and two small divergent distal segments; the glossa

is elongate with a flabellum, a distinct glossal rod, and a narrow

salivary channel margined except distally by short hairs, and lacks

noticeable branched hairs apically (or elsewhere); the ends of the loral

arms are on the cardo-stipital articulation; and the galeal comb is ab-

sent (as is the stipital comb). The galeae are elongate (longer than

the stipites). Other characters suggesting a relationship to long-

tongued bees are the hairs on the propodeal triangle (metapostnotum) and

the lack of a distinct oblique brush on the under side of the middle

tibia. Attributes of Pararhophites unusual among long-tongued bees are

the presence of distinct volsellae (shared with Fideliidae); the short

and scarcely curved subligular process; the long ligular arms, extending

almost to the base of the prementum (shared with Lithurginae); and the

lack of pygidial and basitibial plates (shared with Fideliidae,

Megachilidae, Apidae, and others). At this time, however, there is no

reason to change Popov's (1949) placement of Pararhophites as a tribe

Pararhophitini in the Anthophorinae.

Examination of type material of Andrena melittoides Friese, 1899,

the type species of Melittoides Friese, 1921, verified that this is an

Andrena, not a melittid.

The present study uncovered several genera and subgenera based on

undescribed species. Such species, and a few other taxonomic findings

not related to the higher classification of the family, are described

in the appendices. The Meganomiinae proved so little known, three of

the four genera and four of eight species being new, that its species

are revised in Appendix |. The Sambini was even less well Known, but

since most of its genera and subgenera are monotypic, no treatment is

given in Appendix | except for descriptions of new species.

These comments raise the question of the number of genera recog-

nized. Genera of bees are generally large and easily distinguished. In

an archaic group such as the Melittidae, however, the existence of

various distinctive, small genera is not surprising. There are a few

moderately large genera such as Melitta and Dasypoda; Hesperapis, per-

haps the largest genus, contains nearly 40 species (Stage, 1966).

AUTHORSHIP

Of course the bulk of the work is by Michener. However, Dr. G. |.

Stage of Stafford Springs, Connecticut, has been kind enough to allow

use of his unpublished work (Stage, 1966) in the sections on Hesperapis

and Xeralictoides. The descriptions of the new taxa involved are Stage's

work and the new names are to be attributed to him.

Michener: Classification of Melittidae 2

ACKNOWLEDGEMENTS

It is a pleasure to acknowledge the help of numerous people without

whom this paper would be less complete. Dr. Les Greenberg examined the

mouthparts of many short-tongued bees and thus provided the basis for

my treatment of these structures. Mr. Robert W. Brooks did the computer

work which assisted in preparation of cladograms.

As noted above, Dr. G. |. Stage permitted the inclusion of portions

of an unpublished revision of Hesperapis and Xeralictoides (Stage,

1966); this allowed me to include here a consideration of all genera and

subgenera of Melittidae. Dr. Stage's sections are appropriately marked

and are his work, although imbedded in material by me.

For the loan of important material, including types, | am indebted

especially to the late Dr. E. Konigsmann of the Zoologisches Museum,

Humboldt-Universitaét, Berlin; also to Dr. P. D. Hurd, Smithsonian

Institution, Washington, D.C. and Dr. J. G. Rozen, Jr., American Museum

of Natural History, New York City.

Mrs. Joetta Weaver assisted with editorial work and typed the

manuscript. Mr. H. Harris of the Academic Computer Center of the

University of Kansas assisted greatly in the computer preparation of

camera-ready copy.

This study was supported by National Science Foundation grant DEB

77-23035, which is gratefully acknowledged.

METHODS

| have not sought to examine every species of every genus. For ex-

ample, in genera like Melitta, Macropis, and Dasypoda, | have examined

all available material and this represents a good sample of the

described species. The current generic concepts seem acceptable and

Species not seen by me are probably correctly placed. Unfortunately in

the fauna of southern Africa there exist species not available to me

whose generic placement is uncertain.

In the descriptions, each character (or group of morphologically

associated characters) is lettered. For coordinate, related taxa (e.g.,

the genera of a subfamily), a given variable (or group of variables) has

the same letter in the description of each taxon. This system permits

brevity of discussions and allows users to readily compare a given

variable among all coordinate taxa. The lettering has no other meaning.

In some cases a distinctive character found in only one genus is under-

lined, and to save space the alternative common character is not

repeated for all other genera.

A substantial list (Table 1) of apomorphic characters is numbered

and was used in cladogram-making. Characters may be referred to either

by letters (which represent variables in coordinate taxa) or by numbers

(which represent apomorphies or plesiomorphies relative to other

Melittidae), or by both.

Terms for structures are essentially those used by Michener (1944,

1965), with a few minor and easily understood modifications for sim-

plicity. Certain terms for mouthparts are derived from Winston (1979).

Thus, instead of regarding paraglossae as two segmented, the term is

limited to the distal "segment," the proximal one being the paraglossal

suspensorium.

4 Contrib. Amer.” Ent. Inst, wor. 16, no. 3, 1961

The only new term introduced for a structure is keirotrichia. These

are the small hairs, usually of uniform length, blunt, spatulate, or

bifid, occupying an area on the inner side of the hind tibia of most

bees. The size of this area is a taxonomic character of interest. Oc-

casionally the area is absent, scopal hairs covering the inner as well

as the outer surface of the tibia.

Illustrations of mouthparts, genitalia, and associated sterna were

prepared in a rather standardized way, although figures of certain

structures are omitted for some taxa if similar structures are il-

lustrated for related taxa, or if good illustrations are readily

available elsewhere. For figures of inner views of maxillae, the whole

maxilla was slightly flattened, usually by a gently pressing coverslip,

so that the galea and stipes are ih the same plane. The orientation of

the lacinia was not at all standardized by this procedure, and apparent

differences in the shape of the lacinia and adjacent parts of the galea

are not meaningful. Various conventions used in drawing the anterior

(or upper if the mouthparts are considered to extend forward) surface

of the labium are mostly evident from the drawings. Thus the paraglossa

and its suspensorium are shown on one side, the palpus on the other, and

the position of the subligular process on the posterior surface is shown

by broken lines. The basal fragmentum of the prementum has its distal

end toward the viewer and basal end away; its shape as shown in the

drawings depends so much on orientation that it means little.

Except in certain cases, as noted in figure captions, where dotted

areas represent areas of dark pigmentation, dotted areas are membranous.

Dots representing sensilla are larger and differently located.

CLADOGRAMS

Using a list of variables for which polarity (evolutionary direc-

tion) could be determined with greater or lesser confidence, | have

developed a list of 102 apomorphic characters upon which are based

cladograms for the supraspecific taxa. Polarity was ascertained

(sometimes only tentatively) by consideration of related taxa (out

groups) (see Stevens, 1980). Thus hairy paraglossae occur in the out-

groups that | have used for Melittidae, i.e., the Colletinae (including

the Paracolletini) and the Andreninae. Within the Melittidae,

therefore, this character is considered as plesiomorphic, and reduced

abundance of paraglossal hairs as apomorphic. For the variable,

paraglossal size, there is a trend from large (and hairy) to small,

rudimentary, and even absent. Such a series obviously involves in-

creasingly apomorphic steps since the structure is one that is well

developed in many insect orders and in other families of bees. Similar

considerations apply for many variables having only two characters. The

last antennal segments in most Hymenoptera are subcylindrical. They are

broadly expanded and flat in males of Ceratomonia, slender and hooked

in Meganomia. These are two obvious apomorphies. Frequency (i.e.,

distribution among taxa) of a character within a group is not a suitable

indication of its plesiomorphy or apomorphy. However, a frequent

character in outgroups is likely to be plesiomorphic in the taxon under

study and its alternative, therefore, is likely to be apomorphic. A

character associated with a known plesiomorphy, especially in outgroups,

is also likely to be itself plesiomorphic, the alternative of the

Michener: Classification of Melittidae 5

variable therefore apomorphic.

Table 1 is a list of 102 apomorphic characters, with parenthetic

notes on the distribution of plesiomorphies of the same variables and

on the basis for my judgement as to polarity within each variable. Such

information is frequently abbreviated, but should be sufficient to

suggest reasons for decisions on polarity of each variable. Essentially

similar methods were used by Stage for his section on Hesperapis.

TABLE 1. LIST OF APOMORPHIES (SEE TABLE 3 FOR APOMORPHIES WITHIN

HESPERAPIS GROUP).

1. Head, thorax, and abdomen with yellow integumental markings.

[Such yellow markings, suggestive of the Anthidiini, occur in

various groups of bees but not in the out groups for Melittidae;

they appear to be clearly derived where present among melittids. |

2. Clypeus of male largely yellow. [A yellow clypeus occurs in males

of many bees, including members of all major families except

Megachilidae. However, a black (concolorous) clypeus is more com-

mon, and occurs in the great majority of such probably primitive

groups as Colletinae. It is a weak variable for our purposes not

only because it seems to have arisen repeatedly among bees but

because there is a possibility that the polarity is misinter-

preted. In addition to the genera shown to have this character in

the cladograms, the lower half of the clypeus is yellow in males

of Hesperapis (Zacesta) rufipes. |

3. Mandible of female simple. [This character occurs in a few other

groups of bees (e.g., Panurginae) but is in every case evidently

derived from the widespread, common type of mandible with a

preapical tooth (apex of the pollex, Michener and Fraser, 1978) on

the upper margin. ]

4. Mandible of female modified as long, broad blade. [A unique

structure found only in Dolichochile. |

5. Mandible of female tridentate. [In Melittidae found only in

Haplosamba; a rare and derived feature in the outgroups also. |

6. Mandible of male simple, without preapical tooth, although with

flaring upper edge. [Although a flaring edge, with evidence of

tooth (apex of pollex) remaining, occurs in some species of both

Hesperapis and Capicola, the apomorphy as stated is unique to

Xeralictoides. |

7. Hind femur with subapical pit on inner surface. [A unique feature

found only in Xeralictoides. |

8. Paraglossa with reduced pubescence, largely hairless. [In nearly

all bees, as well as in many other Hymenoptera, the paraglossae

are densely hairy and often annulate like the glossa. They are

smaller in melittids than in most bees, and nearly hairless in

Dasypodinae. |

10.

ike

13.

14.

15.

16.

ear

18.

10.

20.

ce,

23.

Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

Paraglossa much shorter than suspensorium. [The paraglossa is at

least about as long as its suspensorium in most other Hymenoptera

including the out groups. This character involves progression

beyond 8. ] |

Paraglossa short, small, cylindrical. [See character 9.

Character 10 involves progression beyond 9. ]

Paraglossa short, small tapering. [See character 9. Character 11

involves progression beyond 9; probably is not derived from 10. |

Paraglossa virtually absent. [See character 9. Character 12 in-

volves progression beyond 11. |

Galeal blade not notched preapically. [A preapical notch is found

in the out groups as well as many halictids. Its presence is

therefore presumed to be plesiomorphic. However, there is always

the possibility of convergent origin of the notch in different

taxa; in this case the polarity should be reversed. |

Galeal comb reduced to 3-6 teeth. [Outgroups have combs of 9-18

teeth. |

Galeal comb absent. [See character 14. ]

Stipes with flange along posterior margin, at least apically where

it is broad and rounded. [Such a flange unknown except in Sam-

bini. |

Stipes with flange extending the whole length of stipes. [A step

beyond character 16. |

Posterior margin of stipes with large, preapical concavity. [Such

a concavity is very familiar in long-tongued bees, often margined

with a comb, but is not Known in other short-tongued bees except

the ctenoplectrids. |

Maxillary palpus very short, incompletely two-segmented. [This

palpus is six-segmented in other melittids and in the out groups. ]

Distal third of prementum membranous. [This feature is not known

in other bees or wasps. |

Mentum membranous. [This feature is found in no melittids except

Samba. Other bees that have a reduced or membranous mentum show

quite a different and presumably independently evolved reduction. ]

Labial palpus short (as in Dolichochile). [This feature is unique

to Dolichochile. ]

Maxillary palpus with long setae. [The short setae found in all

others (including out groups) are much elongated in Prosamba. |

24.

26.

28.

29.

oY,

ot.

Se.

33.

34.

36.

Michener: Classification of Melittidae 7

Labrum with lateral lobes. [Such lobes are not found in the out-

groups or in melittids other than Melitta and Dolichochile. ]

Clypeus with median ridge. [This is a weak character because it

occurs in scattered groups of bees and wasps, often as a specific

character. It appears to be an apomorphy in the two groups of

melittids that exhibit it. |

Clypeal apex trilobed. [This feature is associated with the very

short clypeal truncation which is the median lobe. Such a clypeus

is not found in outgroups or melittids other than Meganomiinae.

Vertex convex seen from front, elevated well above summits of eyes

(sometimes apparently secondarily concave although elevated. )

[The high, convex vertex is found in certain other bees (e.g., the

Melitomini of the subfamily Anthophorinae) but is a scattered,

rare phenomenon in the out groups. Presumably the plesiomorphic

character of this variable is a rather flat vertex, with the ocel-

lar area somewhat elevated. |

Vertex concave seen from front. [See character 27. ]

Face of female much broader than length of eye. [The extra-

ordinarily wide, low head of Samba is unique among bees. |

Flagellar apex of male expanded. [The plesiomorphic condition is

a simple, rounded last antennal segment, as in most wasps and

bees. |

Flagellar apex of male hooked. [See character 30. ]

Flagellum of male flattened, with long hairs on under side.

[Cylindrical segments with very short hairs are very widespread

and certainly piasiamer aia. t

Mesoscutal hairs scale-like. [Slender, often sparsely and rather

briefly plumose hairs are characteristic of most melittids and the

outgroups, and are doubtless plesiomorphic. |

Hairs extraordinarily densely plumose. [See character 33. ]

Upper metapleural pit difficult to recognize because it is close

and somewhat posterior to lower pit. [Widely separated pits, one

far above the other, characterize the outgroups and most melit-

tids. |

Front coxa with apical hairy spine or lobe. [Similar projections

are found in a few other bees but not in the outgroups. |

Third trochanter of female angulate beneath. [Such modification

of the trochanter of the female is unusual, scarcely found in the

outgroups. |

38.

SU:

40.

Al.

42.

43.

44,

45.

46.

47.

48.

49.

50.

ol.

ave

Contrib. Amer. Ent. -inst.,..vol.-18,.n0:.8, 1981

Anterior femur of male with basal process. [Among melittids such

a process occurs only in Ceratomonia; similar structures are not

known in the outgroups.

Hind femur of male swollen. [Similar enlargement occurs in

various other bees but rarely if at all in the outgroups. |

Basitibial plate of female not defined apically, hidden under

pubescence. [Well defined basitibial plates characterize females

of most burrowing bees including the outgroups and most melit-

tids. |

Basitibial plate of female absent. [See character 40. |

Hind tibia of female with apical bare area on outer surface.

[Such an area is not found in the outgroups. |

Front tibia with several large teeth on outer surface apically and

subapically. [This is a unique feature known only in

Xeralictoides. |

Middle tibial spur of female short, robust. [This spur is slender

and gently curved in nearly all bees. |

Middle tibial spur of female strongly hooked apically. [See

character 44. |

Female with only one hind tibial spur. [Comparable loss of one

spur and enlargement of the other is not known in female bees

other than Samba. |

Male with inner hind tibial spur much elongated, curved backward,

hairy. [This is a unique modification known only in Prosamba. ]

Hind tibia and basitarsus of female broad. [This character is not

found in the outgroups, and may be related to oil transport. |

Scopa plumose with emergent, long, bare hairs. [Comments same as

for character 48. |

Limited plumose hairs under long simple scopal hairs. [This fea-

ture occurs in various bees but in melittids is restricted to a

few groups and the basis of correlated attributes is derived. ]

Apex of hind basitarsus of female with hairs parted. [Such diver-

gence of hairs is not found in the outgroups. This character is

not the same as the parting of hairs along the upper (= posterior)

margin of the basitarsus in females of some Hesperapis. |

Front tarsus of female with dense, short hairs. [This type of

vestiture is not found in outgroups and is presumably related to

oil collecting. |

OS)

34.

cla

56.

oF:

50.

oo.

60.

61.

62:

63:

64.

65.

Michener: Classification of Melittidae 9

Posterior basitarsus of female with small apical projection or

tooth above articulation of second tarsal segment. [Such a tooth

is rather widespread among bees and could be plesiomorphic for

Apoidea, although certainly apomorphic for Aculeata, for it is

lacking in wasps. |

Posterior basitarsus of female with long apical projection or

process above articulation of second tarsal segment. [This

progressive development of character 53 is found in only a few

taxa. |

Hind basitarsus of male with comb and apical process. [Equivalent

structures are not found in other bees. |

Hind basitarsus of male inflated. [This is an unusual character,

not shared by bees in the outgroups. |

Hind basitarsus of male with hairy pocket on outer side near base.

[This character is unknown in bees other than Samba. ]

Claws with inner rami broad and truncated on hind (sometimes

middle) leg. [Claws in the outgroups and most melittids each have

a strong, acutely pointed inner tooth or ramus. |

Claws with inner rami broad and truncated on all legs. [See

character 58. This is a progressive extension of character 58. ]

Claws of female with inner rami represented by strong tooth. [See

character 58. |

Claws of female with inner rami minute or absent. [See character

98. This is a progressive extension of character 60. ]

Arolia absent. [Loss of arolia occurs in a few other scattered

groups of bees but not in bees that resemble melittids. It is

unusual in this case in that associated structures remain rather

well developed. |

Tibia of female with short, blunt, coarse hairs scattered among

scopal hairs. [This feature is not known in the outgroups. ]

Second transverse cubital vein absent, so that there are only two

submarginal cells. [Loss of this vein is a widespread common

apomorphy in wasps and bees. ]

Base of second submarginal cell more or less transverse. [This is

an unsatisfying character since it is rather variable; even in-

dividual variations in positions of vein intersections can in-

fluence the slope of the first transverse cubital vein. Nonethe-

less the character is in general useful. It is uncommon in the

outgroups but is characteristic of the paracolletine genus

Scrapter. |

66.

67.

68.

69.

70.

vag

133

74.

13;

76.

tis

78.

73,

Contrib... Amery -Ent.inst.g. Vol... 18_,0¥ 3, 1961

Mesoscutum as long as or longer than intertegular distance. [This

character, found in Xeralictoides, does not occur in other melit-

tids. |

Stigma very slender. [This character is not uncommon in large

bees; it appears to have arisen independently in various

families. ]

Stigma very broad. [This character is common in minute bees and

must have arisen independently in various families. ]

Prestigma longer than stigma. [Same comment as for character 67. |

Prestigma much shorter than stigma. [Same comment as for

character 68. |

Jugal lobe about half as long as vannal lobe. [In the outgroups,

at least in presumably primitive members of them, the jugal lobe

is long, two thirds or three fourths as long as the vannal lobe.

The long lobe is therefore presumably plesiomorphous in

Melittidae. |

Jugal lobe less than half as long as vannal lobe. [See character

71, of which this is a progressive extension. |

Profile of propodeum all more or less in one plane. [In the

presumably more primitive genera in the outgroups, the anterior

third to fifth of the profile is more nearly horizontal than the

more posterior or declivous part. ]

Profile of propodeum half horizontal. [See comment under

character 73. ]

Propodeal triangle small. [The triangle is rather large in nearly

all bees, including most members of both outgroups. |

_Propodeal triangle partly hairy. [This is a rare feature in

short-tongued bees. As explained in the discussion of

Sinomacropis, this could be morphologically the same as character

Toc]

Metasomal terga without apical hair bands. [Apical hair bands are

widespread among bees including many members of the outgroups.

Their presence is probably plesiomorphic. |

Metasomal terga with basal hair bands. [Basal hair bands sugges-

tive of those of the halictid genus Lasioglossum, are rare or ab-

sent in the outgroups and must be an apomorphy in melittids. ]

Tergal graduli bent posteriorly at each side but terminating

roughly half way from bend to marginal zone of tergum. [This con-

dition is rather common in bees and could be plesiomorphic.

However, it seems that it is not common in the outgroups and it is

tentatively regarded as apomorphic. |

80.

81.

BS.

84.

85.

86.

87.

88.

89.

90.

a1;

Michener: Classification of Melittidae 11

Tergal graduli strongly continuing to rear at each side, reaching

or nearly reaching marginal zone. [This is a progressive exten-

sion of character 79, and its rarity supports the view that it is

apomorphic. |

Male with "stridulatory areas" on bases of certain terga. [Such

areas are unknown in other bees. |

Prepygidial fascia of female absent. [A dense band of hairs on

the fifth tergum is characteristic of nearly all burrowing bees.

Its loss is a remarkable apomorphy of Eremophanta. |

Pygidial plate of female with longitudinal median elevated area,

or at least with ridges marking the edges of such an area. [Such

an elevated area, extending more or less the full length of the

plate, is not found in the outgroups. |

Pygidial plate of male reduced (poorly defined.) [The pygidial

plate is widespread and doubtless plesiomorphic in bees. It is

more often ill-defined or lost in males than in females. ]

Pygidial plate of male absent. [This is a progressive extension

of character 84. Reversal is possible, probably because genes

specifying the plate are retained in females and have only to be

activated in males. |

Sternum V of male with apical comb. [A comparable comb is not

found in similar bees. ]

Latero-apical lobes of sternum VII of male sclerotized, in same

plane as body of sternum, directed laterobasad, one pair only.

[See comments on character 88. ]

Disc of sternum VII of male a broad plate, almost like preceding

sterna. [Widespread among the presumably primitive families of

bees is the type of sternum VII in which the disc is reduced to a

narrow base from which arise two pairs of usually long, almost

membranous, lateroapical lobes, the upper pair with hairs on the

lower surface and the lower pair with hairs on the upper surface.

Such elaborate, similar structures are unlikely to have arisen in-

dependently; hence this condition is presumably plesiomorphic for

bees. A broad plate must therefore be an apomorphy. |

Sternum VII of male with latero-apical lobes reduced. [See com-

ments on character 88. |

Sternum VII of male without distinct latero-apical lobes. [See

comments on character 88. This is a progressive extension of

character 89. | |

Gonocoxite of male abruptly broadened at base. [This is an at-

tribute not found in other bees. |

12 Contrib. ‘Amer. Ent. inst,, vol. 16,-no. 3, 1981

92. Gonocoxite of male with mesoapical, apically produced lobe. [Such

a lobe occurs in some other bees but is weaker; it is virtually

absent in melittids other than Sambini. |

93. Gonostylus short, broadly fused to gonocoxite. [I make the as-

sumption that a more or less articulated, moveable stylus is

primitive. For insects this is obviously true, but it could be

that for bees it is a reversion, for the broadly fused gonostylus

is common. |

94. Gonostylus enlarged and at least biangulate apically. [This fea-

ture is not found in related bees nor in styli in general. It

must be apomorphic. |

95. Gonostylus deeply bifid. [Same comment as for character 94. |

96. Gonoforceps with inner lobe, often cylindrical, usually bearing

long, coarse bristles or hairs. [Such an inner lobe is not known

in other bees. Its presence must be an apomorphy. |

97. Volsella with digitis much elongated, not opposable to cuspis.

[Digitus and cuspis of about the same length and with opposable

denticles are characteristic of most Hymenoptera including the

outgroups. The elongated digitus must be apomorphic. |

98. Volsella much reduced, without recognizable digitis and cuspis.

[See comments under character 97. ]

99. Penis valve grotesque, with both upward and downward projecting

processes. [Such processes are absent or feebly developed in the

outgroups as well as most melittids. |

100. Gonostylus with defined area of dense setae on outer surface.

[Such an area does not occur in other melittids. |

101. First sternum of female highly modified. [No other bee has an

equivalent modification. It must be an apomorphy. |

102. Antenna of male 14-segmented. [Antennae are 13-segmented not only

in other male bees but throughout the aculeata. |

Using the characters listed in Table 1, and a tabulation of

characters of the variables for all taxa, cladograms were prepared

(Figs. 1, 2). A computer program by J. S. Farris (1970) was used to

order the taxa and characters and to prepare preliminary cladograms.

Differences between Figures 1 and 2 result from different considera-

tions, not all of which are worth discussing in detail. In Figure 1

Rediviva and Macropis have a common stem because of characters 48, 49,

51, and 52, all of which are perhaps related to oil collecting and oil

transport. This arrangement separates Rediviva and Redivivoides, but

then these two genera have four convergent apomorphies, 72, 75, 89, and

93. Figure 2 reverses the arrangement, placing Rediviva and

Redivivoides on the same stem, and showing the oil-manipulating

Michener: Classification of Melittidae 13

characters as convergent in Macropis and Rediviva. In spite of the

number and detailed similarity of these characters, | suspect that the

relationships are better shown in Figure 2.

Figures 1 and 2 also differ in the sequence of branches in the

Meganomiinae. The relations among the four very distinctive genera are

indicated by very few synapomorphies, only three in each figure. Al-

though these characters seem strong, they differ in the two figures, and

a conservative cladogram based on present knowledge would probably show

all four meganomiine genera arising from a common point.

The difference in the basal furcation of the cladogram in Figures |

and 2 depends only on the emphasis placed on character 13, loss of the

preapical notch of the galea. Figure 1 assumes it happened twice, which

is likely considering that it has happened in other groups of bees and

that the notch is bridged by transparent material, thus partly lost, in

Ceratomonia, and moreover is completely lost in one species of

Meganomia.

The grouping of apomorphies on certain stems in Figures 1 and 2

gives some idea of the distinctiveness of the various groups. The

autapomorphies could always be multiplied, especially for taxa con-

taining only one species in which one has only to search for more

characters. Synapomorphies are less easily increased in number. It is

impossible to find all the characters, and | believe that those shown

in Figures 1 and 2 are a representative sample which show that the

Meganomiinae, each meganomiine genus, Macropis, Melitta-Dolichochile,

Eremaphanta, Samba, and the Sambini are all strikingly distinctive taxa.

At the opposite extreme, there are some terminal branches with no

characters marked on them, e.g., Macropis s. str. in both Figures 1 and

2. As indicated the taxonomic treatment, there are other differences,

sometimes characters whose polarity is obscure and which therefore could

not be used in the cladistic study. It must be remembered that lack of

characters, e.g., on the Macropis s. str. line, is not really true; the

plesiomorphic alternatives of the apomorphies shown for the sister group

characterize the seemingly character-free taxa. Probably the unmarked

lines represent paraphyletic taxa, i.e., Macropis s. str. is probably

the group from which Sinomacropis evolved. | have no problem with this

idea. Those who wish classifications and cladograms to be completely

redundant and who do not recognize paraphyletic taxa would have to

develop a different classification (based on which cladogram? ).

Comparison of the cladograms shown in Figures 1 and 2 with

cladograms based on larval characters prepared by Rozen (1978) shows

major similarities. Rozen studied larvae of Meganomia, Melitta,

Macropis, Dasypoda, Capicola, and Hesperapis (as well as Ctenoplectra,

now placed in the family Ctenoplectridae). His cladograms are exactly

concordant with Figure 2 except that Ctenoplectra branches from the

Meganomia stem. Rozen notes, however, that "the apparent closeness

between Ctenoplectra on the one hand and Meganomia, Melitta, and

Macropis on the other, is spurious in that Ctenoplectra differs by many

features, the plesiomorphic-apomorphic polarity of which cannot be

determined." The reasons for separating Ctenoplectra into a family

distinct from the Melittidae are adult characters listed by Michener and

Greenberg (1980).

14 Contrib: Amer: ERT. Inst: vel; “16, no. 3, 1937

Xeralictoides Rese enapily Samba

7 ;

Eremaphanta Haplomelitta

Sinomacropis

Macropis

: Atrosamba

Meganomia

45*

es Haplosamba

1aword

—Promelitta

INILLI

Figure 1. Cladogram for the genera and subgenera (except for

Hesperapis) of Melittidae. (Paramacropis is omitted, being insuf-

ficiently known; Haplosamba is placed only on the basis of female

characters.) The numbers represent apomorphies listed in Table 1.

Asterisks indicate characters not appearing elsewhere in the Melittidae.

A dash beside a number indicates that it represents a strong character,

whose polarity is unlikely to be misunderstood. An x after a number in-

dicates a reversion to what looks like the plesiomorphic condition.

Lines without indications of apomorphies lead to presumably paraphyletic

taxa.

Family Melittidae

Diagnosis. Short-tongued bees (i.e., with labial palpal segments

similar and cylindrical, the first two not elongate and sheath-like)

without glossal flabellum; mentum tapering basally and curled over

v-shaped lorum as in long-tongued families; galeal comb usually present;

mesepisternal disc with system of grooves represented only by groove

from scrobe to posterior margin of mesepisternum; scopa limited to

posterior tibia and basitarsis; volsella present.

Description. a. Labrum usually much broader than long, apical

margin in both sexes fringed with bristles. b. One subantennal suture

Michener: Classification of Melittidae 15

Eremaphanta Xeralictoides

6* Samba

27X

43*

66*

Hesperapis (subg.)

Popovapis

Rediviva

Redivivoides 28%

29* Haplo-

46* melitta

Melitta

Dolichochile :

Capicola

100*

Capicoloides Metasamba

Dasypoda

j3aword

—Promelitta

INI LILI

Figure 2. A.second cladogram for genera and subgenera of

Melittidae, using the same characters as for Figure 1, but making dif-

ferent assumptions in various parts of the tree about which are conver-

gent. Explanation as for Figure 1.

below each antenna, its upper end directed toward middle of antennal

socket or toward either inner or outer margin of socket. c. Facial

fovea absent. d. Lower lateral parts of clypeus not much bent

posteriorly on either side of labrum. e. Glossa usually shorter than

prementum. f. Apex of glossa pointed, usually attenuate. g. Apex of

glossa with branched hairs. h. Flabellum absent. i. Posterior sur-

face of glossa with depression or shallow groove; hairs margining

depression or groove similar to nearby glossal hairs, not short as in

forms with a deep groove. j. Glossal rod absent. k. Paraglossa

usually small, sometimes absent, commonly shorter than suspensorium ex-

cept in Meganomiinae, if densely hairy, with branched hairs at apex. |.

Labial palpus with first two segments more or less cylindrical, not

sheath-like; third and forth segments in line with the first two, not:

projecting laterally. m. Subliqular process of prementum not strongly

curved anteriorly, a pair of scarcely sclerotized bands diverging from

its apex and extending toward lower margins of paraglossal suspensoria.

n. Ligular arms of prementum distinct, not or incompletely fused to

margins of premental sclerite, bases near base of prementum. o. Mentum

elongate, tapering basally, curled over the lorum to its articulation

with the latter, usually well sclerotized (not sclerotized in Samba).

16 Contrib... Amer... Ent... |nst.,. vol..i8, no..3, 1961

p. Apex of mentum articulated with prementum via a detached basal frag-

mentum of the prementum; base of fragmentum directed posteriorly so that

fragmentum is at a strong angle to axis of prementum. q. Lorum well

sclerotized, slender, strongly v-shaped, apices of its arms lying on the

cardines only slightly above the cardo-stipital articulations. r.

Galeal blade usually equal to or shorter than stipes, usually much

longer than prepalpal part of galea. s. Galeal comb present (reduced

in Melitta, greatly reduced in Dolichochile, absent or essentially so

in Dasypoda); stipital comb and its concavity absent (concavity distinct

in Eremaphanta). t. Basistipital process somewhat elongate (short in

Macropis). u. Pre-episternal groove absent, as is scrobal groove in

front of pleural scrobe, so that the only strong groove on disc of

mesepisternum is that between scrobe and meso-metapleural intersegmental

suture. v. Mid coxa fully exposed, nearly reaching lower metapleural

pit, shorter than or equal to distance from summit of coxa to summit of

metapleuron. w. Upper metapleural pit above to below midpoint between

summit of mid coxa and hind wing base. x. Metapostnotum (propodeal

triangle) hairless. (A possible exception, with what appears to be a

partially hairy metapostnotum, occurs in Sinomacropis.) y. Oblique

brush on under side of mid tibia of female (tibial comb of Jander, 1976)

present, sometimes weak. z. Basitibial plate of female usually

distinct, of male often distinct. aa. Scopa restricted to hind tibia

and basitarsus. bb. Jugal lobe of hind wing one fourth to three

fourths as long as vannal lobe, both measured from wing base. cc.

Volsella present (much reduced in Meganomiinae). dd. Penis valves

rarely fused dorsally, not forming a distinct spatha. ee. Pygidial

plate of female present; pygidial and prepygidial fimbriae of female

present except in Eremaphanta, prepygidial fimbria reduced to mere hair

band in some other Dasypodini; pygidial plate of male frequently absent.

ff. Mandibular cusp of larva multidentate, apical concavity scoop-like

(Rozen and McGinley, 1974; Rozen, 1977).

Comments. No known character is both unique to the Melittidae and

a feature of all melittids. However, the combination of a short

proboscis (characters e, r, and especially |) with the form of the lorum

and mentum (characters o, q, as in long-tongued bees) separates

Melittidae and Ctenoplectridae from all other bees. The Melittidae

differs from Ctenoplectridae not only in the glossa (characters e, g,

h) and the presence of a galeal and lack of a stipital comb (character

Ss), but also in other characters listed below and in lacking the

numerous special synapomorphies of the Ctenoplectridae listed by

Michener and Greenberg (1980).

As detailed by Michener and Greenberg (1980), characters 0, p, q,

t, u, Vv, W, and aa distinguish the Melittidae from the short-tongued

families Andrenidae, Colletidae, Halictidae, Oxaeidae, and ;

Stenotritidae. Of these, characters o, p, q, t, v, and aa are shared

with the Ctenoplectridae and long-tongued families. Most of these

characters are not found elsewhere in Hymenoptera and are thus

synapomorphies of the group consisting of Melittidae, Ctenoplectridae,

and the long-tongued families of bees. The Melittidae differs from the

Ctenoplectridae in characters g, h, i, m, s, u, cc, and dd; the

Ctenoplectridae shares the alternatives of most of these characters with

the long-tongued bees.

As to synapomorphies of the Melittidae alone, one can only cite

characters k and u, neither of which is very convincing. The paraglos-

Michener: Classification of Melittidae 17

sae do tend to be small in Melittidae, but are larger in the

Meganomiinae than in other subfamilies. They are as small as in most

melittids in certain Andrenidae. Reduction in size could thus have oc-

curred independently, and in any event the difference in paraglossal

size between that of Meganomiinae and that common in other short-tongued

families of bees is not great. The loss of mesepisternal grooves is

more convincing but is duplicated in certain other bees (most

Megachilidae, some Andrenidae), showing that it could be a convergence.

The loss of the grooves, therefore, could have occurred independently

in various melittid groups, although there is no evidence for this idea.

It is thus possible that there is no true melittid synapomorphy, in

which case Melittidae is not the sister group of the Ctenoplectridae

plus long-tongued (C L-T) bees, but is the paraphyletic group from which

C L-T bees arose. The Melittidae is nonetheless a phenetically compact

group, presumably monophyletic (in the usual, not the Hennigian, sense).

It may well be paraphyletic, however, in the sense that pre-

ctenoplectrids arose from bees that, if Known to us, would be placed in

Melittidae. The short-tongued families Colletidae, Halictidae,

Andrenidae, and Stenotritidae may also have arisen from bees that would

be placed in the Melittidae. A cladogram showing the relationships of

the families that fall between the major short-tongued families and the

long-tongued families, assuming that Melittidae and C L-T bees are

sister groups, is given by Michener and Greenberg (1980).

Key to the Subfamilies of Melittidae

1. With yellow or cream markings on head, thorax, and metasoma; apex

of marginal cell rounded, bent away from wing margin.

sleeps aetna hab d cinsiuictlc et re mleihe nigh aged ta nes wee a te aa Meganomiinae

-- Yellow or cream markings, if present at all, limited except in

Eremaphanta to face in male and absent in female; apex of marginal

cell pointed, on or very near wing margin.

2. Paraglossa largely bare, usually markedly shorter than suspen-

sorium, hairs largely limited to apex, or paraglossa absent;

forewing with two submarginal cells, second usually shorter than

first, first transverse cubital (= base of second submarginal cell)

usually more or less at right angles to longitudinal veins (Figs.

5, 6) and usually close to first recurrent vein. Known larvae do

not spin cocoons.

we ee a me ty Sh Gi tt a Dasypodinae

-- Paraglossa densely hairy; forewing with two or three submarginal

cells, second (if only two cells) or second plus third as long as

or longer than first, first transverse cubital (= base of second

submarginal cell) slanting, usually well separated from first

recurrent vein (Figs. 3, 4). Larvae spin cocoons.

edi Sel chi aka ashe ag inhi amy dol eal Cage ont ata ya actassap abvaatee! elec she sacle dexdiatueton Melittinae

18 Contrib...Amer. Ent. inst., vol. 18, no.'3, 1951

Figures 3-6. Wings of Melittidae. 3, Melitta leporina (Panzer). 4,

Macropis labiata (Fabricius). 5, Hesperapis pellucida Cockerell; 6,

Dasypoda panzeri Spinola. Scale lines = 1.0 mm.

Meganomiinae new subfamily

Diagnosis. Differs from other subfamilies, except for minute bees

of the genus Eremaphanta in the Dasypodini, in having yellow or ivory

maculations not only on face but elsewhere on head, thorax, and

metasoma; in the slender, parallel-sided stigma; and in the much reduced

volsella without recognizable digitis and cuspis.

Description. a. Head, thorax, and metasoma with yellow or cream-

colored maculations. b. Mandible of female simple, of male simple or

with extreme apex slightly bidentate because pollex extends nearly to

apex of rutellum or with pollex shorter than rutellum and ending in a

subapical mandibular tooth. c. Paraglossa longer than its suspen-

sorium, densely hairy with annuli similar to those of glossa. d.

Galeal blade with preapical notch or slit on posterior margin, filled

by transparent material in Ceratomonia (Fig, 10), this notch or slit ab-

sent in Meganomia rossi. e. Labrum much broader than long, that of

female except in Ceratomonia with sharp transverse ridge, beyond which

is a concavity extending to the depressed apical part. f. Frontal

carina on a strongly raised, roof-like ridge in the region between an-

tennae. g. Malar area linear. h. Clypeus with a longitudinal median

ridge, at least partly shiny and not or sparsely punctate. i.

Propodeal triangle dull, granular (rather shiny in Meganomia rossi),

rather large, nearly as wide as width of propodeum, declivous but basal

part not so steeply sloping as distal part. j. Forewing with three

submarginal cells, usually second and always third narrowed toward

Michener: Classification of Melittidae 19

costal margin, second and third together shorter than or as long as

first. k. Basal vein over twice (usually over three times) as long as

first abscissa of Rs. |. Stigma extremely slender (Fig. 57), prestigma

two thirds as long as stigma to longer than stigma; vein r arising near

apex of stigma, sides of stigma basal to vein r parallel or even con-

verging apically. m. Apex of marginal cell rounded, bent away from

wing margin. n. Jugal lobe of hind wing about two thirds as long as

vannal lobe or somewhat shorter, but always over half length of vannal

lobe. o. Mid and hind basitarsi of both sexes narrower than tibiae.

p. Hind basitarsus of female tapering distally, apex except in

Ceratomonia giving rise to segment 2 more or less medially and not much

produced distally above base of second segment. q. Metasomal terga

without apical hair bands, except for strong prepygidial and pygidial

fimbriae (terga V and VI). r. Metasomal terga |-IV of females and I|-V

of males with broad, nearly impunctate apical bands, depressed below

level of rest of tergum all the way across. s. Lateral extremities of

the tergal graduli bent posteriorly and continuing as strong

longitudinal carinae nearly to posterior margins of terga. t. Pygidial

plate of male present. u. Seventh sternum of male with a pair of

sclerotized apical lobes arising medially and extending laterally or an-

terolaterally, often in contact with disc of sternum (so that one may

not immediately see that lobes join the sternum only near the midline).

v. Eighth sternum of male without well defined bevelled or truncate

apical plate, although sometimes with an indication of such a plate. w.

Volsella much reduced, not at all chelate. x. Gonostylus in-

distinguishably fused to gonocoxite.

Comments. The Meganomiinae contains bees rich in specialized fea-

tures. Synapomorphies of the subfamily include characters a, b (simple

mandible of female), h, |, gq, r, s, and w. All of these are absent or

nearly so in other Melittidae and related short-tongued bees. On the

other hand, the subfamily retains various ancestral characters, recog-

nized by their widespread occurrence in related families of short-

tongued bees. Such plesiomorphies are characters c, d, j, n, 0, p, and

t. Of these c and d are the most meaningful, as the rest occur also in

certain other melittids.

Rozen (1977, p. 14) suggests subfamilial status for the group on

the basis of larval synapomorphies.

The Meganomiinae contains four genera, all from eastern or southern

Africa, that are very different morphologically. They are Meganomia,

Agemmonia, Uromonia, and Ceratomonia. Each genus seems about equally

different from the other three, largely due to the specialized derived

features of each. Figures 1 and 2 show two different cladograms for the

subfamily, almost equally parsimonious in character changes (44 vs. 43).

The discordance of the distribution of the synapomorphies indicated

shows that some of them either evolved independently in different genera

or, after evolving, were lost in certain descendant genera.

Key to the Genera of Meganomiinae

|. Arolia absent (although long, hairy, yellow planta projects between

the claws); flagellum of male with apical segment curled and at-

tenuate (Figs. 49-52, 55, 56); hind basitarsus of male bent or con-

torted, or at least with curved carina on outer side.

20 Contrib: Amer: Ent. inst..-vol..16,nou3, 1961

“EnTTC SEAS VERT LREP OR TE RTE Pe ee ere ---Meganomia

-- Arolia present, conspicuously black among associated pale in-

terungual structures; flagellum of male with apex simple or ex-

panded as a plate; hind basitarsus of male simple.

2. Upper part of head gently convex seen from front; ocelli much in

front of posterior edge of vertex (Fig. 58), posterior ocellus

separated from that edge by several ocellar diameters and front

edge of median ocellus nearer to antennal bases than to posterior

edge of vertex; inner orbits diverging below; first flagellar seg-

ment over four times as long as broad; glossa as long as prementum.

oe em ee el aye er cera Ae ae, ame dae a yea Agemmonia

-- Upper part of head seen from front flat or with ocellar region

slightly elevated; posterior ocellus separated from posterior edge

of vertex by an ocellar diameter or little more; front edge of

median ocellus much nearer to posterior edge of vertex than to an-

tennal bases; inner orbits converging below, at least in upper

halves; first flagellar segment less than three times as long as

wide; glossa less than half as long as prementum.

3. Basitibial plate of female not defined; flagellum of male simple;

terga IV and V without sublateral stridulating areas; penis valve

of male enormously developed with dorsal and ventral processes.

ee i ge ea maa sag 25 eo ce Be dere ey cone wee, Uromonia

-- Basitibial plate of female well defined; flagellum of male with

apical segment expanded, plate-like, black; terga IV and V with

sublateral stridulating areas hidden under preceding tergal mar-

gins; penis valve of male of ordinary slender form.

ee oe FO CT ST ee ace ey aes ee Ceratomonia

Ceratomonia Michener new genus

(Figs. 4). 10,.J4-15,.4/7, 48, 5/4. 58,, 19¢)

Type species: Ceratomonia rozenorum new species.

Diagnosis. Male flagellum long, flat, hairy beneath; last segment

expanded, discoid; basitibial plate of female well defined, conspicuous;

labrum a transverse strip, not produced medially; arolia present.

Description. a. Moderate sized (length 13-15 mm); body rather

slender; forewing short (about 7 mm long). b. Inner orbits converging

below in upper half of face. c. Clypeus about twice (male) or more

than twice (female) as wide as long. d. Clypeal apex with median trun-

cation over one third as wide as clypeus, in male produced so that

clypeal apex is weakly trilobed. e. Subantennal sutures converging

above, longer than (male) or subequal to (female) maximum (lower) width

of supraclypeal area. f. Summit of head, seen from front, about

horizontal, with ocellar area slightly elevated. g. Ocelli on summit

of head, posterior ocellus separated from posterior edge of vertex by

Michener: Classification of Melittidae 21

Ce es

Figures 7-13. Anterior views of labium and inner views of maxilla

of Meganomiinae. 7, Ceratomonia rozenorum n. sp. 8, Uromonia stagei n.

sp. 9, Agemmonia tsavoensis (Strand). 10, Ceratomonia rozenorum n. sp.

11, Uromonia stagei n. sp. 12, Agemmonia tsavoensis (Strand). 13,

Meganomia andersoni (Meadeswalaby: Scale lines = 0.5 mm; those for

labia apply also to maxillae of the same species.

22 Conthid:.. Amer... BAL. 1st. vol..78, now 3, 1987

less than an ocellar diameter, distance from anterior edge of anterior

ocellus to antennal base nearly twice distance from anterior edge of an-

terior ocellus to posterior edge of vertex. h. Gena little more than

half width of eye seen from side. i. Mandible of male simple, in

repose with apex behind clypeal margin, completely hiding labrum. j.

Labrum of male about six times as wide as long, with distal depressed

part reduced to a mere transverse rim (not produced medially) with a

fringe of long bristles; subapical transverse ridge present; surface

smooth. kK. Labrum of female about ten times as wide as long, distal

depressed part absent, but apical fringe of long bristles present,

transverse ridge absent or not sharp, whole labral surface convex and

punctate. |. Galeal blade short and blunt (Fig. 10). m. Maxillary

palpus extending well beyond apex of galea. n. Galeal comb with

bristles, except for longer basal ones, of approximately uniform length;

preapical notch of galeal blade filled in with transparent material. o.

Cardo much shorter than stipes, thickened distally. p. Ligular arm

short, reaching neither base nor apex of prementum. q. Glossa and

paraglossa about as in Uromonia. r. Scape reaching middle (male) or

upper margin (female) of median ocellus, in male somewhat thickened

(Fig. 58). s. Flagellum of male long, reaching beyond scutellum, seg-

ment | less than twice as long as apical width, apex deeply concave;

segment 2 about as long as broad, segments 3-10 more elongate and

progressively narrower so that 10 is about twice as long as broad; seg-

ments progressively flatter from 1-11; segments 2-11 with long, erect,

simple hairs, curled at tips, on under surfaces, these hairs mostly

about as long as length of segments from which they arise, but shorter

on 10 and 11; segment 11 broader than rest of antenna, bent downward,

discoid. t. Flagellum of female shorter than eye, first segment nearly

twice as long as broad, segments 2 to 9 broader than long, 10 but little

longer than broad. u. Dorsum of thorax with many short, densely

plumose, appressed hairs, almost scale-like in female, among scattered,

long, thick, unbranched, erect hairs. v. Upper half of triangular area

of propodeum (metapostnotum) less steeply sloping than rest of propodeum

in profile. w. Basitibial plate of female well differentiated, with

a covering of dense short hairs that terminates sharply at the raised

edge of the plate. x. Posterior trochanter of female with ventral mar-

gin gently curved. y. Front coxa without posteriorly directed process.

z. Fore femur of male swollen with retrorse, posterior basal

prominence; middle femur and tibia of male somewhat swollen; male legs

otherwise not greatly modified. aa. Inner side of hind tibia of female

covered with short spatulate-truncate keirotrichia, only at sides with

coarse scopal hairs. bb. Scopal hairs coarse, curved, mostly superfi-

cially simple but many of them with one to several barbs on convex side.

cc. Apex of hind tibia of female without bare area on outer surface.

dd. Hind basitarsus of female nearly parallel-sided, second tarsal seg-

ment articulated to lower part of apex, upper apical angle prolonged in-

to shiny, flat leaf-like projection similar to that of Promelitta

_(Dasypodinae) but with only one or two hairs on the inner surface. ee.

Claws with inner rami truncated lobes. ff. Arolium present; orbicula

long and finger-like on middle and hind legs. gg. Prestigma shorter

than stigma. hh. Basal vein distad to cu-v and nearly three times as

long as first abscissa of Rs. ii. Second and third submarginal cells

both narrowed toward anterior (costal) margins, subequal in length on

posterior margins, together slightly longer than first submarginal cell.

Michener: Classification of Melittidae 23

jj. Marginal cell rather short and broad (Fig. 57). kk. Metasomal

terga IV and V with shining, oval, minutely striate, sublateral

pregradular ("stridulating") area, behind which the gradulus is arched

posteriorly. Il. Seventh tergum of male with weakly bilobed and

slightly upturned apex, pygidial plate joining this apex, not produced

beyond it, basally extending half way or more across disc of tergum

toward gradulus. mm. Metasomal sterna of female without scopa. nn.

Metasomal sterna I-V of male with small, shallow median apical emargina-

tions, without conspicuous apical fringes although posterior margins

somewhat more hairy than elsewhere, IV and V rather hairy with some of

the hairs extending beyond the posterior margins. oo. Sternum V of

male otherwise unmodified. pp. Sternum VI of male with broad apical

emargination on either side of which is a crescentic area of stiff erect

setae, and at apex of lobe lateral to emargination, an area of dense

plumose hairs. qq. Sternum VII of male with broad sinus between

lateral, apical lobe and body of sternum and area of long, dense,

-plumose hairs midapically, at bases of lateral apical lobes. rr. Ster-

num VIII of male without spiculum, with only weak lobes as basolateral

apodemes, with rather slender apical process densely hairy below, plate-

like above and flattened, with transverse preapical ridge. ss.

Gonocoxite of male simple, without special broadened base. tt. Penis

valves slender, nearly straight, blunt, not fused dorsally. uu.

Volsella relatively large but of the usual form for Meganomia.

Comments. Striking derived characters (autapomorphies) differen-

tiating this genus from the others in the subfamily are i, j, k, lI, s,

u, ee, and kk (shared with Meganomia). Plesiomorphies which also dif-

ferentiate this genus from certain others include f, g, w, xX, aa, CC,

ff, mm, oo, and ss.

There is only a single known species, C. rozenorum, described in

Appendix |. It is from Namibia (South West Africa). Some of its

behavioral features are indicated by Rozen (1977) under the heading

'Meganomia species B."

Etymology. Ceratos, horn, with reference to the long and peculiar

antennae of the male; and monia, anagram of Nomia.

Uromonia Michener new genus

(Figs, 8, 11," 1e+28,5oa¢ 57, 56; 197)

Type species: Uromonia stagei new species.

Diagnosis. Male flagellum simple, twelve segmented; basitibial

plate of female not defined; labrum over four times as wide as long,

apical margin convex or broadly truncate; arolia present; penis valves

of males large, complex, with both dorsal and ventral projections.

Description. a. Small but moderately robust, body. length 10-12

mm, forewing length 7-8 mm. b. Inner orbits of male distinctly con-

verging below, except diverging at lower extremities; of female paral-

lel except diverging at lower extremities. c. Clypeus about twice

(male) or three times (female) as wide as long. d. Clypeal truncation

not defined (male) or about half as wide as clypeus (female), no emar-

ginations on clypeal apex that produce a trilobed effect. e. Subanten-

24 Contri6.. Amer: Ent. 1Wst.; vol. 18, ne. 3, 1961

Figures 14-30. Meganomiinae, males. 14-18, Ceratomonia rozenorum

n. sp. 14, metasomal tergum VII. 15, sternum VI (ventral). 16, ster-

num VII (dorsal and ventral). 17, sternum VIII (dorsal and ventral).

18, genitalia (dorsal and ventral). 19-24, Uromonia stagei n. sp., same

sequence of structures plus lateral view of genitalia. 25-30, Agemmonia

tsavoensis (Strand). 25, metasomal tergum VII. 26, 27, sterna V and VI

(ventral). 28, sternum VII (dorsal and ventral). 29, genitalia (dorsal

and ventral). 30, sternum VIII (dorsal and ventral). Scale lines = 0.5

mm; the tergum and sterna V and VI are of one scale, genitalia and ster-

na VII and VII! at another, for each species. :

Michener: Classification of Melittidae 25

nal sutures in male diverging above, about three fourths as long as

lower width of supraclypeal area; in female subparallel and little over

half as long as width of supraclypeal area. f, g, h. Summit of head

and genal area as described for Ceratomonia. i. Mandible of male with

inner tooth very near apex, in repose only partly hidden behina clypeal

margin, labrum not exposed. j. Labrum of male over five times as wide

as long; surface smooth, concave; apical margin broadly truncate but no

sharp ends of truncation. k. Labrum of female over four times as wide

as long, distal depressed part short so that transverse ridge is beyond

middle of labrum, area basal to ridge punctate, apical margin of labrum

gently convex. |. Galeal blade rather short with fringe of long hairs

on posterior margin. m. Maxillary palpus not reaching apex of galea,

first segment longer than next two together. n. Galeal comb as

described for Meganomia. 0. Cardo much shorter than stipes. p.

Ligular arms short, ending near middle of prementum, distal third of

prementum membranous except for subligular process which arises from

weakly sclerotized area on posterior surface of prementum, this area

joined by membrane to rest of sclerotized part of prementum. q. Glossa

and paraglossa as in Figure 8. r. Scape as described for Ceratomonia.

s. Flagellum of male twelve segmented (so that antenna is fourteen seg-

mented), almost reaching front of scutellum, segment 1 1.5 times as long

as wide, segment 2 as long as wide, remaining segments a little longer

than wide, simple. t. Flagellum of female slightly shorter than eye,

ten segmented as in other bees, segment 1 twice as long as broad, 2

broader than long, 3 and succeeding segments about as broad as long,

grading to slightly longer than broad apically. u. Dorsum of thorax

with hairs of varying lengths, briefly plumose. v. Propodeal profile

nearly straight, declivous, especially in female, in male more as

described for Ceratomonia. w. Basitibial area of female as in

Agemmonia but basal carina on anterior rather than posterior margin. x.

Posterior trochanter of female with ventral margin gently curved. y.

Front coxa without posteriorly directed process. z. Legs of male

simple except swollen apical part of hind tibia. aa. Inner side of

hind tibia of female entirely covered with long hairs, keirotrichia ab-

sent. bb. Scopal hairs simple. cc. Apical fourth of outer surface

of hind tibia of female slightly concave, bare, but with scopal hairs

arching over it forming a corbicula-like space. dd. Hind basitarsus

of female tapering apically, apex giving rise to second tarsal segment

near middle but upper apical angle, just above base of second segment,

with sharp, apically directed tooth. ee. Claws of foreleg of both

sexes with inner ramus acutely pointed, shorter than outer ramus but

similar in shape; claws of mid and hind legs with inner rami truncated

lobes. ff. Arolium present; orbicula rather long and finger-like. gg.

Prestigma about as long as stigma. hh. Basal vein and cu-v as in

Ceratomonia. ii. Second submarginal cell with ends subparallel, or

slightly narrowed toward anterior (costal) margin; third strongly nar-

rowed toward costal margin and much longer than second; second and

third together slightly shorter than first. jj. Marginal cell as in

Ceratomonia. kk. Metasomal terga without "stridular" areas. Il.

Seventh tergum of male with broadly truncate pygidial plate extending

slightly beyond apex of rest of tergum, basally plate extending less

than half way across disc of tergum toward gradulus. mm. Metasomal

sterna of female without scopa. nn. Metasomal sterna I-IV but not V

of male with small, median, apical emarginations; all these sterna with

26 Contrib. Amer. -ENt..1 Asti; vol, 16) no:'3,.1981

rather long preapical hairs but these forming dense fringe only on V,

to a lesser degree on IV. oo. Sternum V unmodified. pp. Sternum VI

with narrow, almost slitlike median apical emargination and somewhat

sclerotized posterior lateral angle which bears a pair of heavy, an-

teriorly directed pegs, one often larger than the other but sometimes

of equal size; distal margin of sternum except laterally with fringe

similar to that of V. qq. Sternum VI! of male with lateral apical lobe

overlapping body of sternum so that no sinus is evident between lobe and

body until the parts are manipulated. rr. Sternum VIII without

spiculum, with only weak basolateral apodemes, and with broad, flat,

truncate apical process which has an only weakly plate-like beveled

apex. ss. Gonocoxite of male with abruptly broadened base, expanded

and gently lobed apex. tt. Penis valves robust, expanded, grotesquely

lobed, with both dorsal and ventral subapical projections, the two

valves fused to one another from base to apex, where they project as a

flat, median, apical plate. uu. Volsella absent or small and fused to

base of penis valve.

Comments. Apomorphous (derived) characters distinguishing this

genus include the following: | (fringe on galea), p, s (14-segmented

antenna probably unique in the Aculeata), w (shared with Agemmonia), aa,

bb, cc, ss, tt, uu. Plesiomorphous characters include the following:

Boe Oe Ole Se ey RR Nn, 00:

There is only one species, Uromonia stagei, described in Appendix

|. It is from the coast of Kenya. |

Etymology. Uro, tail (with reference to the large and elaborate

male seats, and monia, anagram of Nomia, the halictid genus whose

name is unfortunately in the root of the generic name Meganomia.

Agemmonia Michener new genus

(Pigs 9) 12, 2o=30;'64, SF, 58,192)

Type species: Nomia tsavoensis Strand, 1920.

Diagnosis. Male flagellum simple, slender, segments 1-4 all much

longer than broad; basitibial plate of female not defined; labrum over

three times as wide as long; arolia present. Top of head convex seen

from front and ocelli far in front of posterior margin of vertex.

Description. a. Moderate sized (length 13-15 mm); body rather

robust; forewing length 11 mm. Longer hairs mostly strongly plumose,

those from mandibles, stipes, and hypostomal area forming flat beard on

underside of head of male. b. Face broad; inner orbits of male paral-

lel in upper two thirds, diverging below in lower third; of female en-

tirely diverging below. c. Clypeus about three times as wide as long.

d. Clypeal truncation of male reduced in width to about distance

between antennal sockets, with broad emargination lateral to truncation

so that apical margin of clypeus is strongly trilobed; of female about

as wide as distance between outer margins of antennal sockets and weakly

emarginate lateral to truncation. e. Subantennal sutures parallel or

slightly diverging above, half as long as lower width of supraclypeal

area. f. Summit of head, seen from front, gently convex. g. Ocelli

in front of summit of head; posterior ocellus separated from posterior

margin of vertex by three or more ocellar diameters; anterior edge of

anterior ocellus about midway between antennal base and posterior edge

Michener: Classification of Melittidae 2/

of vertex. h. Gena nearly as wide as eye seen from side. i. Mandible

of male with weak inner tooth near apex, in repose with apex largely

visible from in front, exposing extreme base of labrum. j. Labrum of

male about three times as wide as long, with apical depressed part and

sharp transverse carina at base; apical margin convex. k. Labrum of

female over three times as wide as long, with distal strongly depressed

part, rounded apically, and high carinate ridge across base, basal sur-

face of this ridge punctate. |. Galeal blade rather elongate (Fig.

12). m. Maxillary palpus less than half as long as galeal blade. n.

Galeal comb reduced to a few slender bristles. o. Cardo a little

shorter than stipes. p. Ligular arm long, extending beyond apex of

Ime cia RAN AA Ae

" wy aN

Figures 31-46. Meganomia, males. 31-36, M. binghami (Cockerell).

31. sternum Vii. 32, 33, "sterna V and. V1 (ventral). 34, 35, sterna

Vile NIL dorsal and ventral). 36, genitalia (dorsal and ventral).

37-41, M. andersoni (Meade-Waldo). 37, 38, sterna V and VI (ventral).

39, 40, ~sterna VII and VIII (dorsal and ventral). 41, genitalia (dorsal

and ventral). 42-46, M. rossi)'n. sp.,; same structures and sequence.

Scale lines = 0.5 mm; the tergum and sterna V and VI are at one scale,

genitalia and sterna VII and VIII at another, for each species.

28 Contribs Amer. Ent: inste, vol. 18,-ne.. 3, 1981

Figures 47-56. Flagella of male Meganomiinae. 47, 48, Ceratomonia

rozenorum n. sp. (lateral and ventral). 49, 50, Meganomia rossi n. sp.

(lateral and straightened ventral). 51, 52, M. andersoni (Meade-Waldo)

(same). 53, Uromonia stagei n. sp. 54, Agemmonia tsavoensis (Strand).

55, 56, Meganomia gigas n. sp. (lateral and straightened ventral).

Scale lines = 0.5 mm.

prementum, but distal third only weakly sclerotized. q. Glossa at-

tenuate, about as long as prementum. Paraglossa about as in

Ceratomonia. r. Scape reaching beyond posterior ocelli, slender. s.

Flagellum of male reaching front of scutellum, no thicker than scape,

segment 1 about four times as long as broad, nearly half as long as

scape; segment 2 twice as long as broad; 3 and 4 longer than broad; 5

slightly so; 6 to 10 about as long as broad; 11 longer than broad. t.

Flagellum of female shorter than eye, segment 1 about four times as long

as broad, segment 2 slightly longer than broad, 3 about as long as

broad, 4 to 9 broader than long, 10 longer than broad. u. Dorsum of

thorax with some small, nearly simple hairs among longer, more or less

erect, strongly plumose hairs. v. Upper two thirds of triangular area

of propodeum sloping strongly but less steeply than posterior surface

of propodeum. w. Basitibial plate of female minutely and closely punc-

tate and densely covered by uniform short hairs that grade into those

of more distal part of tibia, but plate not delimited by carina except

near base on posterior margin. x. Posterior trochanter of female with

a preapical ventral tubercle or angle. y. Front coxa without

posteriorly directed process. z. Legs of male not swollen or otherwise

Michener: Classification of Melittidae 29

highly modified; upper side of hind tibia with two well separated

longitudinal carinae between which is a strong concavity. aa. Inner

side of hind tibia of female about as described for Meganomia;

keirotrichia shorter than in that subgenus and the band of such hairs

broader. bb. Scopal hairs long, sparse, superficially appearing simple

but most with several branches on convex sides near bases. cc. Apex

of hind tibia of female without bare area on outer surface. dd. Hind

basitarsus as described for Meganomia. ee. Claws of middle and hind

legs of female with inner rami truncated lobes, of male and forelegs of

female with inner rami acute blades. ff. Arolium present; orbicula

long and finger-like. gg. Prestigma as long as or longer than stigma.

hh. Basal vein distal to cu-v and over four times as long as first ab-

scissa of Rs. ii. Second submarginal cell with ends parallel; third

narrowed toward costal margin. Third submarginal cell longer than

second, the two together shorter than first. jj. Marginal cell long,

not so slender as in Meganomia s. str. (Fig. 57). kk. Metasomal terga

without "stridulatory" areas. Il. Seventh tergum of male truncate with

apex of pygidial plate extending slightly beyond truncation; pygidial

plate broad, extending basally less than half way across disc of tergum.

mm. Metasomal sterna, especially III to VI, of female with long,

briefly plumose hairs apparently forming a metasomal scopa. nn.

Metasomal sternum | of male with small median apical notch, II-IV

without apical emarginations, not fringed. oo. Sternum V of male with

broad median emargination across base of which is a comb of stiff

bristles, interrupted medially; otherwise not greatly modified. pp.

Sternum VI of male as in Figure 27. qq. Sternum VII of male with

lateral apical lobe bent back at apex which overlaps body of sternum.

rr. Sternum VIII of male with spiculum evident as weak angle;

basolateral apodemes strongly projecting anteriorly lateral to spiculum;

apical process broad, truncate, without a plate-like apex. ss.

Gonocoxite simple, without special broadened base, with long plumose

hairs apically. tt. Penis valves slender, nearly straight, broadened

and rounded at apices, fused dorsally near bases. uu. Volsella small,

almost lost as separate sclerite because of fusion with base of penis

valve.

Comments. Apomorphous (derived) characters include the following:

a (strongly plumose hairs), b, d, e, f, g, m, n, q, w (shared with

Uromonia), x (shared with Meganomia), bb (sparse scopa), gg (shared with

Meganomia), mm, oo (comb shared with Meganomia), rr (especially

basolateral apodemes), uu. Associated with these are the following

plesiomorphous characters which differentiate this genus from certain

others: 2z (little modified legs), ff (arolium), kk, nn, ss (simple base

of gonocoxite). None of the plesiomorphous characters distinguishes

this from all other genera of Meganomiinae.

This genus contains only two species, Agemmonia tsavoensis (Strand)

and A. tavetensis Cockerell, both from arid parts of Kenya. As in-

dicated in Appendix |, it is likely that these forms are not

specifically different.

Etymology. Anagram of Meganomia.

30 Contrib. Amer. Ents. lista vol. 18. noe 3; 1951

Figure 57. Forewings of Meganomiinae. Left to right, Ceratomonia

rozenorum n. sp., Uromonia stagei n. sp., Agemmonia tsavoensis (Strand),

and Meganomia gigas n. sp. Scale line = 0.5 mm.

Genus Meganomia Cockerell

(Figs, 13, 31246, 49-52,.55-59; 193-196)

Nomia (Meganomia) Cockerell, 1909: 402.

Meganomia Cockerell, 1931a, p. 201; Stage, 1971, p. 306; Rozen, 1977,

Type species: Nomia (Meganomia) binghami Cockerell, 1909 (monobasic).

Diagnosis. Male flagellum thick, without long hairs beneath,

curled apically, the last one to three segments narrowed; basitibial

plate of female defined at sides but not apically, hairs on and beyond

plate terminating at the same level so that structure of plate is hid-

den; labrum two to three times as wide as long, apical margin convex or

medially emarginate; arolia absent.

Description. a. Large, robust forms (body length 15 to 22 mm);

forewing length 10 to 13 mm. b. Inner orbits slightly converging below

in upper half of face, below that level diverging, or wholly diverging

below in female of M. gigas. c. Clypeal proportions as in Ceratomonia.

d. Clypeal truncation about one third as wide as clypeus (rounded in

male M. andersoni and rossi), in male distinctly produced so that apex

of clypeus is trilobed, in female not or weakly produced. e. Subanten-

nal sutures converging above, slightly longer to slightly shorter than

maximum (lower) width of supraclypeal area. f, g. Summit of head and

ocellar position as in Ceratomonia. h. Genal area of male about half

as wide as eye seen from side, of female well over half eye width. i.

Michener: Classification of Melittidae 31

Figure 58. Faces of Meganomiinae. Top to bottom, males at left,

females at right, Ceratomonia rozenorum n. sp., Uromonia stage! .. 6p. ,

Agemmonia tsavoensis (Strand), and Meganomia binghami (Cockerell). Scale

ine = 0.5 mm.

32 Contrib. Amer. Ent. Inst.; vol. 16, no. 3, 1981

Mandible of male with inner tooth, in repose only partly hidden behind

clypeal margin, extreme base and apex of labrum often exposed. j.

Labrum of male 2 to over 2.5 times as wide as long, smooth, convex to

emarginate apically, without transverse ridge, not divided into elevated

basal and depressed apical portions. k. Labrum of female about 2.5 to

3 times as wide as long with strong transverse median carina, basal to

which surface is usually roughened, distal to which it is depressed and

smooth. |. Galeal blade moderately long (Fig. 13). m. Maxillary pal-

pus approximately reaching apex of galea. n. Galeal comb with basal

bristles curved, much longer than apical bristles (Fig. 13). o. Cardo

nearly as long as stipes. p. Ligular arms long, extending from base

nearly to apex of prementum. q. Glossa and paraglossa about as in

Uromonia. r. Scape reaching ocellar area, in male somewhat broadened,

especially near base. s. Flagellum of male rather short (reaching wing

base), thick, flattened or crenulate beneath, without long hairs, curled

apically, last one to three or four segments attenuated; segment 1

slender basally, more than two to three times as long as broad. t.

Flagellum of female about as long as eye, segment 1 slender basally,

about three times as long as broad, segments 2 to 9 broader than long

to longer than broad (basal ones longer than distal ones which are al-

ways broader than long), 10 longer than broad, but sometimes only

slightly so, in M. rossi about as long as broad. u. Dorsum of thorax

with erect, finely barbed hairs. v. Propodeal profile as in

Ceratomonia. w. Basitibial plate of female well differentiated (i.e.,

elevated) on anterior and posterior margins but apically grading into

general tibial surface; dense short hairs that cover plate grading into

those of outer tibial surface. x. Posterior trochanter of female with

preapical ventral tubercle or angle. y. Front coxa strongly produced

posteriorly as a hairy process mesal to base of trochanters (weakly so -

in male of M. andersoni and M. rossi). z. Middle coxa of male with a

smooth protruding process (weak in M. andersoni and M. rossi) below

trochanteral base; front and middle femora and tibiae of male not

modified; hind femur of male greatly swollen, tibia and basitarsus

somewhat modified and contorted. aa. Inner side of hind tibia of

female with longitudinal strip of keirotrichia with broadened bifid

apices, on either side of which scopal hairs occupy broad marginal

areas. bb. Scopal hairs coarse, curved (especially those of outer side

of tibia), simple (in M. andersoni) to with rather numerous branches on

convex sides of more strongly curved hairs (in M. binghami). cc. Apex

of hind tibia of female without bare area on outer surface. dd. Hind

basitarsus of female gradually tapering toward apex which gives rise to

second segment near middle, upper apical angle scarcely prolonged or in

M. andersoni and M. rossi prolonged as small, dark, flattened, pointed

projection, a miniature of that found in Ceratomonia. ee. Claws of hind

legs with inner ramus a truncated lobe (or in male and one female of M.

rossi a short, pointed, recurved blade), of other legs with inner ramus

an acute blade somewhat shorter than outer ramus but of similar shape.

ff. Arolia absent; orbicula short, much exceeded by elongate, hairy

planta. gg. Prestigma about as long as stigma. hh. Basal vein

slightly distal to, interstitial with, or basal to cu-v and four to six

times as long as first abscissa of Rs. ji. Second and third submar-

ginal cells as described for Ceratomonia, or ends of second subparallel.

jj. Marginal cell elongate, slender (Fig. 57). kk. Metasomal terga

IV and V with specialized "stridulatory" areas as in Ceratomonia. II.

Michener: Classification of Melittidae 33

Figure 59. Faces of Meganomia. Top, M. gigas n. sp., male, female.

Middle, M. andersoni (Meade-Waldo), male, female. Bottom, ME. POSS! fy.

sp.; female, male. Scale line = 0.5 mm.

34 Contribs Amer Ents anst.¢ vol. 382 nev 3, 1981

Seventh tergum of male with well formed, strongly elevated, apically

rounded pygidial plate which protrudes beyond rest of tergum and is

longer than maximum width of plate; basally marginal carinae of plate

extend half way or more across tergum toward gradulus. mm. Metasomal

sterna of female without scopa. nn. Metasomal sterna I-IV of male with

small, shallow median apical emarginations, larger in M. rossi than in

other species; ||| and 1V with dense apical fringes, absent laterally

on IV and weak laterally on Ill in M. rossi. oo. Sternum V of male

with broad apical emargination containing one or two combs of stiff,

black teeth; lateral margin apically with row of long bristles directed

ventrally; base of sternum except in M. rossi with two large, oval, mem-

branous invaginations. pp. Sternum VI of male with narrow apical emar-

gination and strong, lateroapical, laterally projecting process (at-

_ tenuate, deeply bifid, and projecting posterolaterally in M. rossi).

qq. Sternum VII as described for Ceratomonia but without area of long,

plumose hairs. rr. Sternum VIII with weak angle representing spiculum,

with only weak projections as basolateral apodemes, and with apical

process flat, not truncated or plate-like. ss. Gonocoxite of male with

abruptly broadened base as seen in dorsal view, apex expanded and trun-

cate. tt. Penis valves slender, nearly straight, rounded apically but

laterally compressed and therefore pointed as seen from above, fused

dorsally in basal third or half (fusion fully sclerotized in M.

andersoni and M. rossi so that penis valves may best be considered as

not fused). uu. Volsella as described for Ceratomonia, or in M.

andersoni and M. rossi so reduced as to be difficult to recognize.

Comments. Derived characters that differentiate this genus are s,

w, x (shared with Agemmonia), z, aa, ff, gg (shared with Agemmonia), kk

(shared with Ceratomonia), and oo (combs shared with Agemmonia). No

distinctive plesiomorphic generic characters have been recognized; the

recognized plesiomorphies of this genus are all shared with one or more

other genera of Meganomiinae.

This genus contains Meganomia andersoni (Meade-Waldo), binghami

(Cockerell), gigas new species, and rossi new species. All these

species are described in Appendix |. The genus occurs from the sub-

tropical parts of South Africa and Namibia to Kenya, in rather xeric

localities. Behavior and immature stages of M. gigas are described by

Rozen (1977) who used the name M. binghami.

Subfamily Melittinae

Diagnosis. Differs from the other generally non-maculate subfamily

(Dasypodinae) in having (a) paraglossa densely hairy, at least nearly

as long as its suspensorium; (b) three submarginal cells or if two, then

second subequal to or longer than first (true also of some Sambini, only

slightly shorter in Promelitta); (c) base of second submarginal cell

(first transverse cubital vein) oblique, widely separated from first

recurrent vein (true also in some Sambini), and (d) graduli of metasomal

terga bent posteriorly at lateral extremities and extending about half

way to posterior tergal margin (except in Rediviva and Redivivoides ).

Description. a. Body small (7 mm long) to rather large (15 mm

long), having the form and often the appearance of an Andrena, without

yellow maculations or in Macropis face of male with yellow; supraclypeal

Michener: Classification of Melittidae 35

area of female with small yellow spot in Macropis hedini Alfken. b.

Mandible of female (and male) with subapical inner tooth (apex of pol-

lex). c. Paraglossa about as long as its suspensorium, not distinctly

annulate to rather coarsely annulate (in Dolichochile), densely hairy.

d. Galeal blade without preapical notch or slit on posterior margin.

e. Labrum two to six times as broad as long, in some females of

Macropis with strong transverse ridge. f. Frontal carina on a strongly

convex, sometimes roof-like ridge in the region between antennal bases.

g. Malar area linear to as long as broad. h. Clypeus usually without

longitudinal median ridge. i. Propodeal triangle variable. j.

Forewing with two or three submarginal cells, second (if there are two)

or second plus third about as long as or slightly longer than first,

second (if there are two) or third strongly narrowed toward costal mar-

gin, second (if there are three) parallel-sided or narrowed toward

costal margin. Base of second submarginal cell slanting, not at right

angles to the longitudinal veins, well separated from first recurrent

vein. k. Basal vein 1.3 to 3 times as long as first abscissa of Rs,

variable in position relative to vein cu-v. |. Stigma less slender

than in Meganomiinae, prestigma over one third to nearly two thirds as

long as stigma; vein r arising near middle of stigma; sides of stigma

basal to vein r converging slightly toward base of stigma. m. Apex of

marginal cell narrowly rounded or pointed, bent away from costal margin

only to extent of one or two vein widths. n. Jugal lobe of hind wing

one third to over one half length of vannal lobe. o. Mid and hind

basitarsi narrower than tibiae or in some females hind basitarsus fully

as wide as tibiae. p. Hind basitarsus of female parallel-sided or

usually tapering distally, apex giving rise to segment 2 at lower mar-

gin, apex of basitarsus above base of segment 2 extending slightly

distally. gq. Metasomal terga usually with apical bands of pale hair

arising from depressed apical marginal zones, these bands sometimes only

feebly represented at sides (e.g., in Melitta haemorrhoidalis

Fabricius), or sometimes expanded to cover much of terga (e.g., in some

Sinomacropis); prepygidial and pygidial fimbriae strong. r. Metasomal

terga | to |V of females and | to V of males with rather broad usually

depressed apical zones, impunctate or more finely or sparsely punctate

than rest of exposed tergal surfaces, these zones sometimes not dif-

ferentiated except laterally. s. Lateral extremities of graduli of

terga || to IV of females, II to V of males bent posteriorly, then

fading away, not continuing over half way from bends to posterior tergal

margins, in Rediviva and Redivivoides scarcely continuing posteriorly.

t. Pygidial plate of male present or absent. u. Sternum VI! of male

a plate often with modified apical margin and with greatly reduced

lateral apical lobes or disc reduced but with paired apical processes

or lateral apical lobes or both. v. Sternum VIII of male usually with

heavily sclerotized apical process, with or without apical bevelled

plate. w. Volsella well developed. x. Gonostylus variable.

Comments. The bees here included in the Melittinae have been

divided in some previous classifications (Michener, 1944) into the

Melittinae and Macropidinae. Separation at the subfamily level seems

unnecessary; recognition of two tribes, Melittini and Macropidini, would

be a possibility since Macropis is rather distinctive. If the cladogram

shown in Figure 1 is correct, recognition of tribes seems quite un-

desirable while if that shown in Figure 2 is correct, such recognition

seems more appropriate. (For further comments, see below and the sec-

36 Contrib. Amer. Ent. Inst. vol. 16, no. 3, 1981

tion on Cladograms. )

The Melittinae has no known synapomorphies (Fig. 2) or only one,

shared with the Dasypodinae (Fig. 1). Thus it is probably a

paraphyletic group from which the Dasypodinae and possibly also the

Meganomiinae arose. The decision to unite the Macropidinae with the

Melittinae is based especially on similarities between Macropis and

Rediviva (an obvious relative of Melitta). As indicated in Figure | and

the discussion of cladograms, both have the hind tibia and basitarsus

of the female broad, the latter short, the scopa on each with a plumose

understory and emergent simple bristles, the basitarsus with the hairs

of the apex parted to show a narrow apical hairless line. Both also

have the front and sometimes the middle tarsi thickened and covered with

velvety or spatulate hairs. These are synapomorphies not found in any

related bees and are relatively complex, hence perhaps not evolved in-

dependently in Macropis and Rediviva. If it could be shown that this

complex of features was secondarily lost in the Melittinae that lack

them, they would constitute important evidence of the monophyly of the

Melittinae. Figure 2 suggests another alternative, namely that they

arose independently (convergently) in Macropis and Rediviva as adapta-

tions to oil collecting. |

The similarity of Macropis and Melitta in larval characters is em-

phasized by Rozen and McGinley (1974). There is nothing in their study

that justifies tribal or subfamilial separation of these genera.

The Melittinae as here understood exhibit several plesiomorphies

which unite them. Among these are the slanting base of the second sub-

marginal cell (instead of transverse, a feature of most Dasypodinae),

and especially the fact that the larvae spin cocoons and have all the

necessary labial equipment to do so (lacking in Dasypodinae whose larvae

are known). These features are shared with the Meganomiinae, and appear

to be primitive melittid attributes.

The Melittinae is found in the Holarctic Region and in Africa. It

is absent elsewhere.

Key to the Genera of Melittinae

1. Two submarginal cells; pygidial plate of male present; male with

yellow face marks; gonostylus of male long, slender at base, ar-

ticulated to gonocoxite; eighth metasomal sternum of male not

ending in bevelled area suggestive of a pygidial plate (Holarctic

Region).

-- Three submarginal cells; pygidial plate of male absent; yellow face

marks absent; gonostylus of male broadly fused to gonocoxite;

eighth metasomal sternum of male ending in a bevelled area sugges-

tive of a pygidial plate.

2. Propodeal triangle dull (finely granular), large (width at upper

margin usually at least nearly equal to distance between trans-

metanotal sutures); second submarginal cell wider than long or

about as long as wide; seventh metasomal sternum of male a large

plate, truncate or emarginate apically, the lobes or processes

Michener: Classification of Melittidae a7

reduced to small sclerotic structures and sometimes hair patches at

posterior lateral angles of sternum (Holarctic Region and Africa).

Propodeal triangle shiny (usually smooth), often ill defined, small

(width at upper margin not over half distance between trans-

metanotal sutures and often much less); second submarginal cell

usually longer than wide; seventh metasomal sternum with disc

small, apex bifid or with membranous lobes or both (Africa).

Maxillary palpi reduced to two short, fused segments; mandible of

female slightly longer than eye, distal half a long, flat, pointed

blade, at base of which on inner side are two small teeth; surface

of labrum rather uniformly convex, impunctate except near apical

margin (Eastern North America).

Maxillary palpi six-segmented; mandible of female shorter than eye,

of ordinary form with one subapical inner tooth; labrum with wedge-

shaped, slightly depressed, median punctate area partially dividing

smooth area (Holarctic Region and Africa).

Melitta

Scopa of female consisting of simple bristles; hind tibia and

basitarsus of female slender, the latter three or more times as

long as wide; anterior tarsus ordinary, with ordinary vestiture;

seventh metasomal sternum of male with apex weakly emarginate, at

each side of apex a large, flat, vertically expanded process with

long erect hairs on outer surface.

SP aah cag aerate ahaa it nt vba ig a oedagiog gunner lglg 8 geet are gg Redivivoides

Scopa of female with densely plumose understory beneath long simple

bristles; hind tibia and basitarsus of female broad, the latter

about twice as long as broad; anterior tarsus thickened or

elongated, with dense, short vestiture; seventh metasomal sternum

of male with apex deeply bifid or with two long, slender apical

processes, at each side lobes, if present, hairless (although

spiculate), not flattened, but coarsely striate or ribbed.

Rediviva

Genus Melitta Kirby

(Figs. 60-65, 70-76)

Melitta Kirby, 1802, p. 117.

Type species: Melitta trincincta Kirby, 1802, by designation of

Richards, 1330, 0. Iie;

Cilissa Leach, 1815, p. 155.

Type species: Andrena haemorrhoidalis Fabricius, 1775, by designation

of Westwood, 1840, p. 84.

38 Contrib. Amer. Et ainsts} vol. 187 no: 3, 1981

Kirbya Lepeletier, 1841, p. 145 (preoccupied).

Type species: Melitta tricincta Kirby, 1802, by designation of

Sandhouse, 1943, p. 561.

Pseudocilissa Radoszkowski, 1891, p. 241.

Type species: Cilissa robusta Radoszkowski, 1876 (= Melitta dimidiata

Morawitz, 1876), monobasic.

‘Brachycephalapis Viereck, 1909, p. 47.

Type species: Melitta (Brachycephalapis) californica Viereck, 1909, by

original designation and monobasic.

Diagnosis. Melittine bees with three submarginal cells; mouthparts

ordinary for the group; scopa on outer sides of hind tibia and basitar-

sus of female simple, these segments slender; propodeal triangle large,

dull; seventh metasomal sternum of male with large disc and insig-

nificant lateral apical lobes.

Description. a. Face black. b. Blade of galea with or without

long marginal hairs, often intermediate with rather short hairs; galeal

comb short, with less than eight bristles, often only two or three large

ones. c. Ligular arm of labium long, base near base of prementum or

sometimes separated from latter by about width of prementum, distal part

of ligular arm sometimes weakly sclerotized. Labial palpus of ordinary

elongate form. d. Labrum not much if any over three times as wide as

long, not much narrowed laterally, with lateral apical lobe so that

apical labral margin is often concave. Mandible of ordinary form with

preapical tooth. e. Surface of labrum with impunctate, slightly

elevated basal area partially divided medially by triangular, wedge-

shaped, slightly depressed punctate area which is broad apically and

pointed basally. f. Malar space linear to as long as broad. g.

Propodeal triangle dull, granular or minutely rugose; large, extending

at least half way from anterior propodeal margin to metasomal attachment

and width at upper margin at least nearly equal to distance between

transmetanotal sutures, upper margin only slightly less steeply inclined

than the rest. (In M. nigricans Aflken triangle markedly narrower than

distance between transmetanotal sutures.) h. Forewing with three sub-

marginal cells [rarely two as individual variant (Sakagami, 1976)]. i.

Stigma slightly more slender than in Macropis, so that length from base

to base of vein r is at least twice greatest width (scarcely so in M.

dimidiata Morawitz). j. Second submarginal cell wider than long or

about as long as wide [e.g., in M. americana (Smith)]. k. Basal vein

of forewing less than twice length of first abscissa of Rs and distal

to cu-v or interstitial with it. |. Jugal lobe of posterior wing

slightly less than to slightly more than half length of vannal lobe. m.

Hind tibia and basitarsus of female slender, basitarsus three or more

times as long as broad, ending in small process above base of second

tarsal segment. n. Tibial and basitarsal scopa (on outer sides of

these segments) consisting of simple bristles, sometimes marginal ones

branched. o. Hairs of apex of hind basitarsus of female not parted,

the apical process being merely hairy. p. Hind basitarsus of male

slender, without comb; apex not conspicuously truncate, without projec-

tion below base of second segment. q. Fore and mid tarsi of female not

Michener: Classification of Melittidae 39

Cor Zu

Figures 60-69. Mouthparts of Melittinae. 60, 61, Posterior and an-

terior veins of labium of Melitta leporina (Panzer). 62, 63, Lateral

views of portions of labium of same, positioned relative to Figure 61,

anterior surface upward. 64, 65, Outer and inner views of maxilla of

same. 66, 67, Anterior view of labium and inner view of maxilla of

Dolichochile melittoides Viereck. 68, 69, Same, of Macropis europaea

Warncke. Scale lines = 0.5 mm; the scale is the same for all figures of

any one species.

40 Contrib, Amer. Ent. Inst.,-vol. 18, no. 3, 1981

thickened, without dense, velvety hairs; small segments somewhat more

robust than those of hind tarsi. r. Pygidial plate of male absent. s.

Seventh metasomal sternum of male with disc or body of sternum large,

almost as in preceding sterna; apex broadly truncate or medially emar-

ginate; apical lobes reduced to small sclerotic structures and sometimes

hair patches at posterior lateral angles of sternum. t. Eighth

metasomal sternum of male with apical process ending in a well defined

bevelled area, broader than long or as broad as long, suggestive of a

pygidial plate; base of eighth sternum a rounded or pointed spiculum.

u. Gonostylus robust, rather broadly fused to gonocoxite. v. Volsella

with digitis extending much farther posteriorly than cuspis as a

parallel-sided or capitate, blunt process, usually with teeth along

outer margin but these not opposable to cuspis.

Comments. Melitta is probably near the ancestral melittine group

The other genera of the subfamily may have arisen from forms that would

be placed near Melitta if they were extant. Character s is a synapomor-

phy of Melitta (shared with Dolichochile), if the small bodied seventh

sternum of the male with lateroapical lobes (found in most colletids

panurgines, dufoureines, ctenoplectrids, fideliids, and some

anthophorines, as well as in Meganomiinae and other Melittinae) is the

ancestral state. While this seems likely, a broad seventh sternal plate

is characteristic of sphecoid wasps and it is possible that that of

Melitta is a plesiomorphy derived from wasp ancestors. Character v is

more clearly a synapomorphy of Melitta (with Dolichochile), the opposed

or more nearly opposed digitus and cuspis of all other Melittinae being

plesiomorphic relative to the condition in Melitta. The opposed digitus

and cuspis is characteristic of most Hymenoptera from Symphyta to the

short-tongued bees. The reduced galeal comb (part of character b) is

unusual among related groups of bees and is a probable synapomorphy.

The genus Melitta appears to be most abundant and diverse in the

Palearctic Region, within which it ranges from England, Morocco, and the

Canary Islands to Japan. It occurs also in the Nearctic Region, with

only two species, M. americana Smith in the eastern states, and M.

californica Viereck, in the desert of southernmost California and Baja

California. Otherwise the genus is found only in Africa, where M.

capensis Friese (first described as a variety of the Palearctic M.

dimidiata Morawitz), M. albida Cockerell and others are found in the

arid southern part of the continent. Probably the African species

listed by Cockerell (1935) fall in this genus as currently interpreted.

| have not seen material of this genus from the equatorial parts of

Africa.

The genus seems moderately diverse but | see no need to subdivide

it, as have Radoszkowski (1891) and Baker (1965). Both of these authors

have placed Melitta dimidiata Morawitz in a separate genus,

Pseudocilissa. The distinctive features of M. dimidiata Morawitz include

its large size, long bristles on the lacinia and adjacent basal galeal

margin, the dense long hairs on the under surface of the apical process

of sternum VIII of the male (approached in M. capensis), the robust and

complex penis valve with a strong lateral projection, and the shape of

sternum VI! of the male which is broadest apically instead of basally.

Such features do not seem to justify generic recognition for a.single

species, especially since various other species exhibit almost equally

unusual features. Thus M. capensis Friese (= longicornis Friese) has

two apical processes on the gonostylus, one ventral and recurved, one

Michener: Classification of Melittidae 41

lateral and straight, and basal to these processes a large ventral

pocket from which project long plumose hairs. The fringe of the galeal

blade in this species consists of short hairs only. In the similarly

large M. californica Viereck the apex of the gonostylus is attenuate and

down-curved, and the galeal blade almost lacks a fringe. Among smaller

species, M. nigricans Alfken is unusual in the rather small propodeal

triangle. M. haemorrhoidalis (Fabricius) has an unusually sclerotized

and sharply pointed galeal blade without a fringe. As indicated in the

description above, the malar area is variable. Usually it is short,

nearly linear, often longer above the posterior mandibular articulation

than above the anterior one. However, in M. capensis Friese from South

Africa and the otherwise unrelated M. caroli (Cameron) (described as an

Andrena) and M. altissima Cockerell from the Himalayan Region, the malar

area is about as long as broad.

Warncke (1973) divided the west-Palearctic species into two sub-

genera, Melitta and Cilissa, but gave no characters or reasons for this

arrangement. He failed to mention the name Pseudocilissa but included

its type species in Cilissa. Wu (1978) has reviewed the numerous Chinese

species.

Papers on the nesting biology of Melitta include Malyshev (1923)

and Tirgari (1968).

Genus Dolichochile Viereck

(Figs. 66, 67, 77-79, 97)

Dolichochile Viereck, 1909, p. 49.

Type species: Dolichochile melittoides Viereck, 1909.

Diagnosis. As in Melitta, differing most conspicuously by

characters of mouthparts underlined below.

Description. Agrees with the description of Melitta except as fol-

lows: b. Blade of maxilla with short fringe of rather short hairs (as

in some Melitta); maxillary palpus reduced to two fused segments, total

LS TS A ES A LT cree

fold on which well developed combs usually arise. c. Base of ligular

arm separated from base of prementum by about width of latter (as in

some Melitta). Labial palpus short and robust, fourth segment and

XS AS AS A

ES A ES A A I Kn CMACen

uniformly convex, impunctate except near apical margin. f. Malar space

nearly as long as broad (as in some Melitta). j. Second submarginal

Comments. This genus consists of a single species found in the

Atlantic Coast states of North America. It is plainly a specialized

derivative of Melitta, probably of the group of Melitta americana

(Smith) with which it shares many characters. Recognition of

Dolichochile at any taxonomic level (genus or subgenus) makes the

equivalent taxon Melitta (genus or subgenus) paraphyletic. | do not

reject paraphyletic taxa merely because they are paraphyletic. In this

42 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

case | believe recognition of Dolichochile as a genus or subgenus is

justified by the extraordinary characters listed above under b, c, and

d. The short palpi are unlike those of any other melittid. Other

melittids except perhaps Paramacropis have six-segmented maxillary pal-

pi; in Dolichochile they are two-segmented, and much shortened, so that

the two segments together are little if any longer than the first seg-

ment in other melittids. The proboscis as a whole seems somewhat

reduced, as indicated not only by the greatly reduced palpi, but also

by the small mentum and lorum, greatly reduced galeal comb, and the

like. The coarsely annulate paraglossae are another unusual feature;

more species of Melitta should be examined to determine the constancy

of lack of annuli as a character in that genus. The long, blade-like

mandibles of the female are unlike those of any other bee. The labrum

(character e) is similar to that of Rediviva; it is likely that this

resemblance is a result of convergence. :

Redivivoides Michener new genus

(Figs. 80, 82-84, 97)

Type species: Redivivoides simulans new species.

Diagnosis. Melittine bees with three submarginal cells; mouthparts

ordinary for the group; front and middle tarsi of female as in Melitta,

not thickened or densely hairy as in Rediviva; scopa on outer sides of

hind tibia and basitarsus of female short and simple, these segments

slender; propodeal triangle small, ill-defined, shining; seventh

*metasomal sternum of male with reduced disc, non-bifid apex, large and

vertically expanded, hairy, membranous lobe on either side of apex;

seventh metasomal tergum of male with weak pygidial plate.

Description. Agrees with description of Melitta except as follows:

b. Blade of maxilla with fringe of long hairs; galeal comb long, con-

sisting of over 15 bristles. c. Ligular arm of labium short, base

separated from base of prementum by more than width of prementum, apex

not extending into distal third of prementum, thus even more abbreviated

distally than in Macropis. d. Labrum narrowed and rounded laterally,

without lateral apical lobe, apical margin therefore convex. e. Sur-

face of labrum largely smooth, convex, without median, wedge-shaped,

depressed, punctate area. f. Malar space between one third and one

fourth as long as wide. g. Propodeal triangle shining, poorly defined,

small, not extending half way from anterior propodeal margin to

metasomal attachment and width about one third of distance between

transmetanotal sutures. j. Second submarginal cell a little longer

than wide. k. Basal vein of forewing over twice as long as first ab-

scissa of Rs, slightly to distinctly basal to cu-v. |. Jugal lobe of

posterior wing about two fifths as long as vannal lobe. m. Hind tibia

and basitarsus of female slender, basitarsus about or slightly more than

three times as long as broad, apex rounded above and projecting slightly

beyond base of second segment. n. Tibial and tarsal scopa (on outer

side of these segments) consisting of rather short, simple bristles.

(Anterior margin of hind tibia of female shining with only scattered

long hairs arising from it, thus unlike other genera.) r. Seventh

Michener: Classification of Melittidae 43

hee”

Figures 70-92. Melittinae, males. 70-72, Metasomal sterna VI! and

Vill and genitalia (dorsal and ventral), Melitta tricincta Kirby. 73-75,

Same structures and sequence, M. dimidiata Morawitz. 76, Ventral view

of gonoforceps, M. capensis Friese. 77-79, Sterna VII and VIII and

genitalia (dorsal and ventral), Dolichochile melittoides Viereck. 80,

81, Lateral views, sternum VII, Redivivoides simulans n. sp. and

Rediviva peringueyi (Friese). 82-84, Sterna VII and VIII and genitalia

(dorsal and ventral), Redivivoides simulans n. sp. 85-87, Same struc-

tures and sequence, Rediviva peringueyi (Friese). 88-90, Same structures

and sequence, R. colorata n. sp. 91, Lateral view of genitalia of same.

92, Sternum VII (dorsal and ventral), R. rufocincta (Cockerell). Scale

lines = 0.5 mm; the scale is the same for all figures of any one

species.

44 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

metasomal tergum with slightly elevated, bare pygidial plate, obtusely

angulate posteriorly, much broader than long. s. Seventh metasomal

sternum of male with disc or body rather small; apex slightly emar-

ginate, each side with a large, rounded, vertically expanded process

which is concave and covered with long, erect hairs on lateral surface,

this lobe not striate. t. Eighth metasomal sternum of male with apical

process ending in a bevelled area which is longer than broad and not

defined basally, apex not notched. u. Gonostylus slender basally but

fused to gonocoxite. v. Volsella with digitis directed laterally, not

posteriorly as in Melitta.

Comments. This genus is closely related to Rediviva and could

represent the ancestral stock from which that genus arose. Compared to

Rediviva it possesses a series of presumably plesiomorphic characters

that suggest Melitta. These are characters m, n, 0. q, and the lack of

a deeply bifid apex of the seventh sternum of the male (part of s). In

most other features Redivivoides is like Rediviva. The front legs of the

female (character q) are like those of Rediviva rufocincta (Cockerell )

(Fig. 93) except that the small segments of the tarsus are slender,

sparsely haired. Characters m, n. 0. and q as they occur in Rediviva

and Macropis are presumably related to oil collecting. The alternatives

found in Redivivoides suggest that this genus is not an oil collecting

bee. For bees and melittids as a whole, oil collecting is apomorphic—

hence the presumption that these characters are synapomorphous for

Macropis and Rediviva and plesiomorphous for Redivivoides. Another in-

terpretation is possible, however, i.e., that oil collecting and oil

collecting structures have been lost in Redivivoides, as they appear to

have been in some groups of Centris (Anthophoridae) (J. Neff and B.

Simpson, personal communication).

The presence of a weak pygidial plate (character r) differentiates

this genus from both Melitta and Rediviva. It must be plesiomorphic for

bees, but see discussion under Macropis. The form of the volsella

(character v) is also plesiomorphic for bees, with Rediviva and Melitta

both apomorphic. Such characters suggest that Redivivoides may be a

member of a group from which both Rediviva and Melitta are derived.

The large, vertically flattened (i.e., laterally compressed), hairy

lateroapical lobes of sternum VII of the male probably constitute an

apomorphy for Redivivoides. Such lobes are less developed or even absent

in Rediviva and are hairless, ribbed, and spiculate, not vertically

flattened; | know of no other bees in which they are shaped and

positioned as in Redivivoides.

Redivivoides is known only from western Cape Province, South

Africa. | have seen three species, two of them represented by only one

or two female specimens in the collections known to me; the third, for-

tunately, was collected in good numbers and is named in Appendix |.

Etymology. Rediviva, a related melittid genus, plus-oides,

resembling.

Genus Rediviva Friese

(Figs. 81, 85-97)

Rediviva Friese, 1911, p. 671.

Type species: Andrena (Rediviva) peringueyi Friese, 1911, by designa-

tion of Cockerell, 1931b, p. 402.

Michener: Classification of Melittidae 45

Notomelitta Cockerell, 1933b, p. 128 (new synonym).

Type species: Rediviva neliana Cockerell, 1931b, by original designa-

tion and monobasic.

Diagnosis. Melittine bees with three submarginal cells; mouthparts

ordinary for the group; anterior tarsus of female thickened, often also

elongate, with short, dense vestiture; scopa on outer sides of hind

tibia and basitarsus of female densely plumose, hiding surface, with

long, simple emergent hairs; propodeal triangle small, shining; seventh

metasomal sternum of male with reduced disc and usually with

lateroapical lobes, apex deeply bifid.

Description. Agrees with description of Melitta except as follows:

b. Galeal blade with moderately long marginal hairs. c. Ligular arm

of labium short, base variable but apex not extending into distal third

of prementum, thus even more abbreviated distally than in Macropis. d.

Labrum narrowed and rounded laterally, without lateral apical lobe,

apical margin therefore convex. e. Surface of labrum largely smooth

[or in R. rufocincta (Cockerell) distal two thirds punctate], convex,

without median, wedge-shaped, depressed, punctate area. g. Propodeal

triangle shining, usually poorly defined, minute to small, often not ex-

tending half way from anterior propodeal margin to metasomal attachment

and width not over half distance between transmetanotal sutures, often

much less. j. Second submarginal cell usually longer than wide, in R.

colorata new species about as wide as long. k. Basal vein of forewing

usually twice as long as first abscissa of Rs, slightly distal to

strongly basal to cu-v. |. Jugal lobe less than half as long as vannal

lobe, sometimes only one third as long. m. Hind tibia and basitarsus

of female broad, basitarsus about twice as long as broad, ending in a

broadly rounded or subtruncate apex above base of second tarsal segment.

n. Tibial and basitarsal scopa (on outer sides of these segments) with

a mat of strongly plumose hairs obscuring surface, from which emerge

long simple bristles above the plumosity. o. Hairs of apex of hind

basitarsus of female parted as in Macropis. q. Fore tarsi of female

with small segments thickened and largely or entirely covered with short

dense hairs, sometimes spatulate, sometimes plumose; these features

sometimes somewhat developed on mid tarsus; front tarsus of female often

elongated, and entire front leg in that case unusually slender. s.

Seventh metasomal sternum of male with disc or body rather small, apex

deeply bifid, a simple, membranous, lateral lobe before apex except in

R. colorata new species, this lobe with appressed hairs or bristles on

concave outer surface. t. Eighth metasomal tergum of male with apical

process ending in a bevelled area which is longer than broad and not

defined basally, apex often notched. v. Volsella with digitis ex-

tending posterior to cuspis as a tapering projection, shorter than in

Melitta, with teeth along inner margin more or less opposable to those

of cuspis.

Comments. Rediviva was originally described as a South African

subgenus of Andrena. There exist species of Andrena in the Cape Region

of Africa that superficially resemble the type species of Rediviva. As

a result various authors (including Cockerell, 1933b) placed Rediviva

in the Andrenidae, as a synonym or subgenus of Andrena, and Cockerell

named the melittid group as Notomelitta in the belief that Rediviva was

an Andrena. However, | have examined the type specimen of the type

species of Rediviva from the Berlin museum. It is a melittid, clearly

46 Contrib. Amer. Ent. inst., vol. 18, no. 3, 1981

a member of the group described above, and Notomelitta therefore falls

into the synonymy.

The species of Rediviva look like those of Melitta, particularly

resembling superficially the large species of Melitta such. as M.

capensis Friese. The most distinctive generic characters are ed,e,g,

m, n, 0, and s.

ke \a ,

Min,

ON ea

“\

»)

NS iy

nn AVIAN ae,

= 93 ve a

hey : Wyn tp

Vi ae

Figures 93-96. Front legs of female Rediviva. 93, R. rifocincta

rocconggeenssk 94, R. colorata n. sp. 95, Rs _ peringueyi agree ). 96,

longimanus n. sp. Scale line = 1.0 mm.

An interesting character of the genus is the thickening and dense

pubescence of the fore and often mid tarsi of females (character q).

This feature, together with characters m, n, and o, are in general

agreement with Macropis and suggest that Macropis and Rediviva are

closely related; perhaps these characters are synapomorphies related to

oil collecting although nothing is known about that habit in Rediviva.

Whatever the function of the fore tarsal pubescence of females, it must

be exaggerated in females of certain species which have these tarsi and

indeed the whole front leg greatly elongated (Fig. 96). These are forms

in which the dense hairs of the fore tarsi are plumose. Species of this

kind include R. peringueyi (Friese), a species near R. politissima

event and neliana Cockerell, R. colorata new species and R.

longimanus new species, the last showing the most extreme tarsal elonga-

tion. By contrast, some other species have front tarsi of normal

length, with the pads made of dense spatulate rather than plumose hairs.

R. rufescens (Cockerell) is such a species.

Another synapomorphy of Rediviva (with Redivivoides) is character

g, the poorly defined, small propodeal triangle. Since the triangle is

large in most short-tongued bees including most melittids, it is

presumably a derived feature in Rediviva. The character is possibly the

same as the "hairy propodeal triangle" of the subgenus Sinomacropis of

Macropis. In Sinomacropis there is a small hairless basomedian area.

Although undefined, this could be the triangle proper, in which case the

large hairy area is not part of the triangle, and the situation would

Michener: Classification of Melittidae 47

be the same as in Rediviva. | did not interpret the homologies in this

way in describing ‘Sinomacropis s because of the similarities in contours

of the hairy areas of that subgenus to those of the large triangle of

Macropis s. str.

Rediviva consists of a number of species, most of which are found

in temperate South Africa although R. tropicalis from Zaire probably

belongs here. Some of the species are described and keyed by Cockerell

(1934). Species that fall in the genus are: R. neliana Cockerell,

1931b; peringueyi (Friese, 1911); politissima (Cockerell, 1934);

longimanus new species; intermixta (Cockerell, 1934); tropicalis

(Cockerell, 1934); colorata new species, and rufocincta (Cockerell,

1934). 1 have not seen specimens of R. intermixta and tropicalis.

Probably some other African "Melitta" also belong here. | have seen

about eight species of the genus from South Africa, but very few

specimens of each, and some of them not identified.

Genus Macropis Klug

(Figs. 68, 69)

Diagnosis. Melittinae with two submarginal cells; clypeus of male

yellow; pygidial plate of male well developed.

Description. a. Male with clypeus, lower paraocular areas, and

supraclypeal area yellow or partly so. b. Blade of galea margined with

long hairs; galeal comb long, consisting of over 15 bristles. c.

Ligular arm of labium short, base and apex not approaching base and apex

of prementum. d. Labrum of male five to six times as wide as long, of

female about four times, in both sexes narrowed laterally so that apical

margin is convex, ends narrowly rounded, the whole lens-shaped. e.

Surface of labrum impunctate. f. Malar area linear. g. Propodeal

triangle variable (see subgenera). h. Forewing with two submarginal

cells. i. Stigma slightly broader than in most Melittini so that

length from base to base of vein r is less than twice greatest width.

j. Not relevant because there are only two submarginal cells. k.

Basal vein twice length of first abscissa of Rs or more, variable in

relation to cu-v. |. Jugal lobe of posterior wing half as long as van-

nal lobe or longer. m. Hind tibia and basitarsus of female broad,

basitarsus little over twice as long as broad, broadly truncated at

about level of base of second tarsal segment, and with flattened, hair-

less, process projecting posteriorly from upper end of truncation (this

process absent in M. patellata Patton). n. Tibial and basitarsal scopa

of strongly plumose hairs with apices of some emerging as simple

bristles above plumosity. o. Hairs of apical truncation of hind

basitarsus of female, above base of second tarsal segment, parted (i.e.,

diverging) to expose narrow strip of integument along basitarsal apex.

p. Hind basitarsus of male usually thickened, usually with coarse comb

on under surface at least near base; apex subtruncate, often obliquely

so, usually with short, bare projection below base of second segment.

q. Fore and middle tarsi of female somewhat thickened and covered with

dense, velvety hairs; small segments markedly more robust and also more

densely hairy than small segments of hind tarsi. r. Pygidial plate of

48 Contrib. Amer. Ent..tnst., vol. 78) now 3,..1981

Figure 97. Melittinae. Top row, left to right, faces of

Dolichochile melittoides Viereck, female; Redivivoides simulans n. sp.

male, female holotype. Middle row, left to right, faces of Rediviva

colorata n. sp., female holotype, male; R. longimanus n. sp., female

holotype. Bottom row, left to right, forewings of Dolichochile

iva

longimanus

melittoides Viereck, Redivivoides simulans n. sp., Rediv

n. sp., holotype. Scale line = 0.5 mm.

Michener: Classification of Melittidae 49

male strongly developed, greatly elevated above adjacent tergal surface.

s. Seventh metasomal sternum with disc cr body of sternum small; apical

lobes on each side large, complexly folded, setose. t. Eighth

metasomal sternum with apical process not ending in bevelled area sug-

gestive of a pygidial plate; base of eighth sternum truncately produced,

spiculum not projecting. u. Gonostylus slender at base, usually ex-

panded and bilobed or bifid apically, distinctly articulated to gonocox-

ite. v. Volsella with digitis and cuspis of about equal length,

toothed along opposing margins.

Comments. The characters of females, as well as those of mouth-

parts, and the comb mentioned under item p, have not been verified for

the subgenus Paramacropis.

Possible apomorphic characters for this genus include a and h.

Both of these are features that appear in many other bees. A yellow

clypeus (part of character a) even occurs in one genus of Dasypodinae

(Promelitta) and two submarginal cells (character h) characterizes the

Dasypodinae. It is not clear whether these features arose in an an-

cestral Macropis or in a common ancestor of that genus and Dasypodini.

Characters m, n, 0, and q, are unusual features, not found in other

bees except Rediviva. As noted in discussing the subfamily, they are

complex features; it seems unlikely that they arose twice. They could

be synapomorphies for the subfamily, secondarily lost in those that lack

them. On the other hand, they could have arisen within the Melittinae,

one group of which (Rediviva) then gave rise to Macropis. But if this

were true, how does it happen that Macropis possess plesiomorphies like

character r and the articulated gonostyli of u, not possessed by any

other Melittinae? As to character r, the pygidial plate is doubtless

a plesiomorphy in bees, being derived from that of sphecoid wasps. It

is present in all female melittids. It is possible that it was lost in

males, but since genes for it are present it could have been restored

in males of Macropis and feebly in Redivivoides (as well as some

Hesperapis in the Dasypodinae) by activating in the males genes that had

been functioning only in females. For character u (articulated bases

of gonostyli, likewise, one can argue that the (for Hymenoptera)

plesiomorphy of a freely articulated gonostylus may for Macropis be a

reversion and thus a synapomorphy, for there is enormous variation in

this feature among bees, and in fact the condition is found also in some

Dasypodinae which in other features have no special similarity to

Macropis. In short, we have no clear bases for choosing between these

parts of the cladograms shown in Figures 1 and 2, or between those and

a third alternative involving characters 48, 49, 51, and 52 on the stem

for the subfamily with reversals for those characters on the stems for

Melitta (with Dolichochile) and Redivivoides.

An adequate analysis of the subgenera of Macropis is not possible

since Paramacropis is not known to me in the female sex. In couplet 2

of the following key, the female of Paramacropis would presumably run

to Macropis s. str.

Key to the Subgenera of Macropis

1. Gonostylus of male slender, simple; sternum VI of male with broadly

truncate apical process; sternum VII! with apical process much

broader than body of sternum, densely covered with long hairs.

50 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

(Female unknown; found in maritime province of Siberia).

sos waar a pa sia ca Ramet alee ead cl Chas «Rah Ao yo a Paramacropis

-- Gonostylus of male apically expanded and bilobed or bifid; sternum

Vi of male with apex attenuate; sternum VIII of male with apical

process much narrower than body of sternum, apically pointed, with

hairs of moderate length.

2. Propodeal triangle punctate, hairy except for small median area

(China).

sea Ste aE" Bg RR ay hea ew ai oi 7 Nd ao ea take ig Sinomacropis

-- Propodeal triangle impunctate, hairless, either smooth or rugose

(Holarctic).

De ED, erage et my aN Ae De ee eas ae re et mae ng ie Macropis s. str.

Subgenus Macropis Klug s. str.

Megilla Fabricius, 1805, p. 328. |

Type species: Megilla labiata Fabricius, 1805, designated by Westwood,

1840, p. 158. A subsequent designation by Richards (1935, p. 172)

of Apis acervorum Linnaeus, 1758, would make Megilla a synonym of

Anthophora. )

Macropis Klug, in Panzer, 1809, pt. 107, no. 16.

Type species: Megilla labiata Fabricius, 1804, monobasic.

Diagnosis. Propodeal triangle large, hairless; gonostylus of male

expanded apically, bifid or bilobed.

Description. a. Labrum of female with a transverse ridge across

it. b. Maxillary palpus six-segmented. c. Propodeal triangle large,

hairless, smooth or transversely wrinkled, not punctate. d. Basal vein

interstitial with or basal to cu-v. e. Hind basitarsus of male with

comb of stiff bristles on underside, extending nearly the full length

of the segment. f. Hind basitarsus of male less than half as long as

tibia, less than four times as long as broad, with ventroapical projec-

tion bilobed. g. Sixth sternum of male with apex attenuate, down

curved, with retrorse hairs. h. Eighth sternum of male with apex a

narrow, acutely pointed process, somewhat constricted preapically. |.

Male gonostylus narrow at base but broadened apically, strongly bifid.

Comments. This subgenus is a widespread visitor of Lysimachia in

eastern North America, west to eastern Washington State, and across

Eurasia from western Europe to Japan (Popov, 1958). It probably col-

lects oil as well as pollen from this plant (Vogel, 1976). Its nesting

biology was described by Malyshev (1929) and Rozen (1980). The west-

Palearctic species were reviewed by Warncke (1973).

Illustrations of male genitalia and associated structures have been

provided by Saunders (1882), Mitchell (1960), and Michener (1938).

Benson, Ferriére and Richards (1947) and Michener (submitted) have

requested suspension of the rule of priority for the name Megilla and

preservation of the well established name Macropis for this group of

bees.

Michener: Classification of Melittidae 51

Sinomacropis Michener new subgenus

Type species: Macropis hedini Alfken, 1936.

Diagnosis. Propodeal triangle hairy (or greatly reduced in size);

gonostylus of male expanded apically, bilobed or bifid.

Description. Agrees with that of Macropis s. str. except as fol-

lows: a. Labrum of female evenly convex, as in male. c. Propodeal

triangle shining, with well separated punctures and with hairs except

on small, ill-defined median area (which is possibly the triangle

proper). d. Basal vein distal to cu-v. e. Hind basitarsus of male

with comb of stiff bristles on under side of base only. f. Ven-

troapical projection of hind basitarsus of male not bilobed.

Comments. Alfken (1936) as well as Popov and Guiglia (1936) sug-

gested that Macropis hedini could fall in a new genus. The males of the

group, here described for the first time, show the close similarity to

Macropis s. str., but subgeneric rank is justified. This subgenus is

known from at least three species, all from Sichuan (=Szechuen)

Province, China, only one of them described. All are represented by

specimens in the U. S. National Museum. The two japanese species of the

genus fall in Macropis s. str. to judge by the descriptions (Yasumatsu

and Hirashima, 1956).

The most noteworthy subgeneric character of Sinomacropis is the

punctate and hairy propodeal triangle (hairless only in a small median

area). Among bees the hairy vs. hairless propodeal triangle is usually

a conservative variable, characteristic of families or groups of

families; most short-tongued bees have a hairless triangle. A hairy

propodeal triangle is not found in any other melittids. |

Other features of the subgenus, not mentioned in the above descrip-

tion, include more extensive yellow areas than in other Macropis. In M.

hedini the supraclypeal area of the female has a yellow spot, and in

males of all species there are small to extensive yellow areas on the

femora, tibiae, and tarsi. In some males the posterior margins of the

first few metasomal terga are dull yellowish.

The female of M. hedini has the first two metasomal terga largely

red, a feature not found in the male or in the other species of the

genus. M. hedini was originally described from the female only. Popov

(1958) described what he believed to be the male. However, it appears

that he had the male of another species of the subgenus, as shown, for

example, by the long, thin, mediotarsal segments of the middle leg

(Popov, 1958, Fig. 1). Such a middle tarsus is characteristic of

another species but not of the male associated with M. hedini in the U.

S. National Museum (Washington). Popov's illustrations adequately

represent the subgeneric characters of the sterna and genitalia, even

though probably not correctly attributed to the type species of the sub-

genus.

Etymology. Sino- pertaining to China; plus the generic name

Macropis.

Subgenus Paramacropis Popov and Guiglia

Paramacropis Popov and Guiglia, 1936, p. 287.

52 Contrib. Amer... Ent:.:}fist.; vol. 18, no. 3, 1981

Type species: Ctenoplectra ussuriana Popov, 1936, monobasic.

Diagnosis. Male gonostylus simple, not expanded or lobed apically.

(Female unknown to me. )

Description [based entirely on Popov's (1936) description and

figures of the male]. Agrees with Macropis s. str. except as follows:

b. Maxillary palpus five-segmented. c. Propodeal triangle hairless,

smooth. d. Unknown. f. Hind basitarsus of male over half as long as

tibia, over four times as long as broad, probably without comb and ven-

troapical projection. g. Sixth sternum of male narrowed preapically,

then broadened to truncate, hairy apex. h. Eighth sternum of male with

apex greatly broadened behind preapical constriction, much broader than

any other part of sternum, under (?) side of process densely covered

with long hairs. i. Male gonostylus slender, simple.

Comments. This subgenus, from the maritime province of Siberia, is

markedly different from other Macropis in characters g, h, and i.

Character b should be verified, since nearly all melittids have six-

segmented maxillary palpi. |

Subfamily Dasypodinae

Diagnosis. Differs from other subfamilies in having paraglossae

that, if present, are largely bare, with hairs usually present only

apically; except for Promelitta, paraglossae that are much shorter than

their suspensoria (or absent); two submarginal cells, the second usually

shorter than the first; and base of second submarginal cell (first

transverse cubital vein) usually at nearly right angles to the

longitudinal veins on either side and close to the first recurrent vein.

Description. Agrees with description of Melittinae except as fol-

lows: a. Body minute (4mm long) to rather large (14mm long), without

yellow markings except for some Eremaphanta and the yellow clypeus of

male Promelitta and the partly yellow clypeus of the male Hesperapis

rufipes. c. Paraglossa absent or much shorter than suspensorium except

in Promelitta, largely hairless but with some apical hairs, not an-

nulate. e. Labrum frequently with strong transverse ridge (otherwise

as in Melittinae). f. Frontal line frequently a groove on a slightly

to strongly convex region between antennal bases. g. Malar area linear

or nearly so, about as long as width of first flagellar segment in some

Sambini. h. Clypeus usually without longitudinal median ridge, but with

strong ridge in Samba. j. Forewing with two submarginal cells [three

on one side only of a specimen of Eremaphanta convolvuli (Popov) ],

second shorter than first (subequal to first in some Sambini), strongly

narrowed toward costal margin of wing. Base of second submarginal cell

usually nearly at right angles to longitudinal veins, close to first

recurrent vein. k. Basal vein 1.3 to 3.5 times as long as first ab-

scissa of Rs. |. Stigma less slender than in Meganomiinae, large and

broad in Eremaphanta, prestigma one fifth to over two thirds as long as

stigma; vein r arising near middle of stigma or (in some Eremaphanta and

Dasypoda) at apical third; sides of stigma basal to vein r usually con-

verging toward base of stigma, only slightly so in some, margins paral-

lel in a few. m. As in Melittinae or apex of marginal cell not at all

bent away from costal margin of wing (Eremaphanta). n. Jugal lobe of

hind wing one fourth to over three fourths as long as vannal lobe. o.

Michener: Classification of Melittidae 53

Mid and hind basitarsi of female narrower than tibiae. p. Hind

basitarsus of female usually tapering slightly distally, apex usually

narrow and giving rise to segment 2 medially but in Promelitta and the

Sambini apex broader, giving rise to segment 2 at lower margin, apex

above base of segment 2 sometimes produced distally, hairs on apical

margin not parted as in Macropis. q. Metasomal hair bands absent in

some Eremaphanta, some Hesperapis, and in Sambini; prepygidial and pygi-

dial fimbriae weak in some Dasypodini. r. Depressed apical zones of

terga sometimes narrower than in Melittinae. s. Lateral extremities of

tergal graduli not curved posteriorly or only extending a short distance

posteriorly (in Promelitta, about as in Melitta). u. Sternum VII of

male a transverse plate with two apical lobes, not very different from

preceding sterna, or disc reduced with lateroapical lobes.

Comments. The characters listed under c and j above are synapomor-

phies of the subfamily; c is the most reliable. The transverse base of

the second submarginal cell, more or less at right angles to the

longitudinal veins and close to the apex of the second recurrent vein,

is an unusual feature not found in related bees. Unfortunately it is

less than fully satisfying because the difference between transverse (as

in Dasypodinae) and slanting (as in Macropis and the groups with three

submarginal cells) is small and sometimes almost bridged by variation

within genera or even species. Nonetheless, the character is a valuable

one used with care and does in general distinguish most Dasypodinae from

other melittids. Character s also distinguishes Dasypodinae (except

Promelitta) from other Melittidae (except Rediviva and Redivivoides),

but it is not clear whether it is an apomorphy or plesiomorphy. The

other characters listed above do not distinguish all Dasypodinae from

all Melittinae, but are given to indicate ranges of variation.

Larvae of Dasypoda, Capicola s. str. and various subgenera of

Hesperapis are known (Rozen and McGinley, 1974). While Dasypoda is

quite different from the other two genera, all three exhibit several

apomorphies that distinguish them from the Melittinae and Meganomiinae,

and thus support recognition of the subfamily Dasypodinae. Unfortunately

larvae of the tribes Sambini and Promelittini are unknown.

Key to the Tribes of Dasypodinae

1. Vertex seen from front (except in Eremaphanta s. str.) elevated

well above summits of eyes, usually convex; gonostylus of male (ex-

cept in Dasypoda) robust, fused to gonocoxite; sternum VII of male

without or (in some Dasypoda) with a pair of slender, strip-like

lateroapical lobes; paraglossa small, slender, cylindrical.

Siteiaieieaiaieiaiaiaiieiaeieiieet et Dasypodini

-- Vertex seen from front little elevated above summits of eyes,

gently convex to concave; gonostylus of male long, flexibly joined

or articulated to gonocoxite; sternum VII of male with one or two

pairs of broad, lateroapical lobes; paraglossa tapering distally or

absent.

2. Clypeus of male largely yellow; metasomal terga with basal zones of

pale hair; paraglossa nearly as long as its suspensorium; vertex

54 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

weakly, uniformly convex seen from front.

oiheey Seas oe Sate te eet eeu em rite a hae mei oe Promelittini

-- Clypeus concolorous with rest of head; metasomal terga without

basal hair bands, bands if present apical; paraglossa much shorter

than suspensorium or usually not recognizable; vertex straight or

concave seen from front. |

saasianiani aie ean aaa la aoa aaa aatetntnaatetetelatatatataiate tata Sambini

Promelittini new tribe

This tribe, so far as known, contains only a single Egyptian

species which does not readily fall into either of the other tribes.

Another species of doubtful affinities from southern Africa is tenta-

tively included. The principal tribal characteristics are indicated in

Table 2, the apomorphies in Figures 1 and 2. Since Promelitta does not

have the numerous distinctive characters of the Sambini (Figs. 1 and 2),

it is more similar to the Dasypodini than to the Sambini. However, there

are features in which it resembles the Sambini and differs from the

Dasypodini. Some such features are probable plesiomorphies like the

large lateroapical lobes of sternum VII of the male (plesiomorphy for

characters 89 and 90, Table 1); others are apomorphies shown in Figure

In various features Promelitta resembles the Melittinae rather than

other Dasypodinae. The base of the second submarginal cell is equivocal:

as in some Sambini it slopes more than in most Dasypodinae but less than

in Melittinae. The tergal graduli, however, are about as in Melitta; the

free margin of the marginal cell is slightly concave so that the distal

part of the cell is slender, as in Melitta; and the sixth tergum of the

male is not bilobed but has a median, apical, hairy area, as in

Melittinae.

Thus on the basis of similarity, Promelitta could almost be placed

in the Melittinae. The characters indicating it affinities to other

Dasypodinae are indicated in Figures 1 and 2. It is probably an archaic

type, like several other bees of the Palearctic deserts. Perhaps it is

a derivative of Melittinae, survivor of the group from which Dasypodini

and Sambini arose, as suggested by Figure 2.

Genus Promelitta Warncke

(Figs. 98-106; Skétches A-C):

Melitta (Promelitta) Warncke, 1977, p. 59.

Type species: Dufourea alboclypeata Friese, 1900, by original designa-

tion and monotypy.

Diagnosis. Clypeus of male largely yellow; metasomal terga with

basal, not apical, bands of hair; paraglossa nearly as long as its

suspensorium, tapering at apex, largely bare but hairy apically.

Description. a. Body of moderate size (9mm long), moderately

hairy; clypeus of male yellow except for marginal zone all the way

around. b. Head broader than long, nearly as wide as thorax; vertex

convex seen from front. c. Labrum of male over five times as wide

Michener: Classification of Melittidae

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Michener: Classification of Melittidae ws

as long; smooth, shining, convex; hairs only along extreme apical mar-

gin. d. Labrum of female similar to that of male but with indication

of transverse basal convexity. e. Flagellar segments of male mostly

longer than broad but segment 2 much broader than long, 1 shorter than

3, 4, etc. f. Flagellar segments of female mostly broader than long,

but 1 and 10 longer than broad, 2 shortest, about twice as broad as

Figures 98-105. Promelitta alboclypeata (Friese), male. 98, 99,

Inner view of maxilla and anterior view of labium. 100, 101, Sternum

VII (lateral, dorsal, and ventral). 102, Sternum VI (ventral). 103,

104, Genitalia (lateral, dorsal and ventral). 105, Sternum VIII (dorsal

and ventral). Scale lines = 0.5 mm; one scale is for mouthparts, the

other for abdominal structures.

long. g. Maxillary blade little over twice as long as broad, tapering

from near base to apex, with fringe of scattered, rather long hairs;

maxillary palpus extending much beyond apex of blade. h. Galeal comb

well developed. i. Prementum rather robust; ligular arm arising about

a premental width distad to base of prementum, end jagged, about two

thirds of distance from base to apex of prementum. j. Glossa rather

short but with a slender, attenuate apex; labial palpus extending beyond

apex of glossa. k. Paraglossa small, hairy, attenuate apically, as

long as suspensorium. |. Second and third segments of labial palpus

with apices oblique, laterally produced. m. Basal vein 1.5 to 1.7

58 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

times as long as first abscissa of Rs, curved throughout, distal to

cu-v. nn. Stigma of moderate size, less than half as long as costal

margin of marginal cell; vein r arising little beyond middle of stigma;

margins of stigma basal to vein r converging basally; length from base

to base of vein r about twice greatest width of stigma. o. Prestigma

less than half as long as stigma. First recurrent and first transverse

cubital veins well separated. p. Apex of marginal cell pointed,

separated from wing margin by less than a vein width. q. Jugal lobe of

hind wing half as long as vannal lobe. r. Propodeal triangle smooth,

shining, basal zone longer than metanotum, horizontal, curving onto

declivous posterior face of propodeum. s._ Basitibial plate well

defined in both sexes, rather small, rounded apically. t. Scopa rather

dense (but one can see integument through it), of moderately long,

simple hairs, yellow on under side of basitarsus; tibia with scopa-like

hairs except for nearly bare, smooth zone on underside and zone about

half as wide as tibia of keirotrichia with minutely bifid tips on

posterior (inner) surface, these two zones separated by a zone of long

scopal hairs. u. Mid and hind tibial spurs only gently curved

apically, their margins minutely and densely microserrate ("ciliate").

v. Middle and hind basitarsi of female flat, nearly parallel-sided,

slender, over four times as long as broad, somewhat narrower than tibia;

hind basitarsus giving rise to segment 2 below middle of apex, upper

apical angle produced distally as broad, flat blade which is nearly bare

on anterior (outer) surface. w. Middle and hind basitarsi of male

somewhat flattened, slightly tapering; hind basitarsus ending in narrow

truncation with segment 2 arising below middle of apex. x. Distitarsi

somewhat enlarged, distinctly broader than preceding segments. y.

Claws cleft nearly to middle, inner rami much shorter (an inner tooth)

in female than in male. z. Metasomal terga I-1V of females and I-V_ of

males with broad, depressed, finely punctate, minutely hairy apical

zones. aa. Terga without apical or preapical hair bands, but terga II

to V with conspicuous, broad basal bands of white, appressed hair. bb.

Pygidial plate of female well defined, dull, apex rounded, surface with

abruptly elevated longitudinal median zone. cc. Pygidial plate of male

absent. dd. Sixth sternum of male with apex broadly truncate, median

third of apical part slightly elevated and covered by patch of dense

hairs. ee. Seventh sternum of male with rather reduced disc, bifid

apex, and long, pedunculate, lateroapical lobe that is hairy distally.

ff. Eighth sternum of male with strong apical process that is abruptly

truncate to form a plate (like a false pygidial plate) apically. gg.

Gonostylus of male simple, hairy, about as long as gonocoxite, articula-

tion (rather than fusion) to gonocoxite distinct.

Comments. The genus is known from a single Egyptian species,

Promelitta alboclypeata (Friese, 1900), and perhaps from P. piumipes

(Friese, 1912), new combination, a South African species described under

the generic name Rhinochaetula. The latter species (see Sketches A-C)

may represent another (new) genus, perhaps not even in this tribe, but

it is known only from a female specimen in the Berlin Museum; the

characters of the male are therefore unknown. It differs from the

female of P. alboclypeata as described above in the following features:

b. Head even broader, inner orbits diverging below instead of con-

verging. d. Labrum less than four times as long as broad. f. Flagel-

lar segment 1 about 2.5 times as long as broad. g. Maxillary blade

nearly four times as long as broad. j. Apex of glossa not slender, at-

Michener: Classification of Melittidae 59

Figure 106. Promelitta alboclypeata (Friese). Left to right,

faces of female, male type and forewing. Scale line = 0.5 mm, ap-

plicable only to wing.

tenuate. |. Segments of labial palpus not obliquely produced at

apices. p. Apex of marginal cell less sharply pointed than in P.

alboclypeata. t. Keirotrichia absent, the tibia being covered with

scopa-like hairs. Apex of hind basitarsus, on outer side, with oblique

row of close spaced, posteriorly directed coppery hairs extending from

near base of segment 2 to upper margin of basitarsus basal from upper

distal process. y. Claws not so deeply cleft as in P. alboclypeata,

the inner tooth arising beyond middle. z. Basal hair bands restricted

to metasomal terga || and II!. bb. Pygidial plate of female without

elevated median zone.

gO at

ide : LADD

ye ——— :

WAS

Wy S .

NN SW

eh ae

NS <=

ss Z

K\ Ze

Sketches A-C. Promelitta plumipes (Friese). Freehand sketches of

holotype (female). A, Outer view of hind leg. B, Inner hind tibial

spur. C, Face.

60 Contrib. Amer Ent. iist.»-vol.-18,:no0.:3, 1931

It seems quite likely that P. plumipes is not closely related to P.

alboclypeata but until the male 2 of the former can be found, there is no

justification for erecting anew genus for it. It falls better in

Promelitta than in other genera; placing it here at least calls atten-

tion to its lack of close relationship to Capicola, where most of

Friese's other species of Rhinochaetula fall.

Sambini new tribe

This African tribe can be recognized by the characters indicated in

the key to tribes and in Table 2. Its numerous Serena es are also

indicated in Figures 1 and 2.

The short, broad head with a flat or concave vertex, exaggerated in

Samba, tends to be characteristic of the whole tribe. The paraglossae

are greatly reduced, often absent or indistinguishably fused to the

apices of the suspensoria, but in Prosamba they are distinct although

small and tapering apically. The elongate volcellar digitis is sugges-

tive of that of Melitta but presumably arose independently.

The known species of Sambini are all very different from one

another. They may be remnants of a formerly larger fauna.

Key to the Genera of the Tribe Sambini

1. Female with only one hind tibial spur; vertex strongly concave seen

from front; hind basitarsus of male with hairy pocket near base of

outer surface.

-- Female with two hind tibial spurs; vertex not or scarcely concave;

hind basitarsus of male without hairy pocket.

bce el geek ok ie ans (Ree OR ES EG Ne Sek ee Haplomelitta

Genus Haplomelitta Cockerell

Diagnosis. Female with two hind tibial spurs as in most bees; hind

basitarsus of male without large hairy pit on outer surface near base.

Description. a. Body of moderate size (7-14 mm long), not densely

hairy. b. Head about as broad as thorax; clypeus about three times as

wide as long, not or scarcely overhanging base of labrum medially;

minimum distance between eyes about equal to or slightly more than

length of eye in females, slightly to greatly less than length of eye in

males; vertex concave, straight, or feebly convex seen from front. c,

d. Labrum of both sexes about four times as wide as long with a strong,

rounded, transverse ridge; surface basal to ridge smooth, gently convex,

shining, hairless; surface distal to ridge mostly dull, with punctures

and hairs basal to long marginal bristles. e, f. Flagellar segments

variable in relative lengths; see subgeneric descriptions. g. Maxil-

lary blade three to more than four times as long as broad, apex bluntly

pointed, with or without a few long hairs; palpus extending but little

beyond apex of blade. h. Galeal comb present, reduced to three

bristles in Haplomelitta s. str.; flange on posterior side of stipes

Michener: Classification of Melittidae 61

Figures 107-115. Mouthparts of Sambini. 107, 108. Inner view of

maxilla and anterior view of paraglossa of Haplomelitta (Prosamba)

griseonigra n. sp. 109, 110, Inner view of maxilla and anterior view

of labium=(of HH. (Atrosamba) <atra cm. sp. 2.111, «lle, Same, rH.

(Haplomelitta) ogilviei (Cockerell). 113, Lateral view of prementum,

mentum and lorum of Samba calcarata Friese, anterior surface upward.

114, 115, Anterior view of labium and inner view of maxilla of same.

Scale lines = 0.5 mm; scale is the same for all figures of any one

species.

limited to distal part of stipes near base of palpus, in Haplosamba

flange not so evident and perhaps best considered as extending along

posterior side of stipes. i. Prementum not especially slender; ligular

arm full length or moderately shortened, distal part sharp or jagged.

j. Glossa rather attenuate, not or scarcely exceeding labial palpi. k.

Paraglossa absent or unrecognizable, perhaps a robust, tapering struc-

ture at ends of suspensorium, or present as a small tapering projection

in Prosamba. |. Segments of labial palpus with apex not strongly

produced laterally. m. Basal vein two to nearly four times as long as

first abscissa of Rs, weakly curved, basal or distal to cu-v. n. Stig-

ma of moderate size, more than to slightly less than half as long as

costal margin of marginal cell (sometimes so tapered apically that its

62 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

apex and measurements involving it are judgemental); vein r arising at

or a little before middle of stigma; margins of stigma basal to vein r

only slightly converging basally; length from base to base of vein r

slightly more to slightly less than twice greatest stigmal width. o.

Prestigma a little less than to well over one third as long as stigma.

First recurrent vein variable relative to first transverse cubital. p.

As described for Samba. q. Jugal lobe of hind wing one fourth to nearly

one half as long as aie lobe. r. Propodeal triangle shining,

sometimes (Haplomelitta s. str.) roughened by a series of pits or

ridges, defined by weak z strong lines, surface as seen in profile in

about the same plane as rest of propodeum, at a steeper angle relative

to body axis than in Samba. s. Basitibial plate as described for Samba

but surface not always hairy. t. Tibial and basitarsal hairs as

described for Samba but no plumose hairs beneath scopal hairs. u. Hind

tibial spurs two, rather slender and microserrate ("ciliate") except for

modified inner spur of male of Metasamba. Middle tibial spur microser-

rate, sometimes thickened and toward apex strongly curved. v. Middle

and hind basitarsi of female flattened, margins of middle basitarsus

convex so that it is wider medially than at ends; hind basitarsus nar-

rower than middle one, margins nearly straight, parallel, or in

Metasamba hind as broad as middle basitarsus, margins slightly convex

and converging distally. Apices of middle and hind basitarsi rather

narrowly truncate (sometimes obliquely), second segment arising from

lower part of truncation, upper part of truncation usually produced as

an angle or spine above base of second segment on hind leg and sometimes

also on mid leg. w. Middle basitarsus of male somewhat flattened,

variable in form; hind basitarsus of male flattened or inflated, without

hairy pit on outer surface. x. Distitarsi slightly swollen, broader

than preceding segments. y. Claws cleft as in Samba. z. As in Samba

but apical zones less well defined, narrower (so that line across tergum

is much closer to posterior tergal margin than to gradulus), and in

Metasamba punctate and hairy, forming apical hair bands. aa. Terga

without hair bands except for apical ones in Metasamba. bb. Pygidial

plate of female as in Samba, elevated zone broad and weakly differen-

tiated in Haplosamba. cc. Pygidial plate of male absent. dd. Sixth

sternum of male with apex bilobed. ee. Sternum VII of male deeply me-

dially notched at apex, with two pairs of membranous lateroapical lobes,

one often sessile and sometimes not differentiated as a lobe, but hairy

on one surface. ff. Sternum VIII with robust apical process, its upper

apical surface in situ often resembling a pygidial plate. gg.

Gonostylus of male as long as or longer than gonocoxite, well differen-

tiated, slender, with enlarged, angulate apex.

Comments. This genus contains only five species known to me, all

from arid parts of western South Africa and Namibia. These species are

so different in both structure and appearance that | originally planned

to accord each generic rank. However, as suggested by Figures 1 and 2,

the number of differentiating apomorphies is not great, and | hesitantly

reduced each taxon to subgeneric rank. The resulting genus Haplomelitta

is paraphyletic because Samba is derived from it. Such a classification

seems to me more useful than one which slavishly mirrors the cladistic

pattern. The five species of Haplomelitta, however, seem more different

from one another than many apoid genera.

Michener: Classification of Melittidae 63

Key to the Subgenera Haplomelitta

1. With strong, pale apical tergal hair bands; male with deformed hind

and middle basitarsi and inner hind tibial spur.

se: sh nat eel eh a lh ll ahi le SL en Sk Se ie lak MO ----Metasamba

-- Without metasomal hair bands; legs of male not deformed (hind

basitarsus sometimes inflated).

Pall A aR ner Eee 7 Oger BOM FAA gy PS gen Pg PS eh ot PE a gs etn pe tale I SS gp, ng ah ag A Ae Pelee 2

2. Mandible of female tridentate (male unknown).

eS wf lon: 9 it i eo ein A Haplosamba

-- Mandible of female bidentate (inner tooth sometimes reduced to a

shoulder).

i aS a a ee sas king aS Wak a a aia Pa ag a a a al a a a ge ake ae eR a Sil rae nae eee re 3

3. Propodeal triangle strongly differentiated, basal zone (and

laterally entire length of triangle) with short longitudinal rugae;

hind basitarsus of male strongly inflated, wider than tibia.

Sak dw a NS oS OY ee Haplomelitta s. str.

-- Propodeal triangle differentiated only by weak lines, without rugae

on basal zone; hind basitarsus of male slender, much narrower than

tibia.

4. Second submarginal cell about as long as first; first recurrent and

first transverse cubital veins usually well separated; basal vein

interstitial with cu-v or nearly so; sternum VI of male with two

pairs of pedunculate membranous lobes.

sh sho nek in gn i as Se ob nae a sa dl Sc Sade eh chr nis Atrosamba

-- Second submarginal cell distinctly shorter than first; first recur-

rent and first transverse cubital veins close together, intersti-

tial or the former on either side of the latter; basal vein much

basad of cu-v; sternum VI of male with one pair of pedunculate mem-

branous lobes and one pair of hairy areas.

a aaa al al hala latelataletatetalalalalatatatatatatalalatatatatatatatetatetatatatataiatatel Prosamba

Prosamba Michener new subgenus

(Pigs. 107," 108, 116, 121) 147; 148)

Type species: Haplomelitta (Prosamba) griseonigra new species.

, Diagnosis. Rather small, dark, robust, without metasomal hair

bands, with paraglossa, with long hairs on basal segments of maxillary

palpus, and with only one pair of membranous lateroapical processes on

sternum VII of male.

Description. a. Vertex feebly convex seen from front. Eyes

weakly converging below in female, strongly so in male. b. Clypeus of

female convex, without median ridge. c. Paraglossa distinct, small,

64 Contrio. Amer. Ent. Inst., vol. 78, no. 3, 1981

Figures 116-132. Sambini, males. 116, 117, Sterna VII and VIII

(dorsal and ventral), Haplomelitta (Prosamba) Oriseonioran. Spu..bid,

Lateral view of penis valve of same. 119, 120, Genitalia (except penis

valves and penis) of same (dorsal, ventral, and lateral). [Genitalia

of the only male of H. griseonigra had been dismembered by someone and

intact..|

could not be illustrated 121, 122, Sternum VII (lateral, dorsal

and ventral), H. (Atrosamba) atra n. sp. 123, Sternum VIII of same

(dorsal and ventral). 124, 125, Genitalia of same (dorsal, ventral, and

2 ss RR EL SE

Michener: Classification of Melittidae 65

tapering distally. First two segments of maxillary palpus with long

hairs (Fig. 107). d. Flagellar segment 1 of female fully three times

as long as broad, as long as 2-4 together; all flagellar segments except

1 and 10 broader than long. e. Flagellar segment 1 of male less than

three times as long as broad, about as long as next 2 to 3 segments;

segments 2 and 3 broader than long, 4 to 10 about as long as broad. f.

Basal vein nearly straight, nearly four times as long as first abscissa

of Rs, considerably basal to cu-v. g. Stigma tapering into vein R1 so

that any measurement of stigmal length is arbitrary. Second submarginal

cell distinctly shorter than first. First recurrent vein interstitial

with or on either side of first transverse cubital. h. Jugal lobe of

hind wing one third as long as vannal lobe. i. Propodeal triangle

weakly defined, without rugae. j. Middle tibial spur in both sexes

slender, only weakly curved. k. Middle basitarsus of both sexes broad

near base and tapering to apex; hind basitarsus of male flattened,

largely parallel-sided but narrowed at base and tapering at narrow apex,

that of female parallel-sided but tapering in distal third to narrowly

truncate apex, only an obtuse angle and no projection above base of seg-

ment 2. 1. Terga without apical hair bands; margins of terga I-IV in

female, I-VI in male broadly depressed and impunctate. m. Sternum VII

of male with one pair of membranous lateroapical processes and a pair of

lateroapical hairy areas.

Comments. So far as known, this subgenus contains only a single

species from the Namaqualand area of Cape Province, described in Appen-

dix Il. It is rather small, dark, and has the same robust form as

Atrosamba and Metasamba.

Etymology. Pro, a prefix indicating before, plus Samba, the name

of a related taxon.

Atrosamba Michener new subgenus

(Figs. 109, 110.0 1222126, 447. tas)

Type species: Haplomelitta (Atrosamba) atra new species.

Diagnosis. Large, robust, dark forms without metasomal fasciae;

depressed marginal zones of terga narrow and ill-defined; hind basitar-

sus in both sexes with apical spine projecting above base of segment 2.

Description. a. Vertex approximately straight seen from front.

Eyes parallel in female, converging below in male. b. Clypeus of

female convex, without median ridge. Inner subapical mandibular tooth

very small, often worn so that it seems to be a shoulder rather than a

distinct tooth. c. Paraglossa absent or apparently so. d._ Flagellar

segment 1 of female about 2.5 times as long as broad, as long as next

2.3 flagellar segments together; flagellar segment 2 broader than long,

lateral). 126, Sternum VI of same (ventral). 127, Sternum VI of H.

(Haplomelitta) ogilviei (Cockerell). 128, 129, Sternum VII of same (dor-

sal, ventral and lateral). 130, Sternum VIII of same (dorsal and ven-

tral). 131, 132, Genitalia of same (dorsal, ventral and lateral).

Scale lines = 0.5 mm; scale is the same for all figures of any one

species except for Figures 126 and 127.

66 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

3-9 about as long as broad. e. Flagellar segment 1 of male about 1.5

times as long as broad, subequal in length to segment 3; segment 2

broader than long but all other flagellar segments longer than broad.

f. Basal vein distinctly curved basally, about 3.5 times as long as

first abscissa RS, approximately interstitial with cu-v. g. Stigma

tapering into vein Rl; second submarginal cell scarcely shorter than

first; first recurrent vein substantially distal to first transverse

cubital. h. Jugal lobe of hind wing about half as long as vannal lobe.

i. Propodeal triangle weakly defined, without rugae. j. Middle tibial

spur of female rather robust, strongly curved apically, of male slender

and weakly curved. k. Middle and hind basitarsi of both sexes flat-

tened, approximately parallel-sided, apex of each subtruncate, produced

distally as strong hairy spine or angle above base of segment 2; female

with these spines acute, that of middle basitarsus ending at about

middie of segment 2, of hind nearly reaching apex of segment 2; middle

basitarsus of male with only slightly acute apical angle, hind basitar-

sus with acute but blunt apical process reaching level of base of seg-

ment 2. 1. Terga without apical hair bands; margins of terga 1-4 in

female, 1-6 in male, rather narrowly depressed, weakly so mid-dorsally,

and impunctate. m. Seventh sternum of male with two pairs of pedun-

culate lateroapical processes (Fig. 122) (apices of longer pair probably

torn off in the only male specimen).

Comments. This subgenus contains a single large, robust, black

species from western Cape Province. It is described in Appendix II.

Etymology. Atro-, black, plus Samba, the name of a related taxon.

Haplosamba Michener new subgenus

(Fig. 147)

Type species: Haplomelitta (Haplosamba) tridentata new species.

Diagnosis. Shining, coarsely punctate, with partially red, non-

fasciate metasoma like Haplomelitta s. str.; mandible tridentate in the

female; propodeal triangle weakly defined. This subgenus is known only

from the female.

Description. a. Vertex weakly concave seen from front. Eyes

diverging below. b. Clypeus convex, without median ridge. Apex of

bristles); posterior margin of stipes straight, anterior and posterior

margins straight; ligular arms long, arising at base of prementum and

ending nearer apex than in Haplomelitta.] d. Flagellar segment 1 about

twice as long as broad, as long as next two together; all flagellar seg-

ments except 1 and 10 broader than long but 7-9 scarcely so. f. Basal

vein curved near base, about three times as long as first abscissa of

Rs, considerably distal to cu-v. g. As in Prosamba but in the

available specimen first recurrent a short distance distal to first

transverse cubital. h. Jugal lobe of hind wing over two fifths as long

as vannal lobe. i. Propodeal triangle weakly defined, without rugae.

j. Middle tibial spur rather thick (as thick as longer hind spurs) but

only moderately curved apically. k. Middle basitarsus with margins

convex so that segment is widest medially; hind basitarsus narrower than

Michener: Classification of Melittidae 67

middle, parallel-sided; apices of both truncate with a right angular or

slightly obtuse angle above base of segment 2. 1. Terga without apical

hair bands; margins of terga 1-4 rather broadly and_ distinctly

depressed, impunctate.

Comments. This subgenus contains only a single known species from

Namaqualand, Cape Province, described in Appendix ||. It is superfi-

cially like H. (Haplomelitta) ogilviei, having the same rather slender

form and partly red metasoma.

Etymology. Haplo-, single or simple, appropriate because of the

generic name Haplomelitta, plus Samba, the name of a related taxon.

Figures 133-142. Sambini, males. 133-135, Sterna Vil, VIII, and

genitalia, Haplomelitta (Metasamba) fasciata n. sp. (dorsal and ven-

tral). 136, Sternum VI of same (ventral). 137, Sternum VI! of same

(lateral). 138, 139, Genitalia of Samba calcarata Friese (lateral, dor-

sal, and ventral). 140, 141, Sterna VIII and Vil of same (dorsal and

ventral). 142, Sternum VI of same (ventral), reconstructed

posterolaterally where lines are dotted because the sclerite was broken.

Scale lines = 0.5 mm; scale is the same for all figures of either

species.

68 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

Subgenus Haplomelitta Cockerell s. str.

(Figs. lil, Tle, 127-132, 143, 147, 140)

Haplomelitta Cockerell, 1934, p. 446. :

Type species: Rhinochaetula ogilviei Cockerell, 1932, by original

designation.

Diagnosis. Slender, coarsely punctate, sparsely hairy forms super-

ficially similar to Haplosamba, from which it differs by the bidentate

rather than tridentate mandibles and the grotesquely modified first

metasomal sternum of the female. |

Description. a. Vertex of female gently concave seen from front;

of male straight, slightly elevated in area of ocelli. Eyes parallel in

female, converging below in male. b. Clypeus of female convex, without

median ridge. c. Paraglossa absent or apparently so. d. Flagellar

segment 1 of female over twice as long as wide, as long as next two seg-

ments together; segments 2 to 9 much broader than long. e. Flagellar

segment 1 of male about 1.5 times as long as wide, as long as next 1.5

segments; segments 2 and 3 broader than long, subsequent segments

grading to 6 to 10 which are as long as broad. f. Basal vein gently

curved basally, about three times as long as first abscissa of Rs,

slightly distal to cu-v or essentially interstitial. g. Second submar-

ginal cell distinctly shorter than first. First recurrent vein a short

distance distal to first transverse cubital. h. Jugal lobe of hind

wing about one third as long as vannal lobe. i. Propodeal triangle

defined by strong lines, with longitudinal rugae extending all the way

across lateral parts of triangle and across anterior half medially. j.

Middle tibial spur robust, especially in female, apex strongly hooked in

female. k. Middle and hind basitarsi of female flattened; middle

basitarsus broader than hind, its margins convex so that segment is

widest near middle, apex with right angle above base of segment 2; hind

basitarsus parallel-sided, apex truncate with acute projection extending

over base of segment 2. Middle basitarsus of male slender, nearly

parallel-sided, nearly as long as tibia, obliquely truncate at apex.

Hind basitarsus of male inflated, slightly broader than tibia, rounded

at apex, nearly as long as tibia. |. Terga without apical hair bands;

posterior margins of terga I-|V of females and I-VI of male depressed,

shining, impunctate. Metasomal sternum | of female with a broad, deep,

slender lateroapical lobes and one pair of broad sessile lobes, larger

hairs of both coarsely plumose or pectinate.

Comments. As here defined this subgenus includes only a single

species from western Cape Province, South Africa. It is rather slender,

with the metasoma largely red. Cockerell (1934) included Capicola

diversipes Cockerell (1932), described from a single male, in

Haplomelitta. | have not seen this species; it could be the male of the

form here placed in Haplosamba but seems too small; more material and

associated sexes are needed to determine the subgeneric status of

Haplomelitta diversipes (Cockerell). Rozen (1974) described nests of H.

CH.) ogilviei.

Michener: Classification of Melittidae 69

Metasamba Michener new subgenus

(Figs. 133-138, 144, 147, 148)

Type species: Haplomelitta (Metasamba) fasciata new species.

Diagnosis. Rather small, robust, pale haired, with apical hair

bands on metasomal terga arising from depressed, punctate marginal

zones.

Description. a. Vertex faintly convex seen from front. Eyes

weakly converging below in female, strongly so in male. b. Clypeus of

female with longitudinal median ridge. c. Paraglossa absent or ap-

parently so. d. Flagellar segment 1 of female about 1.5 times as long

as broad, as long as next 1.5 segments; all flagellar segments except 1

and 10 broader than long. e. Flagellar segments 1 and 2 of male

slightly longer than broad, remaining segments all longer than broad.

f. Basal vein weakly curved, two (in male) to nearly three times as

long as first abscissa of Rs, slightly distal to cu-v. g. Second sub-

marginal cell shorter than first. First recurrent vein substantially

distal to first transverse cubital. h. Jugal lobe of hind wing two

fifths to nearly half as long as vannal lobe. i. Propodeal triangle

weakly defined, without rugae. j. Middle tibial spur in both sexes

robust, strongly curved apically. Hind tibia of male thickened beyond

143

Figures 143-146. Hind legs of Sambini. 143, Haplomelitta

(Haplomelitta) ogilviei (Cockerell), male. 144, H. (Metasamba) fasciata

n. sp., male. 145, 146, Samba calcarata Friese, male and female. Scale

lines = 0.5 mm.

70 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

long as basitarsus, apparently twisted so that it is curved downward

ormal spurs are curved upward), with sharp bend upward at extreme

segment is widest medially, narrowed at each end, apex truncate, segment

2 arising from lower part of truncation, upper part forming slightly ob-

tuse angle; hind basitarsus similar but margins less convex, straight

and parallel from near base to apical third which tapers to truncation,

angle above base of segment- 2 acute but not much produced apicad.

Middle basitarsus of male broadest at large, right angular premedian

tooth on anterior margin, tapering to obliquely truncate apex. Hind

basitarsus of male flat but somewhat inflated, shape shown in Figure

144. |. Terga I-IV of female, I-VI of male, with moderately broad

depressed apical margins which are rather finely punctate and give rise

to white apical hair bands. m. Sternum VI! of male with two pairs of

lateroapical processes, one short, pedunculate, the other long, slender,

strap-like with a postmedian, anterior projecting angle.

Comments. This subgenus contains only a single species, recently

collected in Namibia (South West Africa). Its robust body is suggestive

of Prosamba and Atrosamba.

Etymology. Meta-, a prefix meaning along with or after, plus

Samba, the name of a related taxon.

Genus Samba Friese

(Figs. 113-115, 139-142, 145-148)

Samba Friese, 1908, p. 568.

Type species: Samba calcarata Friese, 1908 (monobasic).

Diagnosis. Female with only one hind tibial spur; head extra-

ordinarily short and broad, vertex concave seen from front, shortest

distance between eyes of female greater than length of eye. Male with

hind basitarsus swollen with large hairy pit on outer side near base.

Description. a. Body of moderate size (9-10 mm long), not densely

hairy. b. Head much broader than long, as wide as thorax in male,

wider than thorax in female; clypeus over three times as wide as long,

not overhanging base of labrum; minimum distance between eyes much

longer than length of eye in female, about equal to length of eye in

male; vertex distinctly concave seen from front. c, d. Labrum of both

sexes over four times as wide as long with very strong, sharp, trans-

verse premedian ridge which is elevated to form tooth on each side of

small median concavity; surface basal to ridge flat, smooth, shining,

Figure 147. Faces of Sambini. Top row, left to right, ee

(Prosamba) griseonigra n. sp., female, male; H. (Haplomelitta) ogilviei

(Cockerell), female. Second row, H. (Atrosamba) atra n. sp., female,

male. Third row, H. (Metasamba) fasciata n. sp., female, male. Bottom

row, left to right, Samba calcarata Friese, female, male, Haplomelitta

(Haplosamba) tridentata n. sp., female holotype. Scale line = 0.5 mm.

Michener: Classification of Melittidae

d2 Contrib. Amer. Ent..lmst., vol...416,.n0. 3, 198:

hairless; distal surface of ridge also shining and hairless, surface

beyond ridge punctate, with some hairs basal to long marginal bristles.

e. Flagellar segment 1 of male a little longer than broad, 2 broader

than long, others mostly about 1.5 times as long as broad. f. Flagel-

lar segments of female mostly about as long as broad or a little broader

than long, but segments 1 and 10 much longer than broad. g. Maxillary

blade twice as long as broad, apex broadly rounded with a few long

hairs; palpus extending well beyond blade. h. Galeal comb well

developed; flange on posterior side of stipes extending full length of

stipes. i. Prementum slender; ligular arm arising near base of premen-

tum, ending as blunt point rather near to apex. Mentum not sclerotized,

distal end bent strongly anteriorly. j. Glossa rather attenuate, much

exceeding labial palpi. k. Paraglossa not recognizable, perhaps a

robust, tapering, almost hairless structure at the end of the suspen-

sorium. 1. Segments of labial palpus with apices not laterally

produced. m. Basal vein nearly three times as long as first abscissa

of Rs, curved especially basally, interstitial with or slightly distal

to cu-v. n. Stigma of moderate size, more than half as long as_ costal

margin of marginal cell; vein r arising at middle of stigma; margins of

stigma basal to vein r converging basally; length from base to base of

vein r about twice the greatest width of stigma. o. Prestigma about

one third as long as stigma. First recurrent vein well beyond first

transverse cubital. p. Apex of marginal cell bluntly pointed,

separated from costal margin of wing by two or three vein widths. q.

Jugal lobe of hind wing about one third as long as vannal lobe. r.

Propodeal triangle smooth, shining, defined only by lack of pubescence,

surface as seen in profile in same plane as rest of propodeum, at about

a 45 degree angle to main axis of body. s. _ Basitibial plate well

defined and strongly elevated in both sexes, surface hairy. t. Scopal

hairs short, simple, much shorter plumose hairs beneath them in some

areas but tibial surface easily visible; dense fine yellow hairs, very

different from those of scopa, on under side of basitarsus; tibia with

scopa-like hairs (sparser on upper and lower than on outer surfaces) ex-

cept for zone of keirotrichia over half as wide as tibia. u. Female

with only one hind tibial spur which is large, sickle-shaped, with a

thickened base, and with smooth margins; middle tibial spur thickened,

apical part curved, margins densely microserrate ("ciliate"). Male with

tibial spurs only gently curved, rather slender, minutely microserrate.

v. Middle and hind basitarsi of female flat, margins convex so that mid

basitarsus is widest medially, hind near basal third, both roughly three

times as long as broad, somewhat narrower than tibiae, each rather nar-

rowly subtruncate apically and giving rise to segment 2 near lower

apical angle; upper apical angle of middle basitarsus obtuse, of hind

slightly produced but blunt. w. Middle basitarsus of male somewhat

flattened, slightly tapering toward obliquely truncate apex. Hind

basitarsus of male swollen, parallel-sided except narrowed at each end,

a large pit lined with white hairs near base of outer side, apex sub-

truncate, segment 2 arising from lower part of apex. x. Distitarsi

swollen, much broader than preceding segments. y. Claws cleft to

middle or beyond middle, inner ramus shorter than outer. z. Metasomal

terga I-IV of female and I-V of male with broad, depressed, impunctate,

apical zones which are well defined across each tergum, narrow

sublaterally but broad medially, so that each tergum has a broadly

procurved transverse line across it which mid-dorsally is about midway

Michener: Classification of Melittidae 713

between gradulus and posterior tergal margin. aa. Terga without hair

bands. bb. Pygidial plate of female strongly elevated, well defined,

with abruptly elevated longitudinal median zone. cc. Pygidial plate of

male absent. dd. Sixth sternum of male with apical truncation over one

fourth as wide as sternal width. ee. Seventh sternum of male deeply,

medially notched at apex, with two pairs of membranous lateroapical

lobes, one sessile, the other pedunculate, most of hairs on both with

apices wavey. ff. Eighth sternum with strong slender apical process,

dividing apically into laterally directed processes. gg. Gonostylus of

male longer than gonocoxite, well differentiated at base, slender, but

enlarged and angulate at apex, hairs few, short.

Figure 148. Forewings of Sambini. Left to right, Haplomelitta

(Prosamba) griseonigra n. sp., H. (Atrosamba) atra n. sp., H.

aap lamallnta ogilviei (Cockerell), H. (Metasamba) fasciata n. sp., and

Samba calcarata Friese. Scale line = 0.5 mm.

Comments. This genus is known from a single species, Samba

calcarata Friese, known only from Kenya and Tanzania. As is apparent

from Figures 1 and 2, as well as the above description, it has many

unusual features. It clearly belongs in a genus of its own.

Dasypodini new tribe

This large Holarctic and African tribe can be recognized by the

characters indicated in the key to tribes and in Table 2. Its

synapomorphies are also indicated in Figures 1 and 2.

Key to the Genera of the Tribe Dasypodini

1. Yellow integmental markings present at least on face and legs;

stigma large, slightly shorter to slightly longer than costal mar-

74 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

gin of marginal cell; prepygidial and pygidial fimbriae of female

absent (Central Asian deserts).

sa stn ws eo ek ere eb eT i Lee fig Eremaphanta

-- Yellow integumental markings absent (except sometimes on clypeus of

male only); stigma not enlarged, about half as long as costal mar-

gin of marginal cell; prepygidial and pygidial fimbriae of female

well developed, or the former represented by a hair band similar to

those of preceding terga.

2. Basitibial plate absent; profile of propodeum all more or less in

one plane; galeal comb absent or nearly so; male gonostylus well

differentiated, nearly always deeply bifid (Palearctic Region).

Si nine ht ss kgm pe valk pS ict a a SO eg i 0 GAN ke es stro Dasypoda

-- Basitibial plate of female and nearly all males present, rarely

defined only along posterior margin; base of propodeum more nearly

horizontal than rest in profile [except in some Hesperapis

(Carinapis)]; galeal comb present; male gonostylus broad, fully

fused to gonocoxite, not bifid but often with short inner process

or lobe.

3. Pygidial plate of female with longitudinal median elevated area or

at least ridges representing edges of such an area; male gonostylus

without inner lobe (South Africa).

-- Pygidial plate of female without longitudinal median elevated or

differentiated area (sometimes with triangular basal area); male

gonostylus with inner lobe usually bearing long, coarse, branched

bristles or hairs (North America).

4. Mesoscutum as long as or longer than minimum intertegular distance;

front tibia with series of five or more robust, apical and

posteroapical, amber spines on outer surface.

och veil Pal ga nes pliant hona al oc hga seasons laine eee eae aan einem incndn eal a Xeralictoides

-- Mesoscutum considerably shorter than minimum intertegular distance;

front tibia without such spines.

etd er Sk i eras ak al Sot coe Se sineak gle Rach lt lng ei bo en mocnr ease ner Hesperapis

Genus Dasypoda Latreille

(Figs. 149-158)

Dasypoda Latreille, 1802, p. 424.

Type species: Andrena hirtipes Fabricius, 1793, designated by

Blanchard, 1840, p. 414. [Melitta suanmerdasitels Kirby, 1802,

designated as the type species s by Curtis (1831, 367) was not

originally included in Dasypoda but is a synonym of A. hirtipes. ]

Michener: Classification of Melittidae 715

Podasys Rafinesque, 1815, p. 123.

Type species: Andrena hirtipes Fabricius, 1793 (Podasys was proposed

unnecessarily to replace Dasypoda; the type species is therefore

automatically the same as that of Dasypoda. )

Diagnosis. Large bees without yellow integumental markings, with

abundant long hairs. Galeal comb absent or nearly so; basitibial plate

absent in both sexes; propodeal profile slanting, without subhorizontal

basal zone.

Description. a. Body rather large (10-17 mm long), elongate, with

abundant long hair; yellow integumental markings absent. b. Head round

to broader than long, distinctly narrower than thorax; vertex convex as

seen from front. c. Labrum of male less than three to over four times

as wide as long, with basal half smooth, shining, convex to nearly flat,

sometimes with low ridge separating it from hairy, punctate distal zone.

d. Labrum of female similar in shape to that of male, with elevated,

bare, basal area separated by strong ridge from hairy distal zone. e.

Flagellar segments of male all or mostly longer than broad, segment 1

often longest. f. Flagellar segments of female mostly broader than

long or slightly longer than broad but segment 1, which is much longer

than any other, much longer than broad and narrowed toward base. g.

Maxillary blade over four times as long as broad, nearly parallel-sided

except apically; galeal fringe insignificant; maxillary palpus not at-

taining apex of galea. h. Galeal comb absent or reduced to a few

minute, widely separated bristles. i. Prementum long and slender,

lateral margins curled so as to largely or partially hide ligular arms

which arise near base of prementum and end at about four fifths of

distance from base to apex. j. Glossa tapering but not attenuate at

apex. k. Paraglossa minute, hairless or nearly so. 1. Labial palpus

not or scarcely reaching apex of glossa, segments 2 and 3 subcylin-

drical, apices not produced laterally. m. Basal vein less than twice

to over three times as long as first abscissa of Rs, nearly straight,

distal to cu-v. n. Stigma slender, less than half as long as costal

margin of marginal cell; vein r joining stigma at distal third; margins

of stigma basal to vein r only slightly converging basally. o.

Prestigma nearly to more than two thirds as long as stigma. p. Apex of

marginal cell narrowly rounded, separated from costal wing margin by two

or three vein widths. q. Jugal lobe of hind wing less than half as

long as vannal lobe, measured from wing base. r. Propodeal triangle

rather large, dull, not steeply sloping, flat seen in profile (there

being no basal zone more nearly horizontal than rest). s. Basitibial

plate absent in both sexes. t. Scopa consisting of dense, long,

minutely barbed hairs on both inner and outer surfaces of hind tibiae

and basitarsi; keirotrichia absent. u. Mid and hind tibial spurs

nearly straight, their margins minutely and densely microserrate

("ciliate"). v, w. Mid and hind basitarsi of female long, slender,

tapering or in males sometimes parallel-sided, giving rise to segment 2

in middle of the narrow apex. x. Distitarsi not swollen, often no

broader than preceding segments. y. Claws cleft about to middle, inner

ramus shorter in female than in male. z. Metasomal terga! to IV of

females and | to V of males with broad, depressed, impunctate, often

translucent margins and usually with apical or preapical pale hair bands -

which partly cover depressed marginal areas. aa. Prepygidial and pygi-

dial fimbriae of female dense. bb. Pygidial plate of female well

76 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

defined, smooth but often dull, notched at apex, without median elevated

area. cc. Pygidial plate of male absent. dd. Sternum VI of male

strongly bilobed, the lobes with short hairs. ee. Sternum VI! with

rather reduced disc, apex bilobed, each lobe with slender, dorsally

projecting lateroapical lobe that is hairy distally [lateroapical lobe

greatly reduced in some species, e.g., D. argentata (Panzer)]. ff.

Sternum VIII with broad apical process which is not truncate. gg.

Gonostylus usually as long as or longer than gonocoxite, fused to

gonocoxite dorsally but area of separation more or less clear;

gonostylus usually bifid, sometimes to base, and in species such as D.

argentata Panzer apparently deeply trifid although one of the three

distal processes could be of gonocoxal origin.

Comments. This is a well-known genus of rather large, conspicuous

bees that is not closely related to any other genus. The following

characters are probably apomorphies, for they do not occur in any of the

related genera, nor in genera of related subfamilies and families: e,

h, s, and t.

Hh)

Wy;

Sh IRR ae

Noe

ERS

i ~

Z SS

Pp es

aN S

Figures 149-158. Dasvpoda,. males.....149-156, .D,. hirtipes

(Fabricius). 149, Anterior view of labium. 150, Inner view of maxilla.

151, Sternum VI (ventral). 152, 153, Sternum VII (lateral, dorsal, and

ventral). 154, Sternum VIII (dorsal and ventral). 155, 156, Genitalia

(dorsal, ventral, and lateral). 157, Gonostylar area (dorsal) of D.

argentata Panzer. 158, Sternum VII (dorsal and ventral) of same. Scale

line = 0.5 mm and is applicable to all figures.

Michener: Classification of Melittidae a7

The lack of basitibial plates in females (character s) is a sur-

prising feature for such plates are present in most bees that burrow in

the ground, presumably playing a role in locomotion through the burrows.

Lack of such plates among burrowing bees is largely limited to the

genera Dasypoda and Colletes.

Dasypoda is widespread in the Palearctic Region, ranging from

Portugal to Japan. The distribution includes most of Europe and North

Africa, and in Asia extends south to India (according to Quilis, 1928).

The genus is particularly well represented in the Mediterranean Basin. A

review of the west-Palearctic species was given by Warncke (1973), while

the Spanish species were revised by Quilis (1928). The genus has long

been properly recognized, for example by Dalla Torre (1896), and there

is no need to list the species. The larva is described by Rozen and

McGinley (1974); the nesting biology by Malyshev (1927) and Lind (1968).

Genus Eremaphanta Popov

(Figs. 159-167)

Diagnosis. Small bees, commonly with extensive yellow markings but

at least with yellow areas on face and legs; stigma large, transparent,

approximately as long as costal margin of marginal cell and more than

three times as long as prestigma; prepygidial and pygidial fimbriae of

female absent.

Description. a. Minute (4-6.5 mm long), rather slender, sparsely

hairy; yellow integumental markings extensive or limited to mandible,

labrum, clypeus and legs. b. Head transverse to elongate, in latter

case distinctly narrower than thorax; vertex convex as seen from front.

c, d. Labrum less than three to nearly four times as wide as long, con-

vex, smooth and shining except for hairy apical margin. e, f. Flagel-

lar segments except the last broader than long, or in some males

slightly longer than broad; segment 1 not strikingly different in length

from subsequent segments. g. Maxillary blade three or somewhat more

times as long as broad, tapering to narrowly rounded to almost pointed

apex; galeal fringe present, hairs rather short to moderately long; max-

illary palpus extending beyond apex of galea. h. Galeal comb present.

Stipes with large concavity in distal two thirds of posterior margin,

this concavity with short hairs, without teeth such as are found in long

tongued bees. ji. Prementum robust, ligular arms arising near base

[dispar (Morawitz)] or beyond end of basal fourth [fasciata Popov],

ending one third to one fourth of length of prementum from apex. j.

Glossa tapering, attenuate at apex at least in E. dispar (Morawitz). k.

Paraglossa minute, partly hairless. 1. Labial palpus equal to or ex-

ceeding glossa, in E. fasciata Popov but not in E. dispar (Morawitz)

segments 2 and 3 ending obliquely and produced Jaterally. m. _ Basal

vein one to two times as long as first abscissa of Rs, curved, distad to

cu-v. n. Stigma large, transparent, slightly shorter than to slightly

longer than costal margin of marginal cell, vein r joining stigma near

middle, margins of stigma basal to vein r converging basally. o.

Prestigma less than one third as long as stigma. p. Apex of marginal

cell pointed, on or very near wing margin. q. Jugal lobe of hind wing

over two thirds to over three fourths as long as vannal lobe. r.

Propodeal triangle large, minutely sculptured, shining or partly dull,

nearly all horizontal; propodeum in profile about half horizontal and

78 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

half vertical, one surface rounding onto the other. s._ Basitibial

plate present in both sexes, rounded, shining. t. Scopa sparse, con-

sisting of long, simple hairs, largely confined to outer surfaces of

hind tibia and basitarsus; keirotrichia occupying undefined,

longitudinal median band on inner tibial surface. u. Mid and hind

tibial spurs nearly straight, their margins minutely serrate. v, w.

Mid and hind basitarsi somewhat flattened in female, scarcely so in

male, slightly tapering, giving rise to segment 2 in middle of apical

margin. x. Distitarsi not noticeably swollen although somewhat broader

than preceding segments. y. Claws of female with inner tooth simple or

absent; of male, at least in E. dispar (Morawitz), cleft, inner ramus a

little shorter than outer. z. Metasomal terga |-V of females and I-VI

of males with depressed, impunctate, transparent, smooth apical margins,

relatively narrow on anterior terga and progressively broader

posteriorly. aa. Terga with or without apical hair bands. Dense

prepygidial and pygidial fimbriae absent. bb. Pygidial plate of female

well defined, sometimes [in E. fasciata and dispar (Morawitz)] occupying

almost whole dorsal surface of tergum VI. cc. Pygidial plate of male

absent. dd. Sternum VI of male bilobed or medially notched [at least

in E. dispar (Morawitz)]. ee. Sternum VII with rather large disc which

has two, pointed, hairy lobes at apex; lateroapical lobes absent. ff.

Sternum VIII with apical process obliquely truncate. gg. Gonostylus

simple, about as long as gonocoxite, broadly fused to it.

Comments. This is a genus of minute bees which, by their size and

bright coloration, resemble the species of Perdita (Panurginae),

Nomioides and Habralictus (Halictidae), and certain groups’ of

Euryglossinae (Colletidae). Popov (1957) was quite correct in saying

that Eremaphanta is distinct from Hesperapis, a genus into which | had

earlier suggested (Michener, 1944) that Eremaphanta might fall. In 1944

| had not seen specimens of Eremaphanta, and based my suggestion

primarily on Popov's (1940) illustrations of the genitalia and hidden

sterna, which are similar to those of Hesperapis and doubtless indicate

common ancestry. Eremaphanta differs from Hesperapis and other

Dasypodini in a series of apomorphic characters as follows: a, h (con-

cavity in posterior margin), n, 0, r, t, and aa. Another character,

equally distinctive but not known to be apomorphic, is q.

Eremaphanta is Known only from arid parts of Central Asia (the

Turkmenskaja S.S.R. and the Tadzikskaja S.S.R.).

Key to the Subgenera of Eremaphanta

1. Terga without apical hair bands; head broader than long; vertex

gently convex, scarcely above level of summits of eyes.

tite Eremaphanta s. str.

-- Terga with apical hair bands; head longer than broad; vertex ex-

tending far above summits of eyes.

2a al st OE sl yd ha lndiat addon sda haat a imo dad daa en lasadatnrts eat eter Popovapis

Michener: Classification of Melittidae 79

Subgenus Eremaphanta Popov s. str.

Eremaphanta Popov, 1940, p. 53.

Type species: Rhophites vitellinus Morawitz, 1876.

Description. a. Head broader than long, posterior ocelli so

placed that, seen from front, they break the convexity of the vertex.

b. Body with extensive yellow markings. c. Metasomal terga without

apical hair bands. d. Labrum nearly four times as wide as long. e.

Maxillary blade less than three times as long as broad. f. Ligular arm

Figures 159-167. Eremaphanta. 159, 160, Inner view of maxilla and

anterior view of premental area of E. (Eremaphanta a) fasciata Popov.

[Glossa broken off.] 161, 162. Inner view of maxilla and anterior view

of labium of E. (Popova is) dispar (Morawitz). 163, Sternum VI, male,

of same (ventral). 164, 165, Sterna VII and VIII (dorsal and ventral )

of same. 166, ey. Genitalia (lateral, dorsal, and ventral) of same.

Scale line = 0.5 mm and is applicable to all figures.

80 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

arising over one fourth of distance from base to apex of prementum. g.

Labial palpus less than half as long as prementum, segments 2 and 3

ending obliquely and produced laterally. h. Claws of female simple or

each with minute tooth on inner surface.

Comments. This subgenus contains E. convolvuli Popov, fasciata

Popov, minuta Popov, turcomanica Popov, and vitellina Morawitz. The

characters of the mouthparts and claws listed above have been examined

by me only for E. fasciata. The other species are placed here on the

basis of description and figures (Popov, 1940, 1957).

Popovapis Michener new subgenus

Type species: Rhophites dispar Morawitz, 1892.

Description. a. Head much longer than broad, vertex extending

behind ocelli so that, seen from front, summit of vertex is above and

behind ocelli. b. Yellow markings limited to mandibles, labrum,

clypeus, tegula, legs, and the sometimes reddish yellow metasoma. c.

Metasomal terga with distinct apical hair bands. d. Labrum less than

three times as wide as long. e. Maxillary blade over three times as

long as broad. f. Ligular arm arising near base of prementum. g.

Labial palpus longer than prementum, segments more or less cylindrical.

h. Claws of female each with large tooth on inner surface representing

inner ramus.

Comments... 1 nis..subgenus contains .E. dispar (Morawitz) and

zhelochovtsevi Popov. Of these | have seen only the first and the

characters of the mouthparts and claws, not described by Popov, may not

all apply to the second species.

Etymology. Named in honor of V. B. Popov who described the genus

and most of its species.

Genus Capicola Friese

(Figs. 168-177)

Diagnosis. Similar to Hesperapis but pygidial plate of female with

longitudinal median, defined, usually elevated area; male gonostylus (or

possibly distal part of gonocoxite) with inner, basal process or lobe,

usually bearing large setae.

Description. a. Small to middle sized (3-11 mm long), with the

form of a Halictus; yellow integumental markings absent. b. Head

broader than long, about as wide as thorax, vertex convex as seen from

front. c. Labrum of male nearly to more than four times as wide as

long, basal half smooth, shining, convex, no ridge separating it from

distal half which is hairy and punctate. d. Labrum of female similar

in shape to that of male; basal area dull to shiny, convex, biconvex, or

divided by longitudinal grooves into a series of convexities, or other-

wise roughened; distal hairy zone often narrow or limited to margin. e.

Flagellar segments of male mostly about as broad as long or longer than

broad, segment 2 much broader than long, 1 and 3 often broader than

long. f. Flagellar segments of female mostly broader than long or as

long as broad, segment 2 distinctly broader than long, 1 broader than

long to longer than broad. g. Maxillary blade about four times as long

Michener: Classification of Melittidae 81

as broad, shaped about as in Hesperapis pellucida (Fig. 18/7), “more

tapering and pointed in C. braunsiana, blunt in C. nanula; galeal fringe

insignificant to moderately developed. h. Galeal comb usually large

(about 16 spines), only 8 spines in C. nanula; maxillary palpus at-

taining apex of galea. i. Prementum about as in Hesperapis pellucida

(Fig. 186) but ligular arms somewhat shorter, in C. nanula as short as

in H. ilicifoliae (Fig. 188). j. Glossa sometimes more tapering and

more sharply pointed than in Hesperapis pellucida, or in C. nanula

rather blunt. k. Paraglossa minute, with few short apical and

subapical hairs. |. Labial palpus attaining (in C. aliciae) or usually

surpassing apex of glossa, segments 2 and 3 subcylindrical, apices not

produced laterally. m. Basal vein less than to more than twice as long

as first abscissa of Rs, gently curved or rather strongly so near base,

distal to cu-v. n. Stigma more than half as long as costal margin of

marginal cell; vein r joining stigma slightly beyond middle; margins of

stigma basal to vein r converging (sometimes only slightly) basally. o.

Prestigma less than one third to over one half as long as stigma. p.

Apex of marginal cell pointed or narrowly rounded, separated from wing

margin by one or two vein widths. q. Jugal lobe of hind wing over half

to less than two thirds as long as vannal lobe. r. Propodeal triangle

of moderate size to rather short and transverse, minutely roughened and

slightly shining to coarsely rugose striate or pitted; propodeum seen in

profile with basal zone more horizontal than declivous posterior sur-

face. s. Basitibial plate present in both sexes. t. Scopa consisting

of long, simple hairs on outer sides and upper and lower margins of hind

tibia and basitarsus; shorter plumose hairs under long hairs along

posterior (upper) part of outer surface of tibia, from basitibial plate

to apex; inner surfaces of tibia and basitarsus with shorter, finer,

denser hairs, those of tibia (keirotrichia) blunt at apices and forming

a band the length of tibia and over two thirds as wide as tibia. u.

Mid and hind tibial spurs strongly curved or not, slender, margins of

hind spurs and middle spur of males minutely and densely microserrate,

middle spur of female C. nanula similar, of other species with only a

few, large, widely separated teeth. v, w. Mid basitarsus of female

broadest near base or near middle, often broader than hind basitarsus

which is broadest near base or is nearly parallel-sided in C. aliciae

and nanula; these basitarsi giving rise to segment 2 near middle of nar-

row apex. x. Distitarsi not much swollen, not or only moderately

broader than preceding segments. y. Claws cleft much less than _ half

way to base, inner ramus a rather short tooth in female, only somewhat

shorter than outer ramus in male. z. Metasomal terga | to V of female

and | to VI of male with slightly depressed, translucent, marginal

zones, those of | to |V of female and most of those of males usually

largely covered with apical white hair bands. aa. Prepygidial and py-

gidial fimbriae of female present, the former sometimes reduced to a

band almost like apical bands of preceding terga. bb. Pygidial plate

of female well defined, dull, with longitudinal median elevated zone

(represented by pair of subparallel ridges or carinae in C. braunsiana,

in which area between these ridges is depressed almost to level of rest

of plate; similar ridges in c. aliciae close together so that elevated

zone is narrow). cc. Pygidial plate of male absent. dd. Sternum VI

of male strongly bilobed with added pair of lateroapical lobes so that

margin is commonly four-lobed; lobes hairy, often with coarse

specialized hairs. ee. Sternum VII of male variable. ff. Sternum

82 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

VIII of male with broad, subtruncate or rounded apical process which in

situ. often looks like a pygidial plate. gg. Gonostylus in-

distinguishably fused to gonocoxite, probably much shorter than to

nearly as long as gonocoxite, not bifid, without inner process or

projection bearing large hairs.

Comments. This genus, which is restricted to southern Africa, is

closely related to Hesperapis. The species have the aspect of the

middle-sized to small species of Hesperapis. Most of the characters

listed are plesiomorphic but the elevated or at least defined median

zone of the pygidial plate of the female is a synapomorphy (also found

in the Sambini, where it must be of independent origin). Another

probable synapomorphy, not always evident in preparations and not

described above, is the presencé of retrorse, apparently eversible, mem-

branous, dorsal, preapical lobes on the penis valves. They are shown in

Figure J/77 for C. braunsiana; they exist in other species including C.

aliciae although they were not visible in the specimen of that species

illustrated in Figure 171.

The other characters that differentiate Capicola from Hesperapis

are listed under the latter genus; some are synapomorphies for

Hesperapis. Rozen and McGinley (1974) found synapomorphies separating

larvae of Hesperapis from those of Capicola s. str. The groups would

perhaps have been recognized at the subgeneric rather than generic level

except for the enormous geographic distance separating Hesperapis in

North America from Capicola in South Africa. Previous authors have

sometimes united the genera (Cockerell and Ireland, 1933), some of the

African species having been described as Hesperapis (Cockerell, 1934).

Subgenus Capicola s. str.

Capicola Friese, 1911, p. 672.

Type species: Capicola braunsiana Friese, 1911 (monobasic).

Rhinochaetula Friese, 1911, p. 185.

Type species: Capicola (Rhinochaetula) cinctiventris Friese, 1912, by

designation of Cockerell, 1915, p. 343.

Diagnosis. Differs from Capicoloides by the lack of large, blunt,

spine-like hairs in the tibial scopa and the broad, apically bilobed

body of sternum VII of the male.

Description. a. Labial palpus surpassing apex of glossa. b.

Outer surface of hind tibia of female without spine-like hairs inter-

spersed among scopal hairs. c. Sternum VI of male broadly truncate,

apical margin with four lobes bearing thickened setae, median lobes

divergent. d. Sternum VII of male with body rather broad, apex

bilobed. e. Gonostylus of male broad, rather long, blunt, outer sur-

face with well defined area of coarse, dense setae.

Comments. Characters c, e, and perhaps d are synapomorphies

that characterize the subgenus. The hind tibiae of certain males

(C. rufiventris Friese) are greatly expanded apically, triangular,

with the tibial spurs widely separated. The male of C. flavitarsis

Friese has hind tibiae intermediate between the ordinary unmodified

type and that of C. rufiventris.

Michener: Classification of Melittidae 83

The identity of C. cinctiventris is important because it is the

type species of Rhinochaetula and because the name has been used for a

species here placed in the subgenus Capicoloides. This name was proposed

by Friese (1912) as a subgenus of Capicola for three species that sup-

posedly differed from Capicola proper in having the middle tibial spur

of the female minutely serrate ("ciliate") instead of having a few

coarse teeth. Actually, only one of the three species, plumipes Friese,

exhibits this character and it is tentatively placed in Promelitta. The

type species of Rhinochaetula, then, regardless of what species it

really is, has a tibial spur like that of most other Capicola. The type

specimen of C. cinctiventris in Zoologisches Museum, Berlin, is labelled

"Bushmanid./Jackals Water/Lightfoot." It agrees with an identically

labelled specimen in the South African Museum, Cape Town, Friese

(1912), however, gave the locality as "Ookiep," in "Kl. Namaland." |

therefore suspected that the type was erroneous. However, Dr. E.

Kénigsmann of the Berlin museum was doubtful, indicating that Friese

often made errors in citing localities. At my request Dr. Konigsmann

compared the photograph that Friese published in his original descrip- |

tion with the type specimen. The photograph was undoubtedly of that

specimen, as shown by positions of wings, legs, etc. There is nothing

in the original description of C. cinctiventris contrary to this inter-

pretation. This information validates the type specimen and shows that

Rhinochaetula is a synonym of Capicola s. str.

This subgenus includes the following species: braunsiana Friese,

1911 [= aurescens Cockerell, 1932; turneri (Cockerell, 1934)];

cinctiventris Friese, 1912; flavitarsis Friese, 1912; nanula Cockerell,

1936, rufiventris Friese, 1912 (= femorata Friese, 1924; callura

Cockerell, 1932; nivea Cockerell, 1932). The synonymies indicated above

are not always based on type material and should be verified when the

species are revised, although types of Friese's species and cotypes (=

paratypes) of some of Cockerell's were studied.

Nests of C. braunsiana were described by Rozen (1974).

Various other forms that doubtless fall in this subgenus have been

named, for example by Cockerell (1934).

| have examined types of Rhinochaetula (?) armaticeps Friese, 1913,

and R. (?) capensis Friese, 1913. Both are species of the colletid

genus Scrapter, and form the new combinations Scrapter armaticeps

(Friese) and S. capensis (Friese). The last mentioned is not the same as

Polyglossa capensis Friese, 1909, which is presumably also a Scrapter.

Capicoloides Michener new subgenus

Type species: Capicola aliciae Cockerell, 1932.

Diagnosis. Outer surface of hind tibia of female with large,

blunt, spine-like hairs among scopal hairs; sternum VII of male with

body reduced, apex with laterally directed lobes.

Description. a. Labial palpus attaining apex of glossa. b.

Outer surface of hind tibia of female with large, rather short, blunt,

dark tipped spine-like hairs interspersed among the long scopal hairs

from basitibial plate to tibial apex. c. Sternum VI of male with apex

bilobed, extending well beyond small lateral lobes (Fig. 168), median

lobes not divergent. d. Sternum VII of male with body reduced, narrow

84 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

fa i hi y Yo

-Z

‘Z

que

183

Figures 168-185. Capicola and Hesperapis, males. 168, Sternum VI,

center. 169, 170, Sterna VII and

C. (Capicoloides) aliciae Cockerell

Vill (dorsal and ventral) of same. 171, 172, Genitalia (dorsal, ven-

tral, and lateral) of same. 173, Sternum VI, C. (Capicola) braunsiana

Friese (ventral). 174, 175, Sterna VII and VIII (dorsal and ventral

of same. 176, 177, Genitalia (lateral, dorsal, and ventral) of same.

178, Sternum VI, H. (Panurgomia) pellucida Cockerell (ventral). 179,

180, Sterna VII and VII! (dorsal and ventral) of same. 181, 182,

[sean ase ane manner en sa ima eae

Michener: Classification of Melittidae 85

basally, expanded to laterally projecting lobes apically (Fig. 169). e.

Gonostylus slender, apparently rather short (base not defined), without

area of coarse, dense setae.

Comments. This subgenus is especially distinctive because of the

small seventh sternum of the male, which bears lateroapical lobes. For

bees in general this is doubtless a plesiomorphy, for it is a widespread

feature among the short-tongued families. However, if the cladograms

(Figs. 1, 2) are correct, this character of Capicoloides may be a rever-

sion, since Eremaphanta has sternum VII about as in Hesperapis and

Capicola s. str. The gonostyli of Capicoloides lack the specialized

areas of setae which seem to be apomorphous in Capicola proper, again

suggesting a plesiomorphous condition for Capicoloides. The only obvious

apomorphy for this subgenus is character b.

The only included species known to me is C. aliciae Cockerell.

Specimens probably to be included in C. aliciae were identified by

Cockerell as C. cinctiventris Friese. If this identification were cor-

rect, the name Rhinochaetula would be available for this subgenus. (See

notes under the subgenus Capicola. )

Genus Hesperapis Cockerell

(Figs. 178-190)

Diagnosis. Bees usually with a fasciate metasoma and form similar

to that of Halictus, but metasoma of female much flattened and integu-

ment soft. The only close relative except Xeralictoides is Capicola,

from which it differs by the lack of a longitudinal median raised or

defined area on the pygidial plate of the female, and the presence of a

Process or lobe, usually with long hairs, on the inner side of the base

of gonostylus (or perhaps apex of gonocoxite).

Description. a. Small to large (4-15 mm long), with form of a

Halictus; yellow integumental markings absent, or in male of H. rufipes,

lower half of clypeus yellow. b. Head broader than long to about as

broad as long, vertex convex seen from front, sometimes straight, or in

some with "long" head and elevated vertex, slightly concave. c. Labrum

of male less than four times as wide as long, rarely little over twice

as wide as long, two thirds smooth, often shining, convex, sometimes

protuberant mediobasally, no sharp ridge separating basal smooth zone

from distal punctate, hairy zone. d. Labrum of female usually about

four times as wide as long, rarely little over twice as wide as long,

basal convex zone smooth or dull, sometimes forming a rounded ridge

separating it from apical hairy zone along margin, or sometimes (e.g.,

H. ilicifoliae) with sharp transverse ridge near base. e. Flagellar

segments of male mostly longer than broad to broader than long, segment

2 distinctly broader than long, segment 1 sometimes broader than long

even in some species with most segments elongate. f. Flagellar seg-

ments of female as described for Capicola. g. Maxillary blade usually

about as illustrated for H. pellucida (Fig. 187), rarely very short and

Genitalia (dorsal, ventral, and lateral) of same. 183, Sternum VII, H.

(Zacesta) rufipes (Ashmead) (ventral). 184, Ventral view of gonostylar

region of same. 185, Ventral view of gonostylar region of H.

(Carinapis) carinata Stevens. Scale lines = 0.5 mm; scale is the same

for all figures of any one species.

86 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

rounded (H. ilicifoliae), sometimes pointed; galeal fringe insignificant

to moderately developed. h. Galeal comb with four to nine spines; max-

illary palpus not attaining to exceeding apex of galea. i. Prementum

quite variable in length of ligular arms (Figs. 186, 188, 190). j.

Glossa usually rather long and pointed, very short in H._ ilicifoliae

(Fig. 190). k. Paraglossa as described for Capicola. |. Labial palpus

not attaining to exceeding apex of glossa, segments 2 and 3 subcylin-

drical, apices not produced laterally. m. Basal vein as described for

Capicola. n. Stigma as described for Capicola or in largest species of

subgenus Carinapis less than half as long as costal margin of marginal

cell, vein r joining stigma at distal third, and margins of stigma basal

to vein r parallel. o. Prestigma about one third to about one half as

long as stigma. p. Apex of marginal cell as in Capicola. q. Jugal

lobe of hind wing one fourth to three fourths as long as vannal lobe.

r. Propodeal triangle large, smooth, sometimes striate or pitted at

base, or densely punctate throughout; propodeum seen in profile with

basal third or more distinctly more horizontal than posterior declivous

surface except in some species of subgenus Carinapis in which only basal

zone is slightly more horizontal, or entire propodeum declivous as in

Dasypoda. s. Basitibial plate as in Capicola or occasionally only

posterior margin defined. t. Scopa and other hind tibial hairs as in

Capicola or in some species of subgenus Carinapis, scopal hairs with a

few barbs and shorter plumose hairs absent; band of keirotrichia as in

Capicola or narrow, often only one third as wide as tibia, in subgenus

Carinapis. u. Tibial spurs as described for Capicola, most subgenera

having more or less fine adjacent teeth on middle spur of female but

some Carinapis having coarse, well separated teeth. v, w. Mid and hind

basitarsi of female broadest near bases, narrowed apically, giving rise

to segment 2 near middle of narrow apex. x. Distitarsi as described

for Capicola. y. Claws as in Capicola but cleft sometimes reaching

nearly half way to base. z. Tergal bands as described for Capicola,

tergum V_ often with a band like preceding terga. aa. Prepygidial and

pygidial fimbriae present, the former often reduced to a band like those

of preceding terga (see character z). bb. Pygidial plate of female

well defined, without longitudinal median elevated or delimited area.

cc. Pygidial plate of male absent except in some Carinapis. dd. Ster-

num V1 of male rather flat, apically bilobed. ee. Sternum VII of male

broad, short, flat, transverse, apically strongly bilobed. ff. Sternum

VII! of male as described for Capicola but apex rarely exposed enough to

resemble a pygidial plate. gg. Gonostylus indistinguishably fused to

gonocoxite, probably nearly as long as gonocoxite, with an inner basal

lobe or process (perhaps gonocoxal in origin) which, except in the sub-

genus Panurgomia, bears long, coarse, hairs which are barbed or plumose.

Comments. This North American genus contains species very similar

to those of the African Capicola. Much of the above description

describes ranges of variation different from those of Capicola, but not

characters by which the two genera can be distinguished. The following

characters, however, do differentiate these two genera: h (reduced

galeal comb), r (larger propodeal triangle), bb, gg. Of these, h and gg

appear to be synapomorphies for Hesperapis (plus Xeralictoides). There

are also larval synapomorphies of Hesperapis (several subgenera) that

separate it from Capicola s. str. (Rozen and McGinley, 1974). An ad-

ditional character is the lack of the membranous, retrorse, subapical

projections of the penis valves characteristic of Capicola. This

Michener: Classification of Melittidae 87

Figures 186-190. Mouthparts of Hesperapis. 186, 187, Anterior view

of labium and inner view of maxilla of H. (Panur omia) pellucida

Cockerell. 188, Anterior view of labium of H. (Zacesta) rufipes (Ash-

mead). 189, 190, Anterior view of labium and inner view of maxilla of

H. (Amblyapis) ilicifoliae (Cockerell). Scale lines = 0.5 mm.

character as well as r and bb are presumably plesiomorphies, being

shared with various other genera of Dasypodinae.

The characteristic lobe (character ss) on the inner surface of the

gonoforceps is hereafter referred to as a process of the gonostylus, al-

though its morphological origin is not certain.

The nesting biology of several species has been described by Stage

(1966).

The following material on the subgenera of Hesperapis is extracted

from an unpublished thesis by Dr. G. |. Stage 71566) with his permis-

sion. The recognition of most of the subgenera is entirely his.

Paragraphs that are nearly identical to his, or modified only by minor

shortening, are followed by the notation "(Stage)."" Paragraphs based

upon Stage's material but largely reorganized are followed by the nota-

tion "[Stage]". His almost unmodified descriptions of all the new sub-

genera, are therefore described and named by him, as is the genus

Xeralictoides. Aside from shortening, species lists (names extracted

from his work, but not quoted), and comments designed to make his style

more compatible with the rest of this paper, the only significant tax-

onomic differences between his work and what follows are (1) recognition

of Zacesta as a subgenus different from Hesperapis s. str., and (2)

recognition of Xeralictoides as a genus different from. Hesperapis. Both

of these changes are made at the request of Dr. Stage. | have modified

Stage's key to subgenera in accordance with this request, but it uses

88 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

the same characters in essentially the same way as his key.

In each subgeneric description, the diagnostic combination of

characters is indicated by underlines.

Relationships among the Subgenera of Hesperapis. While the fol-

lowing comments do not agree with those of Stage (1966), partly because

mine are less detailed, they would not have been possible without

Stage's study. Table 3 lists the subgeneric variables (as well as

generic characters for Xeralictoides) for which plesiomorphic vs.

apomorphic alternatives are recognizable on the basis of distribution of

the characters in other Melittidae. Figure 191 shows the distribution of

the apomorphies among the subgenera; it is modified from Stage's

cladogram.

TABLE 3... LIST: OF. APOMORPHIES FOR. XERALICTOIDES AND

HESPERAPIS (MODIFIED FROM STAGE, 1966).

1. Males larger than females. [A rare feature of bees, not found in

other melittids, although evident in Anthidium (Megachilidae),

etc. |

2. Vestiture primarily short, appressed, and dense. [In most bees the

pubescence is more erect, more sparse, and commonly longer, also

commonly less densely plumose. ]

3. Oral vestiture of female composed of simple, long, apically wavy

hairs. [In most melittids it is composed of shorter, sparser,

plumose hairs. ]

4. Oral vestiture of female composed of long, dense hair. [See

character 3. ]

2. Head as long as broad or nearly so. [Head markedly broader than

long in most melittids. ]

6. Vertex concave medially seen from front. [The vertex is flat, or

in most Dasypodini/ convex, in most bees. In others where it is

concave, the concavity is broad, while in Xeralictoides it is a

rather small median concavity. ]

7. Clypeus flat. [The clypeus is convex in most melittids. ]

8. Prementum and labial palpi long (Fig. 188). [These structures are

of ordinary length in most melittids. ]

9. Mesoscutum narrow, so that intertegular distance is less than its

length. [The mesoscutum is distinctly shorter than the inter-

tegular distance in most melittids. This is character 66 of Table

10. Propodeal triangle dull, contiguously punctured, in contrast to ad-

jacent areas. [In other melittids, the triangle is either

similarly sculptured to adjacent parts of the propodeum, or is more

shiny than adjacent areas. |

Michener: Classification of Melittidae 89

11. Front tibia with several apical and subapical setae of outer sur-

face greatly thickened, spine-like. [Such modification unknown in

other bees. This is character 43 of Table 1. |]

12. Hind femur with subapical pit on posterior surface. [This feature

is not known in other bees. This is character 7 of Table 1. |

13. Basitarsal scopa parted, i.e., hairs diverging along posterior (=

upper) margin of basitarsus, so that this margin is bare. [In

other melittids the hairs do not diverge in this way and some of

them arise from the margin. |

14. Tergum V of female with apical hair band similar to those of

preceding segments. [In most melittids the prepygidial fimbria is

distinct from the bands of the preceding terga. |

15. Pygidial plate of female with area of plumose hairs medially. [In

other melittids such a hairy area is absent. |

16. Sternum VI of male with apical lobes apically fringed, otherwise

glabrous. [In other Hesperapis the lobes have erect subapical

brushes. |

17. Sternum VII of male with apical lobes long and slender, about four

times as long as greatest breadth. [In other Hesperapis and

related bees these lobes are relatively short and broad. ]

18. Gonostylus with inner lobe bearing two brushes. [In other

Hesperapis the inner lobe bears one brush or is hairless. |

19. Gonostylus with inner lobe cylindrical, glabrous. [In other

Hesperapis the inner lobe has one or two brushes of hair. |

20. Penis valve biramous and strongly angled basolaterally. [In other

Dasypodini the penis valves do not show these features. |

Amblyapis lacks all of the apomorphies listed in Table 3. It could

therefore be a paraphyletic group from which Xeralictoides and other

groups of Hesperapis arose. Zacesta has its own apomorphies, not all

shared with any other group of Hesperapis. All the remaining taxa are

united by character 14, i.e., the prepygidial fimbria of the female is

reduced to an apical band like that of preceding terga. Although some

Capicola show the same attribute, and in Eremaphanta it is carried to an

extreme, there is no evidence that it arose more than once in the

Hesperapis-Xeralictoides group. If it did arise independently, of

course other taxa would arise from the lowest furcation of the

cladogram, which would become a quinquefurcation. Placement of the

remaining taxa is indicated in Figure 191, which suggests that Carinapis

is a paraphyletic group from which Hesperapis s. str. and Panurgomia

arose.

It should be noted that, although far apart in the cladogram (Fig.

IBN. Hesperapis. s. str. and Zacesta are quite similar, so that Stage

(1966) united them under the former name. Two of the three apomorphies

90 Contrib: Amer. Ent. Inst. yovel «Fey ne. 3, 1954

Hesperapis s. str.

Carinapis 3 Panurgomia

2 ;

2p.

ote

Disperapis 4 Xeralictoides

*8 ao

N \ a

Zacesta

: ay 14

Amblyapis

Figure 191. Cladogram for the subgenera of Hesperapis and the

genus Xeralictoides. The numbers represent apomorphies listed in Table

3. The data are from Stage (1966).

for each subgenus shown in the figure are the same. If characters 13

and 14 could be shown to arise independently in different taxa, then

Zacesta-Hesperapis could be viewed as a single taxon near the root of

the cladogram. Independent development of character 14 is not im-

probable; it clearly arose independently in the closely related genus

Capicola. Character 13, however, appears to be a unique development; it

is not Known in any other bees.

Key to the Subgenera of Hesperapis

(modified from G. |. Stage, 1966)

1. Propodeal triangle contiguously punctured throughout and dull, in

contrast to sparsely punctured and shining posterior and lateral

propodeal surfaces; pygidial plate of female more or less flat with

Michener: Classification of Melittidae 91

mesal area bearing patch of appressed, plumose hairs.

a ana a ag ca lag aad catenins teat 0 an als ant es acd os oa se ase neo Panurgomia

Propodeal triangle smooth and shiny, at most anteriorly rugose or

irregularly punctured, or if dull and closely punctured, then

posterior and lateral propodeal surfaces are similar; pygidial

plate of female often strongly convex with elevated basal triangle,

always without mesal hair patch.

sens, em ey omc rs vt” ssa etal gg ma ks uk oa na ms ng cf gad TS csp na ey “hrc en Mims 2

Females

ca nei mek eee ‘mba "een pasa (scence hn ie ys [yas aT: pe pcm cal a ee as ile gt “tse 3

Males

Sh neh ks ih “Sst, et ly Sb, Sl ls ni: “eS nl agi“ octal: ad ads le ll a fj

Hairs along upper (i.e., posterior) margin of hind basitarsus

parted, margin being narrowly hairless and exposed.

i a“ aa nm’ | fi ms mm. ay lh Jl gle pu ag nn Ot nog Ske" sc Oe a a en Sea mel a bi el etm: etn em 4

Hairs along upper (i.e., posterior) margin of hind basitarsus not

parted, arising on as well as on both sides of the margin.

wth inl Se tad ik yy Sep lp oR se rib Dosa on le Ake lk st WG ag 5

Pygidial plate with elevated, triangular basal area margined by

carinae which meet posteromedially (except in H. oliviae which

lacks these carinae); body length 7-15 mm.

aaa i ila Ll, kd ta ae cid roche etn clin Seng Carinapis

Pygidial plate flat with shallow longitudinal sulcus near apex,

without elevated area; body length 6-7 mm.

it ae oe sec tao ee ala eS eye hap hea ag ov ma eo el a oe Hesperapis s. str.

Head length only slightly less than width (9:10), so that head

seems elongate; labial palpus with segment 2 much longer than 1.

TA a A te ie ee Nt i oN ih a eS ih ee Zacesta

Head length much less than width; labial palpus with segment 2

little if any longer than 1.

sic ooh Bl SS age ac ls RCA WA cain eg a a NaM A ae ii. call iid aS es

Flagellar segment 1 at least 1.6 times length of 2

aieetielatatalaalelatatataiahatatatalatatatatatatetatetateteiatetatatatalatetatetalehetatatataeeataatale Disparapis

Flagellar segment 1 equal to or shorter than 2.

aaa aaa lalallala all lal all alata aiatatatatatetetelatetateteatetaatataabataaen Amblyapis

Labial palpus with segment 2 much longer than 1; head about as long

as broad.

aaa arlene aaleiaiahaletatatatatatetatalatetatatatatatetetetetatetatabatatetate Zacesta

Labial palpus with segment 2 little if any longer than 1; head

distinctly broader than long (except in some Hesperapis. s. str.).

Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

eet Amblyapis, Carinapis, Disparapis and Hesperapis s. str.

Good subgeneric characters are difficult to find among many males.

The following key to males that run to the second alternative of couplet

7 in the above key utilizes species-group characters, so that most sub-

genera come out in two or more places.

Key to the Subgenera of Certain Male Hesperapis

(modified from Stage, 1966)

Body length greater than 6 mm or metasomal terga without white

apical hair bands.

Body length 6 mm or less and metasomal terga with distinct, white,

apical hair bands.

Gey Cembe 41k a ls pau) cs ee ee Hesperapis’s.. Str.

Pygidial plate absent, metasomal tergum VII covered with

pubescence.

Bare, subtriangular pygidial plate present, usually elevated and

defined by carinae at least apically.

Posterior lobe of pronotum normal, not produced into spine; galea

either pale, reticulate, heavily punctured or with surface hair or

apical fringe.

Posterior lobe of pronotum produced into stout, weakly recurved

spine; galea dark, very sparsely punctured, otherwise glabrous.

ee oy ee eee OO a al Bo Carinapis

Metasomal sternum VI with apical lobes acute, dull, and bearing

inner, subapical brush of suberect hair; metasomal terga with

apical pale hair bands occasionally very weakly developed or ab-

sent. :

Metasomal sternum VI with apical lobes rounded, shiny and rela-

tively free of hair except for apically directed marginal fringe,

metasomal terga always with distinct pale apical hair bands.

eS ee ee eee Cree Mees Ree eRe Oe e Disparapis

Metasomal terga with uniform sized, well spaced punctures, shiny in

appearance.

Metasomal terga with irregular sized, contiguous punctures, dull in

appearance.

Michener: Classification of Melittidae 93

a a a aaa aaah lal al all oleate Amblyapis

6. Small species, less than 7.5 mm in length. :

-- Large species, more than 7.5 mm in length.

Disiumites f als s et altace 9 ete ied ics age ie ie tg ay oe i Carinapis

7. Apical lobes of metasomal sternum VI with subapical margins nearly

straight and inner margin of each lobe bearing elongate, submar-

ginal brush of suberect hair.

-- Apical lobes of metasomal sternum VI with apices produced laterally

and inner margin of each lobe bearing subovate, submarginal brush

of hairs in a whorl.

Subgenus Amblyapis Cockerell

Halictoides (Amblyapis) Cockerell, 1910, p. 362.

Type species: Halictoides (Amblyapis) ilicifoliae Cockerell, 1910

(monobasic and original designation).

Diagnosis. See characters underlined below.

_ Description. Small bees, slightly over 4 to 9 mm in length. Head:

Integument black or very dark brown with appendages often lighter.

Vestiture variable, generally suberect, sparse and long in females, that

of male longer and more dense, often obscuring integument, particularly

on face around antennal bases and clypeus laterally; oral vestiture nor-

mal, not modified as in female of Hesperapis. s. str. Head broad with

length from slightly less than 0.75 to slightly more than 0.85 width;

inner orbits of eyes nearly parallel to weakly convergent below in

female and weakly to strongly convergent below in male; vertex seen from

in front medially flat or weakly convex in female, more strongly convex

in male; clypeus weakly protuberant and weakly convex; first three

flagellar segments variable but 1 or 2 never longer than 3; mandible of

female generally narrowly rounded apically with well developed inner

subapical tooth, that of male similar but with subapical tooth area

weakly produced into blade. Mesosoma: Integument usually black,

sometimes dark reddish-brown on appendages. Vestiture generally longest

on mesepisternum and lateral faces of propodeum, shorter and sparser on

mesoscutum and very short and sparse or lacking on metepisternum.

Posterior lobe of pronotum not produced into recurved spine; propodeal

enclosure similar to adjacent areas, smooth or at most anteriorly sculp-

tured, but otherwise shiny and not punctured as in Panurgomia (except in

H. arida where it is so punctured but adjacent parts of propodeal in-

tegument are also heavily punctured and dull); jugal lobe between two

thirds and three fourths length of vannal lobe; mesotibial outer vesti-

ture primarily uniformly plumose but often with thick, spinose hairs in-

termixed apically; mesotibial spur translucent white, spiculate over en-

tire length, margins finely serrate; metatibial spurs similar to

mesotibial spurs; tibial scopa usually with plumose decumbant hairs

94 Contrib. Amer: Ent. iMist:> vol. 18;'no. 3, 1961

among long scopal hairs in some areas; basitarsal scopa not parted along

posterior (= dorsal) margin of basitarsus. Metasoma: Integument dark

reddish- brown to black, Araast the latter, terga I-IV of ek Le

REE HSa cubis recent for H. leucura and H. arida where hair bands are

well developed). Pygidial plate (female only) variable but not like

that of Disparapis, ranurgcma or Carinapis; seaniulh Vi of male with

apical lobes acute, ateranly directed directed and bearing brush aring brush or erect hair

in a whorl on inner subapical margin; sternum VII of male with apical

margin medially acutely produced or not between the two major lobes

which are well developed and narrowly rounded apically; male genitalia

with gonostylus bearing inner lobe with wer developed marginal fringe

composed. usually of both simple and plumose long hairs. (Stage)

Comments. This subgenus contains six species, one of which has not

been published. The named species are H. (Amblyapis) ilicifoliae

(Cockerell, 1910), parva Michener, 1937; Tarreae Cockerell, 1907;

leucura Cockerell, 1924; arida Michener, 1936. The range of the sub-

genus is in deserts from west Texas to Baja California, California, and

Nevada and in chaparral regions of southern and central California. At

least some of the species are oligolectic, but on unrelated plants so

that there is no particular host group for the subgenus. [Stage]

Subgenus Hesperapis Cockerell s. str.

Hesperapis Cockerell, 1898, p. 147.

Type species: Hesperapis elegantula Cockerell, 1898 (monobasic).

Diagnosis. See characters underlined below.

Description. Small species, usually less than 6 or 7 mm long.

Head: Integument black except for apical margin of clypeus and appen-

dages which may be reddish. Vestiture variable but mostly short, dense,

almost entirely plumose and often nearly obscuring integument of face face

and vertex; vestiture on inner basal parts of mandibles, on stipes

laterally, on prementum ventrally, and on genae and occiput ‘adjacent to to

proboscidial f fossa c composed of of long, simple, apically wavy hairs in

female; short, plumose in male. Head nearly round seen from in front

with length at least nine-tenths width (except in some males where head

is is usually slightly shorter); inner orbits of eyes slightly converging

below in female, weakly converging in male; vertex eennt ce clypeus

strongly convex and protuberant apically; flagellar seam ments 1 and 2

equal in female, 3 only slightly longer; in male 2 slightly longer than

1 and 3 markedly so so; middle flagellar segments of male much longer than

broad. Mandible with small preapical tooth, not ‘broadened into blade.

Mesosoma: Integument usually shiny, black except legs which may be red-

dish. Vestiture primarily composed of densely plumose hair and

generally short and dense on mesoscutum, longer and less dense but ap-

pressed on pleural areas and longer, more dense and appressed on

propodeum. Posterior lobe of pronotum normal, not produced into

recurved spine; propodeal enclosure sculptured or smooth but always

shiny and never punctured as in Panurgomia; jugal lobe between two-

thirds and three-fourths length of vannal lobe; mesotibial outer

Michener: Classification of Melittidae 95

vestiture dense, primarily uniformly plumose but females with thick,

spinose spinose hairs sp sparsely intermixed apically; mesotibial spur primarily

translucent silvery-white, spiculate except at apex, margins very finely

serrate; metatibial spurs similar to mesotibial spurs; tibial scopa with

short, plumose decumbant hairs among long simple hairs in some areas;

basitarsal scopa parted, i.e., with distinct narrow longitudinal

division exposing posterior | (= dorsal) edge c edge of ‘basitarsus. Metasoma:

Integument largely reddish in female, black or very dark brown, nearly

black in males. Terga bearing apical hair bands, that of tergum V of

female like that on preceding terga. Pygidial plate (female only) —

without | basal pubescence as in Paoueneeaia Ke ig triangle as in

male wit a apical margin Roti Sts two broadly amt lobes with

simple emargination between them; male genitalia with gonostylus bearing

inner lobe with well ¢c:. --loped marginal fringe composed of both simple

and plumose long hairs. (Stage)

Comments. Of the three species in this subgenus as_ here

restricted, “only one; Hesperapisetegantula Cockerell, has been

published. The subgenus occurs in deserts from western Texas to Idaho,

Nevada, and California. For pollen collecting the subgenus may be

restricted to the boraginaceous genus Coldenia. [Stage]

Subgenus Zacesta Ashmead

Zacesta Ashmead, 1899, p. 73.

Type species: Zacesta rufipes Ashmead, 1899 (monobasic).

Diagnosis. See characters underlined below.

Description. Small species, usually less than 6 or 7 mm long. In-

tegument black except that H. rufipes has apical part of clypeus reddish

in female and yellow in male, legs reddish in male of H. rufipes.

Vestiture mostly short, dense, plumose, nearly obscuring integument t of

face and vertex, especially in in male; vestiture of oral region not

specialized as in females of “Hesperapis s $2 str “Head nearly round seen

from front; inner orbits scarcely converging below in female, more

strongly so in male; vertex seen from cer flat or feebly concave me-

dially in female, feebly convex in male; clypeus strongly convex and

protuberant apically: first three flagellar segments not much +h different

in lengths; middle flagellar segments s of male much broader than long long.

Mandible of male male with inner tooth broadened into a blade. ~ Mesosoma:

Integument shiny, black except for legs which may be brown. Vestiture

primarily densely plumose, mostly short and dense on mesoscutum, longer

and less dense on pleural areas, longer, more dense and appressed on

propodeum. Posterior lobe of pronotum normal, not produced into spine;

propodeal enclosure smooth, shiny; jugal lobe 10 to nearly ./75 times

i of vannal lobe; mesotibial outer vestiture dense, mostly uniform-

ly plumose but females with thick, spinose hairs sparsely i intermixed

apically; mesotibial spur translucent silvery white, with very finely

serrate margins, lacking coarse serrations; metatibial spurs similar;

tibial scopa with short, plumose decumbant hairs among long simple

hairs; basitarsal scopa not parted along posterior (= dorsal) edge of

basitarsus. Metasoma: Integument largely red in most females and some

96 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

males. Terga bearing apical hair bands, that of tergum V of female

coarser, slightly darker, and forming broader band (= prepygidial fim-

bria) than on preceding terga. Pygidial plate (female only) without

basal pubescence or triangle. Sternum VI of male as described for

Hesperapis s. str.; sternum VII of male with apical margin produced to

two rounded lobes, emargination between them produced to small median

angle; genitalia as described for Hesperapis s. str.

Comments. Stage (1966) included two species in this group, which

he included in the subgenus Hesperapis s. str. Of these only one, H.

rufipes (Ashmead), has been named. Both species are known only from the

arid parts of central and southern California. For pollen collecting the

subgenus appears to be restricted to a few genera of Polemoniaceae,

especially Eriastrum (= Hugelia). [Stage]

Disparapis Stage new subgenus

Type species: Hesperapis arenicola Crawford, 1917.

Diagnosis. See characters underlined below. The extremely long,

narrow, apical lobes of sternum VII and the peculiar structure of the

inner lobe of the gonostylus with its two hair brushes are unique sub-

generic characteristics of the males. _

Description. Small to moderately large, 8-14 mm long. Head: In-

tegument usually black but may be dark reddish-brown; appendages may

also be dark reddish-brown. Vestiture mostly erect, sparse and long in

females, that of males longer and more dense, often obscuring integument

of face and particularly of clypeus; oral vestiture normal, not modified

as in female of subgenus Hesperapis. Head broad, length 0.75 to 0.85

width; inner orbits of eyes weakly convergent below in female, strongly

convergent below in male; vertex seen from in front medially flat or

weakly convex in female, strongly convex in male; clypeus of female

weakly protuberant apically and primarily flat, becoming convex only

laterally; clypeus of male similar but more convex; flagellar segment 1

in female at least 1.7 times length of 2 and always much longer than 3,

in male 1 subequal to” and 2

et REESE GREER 0 |

subapical inner tooth area weakly developed into blade (except both

sexes of H. arenicola in which mandibles are apically acute and lack

subapical tooth). Mesosoma: Integument usually black, appendages

sometimes dark reddish-brown. Vestiture relatively long, suberect and

Sparse on mesoscutum, scutellum, metanotum, mesepisternum and posterior

and lateral faces of propodeum and nearly lacking on metepisternum.

Posterior lobe of pronotum not produced into spine; propodeal enclosure

smooth or at most sculptured anteriorly, shiny, never punctured as in

Panurgomia; jugal lobe about two-thirds length of vannal lobe;

mesotibial outer pubescence plumose, without or with only several incon-

spicuous, thick, spinose hairs intermixed; mesotibial spur of females

amber colored, spiculate or not, with single inner row of coarse serra-

tions composed of well developed distinct teeth, of male translucent

white, spiculate, finely serrate; metatibial spurs of both sexes similar

to mesotibial spurs of male; tibial scopa with plumose hairs among long

scopal hairs in some areas; basitibial scopa not parted along posterior

Michener: Classification of Melittidae 97

(= dorsal) margin of basitarsis. Metasoma: Integument shiny, dark

reddish-brown to black, usually the latter. Terga bearing apical pale

hair bands, that of tergum V of female similar to those on preceding

terga, , Pygidial plate (female only) uniform, weakly convex pasaly

weakly concave apically, with abruptly reflexed iptly reflexed e edges, Ss, surface finely

papillose, someti sometimes with faint rugae sparsely rsely superimposed; sed; pygidial

plate lacking elevated basal triangle as in Carinapis and basal

pe ea as in Panurgomia; sternum VI of male with apical lobes broad-

ly rounded (laterally subacute in H. arenicola), smooth and shiny, —

without dense suberect brush of hair but a apically 1 fringed; sternum Vil

of male with a apica margin acutely produced medially and with two af apical

lobes e extremely long, narrow, apically somewhat thickened and id rounded;

male gonostylus bearing lol lobe with two distinct brushes, one one dorsally —

directed, one posteriorly directed. (Stage

Comments. This subgenus contains three species only one of which,

Ae (Disparapis s) arenicola Crawford, 1917, has been published. The sub-

genus occurs in deserts from west Texas and Zacatecas to Baja

California, California, and Utah. The species are probably restricted

to the Compositae for pollen collecting. [Stage]

Subgenus Panurgomia Viereck

Panurgomia Viereck, 1909, 48.

Type species: Panurgomia a feickal Viereck, 1909 (monobasic).

Diagnosis. See characters underlined below. The structure of the

inner process on the gonostylus of the male is unique to this subgenus.

Description. Medium sized bees, 7-15 mm long. Head: Integument

black except for occasional reddish-brown clypeus and gena posteriorly;

mouthparts primarily reddish-brown, thick parts often very dark. Vesti-

ture generally suberect, sparse and long in females, that of males

longer and more dense, often nearly obscuring integument, particularly

around antennal bases and on clypeus; oral vestiture not modified as in

female of Hesperapis s. str. Head shape variable, even in different

- populations of a single species, length varying from about 0.75 to 1.01

times width; inner orbits of eyes nearly parallel to weakly convergent

below in female, weakly to strongly convergent below in male; vertex

seen from front weakly rit iui inch clypeus piper lc variously

ay with well developed inner suber tooth; mandible of male

apically rounded with subapical tooth weakly broadened into blade.

Mesosoma: Integument usually black; pleural areas, propodeum and appen-

dages may be reddish-brown. Vestiture short and sparse on mesoscutum,

relatively long, suberect and sparse on mesepisternum and lateral sur-

face of propodeum, gradually becoming shorter towards posterior face of

propodeum; metepisternum nearly bare with only short and sparse plumose

hair. Posterior lobe of pronotum not produced into spine; propodeal en-

closure coarsely, uniformly, contiguously punctured, duller than spar -

sely pu punctured, shiny, adjacent propodea integument; jugal lobe of hind

wing one-fourth to three-fourths length of of vannal lobe; mesotibial outer

vestiture primarily uniformly plumose in male, female similar but with

98 Contrib. Amer. Ent. Inst., VoltetBy ne, 3, 1981

thick, spinose hairs sparsely intermixed apically; mesotibial spur

finely spiculate over almost entire length, finely serrate; metatibial

spurs similar to mesotibial spurs; scopa with short plumose hairs

reduced or lacking; scopa of basitarsus parted, i.e., with narrow

longitudinal division exposing posterior (= dorsal) edge of basitarsus.

Metasoma: Integument red, reddish-brown or black, usually the latter.

Terga bearing pale apical hair bands, that of tergum V in female similar

bearing brushes of suberect plumose hair along apical margins; sternum

Vil of male with apical margin weakly, acutely produced medially and

with pair of well developed, subacute or narrowly rounded lobes; male

genitalia with gonoforceps bearing a short, brushless, subcylindrical

process in place of inner lobe. (Stage)

Comments. Panurgomia, as recognized here, is a small, distinct

subgenus whose species are similar. Species included by Stage (1966)

are Hesperapis regularis (Cresson, 1878); fuchsi (Viereck, 1909);

semirudis Cockerell, 1910; nitidula Cockerell, 1916; pellucida

Cockerell, 1925, and wilmattae Cockerell, 1933. The subgenus ranges

from Arizona, Sonora, and Baja California to northern Nevada and

California, in both deserts and mesic foothill areas. Pollen collecting

is from different families of plants--Onagraceae, Papaveraceae, and per-

haps Compositae. [Stage]

Carinapis Stage new subgenus

Type species: Hesperapis carinata Stevens, 1919.

Diagnosis. See characters underlined below. The structure of the

pygidial plate in the female is a unique character for this subgenus.

Though not universal in Carinapis, the spineform posterior lobe of the

Ppronotum and the well developed male pygidial plate are diagnostic

characters when present since neither occurs in other subgenera.

Description. Medium to moderately large bees, 7 to 16 mm long with

most species near larger end of range. Head: Integument usually black,

rarely dark reddish-brown; appendages black or more often reddish-brown

with antenna sometimes orange below. Pubescence generally sparse and

long in female, that of male longer and more dense, often obscuring in-

tegument, particularly around antennal bases and clypeus; oral vestiture

not modified as in female of Hesperapis s. str. Head broad, length

varying from 0.75 to 0.86 width in female and from 0.72 to 0.81 width in

male; inner orbits of eyes nearly parallel to weakly convergent below in

female, weakly to strongly convergent below in male; vertex seen from

front medially flat or weakly convex in females, usually much more con-

vex in males; clypeus weakly protuberant, usually weakly convex,

sometimes nearly flat or even concave; first three flagellar segments

variable in male, in female segment 1 longer than 3, 3 longer than 2

(except in H. fulvipes female where 2 and 3 are subequal, each shorter

than 1, and in H. macrocephala female where 1 and 3 are subequal, each

longer than 2); mandible of female generally narrowly rounded or

subacute apically with well developed inner subapical tooth, that of

male apically similar to that of female but with inner subapical edge

Michener: Classification of Melittidae 99

weakly produced into blade. Mesosoma: Integument usually black,

sometimes dark reddish-brown or even yellowish-brown on appendages.

Vestiture generally long and dense (except on H. oliviae), longest on

mesepisternum and lateral face of propodeum, shorter and sparser on

mesoscutum and very short and sparse or lacking on metepisternum.

Posterior lobe of pronotum either normal or greatly produced into recur-

ved spine; propodeal enclosure smooth or at most sculptured anteriorly,

otherwise shiny, not punctured as in Panurgomia (except H. macrocephala

where it is similarly punctured but adjacent parts of propodeum are also

heavily punctured and dull); jugal lobe between one-half and two-thirds

length of vannal lobe; mesotibial outer pubescence primarily uniformly

plumose, without thick, spinose hairs intermixed; mesotibial spur

usually spiculate and very finely serrate along inner edge; metatibial

spurs similar to mesotibial spur of male; scopa usually with short

plumose hairs among long hairs; scopa of basitarsus parted, i.e., with

distinct, narrow longitudinal division exposing posterior (= dorsal) ed-

ge of basitarsus. Metasoma: Integument black, dark reddish-brown or

yellowish, usually the former. Terga bearing pale, apical hair bands,

that of tergum V_ in female similar in color, size and composition to

those of preceding terga. Pygidial plate of female with raised, sub-

triangular area, laterally margined by carinae, in basal middle (except

H. oliviae where raised basal area lacks carinate margins); tergum VII

RESON ERE REREURES NREL GRENIER SURES EN RO Le scueciieieimentemenmmrenammmemmeed eooweanera

or strongly produced, with well developed pair of narrowly rounded to

subacute apical lobes; male genitalia with gonostylus bearing inner lobe

with well developed brush or fringe composed of both basally thick, long

plumose hairs and shorter, often simple hairs. (Stage)

Comments. Carinapis contains more species and has a greater total

distribution than any other subgenus. Included are fifteen species, six

previously described, whose combined distributions cover much of North

America from Oregon, North Dakota and Illinois south to Baja California,

Morelos, and northern Florida. Described species are Hesperapis carinata

Stevens, 1919; fulvipes Crawford, 1917; macrocephala Cockerell, 1924;

oliviae (Cockerell, 1897); rhodocerata (Cockerell, 1897); and rodecki

Cockerell, 1934. Most species collect pollen from Compositae, but some

from Malvaceae and perhaps other families. [Stage]

Xeralictoides Stage new genus

Type species: Hesperapis laticeps Crawford, 1917.

Diagnosis. Distinguished from Hesperapis by the narrow mesoscutum,

spinose front tibia, and subapical pit on the posterior surface of hind

femur. Agrees with description of Hesperapis except for the strongly

concave to flat median part of the vertex seen from front, and can be

recognized especially by the characters indicated by underlining below,

in the description which was prepared by Stage (1966) for:comparison

with subgenera of Hesperapis (see Comments under Hesperapis for attribu-

100 Contrib. Amer: Ent. Inst., vol.-18, no: 3, 1981

tion of Stage's material). :

Description. Medium sized bees, 7-10 mm long, with males generally

as_ large as or larger than most females in any given population. Head:

Integument black except for clypeus which may be dark reddish-brown and

appendages which may be reddish or yellowish. Vestiture generally long,

suberect and sparse on posterior vertex and gena, shorter but more dense

and subappressed on face around antennal bases and lateral to clypeus

where integument is nearly obscured; clypeus nearly bare in female,

sparsely covered with short appressed hair in male; female with vesti-

ture of inner basal area of mandible, stipes laterally and ventrally and

areas adjacent to probosidial fossa unusually dense, with hairs long and

minutely-barbed; corresponding vestiture in males short, normal. Head

shape relatively uniform with length usually slightly more than 0.90

width in females and between 0.85 and 0.90 width in males; inner orbits

of eyes -nearly parallel or slightly converging below; vertex seen from

in front strongly concave to flat medially, usually the former; clypeus

moderately convex and protuberant apically; flagellar segments 1 to 3

usually subequal though segment 2 may be slightly shorter; mandible of

female broadly rounded apically with subapical tooth weakly developed;

a 0 CGE

brown. Vestiture short and sparse on mesoscutum, relatively long,

suberect and sparse on mesepisternum, short, appressed, fairly dense and

conspicuous on metepisternum and long, dense and subappressed, obscuring

integument, on posterior and lateral faces of propodeum. Posterior lobe

of pronotum normal, not produced into spine; minimum intertegular

distance less than length of mesoscutum; propodeal enclosure smooth,

finely reticulate or anteriorly striate but always shiny and never punc-

tured as in Panurgomia; jugal lobe of hind wing between seven-tenths and

three-fourths length of vannal lobe; front tibia with outer apical por-

tion spinose; mesotibial outer vestiture plumose, with thick, darker

spinose hairs sparsely intermixed at least apically; mesotibial spur

finely spiculate over basal three-fourths, inner margin with a few

coarse, well separated teeth in female, rather finely serrate in male;

hind femur with shallow, subapical pit on posterior surface; metatibial

spurs rather robust, curved, margins finely serrate; scopa with short

plumose hairs in addition to long scopal hairs; basitarsal scopa not

parted along posterior (= dorsal) margin of basitarus. Metasoma: In-

tegument reddish to black. Terga bearing pale, apical hair bands, that

a prepygidial fimbria, occur middorsally on tergum V.) Pygidial plate

(female only) without basal pubescence as in Panurgomia or elevated

basal triangle as in Carinapis, apical half commonly with longitudinal

median gentle ridge or convexity; sternum VI of male with apical lobes

acute, laterally directed and bearing brush of erect hair on inner

apical margins; sternum VII of male with apical margin produced medially

and with two well developed subacute apical lobes; male genitalia with

gonostylus bearing inner lobe covered with thick brush composed of long

recurved plumose hairs; penis valve strongly laterally angulate near

base and bearing a basal inner ramus (Stage

To complete the characterization of Xeralictoides in terms of

descriptions of other dasypodine genera, | list the following characters

not included in Stage's description. Except for characters h and r,

Michener: Classification of Melittidae 101

these limit the variability expressed in the description of Hesperapis

rather than indicating differences from that description. (Galeal combs

have not been examined for all species of Hesperapis; some may have

large combs like those of Xeralictoides.) c, d. Labrum about three

times as broad as long, convex, smooth and shining except for narrow,

hairy apical zone. e. Flagellar segments of male mostly longer than

broad. f. Flagellar segments of female mostly broader than long. g.

Maxillary blade about as illustrated for Hesperapis pellucida but with

strong marginal fringe. h. Galeal comb well developed, with about 12

spines; maxillary palpus exceeding apex of galea. i. Ligular arms

long, extending nearly to base of prementum. |. Labial palpus not

quite attaining apex of glossa. m. Basal vein a little over 1.5 times

as long as first abscissa of Rs, curved throughout, distal to cu-v. n.

Stigma as described for Capicola. 0. Prestigma less than half as long

as stigma. r. Propodeum seen in profile with subhorizontal basal zone

nearly as long as posterior declivous zone.

Comments. Xeralictoides contains two species, only one of which,

Hesperapis laticeps Crawford, has been named. These species from the

deserts of California and Nevada, are associated with two loasaceous

plants, Eucnide and Mentzelia. Further study of host preference and

floral behavior should be particularly interesting since the oral vesti-

ture of the female is modified apparently to aid in pollen collection

and perhaps even transportation. This vestiture, composed of unusually

long, dense, minutely barbed hairs, frequently is packed heavily with

pollen. [Stage]

As shown in Figure 191, Xeralictoides is a derivative of

Hesperapis; its recognition makes the latter a paraphyletic genus. |

believe that it is reasonable and desirable to recognize paraphyletic

taxa when phenetic considerations justify such action. In this case, |

consider the recognition of Xeralictoides at the generic level on

phenetic grounds debatable. It is clearly the most distinctive of the

groups derived from a proto-Hesperapis ancestor, as shown by the seven

apomorphies on its stem in Figure 191, as well as by characters h andr

above whose directionality is not known. For me these are reasonable

indications of generic distinctness. A decisive factor, however, was

recognition of Xeralictoides at the generic level by its describer, Dr.

G. |. Stage (personal communication), whose knowledge with this group

greatly exceeds my own. Phenetically (i.e., in terms of number of

characters) Xeralictoides is more different from Hesperapis than is

Capicola, but cladistically Xeralictoides obviously falls with

Hesperapis, since it shares with Hesperapis the synapomorphies that

distinguish that genus from Capicola.

Etymology. Named with reference to the halictid genus Xeralictus

which visits the same flowers and is superficially similar in ap-

pearance.

102 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

APPEND IX<ts2 THE-SPECIES:OF MEGANOMINAE

This subfamily, under the generic name Meganomia, was reviewed by

Stage (1971). Keys and descriptions of the genera are in the body of

the present paper. Most of the structural differences among the forms

are incorporated in those descriptions. The taxonomic descriptions

below are therefore largely limited to characters of sculpture and

coloration.

Genus Ceratomonia

This genus, so far as known, contains only the species described

below.

Ceratomonia rozenorum Michener new species

(Figs...7,°10,.. 4e48y 47:48; 87,:58;..198)

Meganomia sp. B, Rozen, 1977, p. 2.

Diagnosis. Last antennal segment of male black, flattened,

discoid; scutum and scutellum of both sexes finely punctate, nearly as

closely so as is possible; basitibial plate of female clearly defined.

Description. Size and structure as indicated in generic descrip-

tion. Male: Coloration. Black, the following parts yellow: labrum;

mandible except red-brown apex; clypeus; paraocular area, extending up

along eye margin to truncated summit half way from antennal base to

ocelli; supraclypeal area extending up to area above antennal bases and

continued as line to small yellow area below median ocellus; band along

posterior orbit, narrowed and separated from eye above, broadened and

joining yellow hypostomal area below; most of stipes; scape except dor-

sal black streak on distal half; pedicel and flagellar segment 1 except

dusky areas on dorsal surfaces; posterior margin of pronotum, both

laterally and dorsally, including much of pronotal lobe; anterior half

of tegula except testaceous translucent margin; axillary sclerites;

posterior margin of scutellum, yellow extending forward sublaterally to

varying extent; axilla; metanotum except laterally; venter of thorax,

extending up to middle of side on anterior surface of mesepisternum;

small spots at lower and upper ends of metepisternum; legs except for

black arolia, distal parts of claws, extreme bases of coxae, broad band

along upper inner surface of hind femur, and band along inner surface of

hind tibia; two large and almost contiguous spots on anterior surface of

metasomal tergum |, bands across terga I-VI, gradually narrowed medially

and broadened laterally, each approximately median on its tergum, being

immediately anterior to broad, depressed, marginal zone; bands of terga

I-11] sometimes with small sublateral emargination or enclosed black

spot on posterior edge sublaterally; tergum VII (reddish posteriorly); .

metasomal sterna (posterior ones reddish). Flagellar segments 2-10 and

base of 11 reddish yellow. Posterior half of tegula translucent

testaceous. Wings slightly dusky, especially apically; veins and stigma

dark brown except for yellow to yellowish-brown costa of forewing.

Michener: Classification of Melittidae 103

Pubescence. White, rather long and dense in vicinity of antennae and

ocelli, on genal and hypostomal areas; stipes; and thorax (dorsum of

last with intermixed very short hairs--see subgeneric description).

Metasomal terga |-V with rather long, well separated hairs arising just

basal to depressed marginal zones, which are hairless. Punctation.

Lower two thirds of face with punctures of various sizes intermixed,

separated by less than a puncture width of shining ground except for

nearly impunctate longitudinal median clypeal line. Upper part of face

and especially vertex with small punctures widely separated by minutely

roughened, dull ground. Scutum and scutellum dull, finely and closely

punctate, with scattered larger punctures from which long hairs arise.

Sides of thorax with rather close irregular punctures separated by very

little, but shining, ground. Propodeal triangle dull, granular. Terga

(except for broad, depressed, minutely roughened posterior marginal

zones) rather finely punctate, with intermixed larger punctures from

which longer hairs arise, punctures progressively coarser on more

posterior terga; ground between punctures dull on first few terga,

becoming more shiny posteriorly. Sterna more finely punctate than ter-

ga, ground between punctures minutely roughened but shining. Female:

Coloration. Black, the following parts yellow: basal two thirds of

mandible; clypeus except light brown apical margin; paraocular area as

in male but part above antennae more broadly yellow; supraclypeal area

(yellow area not reaching antennae); band along posterior orbit, but

slightly separated from orbit, especially above; small spot behind and

mesal from summit of eye; under side of scape; posterior margin of

pronotum (as in male or sometimes with little yellow below pronotal

lobe); tegula, axillary sclerites, scutellum, axilla, metanotum and

metepisternum as in male; posterior margin of propodeal triangle except

medially; apices of femora, on fore and mid legs extending nearly to

bases on under sides; tibiae, except brown hind basitibial plate covered

with short black hair; tarsi, becoming reddish distally (claws and

arolia colored as in male); anterior surface of metasomal tergum |;

bands like those of male on terga I-V, but bands not or little narrowed

medially, sublateral posterior emarginations not isolated as spots; an-

terior metasomal sterna reddish yellow or partly yellow, grading to

black on V and VI. Labrum light brown. Pedicel and flagellar segments

light brown beneath, dusky above, the darker coloration more extensive

on basal halves of most segments. Wing coloration as in male.

Pubescence. Mostly white, areas of long hairs on head less conspicuous

than in male, that of vertex and dorsum of thorax ochraceous, of under

sides of tarsi reddish yellow, of basitibial plate black; prepyigidial

fimbria and hairs on either side of pygidial plate dense, black.

Punctation. As described for male but area anterolateral to posterior

ocellus with few punctures on minutely roughened ground; tergal puncta-

tion coarser and deeper, contrasting even more with posterior marginal

zones; sternal punctures coarser but less dense than tergal.

Type material. Holotype male, South West Africa (= Namibia): 17-19

km east Usakos, March 18, 1976 (J. G. and B. L. Rozen). Allotype female,

same data but taken March 26. Nine male and eight female paratypes,

same data as holotype but two of each sex taken March 29 and two females

on March 22. Rozen (1977) reports that these bees were collecting pol-

len on Indigofera. Two male and nine female paratypes, 13 km west of

Karibib, March 14, 16, 24, and 26, 1979 (J. G. and B. L. Rozen), on

flowers of Petallidium except two females on Indigofera. The type

104 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

material is in the American Museum of Natural History except for two

pairs of paratypes each in the Snow Entomological Museum, University of

Kansas, and the British Museum (Natural History).

Etymology. This species is named for the collectors, Dr. and Mrs.

J. G. Rozen, in recognition of their interesting work on Meganomia, its

allies, and many other bees.

Genus Uromonia

So far as known this genus contains only the species described

below.

Uromonia stagei Michener new species

(Figs. 6, t1yal9724,1938,+5/)-58;-197)

Diagnosis. The smallest known meganomiine (10-12 mm long); antenna

simple; propodeum of female nearly all yellow; blade of galea fringed

with long hairs.

Description. Size and structure as indicated in generic descrip-

tion. Male: Coloration. Head and thorax black, metasoma black to red-

brown (holotype with first and base of last tergum red-brown, others

nearly black), legs often brown, the following parts yellow (often pale

yellow, especially on metasoma and tarsi, sometimes elsewhere): labrum;

mandible except reddish-black apical two fifths; clypeus; paraocular

area extending up along eye margin to truncated summit half way between

antennal bases and ocelli; supraclypeal area extending upward between

and above antennal bases; small spot (often asymmetrical or absent) an-

terolateral to lateral ocellus; scape except for dorsal black streak on

distal half; under side of pedicel and flagellar segment 1; stripe along

under side of flagellar segments 6-11; posterior margin of pronotum,

both laterally and dorsally, broken middorsally and variably at side,

including most of pronotal lobe; anterolateral area on mesoscutum,;

streak along notaulus on anterior part of mesoscutum; pair of small

spots (or in some paratypes a broken line) across posterior margin of

mesoscutum (or this area all black); spot on tegula; outer half of

median axillary sclerite; posterior third (or in some, half) of scutel-

lum; posterior half (or in some, two thirds) of metanotum; sternum

between middle and hind coxae (in some extending forward to front cox-

ae); large areas on ventral surfaces of coxae, trochanters, and femora;

tibiae except dark brown areas on posterior surfaces, largest on hind

tibia where there is more brown than yellow; basitarsi; small segments

of front tarsus, large areas on small segments of middle tarsus, and

small areas on these segments (especially segments 2 and 5) of hind tar-

sus; narrow bands just anterior to depressed marginal zones of terga |

to VI, these bands narrowed at lateral extremities, then continued an-

teroventrally on ventrolateral parts of terga; most of dorsal surface of

tergum VII; sterna | to |V and part of V (remaining exposed sterna red

Figures 192-198. Outer views of hind legs of males of

Meganomiinae. Shaded areas brown or black, clear areas yellow. 192,

—_—_—_——$_$—$_—$——_—$

Michener: Classification of Melittidae 105

495M. rossi nN. sp.

197, Uromonia stagei n. sp. 198, Ceratomonia rozenorum n. sp. Scale

lines = 0.5 mm.

106 Contrib. Amer. Ents Insts; vel. 16, ne. 3, TST

brown). Flagellum dark brown, lighter beneath, except for yellow areas.

Tegula and axillary sclerites light brown except for yellow areas.

Wings slightly dusky, a dark streak beyond apex of marginal cell; veins

and stigma brownish black, black basally, costa mostly brown.

Pubescence. Dull white, fuscous on vertex, and with black or fuscous

hairs intermixed on scutum and more densely on scutellum; metasomal ter-

ga with hairs that mostly look pale in some lights but fuscous in

others, depressed marginal zones hairy. Punctation. Lower half of

clypeus with moderately coarse punctures of variable sizes; rest of

clypeus with small punctures, widely but variably separated by shining

but finely roughened ground; longitudinal median ridge impunctate.

Supraclypeal area more densely punctate than adjacent areas. Rest of

face with small punctures widely separated by shining but finely

roughened ground. Vertex dull, largely without distinct punctures, but

with numerous small punctures in ocellar triangle. Genal area with

punctures separated by about a puncture width. Dorsum of thorax dull,

minutely granular, hair bases distinct but not in conspicuous punctures.

Sides of thorax with small, shallow punctures separated by shining but

minutely roughened ground. Metasomal terga dull, granular, with widely

separated small punctures, larger and more conspicuous on areas just an-

terior to depressed marginal bands; seventh tergum with coarser punc-

tures, particularly in area of pygidial plate. Sterna with large, im-

punctate although finely roughened areas; punctation more prominent on

posterior sterna. Female: Coloration. Head and thorax black and legs

brownish to black, the following parts pale yellow: basal third of

mandible; lateral third and longitudinal median stripe on clypeus, these

areas united along apical clypeal margin; lower paraocular area, not

reaching subantennal suture, truncate at level of antennal base, but ex-

tending well above that level as slender pointed projection along inner

orbit; irregular area on supraclypeal area, extending upward between an-

tennae to level well above antennal bases; under side of scape;

posterior margin of pronotum (less yellow than in male); streak along

notaulus on anterior part of mesoscutum; transverse band across

posterior margin of mesoscutum; spot on tegula; outer half of median ax-

illary sclerite; posterior three fifths of scutellum; most of axilla;

metanotum; metepisternum; propodeum except for propodeal pit; apices of

front and middle femora (extending well toward base on upper surface of

mid femur); anterior and much of under surfaces of front and middle

tibiae; and most of outer surfaces of front and middle basitarsi.

Metasomal terga I-IV yellowish brown with apical depressed zones dark

brown; pale yellow bands as in male but somewhat broader, terga II-IV

with basal yellow bands also, joined to other yellow bands laterally,

and separated from them only by irregular yellowish brown bands; tergum

V yellow with brownish black apical margin; tergum VI brownish yellow,

pygidial plate dark brown; terga III and IV with brownish black blotches

at extreme sides, not visible from above. Sterna I-III yellow, I] and

111 with blackish median blotches; sterna |V and V red brown; sternum VI

brown. Labrum and extreme anterior margin of clypeus reddish brown.

Flagellum dark brown above, light brown beneath. Tegula transparent

testaceous except for yellowish spot. Axillary sclerites light brown

except for yellow area. Tarsi reddish brown except for yellow areas.

Wings as in male. Pubescence. Of head and thorax as described for male

except that on under surface of mesothorax and metathorax and associated

coxae and trochanters, it is stiff and brassy. Metasoma with hairs of

Michener: Classification of Melittidae 107

first tergum nearly all white, of second partly so; remaining terga with

hairs fuscous or blackish, depressed marginal zones I! to IV with

blackish hairs. Prepygidial fimbria and hairs beside pygidial plate

blackish. Pubescence of legs white, yellowish on undersides of tarsi,

patch of black hairs on outer side of middle tibia apically and on base

of hind tibia, including basitibial plate. Punctation. That of head and

thorax similar to that of male; punctures of head coarser than in male

and those of clypeus nowhere widely separated. Metasomal terga with

ground somewhat shining, more so on posterior terga, but depressed mar-

ginal zones dull; punctures minute on marginal zones, larger elsewhere,

finer and more widely separated than in male except on terga V and VI

and sides of |V where they are coarser. Sterna II and III largely

shining, minutely roughened, with only widely separated minute punc-

tures, but posterior margins of each finely and densely punctate; other

sterna nearly wholly finely and densely punctate.

Type material. Holotype male, allotype female, and one female and

14 male paratypes, Kenya: 13 miles (21 km) south of Malindi, 26 May 1967

(C. D. Michener). The locality is approximately at sea level and within

2 km of the coast. The holotype and allotype are in the Snow

Entomological Museum, University of Kansas; paratypes are in the

American Museum of Natural History, and the British Museum (Natural

History).

Etymology. This species is named after Dr. G. |. Stage in recogni-

tion of his work on melittid bees.

Genus Agemmonia

This genus contains two named forms, as indicated below. The male

of A. tsavoensis is here described for the first time.

Agemmonia tsavoensis (Strand)

(Figs.«.9,. 12,°25*80 54,87, 58727492)

Nomia tsavoensis Strand, 1920, p. 93.

Meganomia tsavoensis: Cockerell, 1931a, p. 201; Stage, 1971, p. 312.

Description. Size and structure as indicated in generic descrip-

tion. Male: Coloration. Black, the following parts pale yellow or yel-

lowish white: clypeus except narrow black apical margin; lower para-

ocular area, narrowed abruptly at level of antennal bases, and continued

along eye margin well above that level; lower part of supraclypeal area

(colored area over twice as broad as long); small V-shaped mark or pair

of converging marks in front of ocelli; area on genal area, upper end of

which diverges from posterior orbit; pair of transverse marks on

posterior pronotal margin dorsally (one specimen only); spot on tegula;

outer part of median axillary sclerite; sometimes weak streak along an-

terior part of notaulus; spot on axilla; pair of ventroapical streaks on

hind femur; spot on base of each tibia, often continued as stripes that

run length of outer sides of tibiae or such stripes indicated by

isolated area on each tibia, such white areas beyond basal spots

sometimes absent on front and middle tibiae; narrow bands across

108 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

metasomal terga just anterior to depressed marginal zones, band of ter-

gum | broken medially and sometimes divided into four small spots, bands

of terga II-V with broad sublateral emarginations on anterior margins,

that of || sometimes broken medially, band of tergum VI! broken into

three spots. Labrum, median third of mandible, under side of distal

half of flagellum, distitarsi, and axillary sclerites (except yellow

area) brown. Arolia and apices of claws black. Tegula except for yel-

low spot translucent testaceous, blackish anteriorly. Wings yellowish

brown, more dusky apically; veins brown, stigma infuscated. Pubescence.

Rather short, that of head and thorax strongly plumose, rather long on

under side of head including mandibles, and on stipes and front coxae;

pubescence of legs and metasoma extremely short. Color mostly whitish,

slightly ochraceous on vertex, mesoscutum, and scutellum, strongly

ochraceous on venter of thorax and coxae. Punctation. Lower half of

face with punctures of various sizes intermixed, separated by somewhat

shining, minutely roughened ground; longitudinal clypeal ridge impunc-

tate. Rest of head and thorax with minute punctures widely separated by

dull ground, punctures somewhat coarser and ground more shining on genal

area and lower three fourths of mesepisternum. Metasomal terga almost

without recognizable punctures except for VI and VII, surface dull, more

shining on VI and VII. Sterna a little more conspicuously punctate than

terga. Female: Coloration. Black, legs and metasomal venter red brown,

the following parts pale yellow: small spot on base of mandible (nearly

absent in one specimen); clypeus except black apical margin; paraocular

area as in male but with narrow point along eye margin up to level of

ocelli; supraclypeal and frontal marks as in male; genal mark as in male

but larger; spot on pronotal lobe and pair of transverse marks on

posterior margin of pronotum dorsally; spot on tegula; outer part of me-

dial axillary sclerite; lateral stripe on scutum; stripe along notaulus,

attenuated posteriorly but joining side of triangular spot, of which

there is a pair on posterior scutal margin; transverse streak on each

side of posterior half of scutellum (one specimen); axilla; transverse

band on anterior half or two thirds of metanotum; lateral area of

metanotum; broad arcuate band across base of propodeal triangle; basal

spots on front and middle tibiae; narrow bands across metasomal terga

anterior to depressed marginal zones, those on terga III-V more or less

midway between these zones and graduli, band of tergum | broadly broken,

of tergum || rather narrowly broken, of terga III-V very narrowly broken

or continuous. Labrum and under side of scape brown. Under sides of

flagellar segments 4-10 red brown. Tegula blackish anteriorly, brown

posteriorly (except for yellow spot). Small tarsal segments reddish

brown, claws and arolia colored as in male. Wings as in male.

Pubescence. As in male but not long and dense on undersides of head and

front coxa, and not unusually short on legs and metasoma; hairs of legs

and metasomal scopa ochraceous; hairs of terga ochraceous, fuscous to

black on terga V and VI. Punctation. As described for female, but lower

part of face with fewer coarse punctures.

Type material. The type, from the Tsavo River, Kenya, taken in

May, a female, is in the Musee Royal de I|'Afrique Centrale, Tervuren. |

have not studied it. It was redescribed by Cockerell (1931a). It has

slightly less extensive yellow markings (e.g., more broadly broken

metasomal bands) than the material on which the above description was

based.

Michener: Classification of Melittidae Tes,

Distribution. Known only from Kenya, but probably occurs in arid

lowland parts of adjacent countries. The specimens on which the above

description is based were from the Mombasa-Nairobi highway 80 km

northwest of Mombasa, 31 May 1967 (C. D. Michener).

Agemmonia tavetensis (Cockerell)

Meganomia tavetensis Cockerell, 1934, p. 444; Stage, 1971, p. 311.

Description. Size and structure as indicated in the generic

description. Female: Agrees with description of A. tsavensis except as

follows: Yellow of paraocular area with slender upper orbital point

reduced, not reaching level of ocelli; yellow stripes along notauli not

reaching spots on posterior scutal margin; outer surfaces of front and

middle tibiae largely yellow; and yellow tergal bands broad, only the

first narrowly broken, the others continuous and occupying most of the

space between the gradulus and the anterior margin of the apical

depressed zone on each tergum.

Type material. The holotype, from the Lumi River, Taveta, Kenya,

taken in December, is in the British Museum (Natural History). The

paratype, on which the above comments are based, is in the National

Museum of Natural History, Washington; it is from the type locality.

Comments. This is probably a color form (i.e., synonym) of A.

tsavoensis. The female (male is unknown) seems morphologically identical

to that species. The supposed difference (lack of the median clypeal

ridge in A. tsavoensis) mentioned by Cockerell (1934) and Stage (1971)

does not exist and is the result of an error. Strand (1920) mentioned

such a ridge, but Cockerell (1931a), when he redescribed the type,

failed to note it. When he described A. tavetensis, Cockerell (1934)

evidently read his own earlier description but not Strand's, and

therefore incorrectly concluded that the ridge was absent in A.

tsavoensis. As pointed out by Stage (1971), the only known locality for

M. tavetensis is in the same general area from which A. tsavoensis is

known.

Genus Meganomia

This genus contains four distinct species separable by the keys

below. Certain unusual features of some of the species are mentioned in

the generic description and even in the subfamilial description, but to

Save space are not repeated in specific descriptions.

Key to females

1. Mesoscutum black or with yellow only near anterior margin, narrow

stripes occasionally following notauli up to five sixths of length

of scutum; yellow of clypeus continuous from side to side.

-- Mesoscutum with yellow along lateral margins and yellow stripes

along notauli extending nearly to posterior margin of mesoscutum;

110

Contrib. Amer. Ent. Inst.;, vol. 18, no. 3, 1981

yellow of clypeus divided by median black zone.

Clypeus and scutellum entirely yellow or clypeus with small black |

areas along upper margins; yellow band of first tergum continuous;

forewing length over 13 mm.

Clypeus with pair of basomedian black areas extending more than

half way down clypeus; scutellum with at least anterior margin and

commonly anterior half or more black; yellow band of first tergum

broken medially; forewing length less than 12 mm.

Last antennal segment scarcely longer than broad; mesepisternum and

metepisternum without yellow areas.

Last antennal segment conspicuously longer than broad (as in

binghami and gigas); mesepisternum and metepisternum with yellow

areas.

AER. Ci Agicca MMM ae oc ok. Dee RY aah are evga ese e cuter a ade adel ei oan a ce baie ea andersoni

Key to Males

Flagellum not crenulate beneath, segments 8 to base of 11 of almost

uniform width but apical part of 11 strongly narrowed and forming a

hook; fore coxa produced posteriorly far behind base of trochanter;

fore basitarsus with relatively short yellowish hairs; fifth tergum

with two apical, black combs, one on either side of midline.

Flagellum crenulate beneath, segments 8 to 11 progressively nar-

rowed and commonly forming a hook; fore coxa not produced

posteriorly behind base of trochanter; fore basitarsus with black

brush of long hairs on under side; fifth tergum with a single

median apical black comb. :

Fore wing length 13 mm or more; yellow band of first tergum con-

tinuous; prominence on anterior margin of inner surface of

posterior tibia nearly two thirds of distance from base to apex of

tibia.

Fore wing length under 12 mm; yellow band of first tergum usually

broken; prominence on anterior margin of inner surface of posterior

tibia little more than half way from base to apex of tibia.

Hind tibia with strong emargination in anterior margin; last anten-

nal segment dark in contrast to three preceding segments.

Michener: Classification of Melittidae 111

-- Hind tibia without emargination in anterior margin; last antennal

segment largely yellowish like three preceding segments.

se ho ge, tc hit se en na eg I Re a andersoni

Meganomia binghami (Cockerell)

(Figs.:31-36, 58; 193)

Nomia (Meganomia) binghami Cockerell, 1909, p. 402; Cockerell, 1910, p.

221; Hedicke, 1931, p. 35 |

Nomia flavofasciata Friese, 1909, p. 170; Friese, 1930, p. 14.

Meganomia bin ar Cockerell, 1931a, p. 201; Cockerell, 1933a, p. 376;

Cockerell, , Poe]; Stage, 1971; De OH.

Diagnosis. This species, like M. gigas, has only one slender

apical antennal segment in the male. It differs from that species in

the somewhat less extensive yellow maculations, those of the posterior

part of the thorax and metasoma being bright yellow rather than pale as

in gigas. The hind coxa of the male is thickened apically so that its

under surface is well below that of trochanter, and in side view the

coxa appears to have an apical ventral tooth.

Description. Male: Length 16-18 mm; forewing length 9.5-11.2 mm.

Structure (characters listed are only those that vary within the genus).

1. Produced median lobe (truncation) of clypeal apex over one third as

wide as clypeus, apex straight (truncate). 2. Frontal carina below an-

tennal bases convex as seen in profile. 3. Apex of mandible with two

teeth (apices of rutellum and pollex) of about equal length. 4. Labrum

over 2.5 times as wide as long, flat, apex somewhat emarginate medially.

5. Flagellum not crenulate beneath, segment 1 nearly three times as

long as broad, segment 2 longer than broad, subsequent segments progres-

sively shorter so that 5 is about as broad as long and 11 is markedly

broader than long; segment 12 equally broad at base but thereafter

abruptly narrowed to form flat, strongly curved hook, longer than any

flagellar segment except 1. 6. Front coxa projecting as a flat

process, broadly rounded at apex, extending well behind base of

trochanter. 7. Middle coxa with large, shining process projecting ven-

trad in front of base of trochanter. 8. Hind coxa in profile with

broad apical tooth, its apex well below level of base of trochanter. 9.

Hind femur less than twice as long as broad in side view. 10. Hind

tibia viewed from inner side with prominence on anterior margin scarcely

beyond middle. 11. Apex of hind tibia with anterior half (outer view)

straight, slanting to rounded but acute anterior apical angle. 12.

Front basitarsus with rather long hairs (one third as long as basitar-

sus) on posterior margin, other basitarsal hairs much shorter. 13.

Hind basitarsus with strong outer ridge at base, which is then abruptly

reduced (almost at a right angle) and forms an arcuately excavated under

surface of median part of basitarsus. 14. Basal vein of forewing basal

to cu-v and over five times as long as first abscissa of Rs. 15.

Metasomal sternum IV without pair of strong tubercles on disc; sternum V

with two apical black combs, one on each side of midline; base of ster-

Tle Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

num with two well defined, invaginated, membranous areas. 16. Apical

lobes of sternum VI rounded, much broader than long. 17. Apex of

gonocoxite with diagonal row of bristles on under surface from inner

apical angle toward subapical outer margin. 18. Penis valves broadly

fused basally. 19. Volsella as usual in the genus, with finger-like

process projecting distally beneath basal part of penis. Coloration:

Black, the following parts yellow: mandible except reddish-black apex;

labrum; clypeus; supraclypeal area except lateral margin, yellow ex-

tending up as narrow line between antennae and sometimes half way to

median ocellus; lateral face mark occupying entire lower paraocular area

and above truncated half way between antenna! base and lateral ocellus,

sometimes extending beyond that level as narrow point along eye margin;

small spot lateral to median ocellus; under sides of scape, pedicel, and

flagellar segment 1; narrow line along lower half of posterior orbit;

sometimes yellow area on upper half of genal area away from eye margin;

large area on stipes; posterior margin of pronotum (broken mid-dorsally,

broadened on posterior lobe which may be largely yellow, and extending

down on lateral margin toward front coxa); area along anterior fifth of

notaulus; commonly disjunct area lateral to notaulus near front of

scutum; anterior half of tegula; axilla or large area on it; line along

posterior margin of scutellum or posterior half of scutellum; transverse

band occupying most of dorsal median part of metanotum; venter of thorax

between coxae; under surfaces of coxae (including ventral process of mid

coxa); trochanters to apices of tarsi of front and middle legs except

brownish-black distal halves of claws, large black area on posterior

side of front femur, basal black streak on dorsal side of front femur,

small black spot on under side of front tibia, and black area on under

and posterior side of middle tibia; hind trochanter, femur, tibia, and

basitarsus yellow with irregular black areas on upper side of trochanter

and femur extending onto front and rear surfaces of latter, blackish

areas around femoro-tibial joint, black distal three fifths of inner

surface of tibia extending onto upper and lower surfaces of tibia and as

premedian band nearly across outer surface of tibia, and black posterior

surface of basitarsus; transverse bright yellow bands in front of

depressed posterior margins on metasomal terga | to VI, band on tergum

usually broken medially (sometimes broadly), on terga || to IV or V nar-

rowed medially, that on || sometimes narrowly broken, on terga | to IV,

V, or VI with shallow sublateral emargination on posterior margin, on VI

with median notch in anterior margin; broad pale yellow bands, often

broken medially, on sterna II to |1V, with oval black spot on each side

of each sternum, on IV spot sometimes represented by emargination in an-

terior margin of yellow band; posterior margin of sternum | sometimes

narrowly yellow. Flagellar segments 1-10 blackish dorsally, 2-11 other-

wise yellowish brown. Posterior half of tegula translucent red brown.

Wings slightly brownish, membrane beyond veins dusky, veins and stigma

dark brown to black. Pubescence. Moderately abundant, white, ochreous

(fading to white) on vertex, dorsum of thorax, and metasomal dorsum,

coppery on under sides of tibiae and tarsi. Punctation. Punctures of

whole body mostly separated by less than or only slightly more than a

puncture width of slightly roughened ground, between ocelli small and

dense, lateral to ocelli small and sparse, on scutellum distinctly

sparser than on scutum and more widely separated with shiny ground

between them; tergal margins broadly depressed, smooth, shining, with

only scattered minute punctures and scattered hairs; sterna with punc-

Michener: Classification of Melittidae 113

tures small and inconspicuous. Female: Length 15-17 mm. Forewing

length 11.5 mm. Structure. Labrum about 2.5 times as broad as long,

apex medially emarginate. Inner orbits approximately parallel so that

clypeal width is about equal to upper interorbital distance. Front

coxal process narrowly rounded at apex, about half as long as

trochanter. Wing venation as in male. Coloration. Black, the following

parts yellow: basal half of mandible; clypeus except pair of black

areas descending more than half way across clypeus from upper margin,

sometimes surrounding and isolating yellow area between them; lower

median part of supraclypeal area; lateral face mark as in male but trun-

cation at upper end sloping upward away from eye; small spot lateral to

median ocellus; under side of scape; band along upper half of genal

area, away from eye margin; sometimes area on stipes; pronotal margin as

in male or broken or absent below lobe which is usually largely yellow;

short streak along anterior end of notaulus (in Beitbridge female over

one-third length of scutum) and sometimes a dot (or large area in

Beitbridge and Louis Trichardt specimens) lateral to anterior end of

this streak; anterior half of tegula and streak along inner margin to

posterior end of tegula; axilla or area on it; line along posterior mar-

gin of scutellum (broken medially) or usually posterior half of scutel-

lum, or in Beitbridge specimen whole scutellum except anterior margin;

band on metanotum as in male; band across and sometimes occupying whole

dorsum of propodeal triangle, this band sometimes broken medially;

sometimes pair of irregular spots on posterior surface of propodeum;

dorsal apices of front and middle femora; basal third to half of front

tibia except on under side, this yellow area continued as stripe that

often fuses with distal patch on anterior surface; basal third of middle

tibia except on under side; distal patch on anterior surface of middle

tibia; bands on metasomal terga | to V in front of depressed marginal

zones, that on | sometimes narrowly broken medially, on I! and II!

somewhat narrowed medially, on | to 1V with shallow sublateral emargina-

tion on posterior margin, on V broader than on other terga and with

median notch on anterior margin often hidden by tergum IV. Labrum and

tarsi brown. Flagellum black to brown above, yellow brown beneath.

Tegula and wings as in male. Metasomal sterna and rarely also terga

(between yellow bands) brown or dark brown. Pubescence. As in male but

hairs of undersides of coxae and trochanters, and of thoracic sternum

variably brassy; distal patches of coppery hairs on outer sides of fore

and middle tibiae; hairs of truncate apex of hind femur fulvous, of

basitibial plate red, grading to fuscous apically; prepygidial and pygi-

dial fimbriae and hairs of sterna, especially apically, fulvous.

-Punctation. As described for male but metasomal sterna more con-

spicuously punctate; depressed marginal zones of terga hairless or

nearly so except laterally; tergum V without such a zone, apex densely

hairy.

Type material. The type, a female, of M. binghami from Damarland,

Namibia (South West Africa), is in the British Museum (Natural History ).

The type of M. flavofasciata from Grootfontein, Namibia, was a female

but the specimen so labelled in the Zoologisches Museum, Berlin, is a

male from Otjiverongo, presumably collected in 1920 and described by

Friese in 1930. Obviously it is erroneously labelled as type. The true

type is perhaps lost, but Friese's description clearly shows that it was

a specimen of M. binghami, not M. gigas (e.g., length 15 mm; posterior

half of scutellum yellow). ;

114 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

Distribution. Arid parts of southern Africa north of the Tropic of

Capricorn. South Africa: 40 km north of Louis Trichardt, Transvaal, 610

m altitude, March 24, 1967 (C. D. Michener); 33 km north of Vanzylsrus,

Cape Province, March 3, 1980 (V. B. Whitehead). Zimbabwe (Rhodesia):

Beitbridge, April 12, 1932 (L. Ogilvie, A. Mackie). Namibia (South West

Africa): 73-76 km south of Outjo, March 25, 1979 (J. G. Rozen);

Otjiverongo, January 11, February, 1920; 27 km south Okahandja, February

17, 1977 (J. G. and B. L. Rozen), one female on Gisekia africana;

Grootfontein; Damarland; Seeheim, February 16-19.

Meganomia gigas Michener new species

(Figs. 55-57, 59, 194)

Meganomia binghami: Rozen, 1977, p. 1-14 (misidentification).

Diagnosis. This is the largest known melittid. It closely

resembles M. binghami but is larger, with more extensive, pale yellow

maculations, without a hind coxal tooth in the male.

Description. Agrees with description of M. binghami except as fol-

lows: Male: Length 17-21 mm; forewing length 12-13 mm. Structure. 1.

Produced median lobe (truncation) of clypeal margin less than one third

as wide as clypeus. 3. Apex of mandible with lower tooth (apex of

rutellum) markedly exceeding upper tooth (apex of pollex). 5. Median

flagellar segments somewhat longer than in M. binghami, so that segment

2 is somewhat longer than broad. 6. Front coxal process convex, nar-

rowly rounded at apex. 8. Hind coxa without preapical tooth, its under

surface being at same level as that of trochanter. 9. Hind femur about

twice as long as broad in side view. 10. Hind tibia viewed from inner

side with prominence on anterior margin distinctly beyond middle (this

is the more distal prominence seen from outer side). 13. Hind basitar-

sus with outer basal ridge less high than in M. binghami, abruptly

reduced (at an obtuse angle) to more shallowly excavated median part of

basitarsus. 16. Apical lobes of sternum VI less broadly rounded than

in M. binghami. Coloration. Yellow of scutellum expanded forward on

each side of middle (but not at extreme sides) so that at point of max-

imum, yellow area occupies two thirds to five sixths of scutellar

length; under surface of front coxa without or with only patches of yel-

low; front and middle trochanters with black basal areas, femora with

black on posterior surfaces, tibiae with black on under and commonly

posterior sufaces; hind trochanter largely black, under side of hind

femur black, continued onto entire posterior surface and joining black

of upper side; outer surface of hind tibia yellow with apical and median

dark spot, inner surface with base and distal half black, this black ex-

tending onto lower surface; yellow of metasomal bands pale like that of

rest of body, bands broad, unbroken, gradually narrowed medially except

that on VI which has median notch in anterior margin, only that of first

and sometimes second tergum with shallow sublateral emarginations on

posterior margins. Posterior half of tegula translucent testaceous.

Pubescence. Hairs on under sides of hind tibiae and tarsi fuscous cop-

pery, almost black in certain lights. Female: Length 17-22 mm; forewing

length 13-14 mm. Structure. Inner orbits slightly diverging so that

clypeal width is slightly greater than upper interorbital distance.

Coloration. Clypeus entirely yellow or with only small pair of black in-

Michener: Classification of Melittidae 115

trusions on upper margin; lateral face mark continued upward medially as

slender line often joining small spot lateral to median ocellus, this

spot sometimes connected by a streak across ocellar triangle to mesal

small spot (sometimes absent) on posterior edge of vertex; lateral small

spot on posterior edge of vertex often present; stipes black; yellow of

pronotum broader than in M. binghami and including almost entire lobe;

yellow of notaulus extending back as slender streak one-third to five-

sixths length of scutum, streak sometimes enlarged at posterior end;

large area lateral to anterior end of notaulus and sometimes extending

to tegula yellow; axilla, scutellum, and metanotum entirely yellow; pair

of spots on posterior surface of propodeum usually present and large;

basal yellow of middle tibia occupying nearly half of outer surface;

bands on metasomal terga broad and unbroken, notch in band of tergum V

often absent. :

Type material. Holotype male and allotype female: 62 km west of

Omaruru, Namibia (South West Africa), March 25, 1976 (J. G. and B. L.

Rozen), in the American Museum of Natural History.

Paratypes (all from Namibia): 37 males and 9 females (one pair

pinned together and presumably taken mating), same data as holotype; 6

females and 1 male, 61 km west of Omaruru, March 22, 1979 (J. G. Rozen);

4 females, 60 km west of Omaruru, March 25, 1976 (J. G. and B. L.

Rozen); 2 males, 11 females, 58 km southwest of Omaruru, March 26, 1976

(J. G. and B.L. Rozen); | male, 58 km west of Omaruru, March 15, 1979

(J. G. and B. L. Rozen); 10 females and 5 males, 52 km west of Omaruru,

March 26 and 27, 1976 (J. G. and B. L. Rozen); 5 males and 2 females, 26

km north of Karibib, March 22, 1976, on Crotalaria podocarpa (J. G. and

B. L. Rozen); 7 males and 3 females, 8 km north of Karibib, March 22,

1976, on Crotalaria podocarpa (J. G. and B. L. Rozen); 1 female, 38 km

north of Usakos, March a 1976 (J. G. and B. L. Rozen); 3 females, 24

males, 17-19 km east of Usakos, March 18, 19, and 22, 1976 (J. G. and B.

L. Rozen); 1 male, 11. km west of Usakos, March 6, 1977 (J. G. Rozen).

Seven paratypes are in the Snow Entomological Museum, University of

Kansas, and four are in the British Museum (Natural History). The rest

are in the American Museum of Natural History.

Comments. Many attributes of the biology of this species were

described by Rozen (1977). Of special interest is its sound production.

Etymology. The specific name is the Greek noun in apposition,

gigas, meaning giant, applied because this is the largest melittid.

Meganomia andersoni (Meade-Waldo)

(Figs, 13, 37+41,:S1, 52, 59,196)

Nomia (Meganomia) andersoni Meade-Waldo, 1916, p. 457.

Meganomia andersoni: Cockerell, 1931a, p. 201; Stage, 1971, p. 312.

Diagnosis. This species and M. rossi are easily distinguished from

the others by the male antennae, the last three segments of which are

much more slender than the preceding segments; also the posteroventral

surface of the front tarsus bears a brush of long black hairs. Females

have yellow lateral mesoscutel margins. From M. rossi this species can

be distinguished by the lack of a strong concavity in the anterior mar-

gin of the hind tibia of the male and the only moderately shortened ter-

116 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

minal antennal segment of the female, this segment being conspicuously

longer than broad.

Description. Male: Length 18-19 mm; forewing length 11-11.5 mm.

Structure. 71. Produced median lobe of clypeal apex one third as wide

as clypeus, apex rounded. 2. Frontal carina below antennal bases

straight seen in profile. 3. Apex of mandible with rutellum markedly

exceeding pollex; upper margin of mandible with gentle median convexity.

4. Labrum less than twice as wide as long, surface strongly concave,

apex rounded. 5. Flagellum crenulate beneath, particularly on segments

2 to 8; segment 1 less than 2.5 times as long as broad; segments 2-4

about as broad as long or somewhat broader, segments 5 to 9 progres-

sively shorter; segments 2 to 7 of approximately equal width, 8 to 11

progressively narrowed, the last three often curled, 8 broader than long

and not greatly narrower than 7, segment 10 over 1.5 times as long as

broad, segment 11 curved and pointed (without a broad base as in M.

binghami and gigas). 6. Front coxa not or scarcely projecting behind

base of trochanter. 7. Middle coxa with weakly elevated cap in front

of base of trochanter. 8. Hind coxa unmodified, its lower surface on

level with base of trochanter. 9. Hind femur more than twice as long

as broad in side view. 10. Hind tibia viewed from inner side with

neither strong prominence nor strong emargination on anterior margin,

but with gentle convexity beyond middle. 11. Hind tibia viewed from

outer side without emargination in anterior margin, apex with strongly

projecting lobe (longer than broad, curled mesad at apex) on anterior

half. 12. Front basitarsus with longest hairs on posteroventral sur-

face, these (black) hairs grading into somewhat shorter (white) hairs on

rest of under surface; other basitarsal hairs much shorter. 13. Hind

basitarsus with outer ridge not strongly elevated at base and not

abruptly reduced or excavated; mediotarsal segments less than twice as

long as broad. 14. Basal vein of forewing interstitial with or

slightly basal to cu-v and less than five times as long as first abscis-

sa of Rs. 15. Metasomal sternum IV without tubercles on disc; V with

one short apical black comb borne on apex of a basally broadened median

part of sternum; margin lateral to comb simple; base of sternum with two

well-defined, invaginated, membranous areas. 16. Apical lobes of ster-

num VI about as long as wide, subtruncate at apices; apicolateral angle

sharp, recurved, not bifid. 17. Apex of gonocoxite without diagonal

row of bristles. 18. Penis valves united only by scarcely sclerotized

material. 19. Volsella greatly reduced, scarcely projecting beyond

margin of gonocoxite. Coloration. Black, yellow markings of head and

thorax (pale yellow on head, otherwise bright yellow) differing from

those of M. binghami as follows: vertical yellow streak between anten-

nae sometimes not joined to yellow of supraclypeal area; lateral face

mark extended up as narrow point along eye margin to level of anterior

ocellus; pedicel and sometimes base of flagellar segment 1 black; narrow

yellow line along lower half of posterior orbit commonly absent; yellow

area on upper half of genal area present but often very reduced; stipes

black; pronotal yellow absent below lobe; scutellum and metanotum black.

Legs yellow with black as follows: upper parts of coxae (more extensive

on front coxa), upper side of front trochanter, basal two thirds of

front femur except beneath, spots on anterior and under surfaces of

front tibia and area on posterior and under surfaces of mid tibia,

posterior and upper surfaces of hind femur (extending down on anterior

surface medially), extreme base and distal half of inner surface of hind

Michener: Classification of Melittidae 117

tibia, postmedian area on outer surface of hind tibia, hind tarsus ex-

cept yellow areas on anterior surfaces of all segments, claws except

brown bases. Metasoma with yellow bands as in M. binghami but all in-

terrupted medially except on tergum VI, on | merely a large spot on each

side, on subsequent terga interruption progressively narrower, on V band

sometimes almost continuous; band of tergum VI with median notch on an-

terior margin. Sternum | with large yellow patch. Sterna I! and Ill

largely yellow, the yellow more or less divided by black medially and

enclosing oval black spot on each side of each sternum. Sternum IV with

lateral yellow spot and spot on either side of midline. Flagellar seg-

ments 1-7 blackish above, brownish yellow below; 8-11 yellow, dark along

upper edge and at apex of 11. Posterior half of tegula translucent

testaceous. Wing coloration as in M. binghami. Pubescence. Coloration

as in M. binghami but with rather long, strikingly white hairs on

hypostomal area, front coxa, posterior sides of front trochanter and

femur, and anteroventral part of front basitarsus; front basitarsus with

dense brush of long black hairs along posteroventral surface; other

front tarsal segments with ventral hairs black; small segments of middle

tarsus with dusky hairs ventrally; hind tarsus with ventral hairs dusky

to black, grading to coppery at base of basitarsus. Punctation. Similar

to that of M. binghami but on thorax and metasoma slightly coarser with

ground between punctures smoother and more shining. Female: Length

14-17 mm; forewing length 10.5-12 mm. Structure. Labrum over twice as

wide as long, apex rounded. Last antennal segment longer than broad.

Inner orbits slightly diverging both above and below. Front coxal

process broadly rounded at apex, extending but little behind posterior

margin of base of trochanter. Wing venation as in male. Coloration.

Black, bright yellow markings differing from those of M. ‘binghami as

follows: clypeus with median black area extending to apical margin,

sometimes enclosing small yellow area; supraclypeal area black or with

small yellow spot; lateral face mark with truncation not or but little

sloping upward toward median ocellus; scape black; stipes black; dorsal

part and lobe of pronotum with more yellow but no yellow below lobe;

yellow stripe along notaulus extending almost to posterior margin of

scutum; yellow lateral to front of notaulus extending back along lateral

scutal margin as far as posterior end of tegula; axilla yellow; scutel-

lum with yellow lateral patches joined or nearly so by narrow posterior

marginal band; metanotum yellow at sides and with pair of small spots or

small band medially; broad, unbroken yellow band occupying dorsum of

propodeal triangle; no yellow on posterior surface of propodeum; small

to extensive yellow areas on upper posterior part of mesepisternum and

upper part to almost whole metepisternum; upper side of front tibia

mostly yellow or yellow narrowed medially by black intrusions from an-

terior and posterior margins, underside black; middle tibia with basal

half to fourth of upper side yellow, this area often continued but nar-

rowed toward apex of tibia, also spot on anterior side of apex of mid

tibia; upper side of last anterior tarsal segment yellow; band of

metasomal tergum | broken, that of || sometimes broken; sublateral emar-

ginations on posterior margins of bands usually unrecognizable. Colora-

tion otherwise as described for M. binghami. Pubescence. As described

for M. binghami but prepygidial fimbria reddish dusky; no fuscous_ hairs

around apex of basitibial plate. Punctation. As described for M.

binghami but mesoscutal punctation coarser and sparser medially, large

mesoscutal punctures as large as those of scutellum; depressed marginal

118 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

zones of metasomal terga with scattered small punctures and hairs.

, Type material. The type, a male, from the Masai Reserve in southern

Kenya is in the British Museum (Natural History). |

Distribution. Arid parts of southern Kenya. Aside from the type,

the species is known from a series of both sexes from 8 miles (13 km)

northeast of Magadi, Kenya, 2500 feet (760 m) altitude, June 16, 1967

(C. D. Michener) and one female from 20 miles (32 km) northeast of

Magadi, 3500 feet (1070 m) altitude, June 16, 1967 (C. D. Michener).

Meganomia (Meganomia) rossi Michener new species

(Figs. 42-46, 49, 50, 59, 195)

Diagnosis. This species differs from its close relative, M.

andersoni, in having a strong concavity on the anterior margin of the

hind tibia of the male and in the very short last antennal segment of

the female, this segment being not or scarcely longer than broad.

Description. Agrees with description of M. andersoni except as fol-

lows: Male: Structure. 3. Upper margin of mandible with strong post-

median protruding convexity. 4. Apex of labrum broadly truncate. 5.

Flagellar segment 8 longer than and less than two thirds as wide as

preceding segments, longer than broad, thus part of attenuate apex of

antenna; segment 10 twice as long as broad; segment 11 curved, finger-

shaped and rounded at apex in side view, pointed in ventral view, more

slender than in M. andersoni. 10. Hind tibia viewed from inner side

with large declivity toward emargination on anterior margin, without

gentle convexity. 11. Hind tibia viewed from outer side with large

emargination in apical half of anterior margin; apex with strong

projecting lobe on anterior half, distal part turned mesad and extending

farther in that direction than basal part extends distad from apex of

tibia. 12. Front basitarsus with brassy hairs on anterior margin even

longer than black hairs on posteroventral surface. 13. Hind mediotar-

sal segments about twice as long as broad. 14. Basal vein of forewing

interstitial with or slightly distal to cu-v. 15. Metasomal sternum IV

with high, sharp tubercle rising from disc mediolaterally; V with part

bearing median comb slender, parallel-sided; apical margin lateral to

comb with shoulder near comb so that comb sits in a small emargination;

base of V without invaginated membranous areas; lateral fringes longer

than in M. andersoni. 16. Lobes of sternum VI longer than wide, sub-

truncate at apices; apicolateral angle produced to long slender, deeply

bifid process. Coloration. Yellow of face as bright as that elsewhere;

yellow of supraclypeal area sometimes reduced to spot, connected to

streak between antennae which continues up to median ocellus, narrow

yellow line along lower half of posterior orbit present in both

specimens; scutum with yellow reduced to small streak near anterior end

of notaulus; axilla and extreme lateral area on metanotum yellow; front

femur black except for yellow stripe on under side connected around an-

terior side of apex of femur to large yellow spot on apex of upper side;

large black area on upper side of middle femur; black areas on outer

sides of front and middle tibiae; yellow absent on hind basitarsus and

nearly so on mediotarsal segments; metasomal sterna brown, III and IV

yellowish lateroapically. Flagellar segments 8 and 9 yellowish brown

along upper edges, 10 entirely yellow, 11 entirely brown. Pubescence.

Scutal and scutellar pubescence shorter than in M. andersoni, ochreous;

Michener: Classification of Melittidae 119

small segments of middle and hind tarsi with ventral hairs brown; hind

basitarsus more hairy than in M. andersoni with broad band of black

hairs on underside from base to apex. Female: Structure. Labrum about

three times as wide as long, apex broadly, shallowly emarginate. Apex

of antenna even more shortened than usual in the genus, last antennal

segment only about as long as broad. Coloration. Lateral face marks

fused above antennae and extending up medially as a band surrounding

median ocellus and continued between lateral ocelli to posterior margin

of vertex; yellow spot on margin of vertex mesal to summit of eye; scape

black or partly yellow beneath; yellow lateral patches on scutellum not

joined; metanotum largely yellow; large yellow spot on each side of

posterior surface of propodeum; mesepisternum black; metepisternum black

or with small yellow area along upper end; yellow of legs limited to

small spots on outer bases of fore and mid tibiae and areas on all

distitarsi; bands of metasomal terga | to II! broken medially.

Pubescence. Hairs of scutum and scutellum shorter, sparser, and more

ochreous than in M. andersoni. Hairs on basitibial plate dusky; scopal

hairs beyond basitibial plate along upper tibial margin dusky; prepygi-

dial fimbria except apical margin and pygidial fimbria dusky.

Punctation. Mesoscutum and scutellum except margins and except for an-

terior and lateral parts of mesoscutum shining with only small and

widely separated punctures; depressed marginal zones of metasomal terga

almost without punctures and hairs.

Type material. Holotype male, allotype female, and 1 male and 1

female paratype: Kitani Lodge, Tsavo National Park, Kenya, 2600 feet

(790 m) altitude, December 29, 1969 (M. E. Irwin, E. S. Ross), in the

California Academy of Sciences. :

Etymology. This species is named after Dr. E. S. Ross of the

California Academy of Sciences in recognition of his extensive worldwide

collecting.

Comment. | am indebted to Dr. G. |. Stage who had borrowed the

specimens from the California Academy of Sciences for the opportunity to

study this species.

120 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

APPENDIX Il. NEW AND LITTLE KNOWN SPECIES OF MELITTINAE

AND DASYPODINAE

The following new species are described because the names were

needed in the treatment of genera in the body of this work. Notes are

also added concerning certain previously described species.

Melitta californica Viereck

Melitta (Brachycephalapis) californica Viereck, 1909, p. 47.

Melitta wilmattae Cockerell, 1937, p. 3 (new synonymy).

Melitta maritima Cockerell, 1941, p. 344 (new synonymy).

| have compared the type of M. californica kindly lent by Dr. D. H.

Kavanaugh of the California Academy of Sciences, and the type of M.

maritima lent by Dr. J. Hall of the University of California at

Riverside with specimens from near the type locality of M. wilmattae;

they are the same species, a conclusion earlier and independently

reached by Dr. G:: |... Stage. In describing Ms: wilmattae;.Cockerell em-

phasized propodeal differences which are trivial and vary among

specimens, and the _ lack of black hairs along the sides of the face of

the female. It is true that the type of californica (as well as that of

maritima) has scattered blackish hairs down to the level of the lower

ends of the eyes; such hairs are said to be absent in the type of M.

wilmattae. In a female from Glamis such hairs extend down about to the

middle of the eye. The type of M. maritima lacked pale apical hair

bands on the metasomal terga, except for lateral remnants on the second

tergum. The specimen was, however, extremely worn, with wings badly

tattered, and the hair bands were obviously worn off. |

The range of the species is probably mostly in Mexico and is

separated by perhaps 2000 km from that of its geographically nearest

congener, M. americana Smith of the eastern United States. The known

localities are as follows: Baja California Sur: Isla de Santa

Margarita, March (type of M. californica). Baja California Norte:

Ensenada, April 13, 1941 (type of M. maritima). Arizona: Dublin, near

Yuma, March 8, 1937 (type of M. wilmatt wilmattae). California: 3 miles (5 km)

northwest of Glamis, Imperial County, March 4, 1972 (E. A. Kane, in col-

lection of R. W. Brooks).

The type of M. wilmattae and the specimens from Glamis were taken

on flowers of Sphaeralcea; it is possible that the species is restricted

to that plant as a pollen source.

The holotype of M. california is in the California Academy of

Sciences, San Francisco. The paratype, in the U. S. National Museum ac-

cording to Viereck, is not there (P. D. Hurd, personal communication).

The holotype of M. wilmattae is in the American. Museum of Natural

History, New York. The type of M. maritima was in the Timberlake

Collection at the University of California at Riverside. It was lost in

the mail after | examined it.

Michener: Classification of Melittidae 121

Redivivoides simulans Michener new species

(Figs. 80, 82-84, 97)

Diagnosis. Rather large melittine, usually with reddish-testaceous

areas on basal metasomal terga; hind tibia and basitarsus of female

slender with scopal hairs on outer surfaces short and simple; tergum VII

of male with weak but distinct pygidial plate. Differs from superfi-

cially similar Melitta capensis Friese and Rediviva rufocincta

(Cockerell) by generic characters including those listed above.

Description. Female: Length 13 mm (12.5-15 mm among paratypes);

wing length 9 mm (varying to 8.5 mm). Structure. Inner orbits parallel,

upper parts somewhat converging so that eyes are closest near upper

ends; face broad, minimum interocular distance about equal to length of

eye; malar area short but distinct, over three times as broad as long;

genal area as wide as eye seen from side; distance between posterior

ocelli slightly less than ocellocular distance; ocelloccipital distance

less than ocellar diameter; flagellar segment 1 over half as long as

scape (excluding basal bulb), 2 broader than long, others longer than

broad and subequal in length except for 10 which is over twice as long

as broad. Anterior basitarsus shaped as in Melitta, about as long as

segments 2-5 together which are not unusually robust or densely haired

as in Rediviva. Second submarginal cell considerably longer than broad,

receiving first recurrent vein in middle; third submarginal cell

receiving second recurrent vein beyond middle. Pygidial plate rounded

at apex, discal part coarsely transversely rugose. Coloration. Head and

thorax black, apical portions of mandible, tarsi and to a lesser extent

femora and tibiae, and tegula rufescent; under side of flagellum red

brown, upper side brownish black. Wings light yellowish brown, veins

and stigma brown. Metasoma black; posterior margins of terga I-IV

broadly yellowish transparent, of sterna |-I1V more narrowly reddish

translucent; following other areas pale red or reddish testaceous: ter-

gum |, tergum II except median basal area and lateral spot, broad band

across tergum III just basal to transparent marginal band, sterna |

(blackish basally) and II, basal parts of sterna ||| and 1V. (In some

paratypes these areas variously reduced, the darkest having reddish only

as follows: tergum |, dorsal sublateral areas on tergum Il, preapical

band on sternum II, and weakly developed on bases of terga II! and IV.)

Pubescence. Rather abundant, yellowish white, ochraceous on dorsum of

head and thorax, forming broad bands of somewhat appressed, short,

plumose hairs on posterior transparent margins of terga I-IV, that on |

with hairs rather sparse, the following areas with darker hairs:

clypeus, labrum and mandible with yellowish dusky hairs, some _ hairs

dusky in some lights; lateral margin of paraocular area, next to eye,

with hairs black; vertex with hairs black to dusky; margin of genal area

behind eye with long black hairs intermixed with paler ones; scutum with

intermixed dusky hairs; outer sides of tibiae and basitarsi with hairs

dusky, black in certain lights, grading into the reddish-yellow hairs of

inner sides of tibiae and tarsi; metasomal terga || to IV with short,

erect, black or dusky hairs in front of transparent marginal bands; ter-

ga V and VI with abundant black hair, small area of whitish hairs at ex-

treme sides of V;_ sterna II! and following with hairs of discs dusky

(varying to yellowish), posterior marginal hairs of II1 and IV_ pale.

Punctation. Labrum impunctate, shining, except for broad, punctate,

hairy apical band; clypeus shining, strongly punctate except for broad

122 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

apical, largely nonpunctate band; rest of face shining, punctures finer

than on clypeus and progressively finer upward; genal area still more

finely and rather closely punctate; mesoscutum finely and closely punc-

tate anteriorly, progressively more shining and more coarsely punctate

posteriorly and on scutellum, which has punctures similar in size to

those of middle of face and mesepisternum; metepisternum with lower half

impunctate, upper half with minute punctures; propodeum shining with

punctures finer anteriorly, coarser posteriorly, absent on small, smooth

undefined triangle; metasomal terga and sterna shining with well

separated punctures, finer on posterior marginal zones than elsewhere,

coarser on posterior than on anterior terga, margins proper narrowly im-

punctate. Male: Length 11.5 mm (10.5-12 mm among paratypes); wing

length 8.5 mm (8-9 mm among paratypes).* Agrees with description of

female except as follows: Structure. Inner orbits slightly converging

below. Distance between posterior ocelli equal to ocellocular distance.

Flagellar segment 1 over half as long as scape (excluding basal bulb), 2

about 1.5 times as long as broad, others about as long as 1, nearly

twice as long as broad, except 11 which is over twice as long as_ broad.

Pygidial plate weakly defined, obtusely angulate at apex. Coloration.

Tibiae and tarsi often paler than in female, hind tibia and tarsus often

red brown. Posterior margins of metasomal terga and sterna I-V yel-

lowish transparent, more broadly so on terga. Tergum || with black con-

tinuous across base of tergum, or in some paratypes this tergum almost

entirely yellowish red, in others black limited to median basal area, in

others with black lateral spot also. Sternum II! and sometimes most of

[V pale red. Pubescence. Whiter (i.e., less yellowish) than in female,

hairs of apical tergal bands longer than in female, and on transparent

margins of terga |-V, that on | with hairs rather sparse; hairs of

clypeus white; scutellum with intermixed dusky hairs like scutum; outer

sides of tibiae and basitarsi with hairs whitish; terga II-V with black

hairs in front of marginal bands, VI and VII with hairs dusky; sterna

with hairs yellowish white except for VI which has dusky hairs, sterna

ll-V with apical paler fringes of longer hairs. Punctation. Areas less

distinctive than in female, so that punctation of clypeus is not much

coarser than that of supraclypeal area, that of genal area and front

part of mesoscutum is not so fine as in female, that of posterior part

of mesoscutum and scutellum finer than in female, not much coarser than

that of anterior part of mesoscutum.

Type material. Holotype female, allotype male, and 14 female and 15

male paratypes, South Africa: Clanwilliam, Cape Province, 6 September

1966 (M. and C. D. Michener); 1 female and 3 male paratypes, 20 miles

(32 km) west of Clanwilliam, 16 September 1966 (C. D. Michener). The

holotype and allotype are in the Snow Entomological Museum, University

of Kansas. Paratypes are in that collection, the American Museum of

Natural History (New York), the National Museum of Natural History

(Washington) and the British Museum (Natural History).

Etymology. The specific name refers to the similarity of this

species to other somewhat distantly related Melittinae, as indicated in

the diagnosis.

Michener: Classification of Melittidae 123

Rediviva peringueyi (Friese) —

(Figs. 85-87, 95)

Andrena (Rediviva) peringueyi Friese, 1911, p. 671.

This is the type species of the genus. The female has elongate

front legs, but not nearly so extreme as in R. longimanus, the front

tibia being only about as long as the eye.

One of Friese's original female specimens from Stellenbosch and a

male from Cape Town are in the South African Museum, Cape Town. The

female type from Paarl is in Berlin. An additional record is a single

female from Clanwilliam, 8 September 1966 (M. and C. D. Michener). All

these localities are in western Cape Province, South Africa.

Rediviva longimanus Michener new species

(Figs. 96, 97)

Diagnosis. Black with pubescence mostly black except on legs;

front legs of female extraordinarily long and slender (Fig. 96). The

closest relative is probably R. peringueyi, from which R. longimanus

differs in both features mentioned above.

Description. Female: Length 14 mm; wing length 10.5 mm.

Structure. Inner orbits parallel, upper parts converging above so that

eyes are closest near upper ends; face moderately broad, minimum in-

terocular distance less than length of eye; malar area short, about five

times as broad as long; face not especially protuberant, apex of clypeus

in side view in front of anterior eye margin by about eye width; genal

area as wide as eye seen from side; distance between posterior ocelli

less than ocellocular distance; ocelloccipital distance about equal to

ocellar diameter; flagellar segment 1 fully half as long as scape (ex-

cluding basal bulb), 2 broader than long, 3 as broad as long, others

longer than broad. Anterior legs extremely attenuate (Fig. 96), tibia

about 1.5 times as long as eye; small segments of tarsus, except for

under sides, densely covered with short, dense, plumose hairs. Second

submarginal cell considerably longer than broad, receiving first recur-

rent vein in middle; third submarginal cell receiving second recurrent

vein beyond middle; basal vein over twice as long as first abscissa of

Rs, interstitial with cu-v; wings badly worn but jugal lobe apparently

slightly less than half as long as vannal. Propodeal triangle weakly

defined, about one third as wide as distance between transmetanotal su-

tures. Pygidial plate subtruncate, smooth but dull. Coloration. Black,

tegula and small segments of mid and hind tarsi reddish; under sides of

third and following flagellar segments brown. Wings strongly dusky,

veins and stigma black. Pubescence. Black except as indicated below,

sparse and short on dorsum of metasoma which completely lacks hair

bands. Hair of labrum, lower margin of mandible, under side of thorax,

coxae, trochanters, apical fringe of tergum V, and metasomal sterna |-V

dusky coppery, apical fringes on sterna II-V long and conspicuous; hairs

of tibiae, tarsi, and tergum VI golden. Punctation. Labrum impunctate

except for apical punctate hairy zone; clypeus with moderate sized punc-

tures separated by about a puncture width except for lower third which

is largely impunctate, surface between punctures shining but minutely

roughened; supraclypeal and lower paraocular areas similar to clypeus

124 Contrib. Amer. Ent. Inst.) vol. 18, no. 3, 1981

but punctures slightly smaller; frons with punctures of same size but

denser, surface duller because of minute roughening; upper part of head

and genal areas with punctures small, ground between them rather dull in.

most areas; shiny and nearly impunctate areas below antennal base and

lateral to posterior ocellus; scutum and scutellum with punctures fine,

finest and ground dull on anterior third of scutum; mesepisternum with

punctures shallow, well separated, ground dull; metepisternum shining,

impunctate; propodeum with punctures minute, widely separated, coarser

and closer on anterior lateral portions, ground somewhat shining,

triangle shining, minutely transversely wrinkled; metasomal terga

shining, broad apical zones minutely roughened but ground in front of

apical bands smooth or nearly se, punctures minute and widely separated

dorsally, coarser and closer laterally and on IV and especially V.

Sterna more coarsely punctate, ground between punctures minutely

roughened, especially on posterior segments.

Type material. Holotype female: South Africa: 5 miles (8 km)

north of Nieuwoudtville, Cape Province, September, 1961, in the South

African Museum, Cape Town.

Etymology. From the Latin longus, long, plus manus, hand, with

reference to the long front tarsi. It is a noun in apposition and hence

not in agreement with the feminine generic name.

Comments. This species is described here from the single known

specimen because it represents the extreme in elongation of the front

legs of the female. Probably this relates to a ae habits from

some specific kind of flower.

A single female specimen ile data in the South African Museum

is morphologically like R. longimanus but has paler wings and

pubescence, in general like that of R. peringueyi but without the weak

apical hair bands on the terga found in that species. It may represent

a color morph of longimanus or a related species.

Rediviva colorata Michener new species

(Figs. 88-91, 94, 97)

Diagnosis. Black with red-brown areas on basal metasomal terga;

metasomal terga I-II| of male, !-1V of female, with apical bands of yel-

low hairs; head longer than broad, clypeus strongly produced and

protuberant, malar area over half (female) to about three fourths (male)

as long as broad; second submarginal cell about as long as _ broad.

Superficially similar to R. rufocincta (Cockerell) but differing in long

face, long tongue, slender front legs of the female, dull facial integu-

ment, and many other characters.

‘Description. Female: Length 12.5 mm; wing length 10 mm.

Structure. Inner orbits slightly converging below except for upper parts

which converge above; distance between eyes less than length of eye;

malar area over half as long as wide; face longer than wide, clypeus

protuberant so that in side view anterior end is in front of anterior

eye margin by distance much greater than eye width; glossa slender, more

than half as long as the head and over half as long as prementum; galeal

blade blunt, about 3.5 times as long as broad; genal area as wide as eye

seen from side; distance between posterior ocelli slightly less than

ocellocular distance; ocelloccipital distance greater than ocellar

diameter; flagellar segment 1 less than half as long as scape (excluding

Michener: Classification of Melittidae te

basal bulb), 2 and 3 equal in length, each broader than long, 4 as long

as broad, 5-9 the same or slightly longer than broad, depending on an-

tennal curviture, 10 nearly twice as long as broad. Anterior leg

slender, tibia slightly longer than eye, tarsus not especially elongate

(Fig. 94), small segments densely covered with plumose hairs except on

underside. Second submarginal cell about as long as wide, receiving

first recurrent vein at apical fourth; third submarginal cell receiving

second recurrent vein near beginning of apical third; basal vein much

over twice as long as first abscissa of Rs, fractionally basal to (or

interstitial with) cu-v; jugal lobe little over one-third length of van-

nal lobe. Propodeal triangle minute, less than one sixth as wide as

distance between transmetanotal sutures. Pygidial plate rounded at

apex, disc closely punctate. Coloration. Black, tegula, dorsal surface

of metasomal tergum |, lateral portions of I! and II|, and base of ster-

num || yellowish brown; anterior surface of tergum | dark brown; small

segments of tarsi and under side of flagellum brown; posterior margins

of metasomal terga | to 1V and sterna | to V broadly yellowish tran-

sparent, with some yellow brown at bases of transparent zones; mandible

with distal half rufescent. Wings light brown, veins and stigma dark

brown, vein R of forewing black. Pubescence. Ochraceous, paler on

under side than elsewhere, apical bands of rather long recumbent yellow

hairs on metasomal terga | to 1V, apical bands of more erect, very long,

pallid hairs on sterna || to V; hairs of labrum, mandible, tibiae and

tarsi golden; black hairs intermixed with pale ones on face, vertex,

along posterior orbit, on scutum and scutellum; front tibia and tarsus

with hairs dusky gold on anterior surfaces; mid tibia and basitarsus

with hairs dusky gold on posterior surfaces; apical part of tibial scopa

with long simple hairs of posterior portion dusky gold to black,

basitarsus with similar dark simple hairs on posterior and distal parts;

parted hairs of basitarsal apex dusky; metasomal tergum |V with a few

intermixed black hairs basally; pubescence of terga V and V/I and sternum

VI black. Punctation. Labrum shining impunctate except for apical,

punctate, hairy zone; clypeus coarsely punctate, more coarsely and ir-

regularly so apically, ground between punctures mostly less than a punc-

ture width and rather dull, minutely roughened; rest of face progres-

sively more finely punctate upward to vertex, ground rather dull with

fine roughening; malar area largely shining, impunctate; genal area

shining with scattered punctures; scutum and scutellum dull with punc-

tures mostly coarser than those of vertex and separated by a puncture

width or more, finer and closer on small posterior discal area of

scutum; mesepisternum- shining with coarse, shallow punctures;

metepisternum shining, lower half impunctate, upper half with minute

punctures; propodeum rather dull with irregular and rather shallow punc-

tures except minute, smooth, shining propodeal triangle; metasomal terga

smooth, shining, punctures minute and widely separated, little coarser

on tergum V; apical margins of terga impunctate; sterna minutely

roughened with well separated punctures, coarser than those of terga ex-

cept on sternum VI. Male: Length 12 mm; wing length 10 mm. Similar to

description of female except as follows: Structure. Inner orbits

slightly converging below except for upper extremities which converge

above; malar area nearly three fourths as long as wide; genal area

slightly narrower than eye seen from side; flagellar segment 2 slightly

longer than wide, 3-10 subequal in length, about 1.3-1.4 times as long

as wide, 11 about twice as long as wide. Anterior leg unmodified. Py-

126 Contrib: Amer. Ent.’ inst.) vol-8, no? 3, 1981

gidial plate absent. Tergum VIII without membranous lobe lateral to

bifurcate apex. (Other known Rediviva all have such a_ lobe.)

Coloration. Flagellum entirely blackish; metasomal terga IV-VI with

posterior margins translucent brownish black, only I-III yellowish trans-

parent. Pubescence. Apical bands of rather long yellow hair only on

terga I-II| and extreme sides of |1V; hair of sterna shorter than in

female and not forming conspicuous bands; tibiae and tarsi with yellow

hairs, dark only on apical three fifths of hind basitarsus; metasomal

tergum II! with a few intermixed dusky hairs on disk; terga IV-VII and

sternum VI with hairs black, yellow only on extreme side of IV.

Punctation. Clypeus somewhat more finely punctate than in female, about

like supraclypeal area; scutum and scutellum with punctures small,

widely separated.

Type material. Holotype female, allotype male, and 1 male

paratype: South Africa: Karkloof, Natal, February 15, 1942 (Markley).

The holotype and allotype are placed in the British Museum (Natural

History). The paratype is in the Snow Entomological Museum, University

of Kansas.

Etymology: From coloratus, colored or variegated, from the variety

of integumental and hair colors of this species.

Haplomelitta (Prosamba) griseonigra Michener new species

(Figs. 107,°108;, 116-121 , 147; °148)

Diagnosis. The only known species of Prosamba, therefore recog-

nized by diagnostic characters of that subgenus (or genus). Similar

robust form, dark coloration and lack of metasomal fasciae to H.

(Atrosamba) atra from which it differs in much smaller size as well as

the subgeneric characters. Similar in size and body form to H.

(Metasamba) fasciata from which it differs by darker pubescence and lack

of metasomal fasciae, as well as subgeneric characters.

Description. Female: Length 8 mm; forewing length 5.5 mm.

Structure. Inner orbits scarcely converging below, minimum distance

between eyes slightly greater than length of eye; malar area short, as

long as median width of flagellar segment 1 medially, but anterior man-

dibular articulation well separated from eye margin; genal area wider

than eye seen from side; distance between posterior ocelli scarcely less

than ocellocular distance; ocelloccipital distance slightly greater than

ocellar diameter; flagellar segment 1 distinctly less than half as long

as scape (excluding basal bulb), 2-10 progressively broader, 2-9 broader

than long or sometimes 2 seeming about as long as broad, 10 about 1.5

times as long as broad. Coloration. Black; distal half of mandible red,

legs and tegula reddish black; underside of flagellar segments 3-10

light brown, rest of antenna dark brown to reddish black; posterior

depressed margins of metasomal terga |-IV translucent reddish black; py-

gidial plate partly red in some paratypes. Tibial spurs translucent

whitish brown. Wings brown, veins and stigma dark brown. Pubescence.

Sparse, brownish, dense white patch obscuring surface lateral to anten-

nal base and more white hairs immediately above antennal bases; erect

white hairs at sides of terga I-IV, with some shorter white hairs

arising dorsad on basal parts of terga; dull whitish or pale hairs on

posterior part of vertex, anterior margin of scutum, lateral extremities

of scutellum and metanotum, lateral surface of mesepisternum; black or

Michener: Classification of Melittidae 127

fuscous hairs on mandible, lower clypeal margin, paraocular area along

eye margin, genal area, anterior surface of mesepisternum, lateral sur-

face of propodeum, most of legs (scopa entirely blackish), and metasoma

(prepygidial and pygidial fimbriae dense and black); black or fuscous

hairs intermixed with paler ones on scape, ocellar region, and dorsal

parts of terga II-IV; under sides of tarsi with orange hair; hair bands

of terga I-IV entirely absent. Punctation. Labrum shining, impunctate,

except punctured beyond transverse ridge; clypeus shining, impunctate

except for scattered coarse punctures laterally and near upper margin;

supraclypeal area with longitudinal median impunctate zone; rest of face

strongly and rather densely punctate, a little more finely so upward,

intervals between punctures much less than puncture diameters, shining;

area lateral to ocelli, extending to upper genal area beyond eye, im-

punctate; scutum more coarsely punctate than head, ground smooth and

shining, punctures mostly separated by less than a puncture width an-

teriorly, but posterior median area with interspaces mostly of several

puncture widths; mesepisterna, lateral and posterior surfaces. of

propodeum, and scutellum similar but more finely punctate; propodeal

triangle shining but minutely roughened; metasoma with rather coarse

punctures on shining ground, basal concavity of tergum | broadly impunc-.

tate and shining, terga I-IV each with largely impunctate transverse

ridge basal to apical depressed zones, the latter broad, smooth, impunc-

tate. Male: Length 8 mm; forewing length 5.5 mm. Agrees with descrip-

tion of female except as follows: Structure. Inner orbits strongly con-

verging below, minimum distance between eyes less than eye length; genal

area narrower than eye seen from side; flagellar segment 1 about half as

long as scape, 2-10 of nearly uniform width, 2 broader than long, 3-5

about as broad as long, the remaining segments progressively longer.

Coloration. Mandible with preapical red band; undersides of front femur

and tibia and base of mid femur, small segments of all tarsi, flagellar

segments 3-10 and tegula red. Wings paler than in female, membrane al-

most clear. Pubescence. Somewhat longer than in female, white, somewhat

yellowish on legs, yellowish orange on undersides of tarsi; face densely

covered with long, appressed, white hair hiding surface; metasomal terga

with rather long, erect (but curved apicad near tips) white hair arising

basal to depressed apical zones, these hairs well separated so that sur-

face is easily visible. Punctation. Clypeus closely and rather finely

punctate, like rest of face; impunctate area lateral to ocelli less ex-

tensive than in female.

Type material. Holotype female and 4 female paratypes: South

Africa: Kamieskroon, Namaqualand, Cape Province, September, 1930 (South

African Museum Staff). They are in the South African Museum, Cape Town,

except 1 paratype in the Snow Entomological Museum, University of

Kansas.

The single male specimen may possibly be a different species since

it was taken ata different place. It is therefore not designated as

the allotype. It is from Bowesdorp, Namaqualand, September, 1941 (South

African Museum Staff).

Etymology. The specific name refers to the grayish black general

appearance.

128 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

Haplomelitta (Atrosamba) atra Michener new species

(Figs. 109, 110, 122-126, 147, 148)

Diagnosis. The only known species of Atrosamba, therefore recog-

nized by diagnostic characters of that subgenus (or genus). Similar in

robust form and dark coloration to H. (Prosamba) griseonigra from which

it differs in larger size, more extensive black hairs, as well as sub-

generic characters. The body form is also similar to that of H.

(Metasamba) fasciata from which it differs by large size, dark

pubescence, and lack of metasomal fasciae, as well as subgeneric

characters.

Description. Female: Length 11-11.5 mm; forewing length 9 mm.

Structure. Inner orbits parallel, distance between eyes greater than

length of eye; malar area short, as long as median width of flagellar

segment 1 posteriorly, but anterior mandibular articulation far from eye

margin; genal area slightly wider than eye seen from side; distance

between posterior ocelli less than ocellocular distance; ocelloccipital

distance about twice ocellar diameter; flagellar segment 1 distinctly

less than half as long as scape (excluding basal bulb), 3-10 of about

the same width (flagellum being more slender than in H. griseonigra), 2

broader than long, 3-9 about as long as broad (often appearing either

shorter or longer depending on flagellar curvature), 10 about 1.5 times

as long as broad. Coloration. Black, mandible with dark red median

band, legs extensively dark red, upper surfaces of femora and tibiae

reddish black; flagellar segments 3-10 rather dark brown beneath.

Tibial spurs dark red brown. Wings brown, veins and stigma dark brown.

Pubescence. Not dense, not obscuring surface except for black pygidial

and prepygidial fimbriae, short and erect on dorsal surfaces of terga

ll-IV but elsewhere moderately long, black or brownish black except for

yellowish-white hairs on vertex and occiput behind ocelli, posterior

lobe and upper (posterior) margin of pronotum, anterior margin of scutum

(mixed with black), lateral extremities of scutellum and metanotum,

metasomal tergum |, lateral extremities of tergum ||; undersides of tar-

si, of hind tibia, and of certain areas on other tibiae with orange

hair; hair bands of terga I-IV entirely absent. Punctation. Labrum

shining and impunctate on transverse ridge and basal to it; densely

punctate distal to ridge; clypeus rather sparsely punctate laterally and

near margins but median third except near margins shining, impunctate or

with sparse punctures; rest of face strongly and rather densely punc-

tate, intervals between punctures mostly less than half a puncture width

and shining; area immediately lateral to lateral ocellus and zone on

genal area behind upper end of eye impunctate, but vertex between these

areas with punctures, irregularly placed but separated by about a punc-

ture width of shining ground; punctation of scutum and scutellum about

like that of vertex, of mesepisternum denser, of metepisternum and

propodeum much finer, rather dense on side. of . propodeum; propodeal

triangle polished; metasomal terga with fine, well separated punctures

dorsally, denser and coarser punctures laterally, ground shining; basal

concavity of tergum | broadly impunctate; posterior marginal zones of

terga |-1V depressed, shining, impunctate, not preceded by shining

ridges as in H. griseonigra. Male: Length 10 mm; wing length 7.5 mm.

Agrees with description of female except as follows: Structure: Inner

orbits converging below; minimum distance between eyes subequal to

length of eye; genal area about as wide as eye seen from side; ocelloc-

Michener: Classification of Melittidae 129

cipital distance about 1.5 ocellar diameters; flagellar segments 3-11

longer than broad. Coloration. Femora and front tibia largely black.

Tibial spurs red. Pubescence. Moderately long, including that of

metasomal terga; that of clypeus long, appressed, white, obscuring

clypeal surface. Pubescence largely yellowish white; black to fuscous

on mandible, paraocular area especially laterally, genal area, posterior

part of mesepisternum, metepisternum, propodeum, tergum V-VII, and

metasomal sterna; black mixed with white on scape, vertex, scutum,

scutellum, legs, and tergum 1V. Punctation. Clypeus finely and rather

densely punctate; rest of face more coarsely punctate, as in female;

terga I-VI with shining depressed marginal zones.

Type material. Holotype female, allotype male, and 2 female

paratypes: South Africa: Springbok, Namaqualand, Cape Province, 7

September 1966 (C. D. Michener). Two female paratypes: 5 miles (8 km)

north of Nieuwoudtville, Cape Province, September, 1961 (South African

Museum). One female paratype: Kamieskroon, Namaqualand, Cape Province,

September, 1930 (South African Museum Staff). The holotype and allotype

are in the Snow Entomological Museum; paratypes are in that institution

and the South African Museum, Cape Town.

One female of the same or a closely related species is from

Worcester, Cape Province, 30 September 1966 (C. D. Michener). It is

larger than typical H. atra, the clypeus punctate almost throughout,

tergum | with much intermixed black hair, and tergum || without pale

hair. An apparently typical male of H. atra is from Ladismith, Cape

Province, September 9, 1948 (C. Jacot-Guillarmod).

Etymology. The specific name refers to the largely black color.

Haplomelitta (Haplosamba) tridentata Michener new species

(Fig. 147)

Diagnosis. Sparsely hairy, shiny black with the metasoma largely

red, superficially resembling H. (Haplomelitta) ogilviei (Cockerell),

from which it differs in the subgeneric characters.

Description. Female: Length 11.5 mm; forewing length 7.5 mm.

structure. Inner orbits diverging below, minimum distance between eyes

equal to length of eye; malar area posteriorly as wide as apex of

flagellar segment 1, anterior mandibular articulation far from eye mar-

gin; genal area wider than eye seen from side; distance between

posterior ocelli much less than ocellocular distance; ocelloccipital

distance much greater than ocellar diameter; flagellar segment 1 about

one third as long as scape (excluding basal bulb), 2-3 distinctly

broader than long, 4-9 slightly so to as long as broad, 10 slightly more

than 1.5 times as long as broad, flagellum slightly tapering so that

segment 10 is a little broader than 2. Scutellum biconvex. Coloration.

Black; mandible, clypeus, genal area, and adjacent parts of head dark

red to reddish black; coxae and trochanters largely dark red; rest of

legs and scape dark red; tibial spurs red; antennal segments 2-12 brown;

tegula and wing base orange; metasomal segments |-I!| and base of IV and

base of pygidial plate red; wings dark brown, veins and stigma dark

brown. Pubescence. Short, sparse, nowhere obscuring surface, fuscous to

blackish, yellowish intermixed on frons, vertex (hairs of vertex and oc-

ciput behind ocelli all yellowish), posterior lobe of pronotum, scutum

and scutellum; light fuscous hairs appearing yellowish in some lights on

130 Contrib. Amer. Ent. Inst., vol. 18, no. 3, 1981

face, legs, and metasomal venter; scopa reddish fuscous; hairs of under

sides of tarsi orange. Punctation. Labrum shining impunctate except

distal marginal zone; clypeus shining, impunctate except for punctate

lateral extremities and a few punctures near upper margin; upper part of

supraclypeal area and lower part of frons closely punctate but rest of

head with punctures well separated by shining ground, lower supraclypeal

area, area between ocelli and eye, and genal area with punctures

separated by several diameters, elsewhere closer, coarsest on lower

paraocular area; central part of scutum, from front to rear, with punc-

tures separated by several puncture widths, on scutellum and on scutum

laterally punctures coarser and closer, on sides of thorax still coarser

and closer; ground everywhere smooth and shining; propodeal triangle

smooth and shining; metasomal terga with punctures laterally and

sublaterally coarse and irregular, separated by broad intervals of

shining ground, medially punctures finer and sparser so that mid-dorsum

of terga I-V is almost impunctate; terga I-IV each with depressed

posterior marginal impunctate zone, sharply set off and somewhat nar-

rowed sublaterally by largely impunctate convexity anterior to each

depressed band.

Type material. Holotype female: South Africa: Wallekraal,

Namaqualand, Cape Province, October, 1950 (South African Museum

Expedition). This locality is at 30 25'S, 17 27'E, according to J. G. |

Rozen, Jr. The type is in the South African Museum, Cape Town.

Etymology. The specific name refers to the tridentate mandibles.

Haplomelitta (Metasamba) fasciata Michener new species

(Figs. 133-138, 144, 147, 148)

Diagnosis. The only known species of Metasamba, therefore recog-

nized by the diagnostic characters of that subgenus (or genus). It is

similar in its robust form to H. (Atrosamba) atra and H. (Prosamba)

griseonigra but differs from both in having nearly all of the hair

white, and in having white, pubescent fasciae on the metasomal terga.

Description. Female: Length 8.5 to nearly 9mm; forewing length

nearly 6mm. Structure. Clypeus and supraclypeal area with longitudinal

median, elevated ridge; inner orbits converging below, minimum distance

between eyes about equal to eye length; malar area medially shorter than

basal width of flagellar segment 1, but anterior mandibular articulation

well separated from eye margin; genal area as wide as eye seen from

side; distance between posterior ocelli less than ocellocular distance;

ocelloccipital distance little greater than ocellar diameter; flagellar

segment 1 little over one fourth as long as scape (excluding basal

bulb), segments 2-9 broader than long, 10 about 1.75 times as long as

broad; flagellum of uniform width beyond about segment 3. Coloration.

Black; mandible with preapical red band; tibiae (except outer sides of

fore and middle tibiae and sometimes basitarsi), tibial spurs, tarsi,

undersides of flagellar segments 3 to base of 9, tegula and wing base

light red; upper side of flagellum and whole of segment 10 dark brown;

posterior depressed margins of metasomal terga I-IV transparent

testaceous, black showing through from succeeding terga. Wings light

brown, stigma and most veins brown. Pubescence. Moderately abundant,

long, white, absent on lower two thirds of clypeus except for hairs and

bristles along lower clypeal margin (in worn individuals absent below

Michener: Classification of Melittidae 131

level of upper margin of clypeus); dense and nearly obscuring surface on

much of face above level of upper margin of clypeus; metasomal terga

I-IV with preapical bands of plumose hair arising just basal to apical

depressed marginal zones which are partly obscured by hair, hair bands

broken or weak medially, especially on anterior terga; hairs of tibiae,

tarsi, and metasomal venter yellowish, those of undersides of tarsi

orange; prepygidial and pygidial fimbriae brown or blackish dorsally,

white only laterally. Punctation. Labrum impunctate and shining (dull

in worn individuals) from transverse ridge to base, closely punctate

distally. Lower two thirds of clypeus with coarse punctures (absent on

median ridge) separated by about a puncture width of shining ground

(dull in worn individuals); upper and lateral parts of clypeus with

somewhat finer and closer punctures; supraclypeal and lower paraocular

areas with fine, close punctures, less close higher on face and on genal

area, on vertex lateral to ocelli smooth with only scattered, coarser

punctures; scutum and scutellum with moderately coarse punctures (about

size of those of upper part of clypeus), separated by about a puncture

width of shiny ground on scutum except margins, closer on scutellum and

margins of scutum; mesepisternum more coarsely and closely punctate;

rest of side of thorax and lower part of propodeum with much finer punc-

tures; propodeal triangle polished and shining but minutely, irregularly

roughened. Metasomal terga I-IV finely punctate, shining, with coarser

punctures laterally and just anterior to bases of fasciae, fine and

close at fascial bases, posterior marginal zones smooth, shining, im-

punctate; tergum V coarsely punctate except finely so in area that sup-

ports prepygidial fimbria. Male: Length 9 mm; forewing length 6 mm.

Agrees with description of female except as follows: Structure. Clypeus

and supraclypeal area without longitudinal median ridge; minimum

distance between eyes less than length of eye; genal area wider than eye

seen from side; distance between posterior ocelli equal to ocellocular

distance; ocelloccipital distance about twice ocellar diamter; flagellar

segment 1 short and robust but because of short scape, about one third

as long as scape; flagellar segments all longer than broad, 1 and 2 only

slightly so, 1-4 convex on upper side so that base of flagellum is

crenulate, segments 1-9 of uniform thickness but 10-11 slightly thicker.

Mid and hind leg modifications as noted in description of subgenus.

Coloration. Distal part of mandible red, apical parts of femora red.

Flagellar segments 3-9 light red, upper surfaces of 3 and 4 infuscated;

segments 1-2 partly light red, segments 10-11 black; metasomal terga

I-VI with transparent, depressed margins. Wings nearly clear.

Pubescence. Long white hair obscuring surface of entire face; white

metasomal fasciae on terga I-VI, not weak or broken medially.

Punctation. Clypeus finely and closely punctate. Terga V and VI punc-

tured about like more anterior ones.

Type material. Holotype female and 1 female paratype: Namibia (=

South West Africa): 61 km west of Omaruru, March 21, 1979 (J. G. Rozen).

Allotype male and 2 female paratypes, same ‘data but March 62, 2°°972%.. 1

male paratype, same data but March 15, 1979. Three female paratypes:

32 km west of Omaruru, March 15, 1979, on Indigofera (J. G. and B. L.

Rozen). The holotype and allotype are in the American Museum of Natural

History; paratypes are in that institution except for two in the Snow

Entomological Museum, University of Kansas.

Etymology. The specific name refers to the banded (fasciate) condi-

tion of the metasoma.

132 Contribs. Amer..-Ent.. inst. vol...318,.ne; 3,.1961

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tive study. Univ. Kansas Sci. Bull., 51:631-667.

Wu, Y.-J. 1978. A study of Chinese Melittidae with descriptions of new

species. Acta Ent. Sinica, 21:419-428.

Yasumatsu, K. and Y. Hirashima. 1956. Discoveries of the genus

aero Klug and Melitta Kirby in Japan. Kontyu, 24:247-255, pl.

26.

Contributions

of the

American Entomological Institute

Volume 18, Number 4, 1981

“v"

MEDICAL ENTOMOLOGY STUDIES - XV.

A REVISION OF THE SUBGENUS PARAEDES OF

THE GENUS AEDES

(DIPTERA: CULICIDAE)

John F. Reinert

CONTENTS

Pi AG Ek a We ee i 1

INTRODUCTION 6.3503 6 8 wee ew a a a 1

GENUS AEDES MEIGEN, SUBGENUS PARAEDES EDWARDS ....... 3

KEYS TO SPECIES OF AZDES PARAL OLS) . i. eae kee ees 8

PAE: ok oe a eo are ee ae ee ee oe ee 9

PUPA 6 yobs: ee8 eW e ee ee 10

FOURTH: SIAGE LARVAR ss Cai 6 eee Ge 10

SPECIES TREATMENT. 3 a6 sues eae Se es ee a 10

borvrvraud. AB awarda) ak einer LG Ge ae ee, 10

bonmege Mattingly ci «Vos Cee ee a Bae ie ae 13

chrysosculo (Theobald) s sigs oo ee Vee etd

colless) Matinely oo esce we OR We ee ee 22

menont NMatlINGy: win Ale eee «AOR EW eels 28

osienlatio (Leieestes) Soa Gi ee Ow OE OU Sw eS 31

pooe: (LUlOW) as via Oe ee ee 38

thavlandensts: ROWG?Y : g5 6 ek We OR Oe ee ae eee ee 40

ACKNOWLEDGMENTS. 66536 eee ee ea a Pee eee ee ee 45

BITBRATURE CIIED «53544 ves bata a rgimes Bi ag ae Haas nen 46

LIST OF FIGURES i085 2°05. 6 Seer ee Ee a ea ee) D2

FIGURE ABBREVIATIONS: yor wo Bee ee ee eee SG D3

FPIGUREB 6: 6 sacs (BE Ee POS OL A are PUA es Pea 54

APPENDICES oe aie He eh A ee i ss ea 83

TABLES 1-5 Records of the branching of the setae on the pupae

Of Aedes (Povaedes) Species iii ww wie GW es 85

TABLE 1; Menvede turers Be es ea eee 85

TABLE 24: CRYVOQS CH Ron h Cie OR ee Oe eS 86

TABI. COMES ie GE EAs rR Ww BO 87

TABLE: 4. OSteniai7oe cia eevee Ce LW i A ee 88

DAB Ba a ee ei ee iia a a ew es 89

TABLE 6. Current taxonomic status of the subgenus Parvaedes.. 90

INDEX e ° e e e ee e« e e® ee 6 @

ieee

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ghee

ir sia 7

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ae ae Pa

ie)

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ae a

Wae aie

haRAR Ma POT Ay,

ahs AYR

MEDICAL ENTOMOLOGY STUDIES - XV.

A REVISION OF THE SUBGENUS PARAEDES OF THE GENUS AEDES

(DIPTERA: CULICIDAE)!

John F, Reinert2

ABSTRACT

The subgenus Pavaedes Edwards of Aedes Meigen is revised. Pavaedes

includes 8 currently recognized species that are restricted to the Oriental

region, All known stages of barvaudi (Edwards), bonneae Mattingly, chryso-

scuta (Theobald), collessi Mattingly, menoni Mattingly, ostentatio (Leicester),

pagei (Ludlow) and thailandensis Reinert are described and illustrated, and are

included in identification keys. The range and mode of setal branching of the

pupae and larvae are given. Geographical distribution, bionomics, type-data

and taxonomic discussions are presented for each species. Aedes chrysoscuta

is resurrected from synonymy with ostentatio. Aedes aurotaeniatus Edwards

is excluded from the subgenus Paraedes.

INTRODUCTION

Edwards (in Barraud 1934: 446) described the genus Pavaedes from speci-

mens of 2 species, barrvaudi (Edwards) and argyrurus (Edwards). His associa-

tion of these 2 species and the basis of the new genus was based primarily on

the absence of setae on the upper calypter of the wing. The latter species was

removed from Pavaedes by Thurman (1954: 84) and placed in Udaya, a then new

subgenus of Aedes Meigen. Mattingly (1958a: 4, 19)elevated Udaya to generic

status and reduced Paraedes to subgeneric rank within Aedes. Species current-

ly recognized in the subgenus Pavaedes have previously been assigned to the

following genera and/or subgenera: Aedes Meigen, Aedimorphus Theobald,

Aioretomyia Leicester, Danielsia Theobald, Ochlerotatus Lynch Arribalzaga,

and Pseudohowardina Theobald,

Pavaedes includes 8 distinct species (Table 6) and is distributed throughout the

Oriental zoogeographical region. The present revision redefines the subgenus

and presents characters for distinguishing it from the other subgenera of Aedes.

1 This work was initiated when the author was assigned at the Walter Reed Army

Institute of Research and was supported in part by Research Contract DAMD-

17-74-C -4086 from the U. S. Army Medical Research and Development Com-

mand, Office of the Surgeon General, Ft. Detrick, Frederick, MD.

oT initial Colonel, Medical Service Corps, U. S. Army, Research Liaison

Officer, Armed Forces Pest Management Board, with mailing address: P. O.

Box 14565, USDA, Gainesville, FL 32604; and Research Consultant and Coop-

erator, Medical Entomology Project, Smithsonian Institution, Washington, DC.

Pe Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

Identification keys are given for separating the adult females, male genitalia,

pupae and larvae for the known stages of the included species, All known

stages of each species are described and illustrated. The males, male and

female genitalia, pupae and larvae of chrysoscuia and ostentatio are described

and illustrated for the first time.

Aedes aurotaeniatus Edwards was transferred from the subgenus Stegomyia

Theobald to the subgenus Pavaedes by Huang (1970: 281). This transfer, I

believe, was based on superficial characters that do not reflect phylogenetic

affinities. Huang (page 287) stated ‘‘. . . the male terminalia of aurotaenia-

tus appears to have some rather basic genitalic characters in common with

Pavaedes, suggesting a close affinity.’’ I have examined the 2 male specimens

(456-100, -105) and the other material reported by Huang and found the male

genitalia to be significantly different from all known species of Paraedes in the

phallosome, basal mesal lobe, proctiger, gonocoxite, gonostylus, tergum IX

and sternum IX, Furthermore, many features of the adult and pupae of these

specimens are not equivalent to the species included here in the subgenus

Paraedes, A comparison of the descriptionby Huang (1970) of aurotaeniatus

and that of the subgenus: Pavaedes presented here clearly indicates that auro-

taeniatus does not belong with the other species of Pavaedes, Aedes auro-

taeniatus will be treated in detail in a separate publication,

The abbreviations used in the literature cited section conform to the

BIOSIS List of Serials, Biosciences Information Service of Biological Abstracts,

Philadelphia, 1972. In the synonymy sections, an asterisk following the

abbreviations used (9 = female, o’= male, P = pupa, L = larva) indicates that

at least some portion of that sex or stage is figured. In the distribution sec-

tions, the abbreviations used are the same as in the synonymy sections, but

with the following 2 additions: p = pupal exuvium andl = larval exuvium. In

the pupal descriptions and tables the number of branches of abdominal seta 1-I

was counted on the basal third of the seta; therefore, only primary branches

are recorded. The scale for the illustrations is in millimeters, Distribution

records are indicated as follows: countries are in capital letters, provinces

and primary administrative divisions are in italics, and place names are with

the first letter capitalized, The number of specimens examined from each

province follows the place name of the province in the distribution section. The

spelling of provincial and locality names was taken from the following Official

Standard Names Gazetteers prepared by the Geographic Names Data Base

Division, Defense Mapping Agency and the Office of Geography, U. S, Depart-

ment of the Interior: Malaysia (1970); Philippine Islands (vols. I and II, 1953);

South Vietnam (1971); Sri Lanka (= Ceylon) (1960); and Thailand (1966). Locality

names which did not appear in the gazetteers were spelled according to the

collection data sheets and labels on the specimens,

Information in the bionomics and distribution sections was taken from the

collection data sheets and specimen labels of the specimens that I examined and

from the published literature. In the type-data sections information within

brackets [ | is additional to that included on the original specimen labels,

Nomenclature and chaetotaxy used for the female, male, male genitalia,

pupa and larva follow Knight (1970, 1971), Knight and Laffoon (1970a, 1970b,

1971a, 1971b), Laffoon and Knight (1973) and Harbach and Knight (1978a,

1978b). The terminology of the female genitalia follows Reinert (1974). The

term scape index is defined as the ratio of the length of the larval antennal

scape to its width at the point of attachment of seta 1-A (Reinert 1979: 152),

Reinert: Aedes (Pavaedes) in the Oriental Region 3

GENUS AEDES MEIGEN

SUBGENUS PARAEDES EDWARDS

Type species: Pavaedes barvraudi Edwards

Aioretomyia in part of Leicester 1908: 185.

Pseudohowardina in part of Theobald 1910: 227.

Danielsia of Ludlow 1911: 128,

Ochlerotatus in part of Edwards 1913: 228; Brunetti 1920: 133; Senior-White

1923: 72,

Aédes (Aédes) in part of Dyar and Shannon 1925: 78; Edwards 1928: 53, 1929: 3,

Aedes (?Aedimorphus) of Barraud 1928: 374, 1934: 267; Edwards 1932: 171.

Paraédes Edwards, in Barraud 1934: 446,

Aedes (Aedimorphus) in part of Bohart 1945; 63; Carter 1950: 89; Knight and

Hull 1953: 453; Chow, Thevasagayam and Tharumarajah 1954: 117,

Paraédes of Thurman 1954: 83 (in part); Mattingly 1971: 1.

Aédes (Paraédes) of Mattingly 1958a: 4, 19, 1958b: 76, 1958c: 106.

Aedes (Paraedes) of Stone, Knight and Starcke 1959: 199; Stone and Delfinado

1973: 306; Knight and Stone 1977: 147,

MALE, Head, Antenna 0. 89-1.30 length of proboscis, lightly plumose

with setae of whorls long and evenly spaced around shaft, pedicel with a few

small scales and short fine setae mesally, maxillary palpus dark scaled,

Short, 0.09-0,12 length of proboscis; clypeus bare; proboscis dark scaled,

0.87-1.14 length of femur I; vertex covered with broad decumbent dark scales;

occiput with a few erect forked scales; eyes contiguous or separated. Thorax.

Scutal integument dark; scutum covered with narrow curved dark scales,

narrow curved pale scale patterns present or absent; prescutellar bare space

without scales; scutal setae as follows: 2-5 anterior promontory, scutal fossal

[2-5 anterior, 1-4 lateral, 0-2 (usually 1) posterior], numerous dorsocentral

(anterior and posterior), acrostichal absent (except in barvaudi which has 3,

4 anterior), numerous supraalar, 4-6 posterior medial scutal, and 1 postalar

callar; scutellum with narrow curved and/or broad scales, 4-11 setae on

median lobe and 2-6 setae on lateral lobe; pleural integument usually dark; antepro-

nota widely separated, each with narrowcurved, and/or broad scales, 5-14 setae;

postpronotum with narrow curved, and/or broad scales, 3-7 posterior setae;

propleuron with broad scales, 3-8 setae (thailandensis with 14-22 setae); pro-

sternum, subspiracular area, mesomeron, metameron and mesopostnotum bare;

paratergite usually bare (scaled in thailandensis); postspiracular area with or

without scales, 1-4 setae; mesepisternum with a patch of broad scales on both

upper and lower areas, 1-4 upper and 6-15 posterior setae; prealar knob with-

out scales, 4-12 setae; mesepimeron with a patch of broad pale scales, 6-15

setae dorsad of scales. Legs. Tibiae I-III and tarsi I-III dark scaled; post-

tarsil,II with ungues equal in size, toothed or simple, III with ungues equal in

size and simple. Wing. Alula with a single row of scales on margin; 1, 2

remigial setae; upper calypter with or without setae on margin. Abdomen.

Terga dark scaled, I with a rectangular patch of pale scales on laterotergite °

II-VI with a laterobasal pale scaled patch. Genitalia. Tergum IX covered with

minute spicules, bilobed, with 2-9 setae on each lobe; gonocoxite moderately

long and moderately broad, tergal surface basomesally with a patch of setae ‘

sternal surface apicomesally with a patch of long setae, scales present; gono-

stylus long, attached at apex of gonocoxite, bifid (trifid in bonneae), one or 2

of arms with spicules forming file-like ridges, one arm with short fine setae,

Spiniform absent; basal mesal lobe connected to gonocoxite at base of mesal

4 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

membrane, well developed, with 2 arms, one arm long, narrow, curved and

with 3 stout apical setae (apices usually recurved), other arm broader, with

3-5 flattened apical or subapical setae or with a number of spines (chrysoscuta

and ostentatio), basal mesal lobe connected mesally to its mate by a band that

is covered with short spicules; proctiger short, apex of paraproct curved ter-

gally as a blunt heavily pigmented point, cercus membranous with a moderately

large and moderately pigmented dorsal plate, setae absent; phallosome with

aedeagus divided into 2 lateral plates each with up to 3 stout teeth, paramere

long, 0.80-1.02 length of lateral plate of aedeagus, parameral apodeme long

and narrow; and sternum IX covered with minute spicules, 2-4 (usually 2) long

stout setae.

FEMALE. Essentially as in male but with the following differences. Head.

Antenna 1.14-1.52 length of proboscis; maxillary palpus 0.16-0.23 length of

proboscis; proboscis 0. 75-1.14 length of femur I; vertex with narrow curved

pale scales forming a triangular patch or forming a line along coronal suture,

with or without erect forked scales. Thovax. Pale scale patterns, when

present, usually better developed than in male; paratergite bare (scaled in

thailandensis and pagei); setal differences as follows: 2-6 anterior promontory,

scutal fossal (2-6 anterior and 1-5 lateral), 0-4 (usually absent) anterior

acrostichal, scutellar (5-13 on median lobe and 4-11 on lateral lobe), 7-15 ante-

pronotal, 4-16 propleural, 1-7 postspiracular, 1-4 upper and 7-18 posterior

mesepisternal, 4-13 on prealar knob, and 6-18 upper mesepimeral. Legs.

Posttarsi I-III with ungues equal in size, toothed or simple. Wing. Upper

calypter with setae on margin. Genitalia. Tergum VIII covered with minute

spicules, moderately pigmented, scales usually absent (0-2 scales in

chrysoscuta), basal 0. 7-1.0 retracted into segment VII, base concave mesally,

apex straight or slightly convex, several to numerous short setae scattered over

apical 0.46-0.94, setae along apex short and thin, some species also with

moderately long thin setae apically, basolateral seta present or absent (usually

absent), VIlI-Te index 0.56-1.12, VIII-Te/IX-Te index 1.73-2.65, length 0.15-

0.28 mm, width 0.22-0.34 mm; sternum VIII covered with minute spicules,

moderately pigmented (thailandensis and menoni with a large lightly pigmented

median area), scales usually absent (0-4 scales in pagei), base concave mesal-

ly, apex usually with a moderately deep median indentation (0.11-0.22 of

length) (bonneae and menoni with only a small indentation, 0.06-0.09 of length)

and with a small to medium size lobe on each side of midline, short to moder-

ately long setae on apical 0.71-0.98 of sternum, setae 1-3-S widely separated,

located ina more or less diagonal line, 1-S basomesad of 2-S and 3-S apico-

laterad of 2-S, apical intersegmental fold lightly pigmented, VIII-S index

0.54-0.98, length 0.17-0.32 mm, width 0.30-0.43 mm; tergum IX covered with

minute spicules, moderately pigmented, apex with a small median indentation

or straight and with 1-7 setae on each side of midline, 3-12 total setae, IX-Te

index 0.56-0.98, length 0.06-0.12 mm, width 0.10-0.15 mm; insula covered

with minute spicules, moderately pigmented, long, tongue-like, with 2-7

small tuberculi each with a minute spicule, tuberculi located about 0.5 from

base; lower vaginal lip with numerous minute to long spicules scattered over

surface, moderately pigmented, narrow, lower vaginal sclerite absent; upper

vaginal lip with short spicules, heavily pigmented, narrow, upper vaginal

sclerite well developed, moderately to heavily pigmented, medium to large

size; postgenital lobe covered with short spicules, moderately long, moder-

ately broad, apex with a very small to moderately deep median indentation

(0.03-0.25 of length), with 4-13 setae on each side of midline, 9-26 total

setae, dorsal PGL index 0. 85-1.09, ventral PGL index 1.20-1.73, ventral

Reinert: Aedes (Pavaedes) in the Oriental Region 5

length 0.09-0.15 mm; perianal membrane with scattered short spicules over

entire surface; cercus covered with short spicules, moderately long, apex

sharply rounded with 2,3 long stout setae, dorsal surface with a number of

short to moderately long setae on apical 0.70-0.91, scales usually absent

(0-5 scales in bonneae, chrysoscuta and ostentatio, and 1-8 in menoni), ven-

tral surface with a few short setae on lateral margin, cercus index 2.15-

3.48, cercus/dorsal PGL index 2.41-3.71, cercus length 0.15-0.27 mm; one

large seminal capsule (one large and 2 medium size ones in menoni), heavily

pigmented, spherical, several small seminal capsule pores near orifice, base

of accessory gland duct moderately to heavily pigmented.

PUPA. Cephalothorvax. Ocular plate with cuticular facets poorly developed;

setae 1-3-CT moderately long, subequal in length, multiple branched; 9-CT

longer than 8-CT. Respiratory trumpet. Index 3.21-6.31. Metanotal plate.

Seta 11-CT single, occasionally barbed, longer than 10, 12-CT; 10, 12-CT

multiple branched. Abdomen. Setae 1-II, III multiple branched, 1-II with

0-22 branches; 2-II moderately long to long, stout and only slightly shorter than

3-II; 4-VIII single (rarely 2, 3 branched), noticeably longer than 3-VIII; 5-IV-

VI long, single (occasionally 2 branched in collessi); 9-VIII moderately long

to long, single (rarely 2 branched in bonneae). Paddle. Without fringe of

hair-like spicules; index 1.01-2.22; seta 1-P moderately long to long, single.

A summary of ranges of setal branching for known species (bonneae,

chrysoscuta, collessi, ostentatio and thailandensis) follows. Cephalothorax.

Seta 1-CT with 2-7 branches; 2, 3, 7-CT with 2-5 branches; 4-CT single to

7 branched; 5-CT single to 8 branched; 6-CT single; 8-CT with 2-10 branches;

9-CT single to 6 branched. Metanotal plate. Setae 10, 12-CT with 2-8

branches; 11-CT single and occasionally barbed. Abdomen. Seta 1-I with

15-53 branches in basal 0.33; 2, 6-I single or 2 branched; 3-I single to 3

branched; 4-I with 4-10 branches; 5-I with 2-6 branches; 7-I single to 5

branched; 9, 10-I single; 0, 3-II single (occasionally 2 branched in thailanden-

sis); 1-II with 5-23 branches; 2, 6, 9-II singleor2 branched; 4-II with 3-11

branches; 5-II with 2-9 branches; 7-II single to 6 branched; 0, 2, 3, 9, 11,

14-III single; 1-III with 2-10 branches; 4-III single to 5 branched; 5-III with

2-11 branches; 6-III single to 8 branched; 7, 10-III single to 5 branched; 8-III

with 2-6 branches; 0, 2, 9, 11, 14-IV single; 1-IV single to 9 branched; 3-IV

single to 8 branched; 4, 7, 8-IV single to 4 branched; 5-IV single or 2 branched;

6-IV single to 6 branched; 7-IV single to 3 branched; 10-IV single to 5 branched;

0, 2, 5, 9, 11, 14-V single; 1, 6-V single to 6 branched; 3-V single to 4

branched; 4-V with 2-7 branches; 7-V single to 8 branched; 8-V with 2-5

branches; 0, 2, 9, 11, 14-VI single; 1, 4, 6-VI single to 6 branched; 3-VI

single to 3 branched; 5, 7, 10-VI single or 2 branched; 8-VI single to

2 branched; 0, 2, 7, 11, 14-VII single; 1, 4, 10-VII single to 4 branched;

3-VIII single to 5 branched; 5-VII single or 2 branched; 6-VII single to 7

branched; 8-VII single to 6 branched; 9-VII single to 3 branched; 0-VIII single;

4, 14-VIII single to 3 branched. Paddle. Seta 1-P single.

LARVA. Head. Moderately to heavily pigmented; seta 1-C long, stout;

4-C multiple branched, short, mesad and slightly to moderately caudad of 5-C;

0, 6-C multiple branched, long to very long, stout, barbed; 6-C laterad and

slightly cephalad of 5-C; insertions of 4-C and 6-C each approximately equal

distance from 5-C; 7-C multiple branched, long, stout, barbed, caudomesad

of base of antenna; 8-10, 12-C short; 13-C multiple branched, moderately long,

occasionally barbed; dorsomentum with 26-34 teeth; ventromedian cervical

sclerite moderately pigmented. Antenna. Scape moderately long to long,

numerous short spicules scattered along entire length, moderately pigmented,

6 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

index 9. 84-16.90; seta 1-A multiple branched, moderately long, stout, barbed,

inserted 0.45-0.61 from base; 2-6-A inserted at or near apex. Thorax. Seta

8-P single (2 branched in 2 specimens of chrysoscuta), long, stout, barbed.

Abdomen. Seta 6-I-V long, stout, barbed; comb on VIII with 7-18 scales ina

single curved row, each scale with a median apical spine and basally with

short denticles along lateral margins; 4-VIII longer (usually noticeably longer)

than 3-VIII; 1-X short; 2-X multiple branched, moderately long; 3-X very long;

saddle moderately pigmented, incompletely rings segment X, acus absent; 4

anal papillae, moderately long. Siphon. Pigmentation variable; acus well

developed; index 1.77-4.20; pecten on basal 0. 42-0.66 of siphon, with 11-24

teeth, distal one to 3 teeth more widely spaced than remainder; 1-S short,

inserted in basal 0.57-0.69 of siphon distad of last pecten tooth, 2-S inserted

in or near caudal margin of siphon. A summary of ranges of setal branching

for known species (bonneae, chrysoscuta, collessi, ostentatio and thailanden-

sis) follows.. Head. Setae 0, 1, 3, 14, 18, 20-C single; 4-C with 5-13 bran-

ches; 5-C with 3-8 branches; 6, 10-C with 2-6 branches; 7-C with 5-14

branches; 8-C single to 8 branched; 9-C with 2-5 branches; 11-C with 2-8

branches; 12-C with 3-11 branches; 13-C with 4-9 branches; 15-C with 2-5

branches. Antenna. Seta 1-A with 3-6 branches. Thorax. Seta 0-P with

7-20 branches; 1-P single to 4 branched; 2, 9-P single to 3 branched; 3-P with |

2-6 branches; 4, 11-P single to 6 branched; 5, 6, 10, 12-P single; 7-P with

2-4 branches; 8-P single or 2 branched; 14-P with 2, 3 branches; 1-M with

2-5 branches; 2-M single to 5 branched; 3-M single to 3 branched; 4-M with

2-7 branches; 5, 7, 10, 12-M single; 6-M with 3-7 branches; 8-M with 4-9

branches; 9-M with 4-8 branches; 11-M single or 2 branched; 13-M with 9-27

branches; 14-M with 5-20 branches; 1-T single to 4 branched; 2-T single to 6

branched; 3-T with 5-17 branches; 4-T with 2-6 branches; 5, 10-T single; 6,

11-T single to 3 branched; 7-T with 4-12 branches; 8-T with 3-14 branches;

9-T with 2-4 branches; 12-T single or 2 branched; 13-T with 7-20 branches.

Abdomen. Setae 0, 14-VIII single; 1-VIII with 3-7 branches; 2-VIII single to

4 branched; 3-VIII with 3-13 branches; 4-VIII single to 3 branched; 5-VIII with

3-10 branches; 1-X single to 3 branched; 2-X with 4-13 branches; 3-X single;

4-X with 8-10 setae on grid, each with 2-16 branches (some setae of each

specimen always with 8 or more branches), and usually one short precratal

seta. Siphon. Seta 1-S with 2-6 branches; 2, 6, 7, 9-S single; 8-S single to

4 branched.

EGG. Not known.

MEDICAL IMPORTANCE. Females of several species (collessi, ostentatio

and thailandensis) have been collected feeding on man. Leicester (1908: 194)

reports that ostentatio is a vicious biter during the day. Even though the

females of several species feed on man, few investigations have been conducted

for pathogen isolation from members of this subgenus. Four specimens of

ostentatio from The Philippines were dissected and found to be negative for

Brugia malayi (Brug) parasites (Rozeboom and Cabrera 1965: 208). Ramachan-

dran et al. (1970: 511) examined 10 specimens of osteniatio collected from

Trengganu, Malaysia, and found they were negative for Brugia malayi and non-

filarial nematodes. Small numbers of specimens of both collessi and ostentatio

from Malaysia were dissected and found negative for filaria parasites (Ann.

Rep. Inst. Med. Res. for 1963-1965, 1968, Fed. Malaya and Malaysia, 1968:

63, 89, 1968: 27, 1968: 23, 32, 1968: 112).

DISTRIBUTION. The subgenus Pavaedes occurs throughout most of the

Oriental region. Species of the subgenus have been collected from India,

Indonesia, Malaysia, The Philippines, Sri Lanka, Thailand and Vietnam.

Reinert: Aedes (Paraedes) in the Oriental Region 7

DISCUSSION. The subgenus Paraedes is characterized and distinguished

from the other subgenera of Aedes by the following: in the adults by the com-

bination of (1) numerous anterior and posterior dorsocentral setae; (2) acro-

stichal setae usually absent (3, 4 in male of barraudi, 0-3 in female of

collessi and 2-4 in female of thailandensis); (3) lower mesepimeral seta absent;

(4) prescutellar bare space without scales; (5) paratergite usually bare

(scaled in thailandensis and female of pagei); and (6) male palpus short, less

than 0.12 length of proboscis; in the male genitalia by the combination of (1)

aedeagus with 2 approximated lateral plates each with up to 3 lateral teeth;

(2) basal mesal lobe with 2 arms, one arm long, narrow, curved and with 3

stout apical setae, other arm broader, with 3-5 flattened apical setae or with

long narrow spines; and (3) gonostylus moderately long, bifid or trifid, one or

2 arms with spicules forming file-like ridges, one arm with short fine setae,

spiniform. absent; in the female genitalia by the combination of (1) only a

single large seminal capsule (one large and 2 medium size ones in menoni);

(2) insula tongue-like, with 2-7 small tuberculi each with a minute spicule,

tuberculi located approximately 0.5 from base; (3) upper vaginal sclerite well

developed, narrow; (4) tergum VIII and sternum VIII usually without scales;

and (5) cercus moderately long, index 2.15-3.48, usually without scales; in

the pupa by the combination of (1) seta 9-CT longer than 8-CT; (2) seta 1-II

with 5-22 branches; (3) seta 9-VIII moderately long to long, single (2 branched

in 2 specimens of bonneae); (4) seta 1-P moderately long to long, single; (5)

paddle shape (see figures) and without a fringe of hair-like spicules; and (6)

seta 2-II moderately long to long, stout and only slightly shorter than 3-II;

and in the larva by the combination of (1) position and development of head

setae 4-7-C; (2) antennal scape spiculate, seta 1-A moderately long, stout,

multiple branched; (3) 6-I-V long, stout, barbed; (4) 4-VIII single (rarely 2 or

3 branched in chrysoscuta), noticeably longer than 3-VIII; (5) comb on VIII

with 7-18 scales, in a single curved row, each scale with a median apical

spine; (6) segment X with saddle incomplete, acus absent, 2-XK moderately

long and multiple branched, 3-X very long and single; and (7) pecten with dis-

tal one to 3 teeth more widely spaced than basal teeth.

The short maxillary palpi of the males of Pavaedes are similar to those

of the following subgenera of Aedes: Aedes, Bothaella Reinert, Cancraedes

Edwards, Christophersiomyia Barraud, Huaedes Huang, Leptosomatomyia

Theobald, Nothoskusea Dumbleton, Rhinoskusea Edwards, Vervallina Theo-

bald and most species of Geoskusea Edwards.

The absence or reduction of the setae on the margin of the upper calypter

of the male wing is similar to the reduced number found in the males of the

subgenus Cancvaedes, members of the subgenus Rhinoskusea, and the genus

Udaya.

Male genitalia of Pavaedes are unique and easily separated from those of

other subgenera of Aedes by the development of the basal mesal lobe, gono-

stylus and phallosome. Species of the subgenus Pavaedes, however, show

some resemblance in the male genitalia to those of the subgenus Aedes, but

they can be segregated by the above features. Reinert (1976a: 4) pointed out

the similarity of the file-like ridges on the gonostylus between Pavaedes and

the subgenus Jndusius Edwards.

The female genitalia of Pavaedes are most similar to those of the sub-

genus Aedes. Differences between the 2 subgenera are: Pavaedes has the

insula with small tuberculi near middle of length, tergum VIII without scales,

one large seminal capsule (menoni with one large and 2 smaller ones), and

tergum IX with one to 7 setae on each lobe; while Aedes has the insula with

8 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

small tuberculi on apical 0.5, tergum VIII with a number of scales, one large

and 2 smaller seminal capsules, and tergum IX with 12 or more setae on each

lobe. Other subgenera of Aedes with a single large seminal capsule are

Cancraedes, Rhinoskusea, Niveus Group of Finlaya Theobald and several

species of Aedimorphus. These subgenera are easily distinguished from

Pavaedes by: Cancraedes and Aedimorphus have different shaped cerci and

terga VIII and IX, and the tuberculi are located on apical 0.3 of the insula,

and Rhinoskusea and Finlaya have a lip-like insula with long setae and the

upper vaginal sclerite is absent.

Larvae of the species of Paraedes are uniform in external morphology

and difficult to separate with the exception of thailandensis, which has several

unique characters. Pavaedes larvae are very similar morphologically to

those of the subgenus Aedes and Ae. (Neomelaniconion) lineatopennis (Ludlow).

The following features are distinctive for each. Pavaedes larvae have: seta

4-C inserted caudad of 5-C; 5, 6-C very long; 13-C usually 6-8 (4-9) branched;

4-VIII noticeably longer than 3-VIII; siphon without accessory setae; and crab

holes as their usual immature habitat; Aedes larvae have: seta 4-C inserted

cephalad of 5-C; 5, 6-C long; 13-C usually 2-4 (2-5) branched; 4-VIII shorter

than or approximately equal to length of 3-VIII; siphon with accessory setae

inserted usually subdorsally and/or subapically; and fresh water ground pools

as their immature habitat; and Ae. (Neo.) lineatopennis larvae possess the

features listed for the subgenus Aedes except for the accessory setae of the

siphon which are absent and 4-VIII longer than 3-VIII. The latter species also

has setae 1, 2-VIII inserted on a common sclerotized plate, and the saddle

nearly encircles segment X.

The subgenus Pavaedes appears to be most similar morphologically to the

subgenus Aedes as noted above in the characters of the adult, male genitalia,

female genitalia and larva. These 2 subgenera, however, can be separated by

the features previously outlined. These 2 subgenera also have allopatric dis-

tributions.

BIONOMICS. Adults of most species of Paraedes have been collected feed-

ing on man, and at times they have been reported as vicious biters. Biting

activity has occurred in partially and heavily shaded areas of forests, bamboo

groves and coastal villages. Some species have been taken in light and Malaise

traps. Aedes collessi has been collected from human, monkey, Berok and

calf baited traps.

The usual immature habitat is fresh water in small and large crab holes;

however, they have also been collected from water in a small deep hole near

a stream, elephant footprints, nipah palm fronds, ground pools among nipah

palm trees, a small pool ina jungle, anda small wheel track. These habitats

were located in valleys and in hilly and mountainous terrain, in partially or

heavily shaded areas of secondary rain forests, secondary scrub, bamboo

groves and a tapioca plantation.

KEYS TO SPECIES OF AEDES (PARAEDES)

FEMALES

1. Scutellum with broad dark scales; scutum without pale scales on margins

oF prescutehar bare Gpaces ieee Si OO Ei ae es bonneae

Scutellum with narrow curved scales; scutum with pale scales on margins

Of prebeutellar Dare Space eo EG ok es SR AO ee we os 2

2(1).

3(2).

(3).

5(2).

6(5).

4(3).

5(4).

6(2).

Reinert: Aedes (Paraedes) in the Oriental Region 9

Antepronotum with only narrow curved scales. ..........e..-. 3

Antepronotum with broad scales (may also have narrow curved

SOAICB ON ae ng ae a eee ae es 5)

Posttars? 1-H wit toothed: Unsues se rr a es Oe paget

Posttarsi I+ Ul with Simple ungues. 650s Vee es Oe as 4

Genitalia with a single large seminal capsule; antepronotum with only

GaP BCAlGES eee rr ee a a a ee collessi

Genitalia with one large and 2 medium size seminal capsules;

antepronotum with both dark and pale scales. ........ menoni

Paratergite with narrow pale scales; antepronotum with both narrow

curved and broad pale scales; sternum VIII index 0. 86-0.96.

thailandensis

Paratergite without scales; antepronotum with only broad dark and/or

pale scales; sternum VIII index 0.59-0.73. ............ 6

Pleural integument and antennal pedicel pale golden-brown; posttarsus

fil ‘with toothed tngues. FOP Per es . «6 Chrysoscuta

Pleural integument and antennal pedicel dark brown; posttarsus III with

SIM ple VNBUER OI yas SAE Ne Res Oy ostentatio

Gonpsty lus ities PE Os We CO a eee oe bonneae

Golmetylls Dit ee a ee Ee ae aN ees eee 2

Basal mesal lobe with one arm bearing 3-5 flattened apical setae. . 3

Basal mesal lobe with one arm bearing long thin spines. ....... 6

Basal mesal lobe with one arm bearing 3 flattened apical setae.

collessi

Basal mesal lobe with one arm bearing 4 or 5 flattened apical setae.

Basal mesal lobe with one arm bearing 5 flattened apical setae.

thailandensis

Basal mesal lobe with one arm bearing 4 flattened apical setae. .. 5

Gonostylus bifid with longer arm greatly expanded and with 6-8 short

SOUAG ON MUR OUR tMOe eek eae eee 8 eo we al barraudi

Gonostylus bifid with longer arm only slightly expanded and with 3, 4

ST Oe ee ee GP gy ae eg are 6 menoni

Gonocoxite with a subapical patch of 35-45 setae on sternal surface;

basal mesal lobe with one arm gently tapered from base to a pointed

apex and with long dendritic spines on apical 0.4. . . chrysoscuta

Gonocoxite with a subapical patch of 20-25 setae on sternal surface;

basal mesal lobe with one arm swollen near middle and with short

simple spines near middle and on apical 0.25. ...... ostentatio

10 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

PUPAE

1. Seta 1-CT with 2 branches; seta 1-IV single or 2 branched. ._ collessi

Seta 1-CT with 3-7 branches; seta 1-IV with 4-9 branches. ..... 2

2(1). Seta 6-V with 5, 6 branches; seta 6-VI with 2-6 branches; paddle index

iO de bl Od cae Cu wie ee ew ee thailandensis

Seta 6-V single to 3 branched; seta 6-VI single; paddle index

Di Bae obs et es a es a ee. 3

3(2). Insertion of seta 2-II laterad of insertion of seta 3-II; insertion of seta

3-III far laterad of insertion of seta 1-II]......... bonneae

Insertion of seta 2-II mesad of insertion of seta 3-II; insertion of seta

3-III slightly laterad of insertion of seta 1-II]............ 4

4(3). Seta 10-III usually single or 2 branched, rarely 3 branched; seta 10-IV

usually 2 branched, rarely 3,4 branched.......... ostentatio

Seta 10-III usually 4,5 branched, rarely 3 branched; seta 10-IV usually

3,4 branched, rarely 2 branched... 6:6 6 sie heli ios chrysoscuta

FOURTH STAGE LARVAE

Le Pecten with 18-24 teeth; seta 5-C with 3 branches. ... thailandensis

Pecten with 11-16 teeth; seta 5-C with 4-8 branches. ........ 2

2(1). Siphon with basal 0.4 more darkly pigmented than apical 0.4. .collessi

Siphon with apical 0.4 more darkly pigmented than basal0.4.... 3

3(2). Siphon with apical 0.4 darkly pigmented, brown. ....... bonneae

Siphon with apical 0.4 very darkly pigmented, dark blackish-brown .

4(3). Seta 1-P single; seta 8-M with 5,6 branches; pecten usually with 14-16

TOR a a a ee ae ee chrysoscuta

Seta 1-P with 2,3 branches; seta 8-M with 7-9 branches; pecten usually

WED abe FOC. es hee tad Miao Vio ostentatio

SPECIES TREATMENT

AEDES (PARAEDES) BARRA UDI (EDWARDS)

(Pigs..1, 3, 27)

Pavaédes barraudi Edwards, in Barraud 1934: 447 (o*).

Aédes (Pavaédes) barraudi of Mattingly 1958a: 24 (key), 1958b: 78 (“*),

1958c: 106 (lectotype designation).

Aedes (Pavraedes) barvaudi of Stone, Knight and Starcke 1959: 199; Stone and

Delfinado 1973: 306; Knight and Stone 1977: 148.

MALE (Fig. 1). Head. Antenna dark brown, 0. 89-0.94 length of pro-

boscis, pedicel dark brown with a few small brown scales and with short fine

Reinert: Aedes (Paraedes) in the Oriental Region 11

brown setae mesally; maxillary palpus 0.11 length of proboscis; clypeus dark

brown; proboscis 1.01-1.10 length of femur I; vertex covered with broad de-

cumbent dark brown scales, a few narrow curved white scales on interocular

space, ocular line with small narrow curved white scales; lateral surface

covered with broad dark brown scales and with a small patch of broad white

scales anterior to antepronotum and posterior to eye margin; occiput with

narrow curved decumbent white scales and a few moderately long erect forked

golden-white and brown scales; interocular and ocular setae dark brown and

well developed; eyes contiguous anteriorly, but slightly separated dorsally.

Thorax. Scutal integument reddish-brown; scutum covered with narrow curved

reddish-brown scales except for narrow curved white scales on following areas:

a small patch on anterior promontory area, a small patch on scutal fossal

areas extending anteriorly onto lateral area above postpronotum, a small

patch at scutal angle and a few similar scales along posterior margin of scutal

ridge, a small patch on posterior scutal fossal area, a small patch on supra-

alar area anterior to base of wing, a short narrow stripe on anterior portion

of posterior dorsocentral area, a small patch on posteriolateral margin of

prescutellar space and 3 small indistinct patches on anterior dorsocentral and

acrostichal areas each of 3,4 scales; scutum with reddish-brown setae as

follows: 2 anterior promontory, 3,4 anterior acrostichal, numerous dorso-

central (anterior and posterior), scutal fossal (3,4 anterior, 1,2 lateral, and

1,2 posterior), 30-36 supraalar, 3,4 posterior medial scutal; scutellum with a

patch of narrow curved white scales on each lobe, 4,5 long and 1,2 short setae

on median lobe, 3 long and 1-3 short setae on lateral lobe; pleural integument

dark brown; antepronotum with broad white scales, 7-9 brown setae; postpro-

notum with a few narrow curved reddish-brown scales dorsally and a small

patch of 10-12 moderately broad curved pale scales on posterior area anterior

to setae, 4,5 brown posterior setae; postspiracular area with a few broad

white scales, 3,4 golden setae; propleuron with a patch of broad white scales,

4,9 golden-brown setae; paratergite bare; mesepisternum with an upper and a

lower posterior patch of broad white scales, 8-13 golden setae on upper and

posterior areas, lower setae shorter and white; prealar knob with 5,6 golden

setae; mesepimeron with a patch of broad white scales, 9,10 pale golden setae

dorsad of scales; other pleural areas bare. Legs. Coxae I-III with several

brown or golden setae, I with anterior surface with broad brown scales and a

dorsal and a ventral patch of white scales, II and III with a patch of broad white

scales on anterior surface; trochanters I-III with white scales and a few short

setae; femora I and II with anterior surface brown scaled, III with a broad ven-

tral longitudinal stripe of broad white scales from base to near apex on both

anterior and posterior surfaces, a similar stripe on posterior surface of II, I

with a broad dorsal longitudinal white scaled stripe from base to near apex of

posterior surface, I-III with a small dorsoapical white scaled patch; posttarsi

I-III (Fig. 27) with 2 ungues, I and II with ungues equal in size and with a

tooth, IIT with ungues equal in size, both simple. Wzng. Dorsal and ventral

veins brown scaled; costa with a small white scaled patch on base of anterior

surface; alula with narrow pale brown scales along margin; 1,2 golden remigial

setae; upper calypter bare. Halter. Pedicel pale; capitellum white scaled

with a few pale brown scales at base. Abdomen. Terga II-VI dark brown

scaled with a laterobasal white scaled patch (remainder of abdomen removed

with genitalia); sterna brown scaled; terga and sterna with numerous short

golden setae, mostly along posterior margins. Genitalia (Fig. 3). Tergum IX

moderately to heavily pigmented, strongly bilobed, with 7,8 short setae on

each lobe; gonocoxite moderately pigmented, tergal surface with 5-7 long stout

12 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

and 11-13 short to moderately long setae scattered over surface and with a

short double row of short setae on basal 0.2 of mesal margin, lateral surface

with several moderately long setae, sternal surface with 6-9 long stout setae

on lateroapical 0.25 and a number of short setae scattered over remainder of

area, a subapical circular patch of 49-54 short and moderately long setae on

mesal margin, lateral surface and lateral portions of tergal and sternal sur-

faces with scales; gonostylus approximately 0.71 length of gonocoxite, pedicel

short, broad and 0.3 of overall length, distal 0.7 bifid and divided into a long

narrow mesally curved arm with short ridges of spicules on mesal surface,

inserted 0.3 from base of gonostylus, and a mesal longer flattened arm with

apical portion with spicules forming file-like ridges, a row of 6-8 short setae

and several short thin setae on sternal surface, several minute setae in a row

on tergal surface; basal mesal lobe divided at base into 2 long arms, sterno-

mesal arm cylindrical with a slight bend near middle and with 3 moderately

long hook-like setae at apex, outer arm somewhat flattened and forked in

distal 0.36 into an outer extension with 4 short stout setae and a similarly

shaped bare mesal extension, also a ventral sclerotized structure near 0.4

from base of arm; phallosome with aedeagus divided into 2 pigmented narrow

lateral plates connected basally, apical 0.3 with a lightly pigmented sternal

connection, distal portion of each plate bluntly rounded with apex pointed and

projected mesally, paramere approximately 0.88 length of lateral plate, para-

meral apodeme long and narrow; sternum IX large, with 2 long stout setae

near center.

FEMALE, PUPA and LARVA. Not known.

TYPE-DATA. The lectotype male of barvaudi, deposited in the British

Museum (Natural History) (BMNH), possesses the following data on the adult

labels: Coorg, Virajpet, S. India, VI.1927, J. D. Baily | collector]; 2673 | collec-

tion number]; Pavaédes| barraudi Edwards, HOLOLECTOTYPE, P. F. Mattingly,

14: ii: 58; Type | circular label witha red border];andSEAMP Acc. No. 295. The

lectotype is in good condition and the genitalia are mounted in balsam. A

single male paratype has the same label data as the lectotype except for the

collection number, 2672, and a paratype label [circular label with a yellow

border]. The paratype is in fair condition and the genitalia are mounted in

balsam on a small cover glass on a card attached to the pin. Aedes barvaudi

is still known only from the 2 males which Edwards (in Barraud 1934: 447)

designated as cotypes. Mattingly (1958c: 106) selected a lectotype from the

cotypes.

DISTRIBUTION. Two specimens examined: 2°.

INDIA. Virajpet, Coorg, June 1927, J. D. Baily, 2673 (lectotype), 1°,

2672 (paratype), 1°.

Distribution from literature.

INDIA. Virajpet, Coorg, South India (Edwards in Barraud 1934: 447;

Mattingly 1958c: 106).

DISCUSSION. The adult habitus of barraudi is distinctive in that the scale

pattern of the scutum (Fig. 1) easily separates this species from the others of

the subgenus. The presence of 3, 4 anterior acrostichal setae in the male of

barraudi differs from the males of other species of the subgenus which have

these setae absent. However, females of collessi and thailandensis have 0-3

and 2-4 anterior acrostichal setae, respectively.

Male genitalia of barvaudi are most similar to those of collessi especially

in the development of the gonostylus. Features of the basal mesal lobe, gono-

coxite and tergum IX (Figs. 3, 6) are distinctive for these 2 species.

BIONOMICS. Edwards, in Barraud (1934: 447), states that the type was

Reinert: Aedes (Paraedes) in the Oriental Region 13

collected in June.

AEDES (PARAEDES) BONNEAE MATTINGLY

(Figs, 2° 4° 10. 17, 22°93. OY oe)

Aédes (Paraédes) bonneae Mattingly 1958a: 8, 11, 13, 21, 22, 24, 25, 27,

BU, Oy: Des, (Fy on Fy ee els

Aedes (Paraedes) bonneae of Stone, Knight and Starcke 1959: 199; Stone and

Delfinado 1973: 306; Knight and Stone 1977: 148.

MALE. Head. Antenna dark brown, 1.15-1.29 length of proboscis,

pedicel dark brown with a few short fine setae mesally; maxillary palpus

0.11-0.12 length of proboscis; clypeus dark brown; proboscis 1.02-1.10

length of femur I; vertex completely covered with broad decumbent dark

brown scales; lateral surface with broad dark brown scales and with a small

patch of broad pale scales anterior to antepronotum and posterior to eye

margin; occiput with a few short erect forked dark brown scales; interocular

and ocular setae dark brown and well developed; eyes contiguous anteriorly.

Thorax. Scutal integument dark brown; scutum covered with narrow curved

dark reddish-brown scales; scutum with dark setae as follows: 2 anterior

promontory, numerous dorsocentral (anterior and posterior), scutal fossal

(3-5 anterior, 2-5 lateral and 1 posterior), numerous supraalar, 4 posterior

medial scutal; scutellum with a patch of broad scales on each lobe, 6,7 setae

on median lobe and 4 setae on lateral lobe; pleural integument dark brown;

antepronotum with a patch of broad dark brown scales on posterior surface and

a few narrow dark brown scales on anterior area, 5-7 dark setae; postpronotum

with narrow curved dark reddish-brown scales, 3 dark posterior setae; pro-

pleuron with a patch of broad silvery-white scales, 5,6 dark or golden setae;

paratergite bare; postspiracular area with a few broad translucent white

scales posterior of setae (scales rubbed off in some specimens), 3 dark

setae; mesepisternum with a large patch of broad silvery-white scales on

upper area and a small patch of similar scales on lower area, 2,3 dark setae

on upper area and 7-9 setae on posterior area, lower setae short and pale;

prealar knob with 4-6 dark setae; mesepimeron with a large patch of broad

silvery-white scales, 10-12 short dark setae dorsad of scales. Legs. Coxae

I-III with several setae, I with anterior surface with broad brown scales and

with a small patch of silvery-white scales dorsally, II and III with a small

patch of broad silvery-white scales on anterior surface; trochanters I-III

pale scaled and with a few short setae; femora I-III with anterior surface

brown scaled, II and III with posterior surface brown with a ventrobasal pale

scaled area; posttarsi I-III (Fig. 27) with 2 ungues, I and II with ungues equal

in size and with a tooth, III with ungues equal in size, both simple. Wing.

Dorsal and ventral veins with dark brown scales; costa without a patch of white

scales; alula with moderately broad dark brown scales on margin and a second

row of moderately broad to broad scales parallel to margin; 1,2 dark remigial

setae; upper calypter with a number of narrow dark setae on margin. AHalter.

Pedicel pale; capitellum covered with broad dark brown scales. Abdomen.

Terga II-VII dark brown scaled and with a small patch of pale scales on latero-

basal area; sterna dark brown scaled with a few pale scales on basal areas;

terga with a few short setae on lateral and posterior margins. Genitalia (Fig.

4). Tergum IX heavily pigmented, bilobed, with 3-5 short stout setae on each

lobe, cephalic margin emarginate, lateral margin narrowly connected to

14 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

sternum IX; gonocoxite heavily pigmented, tergal surface with 5-7 moderately

long and long stout setae on mediolateral area, 2 moderately long setae on

mesal margin approximately 0.3 from base, sternal surface with several very

long stout setae on apical 0.4, mesal margin with a widely spaced row of stout

setae, apical setae very long and basal setae short, lateral and sternal sur-

faces with numerous broad scales, tergal, sternal and lateral surfaces covered

with minute spicules; gonostylus approximately 1.07 length of gonocoxite, tri-

fid, outer arm long, narrow, with apex pointed and slightly curved, 6,7 short

setae along length, inner arm with a sharp bend at approximately 0.22 from

base, long, narrow, apical 0.25 curved outward, apex with a small patch of

spiculate rows, third arm arising from inner arm near basal bend, arm long,

very narrow and with apex expanded into a hemispherical club bearing 12,13

short setae; basal mesal lobe divided at base into 2 long caudally projecting

arms, dorsomesal arm narrow at base and expanding into a broad apical area

that is covered with moderately long spicules, 3 moderately long, broad, flat

setae at approximately 0.32 from apex, ventral arm long, narrow, covered

with minute spicules, apical area with 3 long curved setae; phallosome with

aedeagus divided into 2 heavily pigmented lateral plates that are connected

basally, each plate curved tergomesad and with 1,2 long basomesal teeth, a

lightly pigmented dorsal flap covering lateral plates, paramere approximately

0.86 length of aedeagus, parameral apodeme moderately broad, approximately

1.65 length of paramere; sternum IX moderately to heavily pigmented, apical

margin rounded, 2 long stout setae on caudomesal area.

FEMALE. Essentially as in male but with the following differences. Head.

Antenna 1.17-1.26 length of proboscis, pedicel also with a few small dark

scales mesally; maxillary palpus 0.18-0.19 length of proboscis; proboscis

1.02-1.07 length of femur I; vertex also with a few narrow pale scales on

coronal suture; lateral surface with pale scaled patch larger; occiput with a

few narrow pale scales in some specimens. Thovax (Fig. 2). Scaling as in

male but also with a few narrow curved pale scales on anterior promontory

area, anterior scutal fossal area, and a few on acrostichal area; scutellum with

a few broad pale scales basad of dark brown scales of median lobe, 8-11 dark

setae on median lobe and 6-8 setae on lateral lobe; antepronotum with narrow

curved dark brown scales on posterior surface and a few broad brown scales

on anterior area (some specimens with broad scales rubbed off), 7-9 dark

setae; postpronotum with 3,4 posterior dark setae; propleuron with 6-8 setae;

postspiracular area with 3,4 setae; mesepisternum with 3, 4 upper and 10-14

posterior setae; prealar knob with 7-9 dark setae; mesepimeron with 15-18

setae dorsad of scale patch. Legs. Posttarsi I-III (Fig. 28) with 2 ungues, I

and IIT with ungues equal in size and with a tooth, III with ungues equal in size,

both simple. Abdomen. Terga with medium size laterobasal pale scaled

patches. Genitalia (Fig. 10). Tergum VIII moderately pigmented, scales

absent, basal 0.75-0.90 retracted into segment VII, base concave mesally,

apex straight, several short setae scattered over apical 0.46-0.61, setae

along apex short and thin with the exception of 2,3 moderately long thin ones,

basolateral seta absent, VIII-Te index 0.56-0.57, VIII-Te/IX-Te index 2. 54-

2.65, length 0.17-0.19 mm, width 0.28-0.32 mm; sternum VIII moderately

pigmented, scales absent, base concave mesally, apex with a small median

indentation (0.06-0.08 of length) and with a very small lobe on each side of

midline, short to moderately long setae on apical 0.71-0.79, seta 2-S approxi-

mately 0.58 from 1-S, seta 3-S approximately 0.42 from 2-S, apical inter-

segmental fold lightly pigmented, VIII-S index 0.54-0.60, length 0.17-0. 20

mm, width 0.30-0.32 mm; tergum IX moderately pigmented, apex nearly

Reinert: Aedes (Paraedes) in the Oriental Region 15

straight, base with a small median indentation, 1,2 setae on each side of mid-

line at apex, 3,4 total setae, IX-Te index 0.56-0.63, length 0.06-0.07 mm,

width 0.11 mm; insula with 4,5 small tuberculi; lower vaginal lip with numer-

ous minute spicules scattered over entire surface, lower vaginal sclerite ab-

sent; upper vaginal lip heavily pigmented, narrow, upper vaginal sclerite

moderately pigmented, large size; postgenital lobe with a moderately deep

median indentation (0.11-0.17 of length) apically, 6-9 (usually 8) setae on

each side of midline, 15, 16 total setae, dorsal PGL index 0. 88-0.96, ventral

PGL index 1.46-1.70, ventral length 0.09-0.10 mm; cercus with 2,3 long

stout setae apically, dorsal surface with a number of short to moderately long

setae on apical 0. 84-0. 88, 2-4 broad scales, ventral surface with a few short

setae on lateral margin, cercus index 2.41-2.58, cercus/dorsal PGL index

3.04-3.13, cercus length 0.16-0.19 mm; one large seminal capsule, base of

accessory gland duct heavily pigmented.

PUPA (Figs. 17, 22). Chaetotaxy as figured and recorded (Table 1).

Cephalothorax. Seta 1-CT with 3-5 branches. Respiratory trumpet. Index

3.73-5.25, mean 4.53. Abdomen. Seta 1-II with 14-22 branches; 2-II inserted

laterad of 3-II; 1-III with 4-10 branches; 1-IV with 4-9 branches; 1-V with 3-6

branches; 6-VI single. Paddle. With minute spicules on apical 0.31-0. 46 of

lateral and apical 0.08-0.16 of mesal margins; index 1.39-2.03, mean 1. 86.

LARVA (Fig. 23). Description based on 10 larval exuviae with associated

adults. Head. Moderately to heavily pigmented; setae 0, 1,3, 14,18, 20-C

single; 4-C with 6-11 (8) branches; 5-C with 5-7(5) branches; 6-C with 4-6 (5)

branches; 7-C with 9-14 (10) branches; 8-C with 2,3 (2) branches; 9-C with 2,3

(3) branches; 10-C with 2-4 (2) branches; 11-C with 4-7 (4) branches; 12-C with

3-10 (5) branches; 13-C with 4-9 (6) branches; 15-C with 2-4 (3) branches;

dorsomentum with 29-33 (usually 32) teeth, heavily pigmented. Antenna.

Scape moderately long, index 9. 84-10.97, mean 10.58; seta 1-A with 3-6 (4)

branches, inserted 0.52-0.55 from base. Thorax. Seta 0-P with 7-14 (8)

branches; 1, 9-P single or 2 (1) branched; 2-P with 2 branches; 3-P with 2-4

(3) branches; 4, 11-P single to 3 (2) branched; 5, 6, 8,10, 12-P single; 7-P with

2,3 (3) branches; 14-P with 2,3 (2) branches; 1-M with 2-4 (3) branches; 2-M

single to 3(2) branched; 3,5, 7,10,12-M single; 4-M with 3, 4 (3) branches; 6-M

with 4-6 (6) branches; 8-M with 5-8 (6) branches; 9-M with 5-7 (7) branches;

11-M single or 2 (1) branched; 13-M with 9-17 (12) branches; 14-M with 10, 11

branches; 1, 6-T single to 3 (2) branched; 2-T with 3,4 (3) branches; 3-T with

6-10 (9) branches; 4-T with 2-5 (3) branches; 5, 10-T single; 7-T with 5-9 (8)

branches; 8-T with 5,6 (6) branches; 9-T with 3, 4 (3) branches; 11, 12-T

single or 2 (1) branched; 13-T with 7, 8 (8) branches. Abdomen. Setae 0,

14-VIII single; 1-VIII with 3, 4 (3) branches; 2-VIII single to 3(2) branched;

3-VIII with 3-7 (5) branches; 4-VIII single or 2 (1) branched; 5-VIII with 4-7

(6) branches; comb on VIII with 11-16 (usually 12-14) scales; 1-X single or

2 (2) branched; 2-X with 6-10 (7) branches; 3-X single; ventral brush with 10

setae on grid, each with 2-13 branches, some setae always with 10 or more

branches, anterior setae shorter and with fewer branches than posterior ones.

Siphon. Moderately pigmented with apical 0.4 more heavily pigmented than

basal 0.4; index 2.67-3.92, mean 3.17; pecten on basal 0.45-0.51, with 11-15

(usually 12,13) teeth, each tooth with one or 2 small ventral denticles near

base, apex of teeth long, narrow and pointed, distal 1,2 teeth spine-Like,

without denticles and wider spaced than remainder; 1-S with 2-4 (2) branches,

inserted on basal 0. 60-0. 64 of siphon distad of last pecten tooth; 2-S single,

short; 6,7,9-S single; 8-S single or 2 (2) branched.

TYPE-DATA. The holotype male, deposited in the BMNH, possesses the

16 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

following data on the adult labels: SELANGOR, Rantau Panjang, 4.3. 1953,

J. A. Reid[on upper surface of label], Ex. Coll. I.M.R. Malaya [on under sur-

face of label]; 598/4 [collection number]; A. (Skusea) near curtipes Det. J. A.

Reid; Aé. (Paraédes) bonneae Mattingly, HOLOTYPE, P. F. Mattingly, 20:

viii: 1957; andTYPE [circular label with a red border]. The holotype is in

good condition and the genitalia are intact. The specimen has the left fore-

and midlegs missing. The pupal and larval exuviae associated with the holo-

type are mounted in balsam ona microscope slide. The allotype is in excel-

lent condition and possesses the same collection data as the holotype except

the collection number, 596/7. The pupal and larval exuviae associated with

the allotype are mounted in balsam on a microscope slide. Paratype labels

are on 12 female and 6 male adult specimens (pupal and larval exuviae are

also indicated) and are as follows: 592 1c, 592/12 Opl, 593 19 and 1c’, 594

lo’, 594/9 Qpl, 596 29 and 10, 596/4 Qpl, 596/6 cpl, 598 29 and 1o’, 598/1

Cpl, 598/5 9pl, 598/6 @pl, and 598/9 9pl. All these paratypes have the same

collection data as the holotype except for the following: 592, 592/12, 593

(10") larvae in ground pool among nipah palms; 594, 596 (19) larvae in crab

holes; 596 (19) larva in water between nipah palm fronds; and 598/1 larva in

nipah palm frond. Mattingly (1958a: 32) reported the type-series was collec-

ted on 4: v : 1953 and consisted of the holotype, allotype and 6 male and 14

female paratypes. All the above specimens I examined were dated 4.3.53.

DISTRIBUTION. Fifty-six specimens examined: 3 Cpl, 7%, 10 Spl and

10°.

MALAYSIA. Selangor, Rantau Panjang, 4 Mar. 1953, 598/4 (holotype);

596/7 (allotype); paratypes: 592, /12; 593; 594, /9; 596, /4, /6; 598, /1, /5,

/6, /9, 2 Spl, 50, 8 Qpl, 42, and May 1957, 037, 1% and 2; other than type-

series, same location, 22 Aug. 1968, 1679, S. W. James, C. Y. Wang and

S. bin Oman, 22; Kg. Rantau Panjang, Aug. 1957, J. A. Reid, 0275/1, /2,

O277/1, 0278/1, 4 Spl, 18, Pepband 19;

Distribution from literature.

MALAYSIA. Selangor, Rantau Panjang (Mattingly 1958a: 32).

DISCUSSION. Adults of bDonneae are easily identified by the scutum which

is dark scaled and without pale scaled areas. This species also has broad

dark scales on all lobes of the scutellum, paratergite bare, femora I-III with-

out dorsoapical white scales, and antepronotum with both narrow curved and

broad brown scales.

The male genitalia of bonneae are distinctive in the trifid gonostylus and

the development of the basal mesal lobe, which has only 3 flattened setae on

the dorsomesal arm (Fig. 4).

Female genitalia of bonneae are separated from those of the other species

by the shape of tergum IX and sternum VIII and the setae being restricted to

the apical area of tergum VIII.

| The shape of the male genital lobe (Fig. 22) of the pupa of bonneae is unique

in that the apex is broad and nearly truncate. Other features of the pupae are:

base of 2-II inserted slightly laterad and cephalad of 3-II; 3-III inserted far

laterad of 1-III; and 6-CT greater than 0.5 length of 7-CT. The latter feature,

6-CT, is shared by pupae of chrysoscuta and ostentatio, but differs from those

of collessi and thailandensis in which 6-CT is less than 0.5 of the length of

7-CT.

The larva of bonneae can be identified by the pigmentation of the siphon

which has the apical 0.4 more darkly pigmented than the remainder. Aedes

chrysoscuta and ostentatio also have the apical portion of the siphon more

heavily pigmented than the remainder; however, the apical 0.4 is very dark

Reinert: Aedes (Paraedes) in the Oriental Region 17

colored while in bonneae it is more lightly colored. The antennal scape index

is 9.84-10.97 for bonneae, but 11.75-12.50 for ostentatio and 15. 31-16. 90 for

thailandensis.

BIONOMICS. In Malaysia larvae have been collected from water in crab

holes, in ground pools among nipah palm trees and in nipah palm fronds during

March, May and August. Females were taken in Malaysia biting man during

August at a coastal village, at sea level, 10-99 m from houses, and in partial

shade.

Mattingly (1958a: 34) records the immature habitats in Malaysia as ground

pools, crab holes and nipah palm fronds.

AEDES (PARAEDES) CHRYSOSCUTA (THEOBALD)

(Figs. 5,11, 18,94; 97 98)

Pseudohowardina chrysoscuta Theobald 1910: 228 (9).

Ochlerotatus ostentatio of Edwards 1913: 228 (in part); Brunetti 1920: 140

(in part); Senior-White 1923: 81 (in part), 1927: 62.

Aedes (?) ostentatio of Edwards 1922b: 468 (in part); Brug and Bonne-Wepster

1947: 185 (in part).

Aedes (? Aedimorphus) ostentatio of Barraud 1928: 374 (2), 1934: 267 (2, in

part); Edwards 1932: 171 (in part); Macdonald 1957: 21 (in part),

Aedes (Aedimorphus) ostentatio of Knight and Hull 1953: 463 @, in part); Chow,

Thevasagayam and Tharumarajah 1954: 117; Horsfall 1955: 518 (in part).

Aédes (Aédimorphus) ostentatio of Carter 1950: 89, 106. 3

Aédes (Paraédes) ostentatio of Mattingly 1958a: 7,9, 22,24,25 (2, key, in pard;

Qutubuddin 1960: 138.

Aedes (Paraedes) ostentatio of Stone, Knight and Starcke 1959: 200 (in pari);

Basio 1971: 26 (in part); Stone and Delfinado 1973: 307 (in part);

Harrison et al. 1974: 153; Knight and Stone 1977: 148 (in part).

MALE. Head, Antenna dark brown, 1.28-1.30 length of proboscis, pedi-

cel pale golden-brown with a few short fine brown setae and small dark scales

mesally; maxillary palpus 0.10 length of proboscis; clypeus pale golden-brown;

proboscis 0.87-0.94 length of femur I; vertex covered with broad decumbent

dark brown scales, a few narrow curved golden-white scales on interocular

space, ocular line with narrow curved golden scales; lateral surface covered

with broad dark brown scales and with a patch of broad white scales anterior

to antepronotum and posterior to eye margin; occiput with a few short erect

forked brown scales and a few broad decumbent white scales; interocular and

ocular setae dark brown and well developed; eyes slightly separated anteriorly.

Thorax. Scutal integument moderately brown; scutum covered with narrow

curved reddish-black scales, except for narrow curved golden scales on

anterior promontory area, anterior scutal fossal area (some specimens also

with golden scales on lateral scutal fossal area and a very narrow line on

acrostichal area, a few along lateral margins of prescutellar bare space, and

a small patch on supraalar area); scutum with reddish-black setae as follows:

2,3 anterior promontory, numerous dorsocentral (anterior and posterior),

scutal fossal (2-4 anterior, 1,2 lateral and 1 posterior), numerous supraalar,

4 short posterior medial scutal; scutellum with a small patch of narrow curved

reddish-black scales on each lobe (some specimens also with a few narrow

curved golden scales on median lobe, 6-8 setae on median lobe, 3, 4 setae on

lateral lobe; pleural integument pale golden-brown; antepronotum covered with

18 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

broad dark brown scales, 6-9 reddish-black setae; postpronotum with narrow

curved reddish-black scales along dorsal margin and a small patch of 3-8

broad dark brown scales on posterior area in front of setae, 3,4 reddish-black

posterior setae; propleuron covered with broad white scales, 5-8 golden setae;

paratergite bare; postspiracular area with 2-5 broad white scales (some speci-

mens with scales rubbed off), 2-4 dark brown setae; mesepisternum with a

large patch of broad white scales on upper area and a small patch of similar

scales on lower area, 2,3 setae on upper area and 8-12 setae on posterior

area, lower setae short and pale; prealar knob with 5-8 dark brown setae;

mesepimeron with a large patch of broad white scales, 8-15 short golden

setae dorsad of scales. Legs. Coxae I-III with several dark brown setae, I

with anterior surface with broad white scales, II and III with a small anterior

patch of broad white scales; trochanters I-III with a few broad pale scales and

a few short golden setae; femur I with anterior and posterior surfaces covered

with dark brown scales, I also with a posteriodorsal pale scaled stripe from

base to near apex, femur II with anterior and posterior surfaces dark brown

scaled and a posteroventral longitudinal white scaled stripe on basal 0.7,

stripe broader proximally, femur III dark brown scaled with an anteroventral

and a posteroventral longitudinal white scaled stripe on basal 0. 80-0. 92,

stripe broader proximally; posttarsi I-III (Fig. 27) with 2 ungues, I and II with

ungues equal in size and with a tooth, III with ungues equal in size, both

simple. Wing. Dorsal and ventral veins dark brown scaled; costa without a

patch of white scales; alula with narrow brown scales along margin; 1,2 short

brown remigial setae; upper calypter bare. Halter. Pedicel pale; capitellum

dark brown scaled. Abdomen. Terga dark brown scaled, II-VII with a small

laterobasal patch of white scales, VII white scaled with a few brown scales

on posterior margin; sterna white scaled, III-VII with a narrow apical band of

brown scales, VIII brown scaled, terga and sterna with short setae, mostly

along posterior and lateral margins. Genitalia (Fig. 5). Tergum IX strongly

bilobed, with 2-5 short setae on each lobe; gonocoxite with an apical sterno-

mesal lobe, tergal surface with 3, 4 long stout and 9-12 moderately long setae

and 3 irregular rows of setae on basal 0.75 of tergomesal margin, setae short

distally and increase in length proximally until they become long at base,

lateral surface with a few long and moderately long setae, sternal surface

with a number of long stout and a few short setae on apical 0.25 and a sub-

apical circular patch of approximately 35-45 moderately long and a few short

setae on mesal margin, a single short seta at base near middle, lateral

surface and lateral areas of tergal and sternal surfaces with scales; gonostylus

with pedicel narrow and distal portion bifid and divided into a narrow club-

shaped tergal arm bearing 11-17 short setae and a file-like area of spicules,

and a flattened rectangular shaped sternal arm with small hair-like spicules

on distal 0.55; basal mesal lobe divided at base into 2 long arms, mesal arm

with distal 0.4 narrower, curved tergally and with 3 moderately long hook-like

setae at apex, proximal 0.5 somewhat flattened, expanded distally, and

covered with short hair-like spicules, outer arm very long, narrow and gently

tapering from near base to apex, distal 0.4 narrower and covered with long

dendritic spines, becoming more numerous distally, basal 0.5 cylindrical

and with a short finger-like projection near base which bears 2,3 short setae,

one or 2 apically and one subapically; phallosome with aedeagus divided into

2 heavily pigmented lateral plates connected basally, each plate with one or

2 stout teeth, apical 0.3 with a lightly pigmented sternal connection, distal

portion of each plate bluntly rounded with apex pointed and projected mesally,

paramere approximately 0.96 length of lateral plate, parameral apodeme

Reinert: Aedes (Paraedes) in the Oriental Region 19

moderately long and narrow; sternum IX large, with 2,3 long stout setae near

center.

FEMALE. The female differs markedly from the male in scale patterns

of the head, thorax and legs and the number of thoracic setae. The following

features differ from the male. Head. Antenna 1.41-1.52 length of proboscis;

maxillary palpus 0.21-0.23 length of proboscis; proboscis 0.77-0.85 length of

femur I; vertex covered with broad decumbent dark blackish-brown scales and

a median triangular patch of narrow curved golden ones, numerous erect

forked golden scales scattered throughout golden scale patch from occiput to

interocular setae, a few erect forked dark scales also on lateral areas of

occiput; interocular setae golden, well developed. Thorax (Fig. 24). Scutum

covered with narrow curved reddish-black scales, narrow curved golden

scales on the following: anterior promontory area, extending from anterior

scutal fossal area over lateral scutal fossal area to scutal angle, a few on

scutal ridge, a broad stripe on acrostichal area extending from anterior

promontory area over posterior medial scutal area and along lateral margins

of prescutellar bare space, a distinct narrow stripe on anterior dorsocentral

area to approximately scutal ridge, and a small patch on supraalar area

cephalad of wing base; scutellum with curved golden scales on all lobes, also

usually a few narrow curved dark scales on posterior margins; setal differ-

ences are: 2-4 anterior promontory, 4-6 anterior scutal fossal, 6 posterior

medial scutal, scutellum with 6,7 lateral and 7-9 median, 8-10 antepronotal,

4-6 postpronotal, 7-10 propleural, 2,3 upper and 8-18 posterior mesepisternal,

6-11 on prealar knob, and 10-16 mesepimeral; postspiracular area with a

patch of broad white scales. Legs. Femora I-III with a small dorsoapical

patch of white scales; posttarsi I-III (Fig. 28) with 2 ungues equal in size and

with a tooth. Wing. Costa with a few broad pale scales at base; upper calypter

with dark setae on margin. Halter. Capitellum with dark scales at base and

white scales apically. Abdomen. Tergum VIII with a laterobasal patch of

white scales. Genitalia (Fig. 11). Tergum VIII with 0-2 scales, basal 0. 85-

0.95 retracted into segment VII, base concave mesally, apex straight, numer-

ous short setae scattered over apical 0.67-0.82, setae along apex short to

moderately long, basolateral seta absent, VIII-Te index 0.74-0.82, VIII- Te/

IX-Te index 2.19-2.31, length 0.25-0.27mm, width 0.32-0.34mm; sternum VII

with scales absent, base slightly concave mesally, apex with a moderately

deep median indentation (0.11-0.13 of length) and a small lobe on each side of

midline, short to moderately long setae on apical 0.92-0.98, apical interseg-

mental fold lightly pigmented, VIII-S index 0.66-0.73, length 0.27-0.30 mm,

width 0.40-0.43 mm; tergum IX with apex with a small median indentation and

with 3-5 short thin setae on each side of midline, 6-10 total setae, IX-Te index

0. 79-0. 84, length 0.11-0.12 mm, width 0.14-0.15 mm; insula with 2-4 small

tuberculi; lower vaginal lip with numerous minute spicules; upper vaginal lip

with vaginal sclerite moderately large, narrow; postgenital lobe with apex with

a very small to small median indentation (0.04-0.11 of length), 11-13 (usually

11,12) setae on each side of midline, 21-26 total setae, dorsal PGL index

0.90-1.07, ventral PGL index 1.45-1.73, ventral length 0.13-0.15 mm; cer-

cus with 2 long stout setae at apex, dorsal surface with a number of short to

moderately long setae on apical 0. 73-0.83, 1-5 scales (usually 1-3), ventral

surface with a few short setae on lateral margin, cercus index 2. 81-3. 32,

cercus/dorsal PGL index 2.64-2.97, cercus length 0.25 mm; one large

seminal capsule, base of accessory gland duct moderately to heavily pigmented.

PUPA (Fig. 18). Chaetotaxy as figured and recorded (Table 2). Cephalo-

thovax. Seta 1-CT with 2-5 branches; 5-CT with 3-6 branches. Respiratory

20 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

trumpet. Index 5.51-6.31, mean 5.92. Abdomen. Seta 4-I with 4-8 (usually

6,7) branches; 5-I with 2-4 (usually 2,3) branches; 1-II with 10-23 branches;

2-II inserted mesad of 3-II; 10-III usually 4,5 (8-5) branched; 7-IV single to

3 branched; 10-IV usually 3, 4 (2-5) branched; 1-V with 2-5 branches; 6-VI

single. Paddle. With minute spicules on apical 0.52-0.956 of lateral and

apical 0.13-0.14 of mesal margins; index 1.32-1.46, mean 1.41.

LARVA (Fig. 24). Description based on 5 larval exuviae with associated

adults and 5 whole larvae. Head. Moderately pigmented; setae 0,1, 3,14, 18,

20-C single; 4-C with 9-13 (12) branches; short; 5-C with 4-6 (5) branches,

very long; 6-C with 4,5 (4) branches, very long; 7-C with 7-13 (9) branches,

long; 8-C with 2,3 (3) branches; 9-C with 2-5 (4) branches; 10-C with 2-6 (4)

branches; 11-C with 4-6 (4) branches; 12-C with 5-10 (8) branches; 13-C with

5-9 (7) branches; 15-C with 3-5 (4) branches; dorsomentum with 30-34 (30)

teeth, heavily pigmented. Antenna. Scape long, index 10.22-10.98, mean

10.53; seta 1-A with 4,5 (4) branches, inserted 0.53-0.57 from base. Thorax.

Seta 0-P with 8-19 (12) branches; 1-P with 2-4 (2) branches; 2-P single to

3 (2) branched; 3-P with 2-6 (3) branches; 4-P with 3-6 (4) branches; 5, 6, 10,

12-P single; 7-P with 2-4 (3) branches; 8-P single or 2 (1) branched, 9-P

single to 3 (3) branched; 11-P with 2-5 (3) branches; 14-P with 2 branches;

1-M with 2-5 (4) branches; 2-M with 2-4 (3) branches; 3-M single or 2 (1)

branched; 4-M with 3-7 (5) branches; 5,7,10-12-M single; 6-M with 4-6 (5)

branches; 8-M with 5, 6 (5) branches; 9-M with 5-8 (7) branches; 13-M with

10-27 (16) branches; 14-M with 8,9 branches; 1-T with 2-5 (3) branches; 2-T

single to 6 (5) branched; 3-T with 7-17 (17) branches; 4, 9-T with 2-4 (3)

branches; 5,10,11-T single; 6-T with 2,3 (3) branches; 7-T with 5-9 (6)

branches; 8-T with 5-10 (8) branches; 12-T with 2 branches; 13-T with 8-10

branches. Abdomen. Setae 0, 2,7,13-I single; 1, 6-I with 2,3 (3) branches;

3-I with 4-8 (5) branches; 4-I with 9-17 (9) branches; 5-I with 3-5 (4)

branches; 9-I with 2-4 (3) branches; 10-I with 2 branches; 11-I single or 2 (1)

branched; 0, 2, 9, 14-II single; 1, 8-II with 2,3 (3) branches; 3-II with 3-7 (7)

branches; 4-II with 5-12 (10) branches; 5-II with 2,3 (2) branches; 6-II with 2

branches; 7-II with 3-7 (4) branches; 10-II single to 4 (2) branched; 11-II single

to 3 (2) branched; 12-II with 2-5 (4) branches; 13-II with 9-11 (11) branches;

0, 2,9, 11, 14-III single; 1, 13-II with 5,6 (6) branches; 3-III with 3-6 (6)

branches; 4-III with 2-4 (2) branches; 5, 8-III single to 3 (2) branched; 6-III

single or 2 (2) branched; 7-III with 4-9 (6) branches; 10-III with 2,3 (3)

branches; 11-III single or 2 (1) branched; 12-III with 2 branches; 0, 2,9, 14-IV

single; 1-IV with 5,6 (6) branches; 3-IV with 5,6 (5) branches; 4-IV with 2,3

(2) branches; 5, 8-IV single to 2 (2) branched; 6, 11, 12-IV with 2 branches;

7-IV with 2-4 branches; 10-IV single to 3 (1) branched; 13-IV with 4 branches;

0,2,9,14-V single; 1-V with 3-7 (5) branches; 3, 10, 13-V with 3,4 (3) branches;

4-V with 6-9 (8) branches; 5, 6, 12-V single to 2 (2) branched; 7-V with 4-13

(6) branches; 8-V with 2,3 (2) branches; 11-V with 2 branches; 0, 2,9, 10, 14-VI

single; 7-VI with 7 branches; 3-VI with 3-8 branches; 4-VI with 3-6 branches;

5-VI with 2-4 (2) branches; 6-VI single to 4 branched; 7-VI with 3-7 (4)

branches; 8-VI single to 2 (1) branched; 11, 12-VI with 2 branches; 13-VI with

12-20 (12) branches; 0, 2, 14-VII single; 1-VII with 2-5 (3) branches; 3-VII with

5-7 (7) branches; 4, 5-VII with 4,5 (5) branches; 6-VII with 10-14 branches;

7-VII with 2-4 (4) branches; 8-VII with 9-13 branches; 9-VII with 3 branches;

10-VII with 3-5 branches; 11-VII single or 2 (2) branched; 12-VII with 3, 4 (3)

branches; 13-VII with 7-11 branches; 0, 14-VIII single; 1-VIII with 3-7 (4)

branches; 2-VIII with 2-4 (3) branches; 3-VII with 3-9 (7) branches; 4-VII

single to 3 (1) branched; 5-VIII with 4-7 (5) branches; comb on VIII with 12-16

Reinert: Aedes (Paraedes) in the Oriental Region 21

(usually 14-16) scales; 1-X single to 3 (2) branched; 2-X with 4-7 (6) branches,

moderately long; 3-X single, long; 4-X with 8,9 setae on grid, each with 8-16

branches and one short precratal seta. Siphon. Moderately to heavily pig-

mented, apical 0.4 very heavily pigmented, dark brownish-black; index 3.37-

4.20, mean 3.76; pecten on basal 0.51-0.56, with 12-16 (usually 14-16) teeth,

each tooth with one or 2 small ventral denticles near base, apex long, narrow

and pointed, distal 1-3 teeth spine-like, without denticles and wider spaced than

remainder; 1-S with 2-6 (5) branches, inserted on basal 0. 68-0. 69 of siphonand

distad of last pecten tooth; 2-S single, short; 6, 7, 9-S single; 8-S with 2 branches.

TYPE-DATA. The holotype female of Pseudohowardina chrysoscuta

Theobald is in the BMNH and possesses the following data: Peradeniya, Cey-

lon, 4-07, Green; Pseudohowardina chrysoscuta Type 2, F. V. T.; Type.

DISTRIBUTION. Ninety specimens examined: 10 %pl, 5p, 2%, 12 2pl,

1 2p, 32, 3 larval exuviae, 3 larvae and 1 pupa.

SRI LANKA. Central, Kandy District, Kandy, Peak View Motel, 15-24

Jan. 1970, D. R. Davis and W. H. Rowe, 1° and 2°; same province and dis-

trict, Wakarwatte, 23-24 Jun. 1975, 75-2; 76-1, -2, -3, -4; 78-1, -2, -3;

79-1; 83-1, -2, -3, -4, -5, -100, -101, -103, -104, -105, -106; 84, 84-1,

-2, -3, -4; 85, 85-2, -3, E. L. Peyton and Y-M. Huang, 10 Cpl, 5 Op,

1°, 10 Spl, 1 2p, 12, 1P, 3L, 31; same except Udawattekele, 25 Jun. 1975,

109-1, 1 2pl. Western, Alutgama, 26 Oct. 1978, No. 56, N. Jayasekera, 1

Ppl.

Distribution from literature.

INDIA (Bohart 1945: 63; Brug and Bonne-Wepster 1947: 185); Malabar

Coast (Barraud 1928: 374, 1934: 268; Mattingly 1958a: 28, questions the

records from India).

SRI LANKA (Bohart 1945: 63; Brug and Bonne-Wepster 1947: 185);

Peradeniya (Theobald 1910: 224), Bibile (Barraud 1934: 268); Central, Kandy

District, Kandy, Peak View Motel (Harrison et al. 1974: 153); Bibile

(Mattingly 1958a: 26).

DISCUSSION. Adults of chrysoscuta are similar in habitus to those of

ostentatio. Prime differences useful in distinguishing the 2 species follow,

for chrysoscuta: pleural integument, antennal pedicel, and clypeus each pale

golden-brown; antepronotum of female covered with broad dark brown scales;

costa of female wing with a few pale scales at base; female posttarsus III with

both ungues toothed; female antenna length 1.41-1.52 of proboscis; female pal-

pus length 0.21-0.23 of proboscis; female proboscis length 0.77-0.85 of femur

I; and male antennal length 1.28-1.30 of proboscis; and for ostentatio: pleural

integument, antennal pedicel, and clypeus each dark brown; antepronotum of

female covered with broad dark brown scales and a small patch of broad white

scales; costa of female wing completely brown scaled; female posttarsus III

with both ungues simple; female antennal length 1.27-1.35 of proboscis; female

palpus length 0.18-0.19 of proboscis; female proboscis length 0. 91-0. 99 of

femur I; and male antennal length 1.17-1.21 of proboscis.

The sexes of chrysoscuta are dimorphic in the pale scaled patterns of the

head and scutum. These pale scaled patterns are also variable between indi-

viduals from the same collection.

Male genitalia of chrysoscuta closely resemble those of ostentatio, how-

ever, they can be separated by the following, in chvysoscuta: sternal surface

of gonocoxite with a subapical patch of 35-45 setae; tergal surface of gono-

coxite with 3 irregular rows of setae on basomesal area, setae short distally

and long basally; sternal surface of gonocoxite with a single long stout seta on

median basal area; and basal mesal lobe bifid with one arm gently tapering

22 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

from base to a pointed apex and with long dendritic spines on apical 0.4; and

in ostentatio: sternal surface of gonocoxite with a subapical patch of 20-25

setae; tergal surface of gonocoxite with one or 2 irregular rows of setae on

basomesal area, setae short distally and moderately long basally; sternal sur-

face of gonocoxite almost always without a single long stout seta on median

basal area; and basal mesal lobe bifid with one arm swollen near middle and

with short simple spines near middle and on apical 0.25. The male genitalia

of both chrysoscuta and ostentatio are unique and easily separated from those

of the other species of the subgenus in the development of the basal mesal lobe

in that one of the arms does not have stout flattened setae.

Female genitalia of chrysoscuta are similar to those of ostentatio. They

can be separated by the following, in chrysoscuta: sternum VIII length 0. 40-

0.43 mm; tergum VIII length 0.32-0.34 mm; postgenital lobe with 21-26 setae;

cercus length 0.25 mm; cercus index 2. 81-3.32; and tergum IX index 0.79-

0. 84; and in osteniatio: sternum VIII length 0.35-0.40 mm; tergum VIII length

0.25-0.31 mm; postgenital lobe with 14-17 setae; cercus length 0.17-0.19 mm;

cercus index 2.20-2.58; and tergum IX index 0.67-0.75.

The pupa of chrysoscuta is very similar to that of ostentatio, but can be

distinguished by the usual branching of setae 4, 5-I and 10-III, IV.

Larvae of chrysoscuta and oStentatio are very similar. The 2 species can

be separated by the following, in chrysoscuta: seta 1-P with 2-4 branches;

8-M with 5,6 branches; 7-IV with 2-4 branches; 8-VI single or 2 branched;

and scape index 10. 22-10.98; and in osteniatio: seta 1-P single; 8-M with 7-9

branches; 7-IV with 7-10 branches; 8-VI with 3-5 branches; and scape index

11.75-12.50. Also, larvae of chrysoscuta usually have 14-16 (12-16) comb

scales while those of ostentatio usually have 9-12 (9-14) comb scales.

BIONOMICS. Adults were taken in Sri Lanka in a Malaise trap at an

elevation of 549 m.

In Sri Lanka immatures were collected 9 times from fresh water in both

small and large crab holes located in partial shade of a secondary rain forest

in mountainous and valley terrain, and at 600 m elevation during June and

October.

Adults from Sri Lanka were collected in April (Theobald 1910: 229) and

January at an elevation of 1800 ft (Harrison et al. 1974: 153).

AEDES (PARAEDES) COLLESSI MATTINGLY

(Pies 2 6, 12. 10,°22, 25, 27° 38)

Aédes (Paraédes) collessi Mattingly 1958a: 13, 18, 21, 22, 24, 25, 27, 28

(oy, OF, PY Lt, keyel.

Aedes (Paraedes) collessi of Stone, Knight and Starcke 1959: 199; Stone and

Delfinado 1973: 306; Knight and Stone 1977: 148.

MALE. Description based on holotype and 2 other males. Head. Antenna

dark brown, 1.20-1.21 length of proboscis, pedicel golden-brown with a few

short fine setae mesally; maxillary palpus 0.09 length of proboscis; clypeus

golden-brown; proboscis 0.96-1.07 length of femur I; vertex and lateral sur-

faces covered with broad decumbent dark blackish-brown scales and with a

very small patch of broad pale scales anterior to antepronotum and posterior

to eye margin; occiput with a few short erect forked dark brown scales; inter-

ocular space covered with broad dark blackish-brown scales; ocular and inter-

ocular setae dark brown and well developed; eyes separated anteriorly. Thorax.

Reinert: Aedes (Paraedes) in the Oriental Region 23

Scutal integument dark brown; scutum covered with narrow curved dark

reddish-brown scales except for a very small patch of narrow curved pale

scales on anterior promontory area; scutum with dark setae as follows: 3,4

anterior promontory, numerous dorsocentral (anterior and posterior), scutal

fossal (3 anterior, 2,3 lateral and 1 posterior), numerous supraalar, 4 posteri-

or medial scutal; scutellum with narrow curved reddish-brown scales on each

lobe, 4 setae on median lobe, and 2,3 setae on lateral lobe; pleural integument

light brown; antepronotum with a few narrow curved dark brown scales (one

specimen also with 2,3 broad dark brown scales), 6-8 dark setae; postpronotum

with a few narrow curved dark reddish-brown scales on upper area, 3 dark

posterior setae; propleuron with a few broad white scales, 3,4 setae; para-

tergite bare; postspiracular area with 1,2 dark setae; mesepisternum with a

moderately large patch of broad white scales on upper area and a small patch

of similar scales on lower area, 1,2 dark setae on upper area and 6 setae on

posterior area, lower setae short and pale; prealar knob with 5, 6 dark setae;

mesepimeron with a large patch of broad white scales, 6-8 short setae dorsad

of scales. Legs. Coxae I-III with several setae, I with anterior surface with

broad scales, white scaled on upper area and pale brown scales on lower area,

II and III with a small patch of broad white scales on anterior surface; trochan-

ters I-III with a few pale brown and white broad scales and a few short setae;

femora I-III with anterior surface brown scaled, I-III with posterior surface

brown scaled, and II and III also with a pale scaled ventral stripe from base to

near apex; posttarsi I-III (Fig. 27) with 2 ungues equal in size and with a tooth.

Wing. Dorsal and ventral veins dark brown scaled; costa without a patch of

white scales; alula with narrow dark brown scales on margin; one dark remigial

seta; upper calypter apparently without setae. Halter. Pedicel pale, capitel-

lum dark brown scaled. Abdomen. Terga II-VII dark brown scaled and with

a moderately large patch of broad white scales on laterobasal area, VIII dark

brown scaled with 1-3 white scales on laterobasal area; sterna white scaled

with a posterior band of brown scales, VIII brown scaled; terga with a few

short brown setae on lateral and posterior margins. Genitalia (Fig. 6). Ter-

gum IX moderately pigmented, bilobed, with 3,4 short setae on each lobe,

cephalic margin emarginate, lateral margin narrowly connected to sternum

IX; gonocoxite heavily pigmented, tergal surface with long setae on apical 0.5,

a row of 9,10 short to moderately long setae on mesal area extending from

near base to approximately 0.3 from apex, 1-3 broad scales, a few very long

stout setae along lateroapical surface, sternal surface with an apical patch of

long stout setae extending along mesal margin as a closely set row to

approximately 0.5 from base, a few scattered setae on basal and lateroapical

areas, a number of broad scales on sternal and lateral surfaces, tergal,

sternal and lateral surfaces covered with minute spicules; gonostylus approxi-

mately 0.73 length of gonocoxite, bifid, with inner arm large and with apex

bluntly rounded, a large patch of spicules forming file-like ridges, a row of

8,9 short setae along inner margin of this patch, 4,5 short setae along outer

margin, other arm short, curved sternomesally, apex pointed and slightly re-

curved, outer margin with short ridges of spicules on apical area; basal mesal

lobe divided at base into 2 long caudally projecting arms, dorsomesal arm

with base broad, flattened, mesal area expanded into a point, covered with

numerous long spicules, apical area narrow and terminating with 3 short

dark blunt flattened setae, ventral arm long, narrow, basal portion curved

ventrad, entire surface covered with minute spicules, apex with 3 moderately

long stout setae; phallosome with aedeagus divided into 2 heavily pigmented

lateral plates connected basally, each plate curved tergomesad and with 1, 2

24 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

long basomesal teeth, a lightly pigmented dorsal flap covering lateral plates,

paramere approximately 1.02 length of aedeagus, parameral apodeme moder-

ately broad, approximately 1.17 length of paramere; sternum IX moderately

to heavily pigmented, apical margin rounded and extended far cephalad, 2 long

stout setae on caudomesal area.

FEMALE. The female differs markedly from the male in the scale pat-

terns of the head and thorax and the number of thoracic setae. The following

features differ from the male. Head, Antenna 1.15-1.30 length of proboscis,

flagellomere 1 approximately 0.75 length of combined lengths of 2 and 3, 1

also with several small broad dark scales on basal 0.5, pedicel with a few

small broad pale brown scales mesally in addition to short fine hairs; maxil-

lary palpus 0.17-0.20 length of proboscis; clypeus dark brown; proboscis 0. 99-

1.03 length of femur I; vertex with broad decumbent dark blackish-brown

scales and a large triangular patch of narrow curved golden scales extending

from occiput anteriorly over interocular space, similar golden scales on ocu-

lar line; lateral surface with a larger patch of broad white scales anterior to

antepronotum; occiput with numerous erect forked golden scales and similar

scales extending onto posterior portion of vertex which is golden scaled.

Thorax (Fig. 2). Scutum with narrow curved reddish-brown scales, but with

narrow curved golden scales on following areas: broad stripe extending from

anterior promontory area over anterior and posterior acrostichal areas and

onto posterior medial scutal area at which point it forks and forms a narrow

stripe along lateral margins of prescutellar bare space, similar scales cover-

ing anterior scutal fossal area and anterior dorsocentral area to just posterior

of scutal ridge, golden scales on dorsocentral and acrostichal areas combined

cover most of anterior median area of scutum, golden scales also on anterior

and anterior portion of lateral scutal fossal areas, a small patch at scutal

angle and a medium size patch on supraalar area (golden scaled areas some-

what variable, some specimens with a few narrow curved reddish-brown

scales on areas between anterior acrostichal and anterior dorsocentral areas);

setal differences are as follows: 3-5 anterior promontory, scutal fossal (3, 4

anterior, 2-4 lateral and 1 posterior), 0-3 anterior acrostichal; scutellum with

narrow curved golden scales on each lobe, lateral lobes of some specimens

also with a few narrow curved reddish-brown scales, 4-6 setae on median

lobe, 4-6 setae on lateral lobe; pleural integument dark brown; antepronotum

with numerous narrow curved dark reddish-brown scales, 8-11 dark setae;

postpronotum with narrow curved dark reddish-brown scales on dorsal and

posterior areas, 4-6 dark posterior setae; propleuron with a large patch of

broad white scales, 5-9 setae; paratergite bare (allotype with a few narrow

curved reddish-brown scales on posteriolateral area); postspiracular area

with several broad white scales posterior to setae, 2-4 dark setae; mese-

pisternum with a large patch of broad white scales on upper area and a medi-

um size patch of similar scales on lower area, 2,3 upper and 7-11 posterior

setae; prealar knob with 4-8 dark setae; mesepimeron with 9-11 setae dorsad

of large patch of broad white scales. Legs. Coxae I with anterior surface

dark brown scaled with a small patch of white scales on dorsal area; femur III

with anterior surface brown scaled with a broad ventral white scaled stripe

from base to near apex, I-III with posterior surface brown scaled with a white

scaled stripe from base to near apex, stripe dorsal on I and ventral on II and

III, II and III also with a small white scaled dorsoapical patch, white scales

also extending onto dorsal areas of anterior and posterior surfaces (some

specimens also with a few dorsoapical white scales on I); posttarsi I-III (Fig.

28) with 2 ungues equal in size and simple. Wing. Costa with a small white

Reinert: Aedes (Paraedes) in the Oriental Region 29

scaled patch at base; upper calypter with a number of narrow dark setae on

margin. Abdomen. Terga II-VII dark brown scaled with a large laterobasal

white scaled patch. Genitalia (Fig. 12). Tergum VIII with scales absent,

basal 0.75-0.90 retracted into segment VII, base concave, apex straight,

numerous short setae scattered over apical 0.79-0.83, setae along apex

short and thin, basolateral seta absent, VIII-Te index 0.64-0.68, VIII-Te/Ix-

Te index 1. 73-1. 89, length 0.15-0.17 mm, width 0.23-0.26 mm; sternum VII

with scales absent, base slightly concave mesally, apex with a moderately

deep to deep median indentation (0.17-0.20 of length) and a small lobe on each

side of midline, short to moderately long setae on apical 0.77-0.94, seta 2-S

approximately 0.38 from 1-S, seta 3-S approximately 0.62 from 2-S, apical

intersegmental fold lightly pigmented, VIII-S index 0.60-0.65, length 0.19-

0.21 mm, width 0.31-0.32 mm; tergum IX with apex with a small median

indentation and with 3-5 (usually 3, 4) short thin setae on each side of midline,

6-9 total setae, IX-Te index 0.76-0.85, length 0.09-0.10 mm, width 0.11-

0.12 mm; insula with 5,6 small tuberculi; lower vaginal lip with numerous

short spicules scattered over entire surface; upper vaginal lip with upper

vaginal sclerite moderately pigmented, moderately large size; postgenital

lobe with apex with a very small median indentation (0.07-0.08 of length),

6-9 (usually 7,8) setae on each side of midline, 14-17 total setae, dorsal PGL

index 0.90-0.96, ventral PGL index 1.52-1.56, ventral length 0.09-0.10 mm;

cercus with 2,3 long stout setae at apex, dorsal surface with a number of

short to moderately long setae on apical 0.70-0.88, scales absent, ventral

surface with a few short setae on lateral margin, cercus index 2.51-2. 54,

cercus/dorsal PGL index 2. 78-3.08, cercus length 0.17 mm; one large

seminal capsule, base of accessory gland duct heavily pigmented.

PUPA (Figs. 19, 22). Chaetotaxy as figured and recorded (Table 3).

Cephalothorax. Setae 1, 5-CT with 2 branches; 9-CT single or 2 branched.

Respiratory trumpet. Somewhat swollen at middle; index 3.33-4.46, mean

3.83. Abdomen. Seta 1-II with 5-14 branches; 2-II inserted laterad of 3-1;

1-III with 2,3 branches; 6-III single; 1-IV-VI single or 2 branched; 3, 6-V

single; 4-V with 2-4 branches; 1-VII single. Paddle. With minute spicules

on apical 0.42-0.48 of lateral and apical 0.16-0.18 of mesal margins; index

1.94-2.22, mean 2.04.

LARVAE (Fig. 25). Description based on 6 larval exuviae with associated

adults. Head. Moderately to heavily pigmented; setae 0,1,3,14,18, 20-C

single; 4-C with 5-7 (7) branches; 5-C with 5-7 (6) branches; 6-C with 4, 5 (4)

branches; 7-C with 9-13 (11) branches; 8-C with 2,3 (2) branches; 9-C with

2-5 (4) branches; 10-C with 3,4 (3) branches; 11-C with 2-6 (5) branches;

12-C with 6-10 (7) branches; 13-C with 5-8 (6) branches; 15-C with 2, 3 (3)

branches; dorsomentum with 29-33 teeth, heavily pigmented. Antenna. Scape

moderately long, index 10.00-11.61, mean 10.80; seta 1-A with 4-6 (5)

branches, inserted 0.54-0.61 from base. Thorax, Seta 0-P with 12-20 (17)

branches; 1,5,6,8,10, 12-P single; 2, 14-P with 2,3 (2) branches; 3-P with

3,4 (4) branches; 4-P with 3,4 (3) branches; 7-P with 2,3 (3) branches; 9-P

single or 2 (2) branched; 11-P with 2-6 (3) branches; 1-M with 4 branches;

2-M with 3 branches; 3,5,7,10, 12-M single; 4-M with 2-5 (4) branches; 6-M

with 5-7 (6) branches; 8-M with 5-9 (6) branches; 9-M with 6-8 (8) branches;

11-M single or 2 (1) branched; 13-M with 22-25 (22) branches; 14-M with

13-20 (18) branches; 1-T single to 4 (4) branched; 2-T with 2-4 (4) branches;

3-T with 8-14 (11) branches; 4-T with 3-5 (5) branches; 5, 10-T single; 6-T

with 2 branches; 7-T with 4-8 (5) branches; 8-T with 7-14 (10) branches; 9-T

with 3,4 (3) branches; 11, 12-T single or 2 (1) branched; 13-T with 12-20 (18)

26 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

branches. Abdomen. Setae 0,4, 14-VIII single; 1-VIII with 5-7 (6) branches;

2-VIII single or 2 (2) branched; 3-VIII with 6-10 (8) branches; 5-VIII with 5-8

(5) branches; comb on VIII with 7-10 (usually 8,9) scales; 1-X single or 2 (2)

branched; 2-X with 7-12 (9) branches; 3-X single; ventral brush with 10 setae

on grid, each with 2-14 branches, some setae always with 8 or more branches,

cephalic setae shorter and with fewer branches. Siphon. Moderately pigmented

with basal 0.4 heavier pigmented; index 3.13-3.91, mean 3.57; pecten on

basal 0.42-0.53, with 12-16 (usually 13, 14) teeth, each tooth with one or 2

small ventral denticles near base, apex long, narrow and pointed, distal 1-3

teeth spine-like, without denticles and wider spaced than remainder; 1-S with

2-5 (2) branches, inserted on basal 0.57-0.60 of siphon distad of last pecten

tooth; 2-S single, short; 6,7,8, 9-S single.

TYPE-DATA. The holotype male, deposited in the BMNH, possesses the

following data on the adult labels: SELANGOR, Rantau Panjang, Mar. 1956

[on upper surface of label]; Ex. Coll. I.M.R. Malaya [on lower surface of

label]; No. 013/1, Larvae in progeny raised in lab., Mother caught attacking

man; Aédes (Aedi.) near ostentatio; Ae. (Pavraédes) collessi Mattingly

HOLOTYPE, P. F. Mattingly 20: viii: 1957; and Type [circular label with a

red border]. The holotype adult is in very poor condition and has only the

following structures remaining on the pin; fragment of the thorax, both wings,

left midleg, and most of the abdomen. The genitalia are mounted in balsam

on a small cover glass on a card attached to the pin and one leg is

mounted on a second small cover glass on a card attached to the pin. Associ-

ated larval and pupal exuviae are mounted in balsam under separate cover

glasses on a microscope slide and are in good condition. The allotype is in

excellent condition and possesses the same collection data as the holotype

except the number 013/2. One paratype female, number 013/3, has the

same data as the holotype. The allotype and paratype each have associated

larval and pupal exuviae mounted in balsam on microscope slides. Another

female specimen has a paratype label and the same data as the holotype

except for the following: 1953, H.B.T. [human bait trap], and J. A. Reid

collector. This specimen was not mentioned by Mattingly (1958a: 28).

DISTRIBUTION. Fifty-five specimens examined: 1 “pl, 1p, 1°, 5 ¥pl

and 34%.

MALAYSIA. Sabah, Tenom, Mandalom Forest Reserve, 9 Apr. 1970,

S-405, -440, S. bin Omar, C. Y. Wang and Ho, 20%; Kudat, 20 Aug. 1966,

459, 22; Lingan Patpar (? spelling), 25 Jun. 1966, 455, 1°; Timbang Kota

Belud (? spelling), 24 Jun. 1966, 460, 1%. Selangor, Kula Selangor, 1968,

S-315, -111, 1%p, 1°; Puchong, May 1956, 0.27, 12; Rantau Panjang, Mar.

1953, 013/1 (holotype), 013/2 (allotype), 013/3 (paratype), 1 “pl and 2 pl;

11 Feb, 1953, 593/1, J. A. Reid, ? Spl; Mar. 1953, 593/2, /3,) 2 Ypl; 1953,

J. A. Reid, 12 paratype and 19; 21 Nov. 1952, J. A. Reid, 12; 19 Mar. 1953,

J. A. Reid, 22; Apr. 1956, 12; Kuala Lumpur, Gombak Rd., 28 Feb. 1904,

G. F. Leicester, 2°.

Distribution from literature.

MALAYSIA. Selangor, Rantau Panjang (Mattingly 1958a: 28; Macdonald

1957: 21; Wharton et al. 1964: 68); Selangor, Sabak Bernam Dist., Torking-

ton Estate (Univ. Calif. Internat. Center Med. Res. Ann. Prog. Rep. 1979:

29); Perak, Parit, Mukim Kampong Gajah (Ann. Rep. Inst. Med. Res. for

1965, Malaysia, 1968: 32); Puchong (Macdonald 1957: 21).

DISCUSSION. Adults of collessi are similar to those of menoni and pagei.

The 3 species, however, can be separated by the following features in the

female: scutal pale scale patterns different (see Fig. 2); collessi with ante-

Reinert: Aedes (Paraedes) in the Oriental Region 27

pronotum with narrow curved reddish-brown scales, postpronotum with 3

setae and narrow curved reddish-brown scales, 0-3 (usually 2,3) anterior

acrostichal setae, 4 posterior medial scutal setae, and with narrow curved

golden scales on interocular space; menoni with antepronotum with narrow

curved reddish-brown scales on inner area and narrow curved golden scales

on outer surface, postpronotum with 4-6 setae and a few narrow curved golden

scales, acrostichal setae absent, 4 posterior medial scutal setae, and with

narrow curved white scales on interocular space; and pagez with antepronotum

with narrow curved reddish-brown scales on inner area and narrow curved

golden scales on outer surface, postpronotum usually with 3 (some specimens

with 4) setae and narrow curved reddish-brown scales, 6 posterior medial

scutal setae, and with narrow curved golden scales on interocular space.

The following combination of features distinguishes females of collessi

from other species of Pavaedes: eyes separated in front; scutal pale scaled

pattern (see Fig. 2); antepronotum with narrow curved reddish-brown scales;

scutellum with narrow curved golden scales on each lobe (some specimens

also with narrow curved reddish-brown scales on median lobe); costa with a

small white scaled patch at base; and femora II and III with a small dorso-

apical patch of white scales.

Males of collessi can be distinguished from those of menoni by the follow-

ing: collessi with pale scales on anterior promontory area; maxillary palpus

0.09 length of proboscis; antepronotum with narrow curved reddish-brown

scales (one specimen also with a few broad brown scales) and 6-8 setae;

postspiracular area with 1,2 setae; prealar knob with 5, 6 setae; mesepimeron

with 6-8 setae dorsad of scale patch; and characters of the genitalia (see

Fig. 6); and menoni possesses: pale scales on anterior promontory, anterior

and lateral scutal fossal areas and scutal angle; maxillary palpus 0.12 length

of proboscis; antepronotum with broad brown scales and 11,12 setae; post-

spiracular area with 3,4 setae; mesepimeron with 9,10 setae dorsad of scale

patch; and characters of the genitalia (see Fig. 7). :

Pupae of collessi are distinguished by: seta 1-CT with 2 branches, 5-CT

with 2 branches, 1-III with 2,3 branches, 6-III single, 1-IV-VI single or 2

branched, paddle index 1.94-2.22, and shape of the male genital lobe (see

Fig. 22).

Aedes collessi larval features of importance in separating this species

from the others of the subgenus are: seta 8-S single, 1-S inserted on basal

0.57-0.60 of siphon, and siphon with basal 0.4 darker pigmented than apical

0.4.

BIONOMICS. Females in Malaysia have been collected in both human and

calf baited traps. Adults were taken during the months of February through

June and August. Larvae were collected from water in a small pool in the

jungle and in a ground pool among nipah palm trees during February and

March.

In Malaysia Wharton et al. (1964: 68) obtained females from a net trap

baited with a man at ground level and from net traps baited with Macaca

monkeys at ground level and on platforms in trees. Mattingly (1958a: 29,

31) reported that females were taken biting man or in human baited traps and

larvae were collected from ground pools and, in one case, in a ground pool

located in nipah palms. In Malaysia collessi was also collected from Berok

baited traps, dissected and found negative for filaria parasites (Ann. Rep.

Inst. Med. Res. for 1963, Fed. Malaya, 1968: 48, 63).

28 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

AEDES (PARAEDES) MENONI MATTINGLY

(Figs. 2, 7, 13, 28)

Aédes (Paraédes) menoni Mattingly 1958b: 77 (¢*, &*); of Qutubuddin 1960: 135.

Aédes (Pavaédes) Travancore sp. of Mattingly 1958a: 24 (keys).

Aedes (Paraedes) menoni of Stone, Knight and Starcke 1959: 200; Stone and

Delfinado 1973: 207; Knight and Stone 1977: 148.

MALE. The following description is of the holotype. Head, Antenna

brown, 1.16 length of proboscis, pedicel golden-prown with a few short fine

setae and a few small brown scales mesally; maxillary palpus 0.12 length of

proboscis; clypeus golden-brown; proboscis 0.94 length of femur I; vertex

covered with broad decumbent dark brown scales except for a few erect forked

golden scales on posterior area, narrow curved golden-white scales on ocular

line, lateral surface with broad dark brown scales and a small patch of broad

white scales on median anterior margin at level with antepronotum; occiput

with a few narrow decumbent white scales and numerous erect forked golden-

brown scales; interocular space with narrow curved golden-white scales, 2

pairs of brown setae; several dark brownish-black ocular setae; eyes slightly

separated anteriorly. Thovax. Scutal integument dark golden-brown; scutum

badly rubbed but remaining scales indicate that entire surface covered with

narrow curved reddish-brown scales except for narrow curved white scales

on the following: 4 on anterior promontory area, a small patch on anterior

and extending onto lateral scutal fossal areas, and a small patch on scutal

angle; scutum with dark brown setae as follows: 4 anterior promontory, numer-

ous dorsocentrals (anterior and posterior), scutal fossal (4 anterior, 2 lateral

and 1 posterior), numerous supraalar, 4 posterior medial scutal; scutellum

with a patch of narrow curved reddish-brown scales and a few narrow curved

white scales on basal area of lateral lobe, median lobe with only 3 narrow

reddish-brown scales remaining, 4 long and 3 short setae on median lobe, 2

long and 2 short setae on lateral lobe; pleural integument golden-brown; ante-

pronotum with broad dark brown scales, 11,12 dark setae; postpronotum with

several narrow curved reddish-brown scales on dorsal area, 3, 4 dark posteri-

or setae; propleuron with a patch of broad silvery-white scales, several

golden-brown setae; paratergite bare; postspiracular area with 3,4 golden-

brown setae; mesepisternum with broad silvery-white scales, a few remaining

on upper area and a small patch on lower area, 3 upper and 9 (?) posterior

brown setae; prealar knob with 7,8 golden-brown setae; mesepimeron with a

patch of broad silvery-white scales on upper area, 9,10 golden setae dorsad

of scale patch. Legs. Coxae I-III with golden-brown setae, I with broad white

scales and a small median patch of broad brown scales on anterior surface, II

and III with a patch of broad white scales on lateral surface; trochanters I-III

with a few pale scales and a few short setae; femora I-III with anterior surface

brown scaled, I-III with posterior surface brown scaled except for pale scaled

areas as follows: I with a narrow dorsal longitudinal stripe from base to apex,

II with a similar stripe on ventral area, and III with a large ventrobasal patch

which narrows apically; posttarsi I-III with 2 ungues, I and II with ungues

unequal in size, both simple, II with ungues equal in size, both simple. Wing.

Dorsal and ventral veins with dark brown scales; costa without a patch of

white scales; alula with several narrow brown scales on margin; no remigial

setae seen; upper calypter apparently without setae. Halter. Pedicel pale;

capitellum brown scaled with a few pale scales at base. Abdomen. Only basal

Reinert: Aedes (Paraedes) in the Oriental Region 29

2 segments remain on specimen, remainder mounted on slide with genitalia.

Tergum II brown scaled with a small laterobasal patch of broad white scales;

sternum II with a few pale brown scales. Genitalia (Fig. 7). Genitalia of

holotype damaged and in poor condition. Tergum IX moderately to heavily

pigmented, bilobed, with 8,9 short thin setae on each lobe, cephalic margin

slightly emarginate, gonocoxite moderately to heavily pigmented, moderately

long, moderately broad, mostly denuded of setae and scales but appears to

have 4 short and 2 longer setae on basomesal area of tergal surface, a few

scattered moderately long setae on median lateral area and a group of long

stout setae on apical area of tergal surface, sternal surface with a patch of

moderately long and long setae on apicomesal area, a few moderately long

and long setae scattered over remainder of sternal surface, several broad

scales on lateral and sternal surfaces, tergal, sternal and lateral surfaces

covered with minute spicules; gonostylus approximately 0.69 length of gono-

coxite, bifid at approximately 0.61 from base, basal area moderately broad,

inner arm narrow with a row of short stout recurved spicules on apicolateral

area and 3,4 minute setae basad of spicules, outer arm curved mesad, short,

narrow but with apical area somewhat broader and covered with short rows of

spicules forming file-like ridges; basal mesal lobe divided at base into 2 long

caudally projecting arms, dorsal arm with basal area narrow and apical por-

tion broader and flat with 4 short flattened setae on apical margin, ventral

arm long, narrow, curved, basal portion covered with short spicules, apex

with 3 long setae; phallosome with aedeagus divided into 2 heavily pigmented

lateral plates which are connected basally, each plate curved tergomesad with

2,3 basomesal teeth, paramere approximately 0.98 length of aedeagus, para-

meral apodeme narrow; sternum IX moderately to heavily pigmented with 2

long stout setae on caudomesal area.

FEMALE. Features differing from male are listed below. Description

based on allotype and 2 paratypes. Head. Antenna 1.31-1.39 length of pro-

boscis; maxillary palpus 0.19-0.21 length of proboscis; proboscis 0.89-0.97

length of femur I; vertex with larger area of narrow curved golden-white

scales, lateral surface with white scaled patch larger; occiput with erect

forked scales more numerous. Thorax (Fig. 2). Scutum covered with narrow

curved reddish-brown scales except for narrow curved golden scales as

follows: a small patch on anterior promontory area, a large moderately broad

patch on anterior and lateral scutal fossal area extending onto scutal angle, a

narrow patch along scutal ridge and onto posterior dorsocentral area, a few

scales forming indistinct lines on posterior dorsocentral and acrostichal areas

(one specimen with a few scales on anterior acrostichal area) and along lateral

margins of prescutellar bare space, and a small patch on supraalar area;

scutellum with narrow curved golden-white scales on all 3 lobes; setal differ-

ences are as follows: 3,4 anterior promontory, scutal fossal (4, 5 anterior

and 2-4 lateral), scutellar (4-6 long and 3-5 short setae on median lobe, 2-4

long and 2-4 short setae on lateral lobe); 8-12 antepronotal, 4-6 postpronotal,

1-4 postspiracular, 2,3 upper and 7-10 posterior mesepisternal, and 6-18

mesepimeral; antepronotum with narrow curved reddish-brown scales on dor-

sal half and narrow curved golden-white scales on ventral half; postpronotum

with a few golden-white scales dorsad of narrow curved reddish-brown scales.

Legs. Femur III with an indistinct basoventral patch of pale scales; posttarsi

I-III (Fig. 28) with 2 ungues equal in size and simple. Wing. Costa with a

small patch of white scales at base; 1,2 remigial setae; upper calypter with

several dark setae on margin. Abdomen. Terga II-VI brown scaled with a

large laterobasal patch of white scales, II-VII also with a narrow dorsobasal

30 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

band of white scales connected with laterobasal patches of white scales, white

scaled band on VII broad; sterna II-VII white scaled, III-VII also with a narrow

apical band of brown scales, brown scaled band on VII covering apical 0. 4.

Genitalia (Fig. 13). Tergum VIII with scales absent, basal 0.8 retracted into

segment VII, base concave mesally, apex straight, numerous short setae

scattered over apical 0.94, setae along apex short and thin, basolateral seta

absent, VIII-Te index 1.12, VII-Te/IX-Te index 2.53, length 0.28 mm, width

0.25 mm; sternum VIII with a large lightly pigmented median area, scales

absent, base slightly concave mesally, apex with a small median indentation

(0.09 of length) and a medium size lobe on each side of midline, short to mod-

erately long setae on apical 0.91, apical intersegmental fold lightly pigmented,

VIlI-S index 0.98, length 0.32 mm, width 0.33 mm; tergum IX with mesal

area lightly pigmented, apex with a small median indentation and with 4-6

setae on each side of midline, 9-11 total setae, IX-Te index 0.97-0.98, length

0.10-0.11 mm, width 0.11 mm; insula with 7 small tuberculi; lower vaginal

lip with numerous long spicules scattered over surface, lower vaginal sclerite

absent; upper vaginal lip with upper vaginal sclerite moderately pigmented,

medium size; postgenital lobe with apex with a moderately deep median inden-

tation (0.22-0.23 of length), 6-9 setae on each side of midline, 13-17 total

setae, dorsal PGL index 0.93-0.94, ventral PGL index 1.66-1.73, ventral

length 0.11-0.13 mm; cercus with 2,3 long setae at apex, dorsal surface with

a number of short to moderately long setae on apical 0.89-0.91, 1-8 scales,

ventral surface with a few short setae on lateral margin, cercus index 3.39-

3.48, cercus/dorsal PGL index 3.52-3.71, cercus length 0.22-0.27 mm; one

large and 2 medium size seminal capsules, base of accessory gland duct heavily

pigmented.

PUPA and LARVA. Not known.

TYPE-DATA. The holotype male, deposited in the BMNH, possesses the

following data on the adult labels: S. India, Travancore, M.O.T. Iyengar;

Pres. by I. M. Puri, XII. 1936 [Mattingly 1958b: 77, erroneously reported

this as Dec. 1935]; Hypopygium on slide; Aédes (Paraédes) menoni Mattingly,

HOLOTYPE, P. F. Mattingly, 31: i: 58, Terminalia on slide; and Type

[circular label with a red border]. The holotype is in poor condition and is

as follows: a minuten pin extends through the mesepisternum of both sides,

notum badly rubbed, right antenna missing, right wing partially broken, tarso-

mere 9 of left foreleg missing, right mid- and hindlegs missing, and abdomen

removed and mounted on a slide. The genitalia are mounted in balsam

on a microscope slide and are badly damaged. The allotype and 2 female

paratypes possess the same label data as the holotype and are in the BMNH.

These 3 female specimens are in good condition with the exception of rubbed

areas on their thoraces.

DISTRIBUTION. Four specimens examined: 1° and 39.

INDIA. Travancore, Dec. 1936, M.O.T. Iyengar, holotype 1°, allotype

1°, paratypes 2°.

Distribution from literature.

INDIA. Travancore, Southwest India (Mattingly 1958b: 77).

DISCUSSION. The habitus of the adults of menoni, especially the scale

pattern of the scutum, set this species apart from those of other species of

the subgenus as shown in Figure 2.

Female genitalia of menoni differ from all other species of the subgenus

by the presence of 3 well developed seminal capsules while the others each

have a single large one. Other differences of menoni are sternum VIII index

0.98, tergum VIII index 1.12, cercus index 3.39-3.48, tergum IX index 0.97-

Reinert: Aedes (Pavaedes) in the Oriental Region 31

0.98, and cercus/dorsal PGL index 2.52-3.71. Even though the 3 females

(allotype and 2 paratypes) possess the same collection data as the holotype

male and share a number of habitus features, the differences in the female

genitalia, especially the presence of 3 seminal capsules, suggest that positive

association of the sexes of this species needs to be confirmed by progeny

reared specimens, both male and female, with associated immature exuviae.

The male genitalia, even though the single specimen is badly damaged,

are distinct and easily distinguished by the development of the basal mesal lobe

and gonostylus from those of the other species of the subgenus (see Fig. 7).

BIONOMICS. Nothing is known about the bionomics or medical importance

of this species.

AEDES (PARAEDES) OSTENTATIO (LEICESTER)

(Figs. 2, 8, 14, 20, 22, 26-28)

Aioretomyia Ostentatio Leicester 1908: 193 (¢).

Aioretomyia ostentatio of Brunetti 1912: 490.

Ochlerotatus ostentatio of Edwards 1913: 228 (in part); Brunetti 1920: 140

(in part); Senior-White 1923: 81 (27 part).

Aedes ostentatio of Edwards 1922a: 263 (key); Stojanovich and Scott 1965: 20

(2*, key), 1966: 110 (2*, key).

Aedes (?) ostentatio of Edwards 1922b: 468 (in part); Brug and Bonne-Wepster

1947: 185 (in pari).

Aédes (Aédes) ostentatio of Edwards 1928: 53, 1929: 3.

Aedes (? Aedimorphus) ostentatio of Edwards 1932: 171 (in part); Barraud 1934:

267 @, in part); Macdonald 1957: 21 (in part).

Aédes ostentatio of Brug and Edwards 1931: 259.

Aedes (Aedimorphus) ostentatio of Bohart 1945: 52, 63 (2, key, in part); Knight

and Hull 1953: 463 (2, iz part); Horsfall 1955: 518 (in part).

Aédes (Paraédes) ostentatio of Mattingly 1958a: 7, 9, 22, 24, 25 (2*, key,

in part), 1958c: 106 (lectotype designation).

Aedes (Paraedes) ostentatio of Stone, Knight and Starcke 1959: 200 (in part);

Army Mosquito Project 1965: 29 (key); Stone et al. 1966: 50 (key);

Basio 1971: 26 (in part); Stone and Delfinado 1973: 307 (in part); Knight

and Stone 1977: 148 (” part).

MALE. Head, Antenna dark brown, 1.17-1.21 length of proboscis, pedi-

cel dark brown with a few short fine brown setae and small dark scales mesal-

ly; maxillary palpus 0.09-0.10 length of proboscis; clypeus dark brown; pro-

boscis 0.87-0.97 length of femur I; vertex covered with broad decumbent

dark brown scales, a few narrow curved golden-white scales on interocular

space, ocular line with narrow curved golden scales; lateral surface covered

with broad dark brown scales and with a patch of broad white scales anterior

to antepronotum and posterior to eye margin; occiput with a few short erect

forked brown scales and a few broad decumbent white scales; interocular and

ocular setae dark brown and well developed; eyes slightly separated anteriorly.

Thorax. Scutal integument dark brown; scutum covered with narrow curved

reddish-black scales except for narrow curved golden scales on anterior

promontory area and anterior scutal fossal area; scutum with reddish-black

setae as follows: 2-4 anterior promontory, numerous dorsocentral (anterior

and posterior), scutal fossal (3,4 anterior, 2,3 lateral and 1,2 posterior),

32 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

numerous supraalar, 4 short posterior medial scutal; scutellum with a small

patch of narrow curved reddish-brown scales on each lobe, 5-8 setae on medi-

an lobe, 3-5 setae on each lateral lobe; pleural integument dark brown; ante-

pronotum covered with broad dark brown scales, 7-9 reddish-black setae;

postpronotum with narrow curved reddish-black scales along dorsal margin

and a small patch of 4-8 broad dark brown scales on posterior area anterior

of setae, 3,4 reddish-black posterior setae; propleuron covered with broad

white scales, 5,6 setae; paratergite bare; postspiracular area with 2,3 broad

white scales (Some specimens with scales rubbed off), 3,4 dark brown setae;

mesepisternum with a large patch of broad white scales on upper area anda

small patch of similar scales on lower area, 2,3 setae on upper area and 7-9

setae on posterior area, lower setae short and pale; prealar knob with 6-8

dark brown setae; mesepimeron with a large patch of broad white scales, 8-11

short golden setae dorsad of scales. Legs. Coxae I-III with several dark

brown setae, I with anterior surface with broad white scales and a small

anterior median patch of broad brown scales, II and III with a small anterior

patch of broad white scales; trochanters I-III with a few broad pale scales

and a few short golden setae; femur I with anterior and posterior surfaces

covered with dark brown scales, I also with a posteriodorsal pale stripe from

base to near apex, femur II with anterior and posterior surfaces dark brown

scaled and a posteroventral longitudinal dusty-white scaled stripe on basal 0.7,

stripe broader proximally, femur III dark brown scaled with an anteroventral

and a posteroventral longitudinal dusty-white scaled stripe on basal 0. 70-0. 85,

stripe broader proximally; posttarsi I-III (Fig. 27) with 2 ungues, I and II with

unques equal in size and with a tooth, III with ungues equal in size, both simple.

Wing. Dorsal and ventral veins dark brown scaled; costa without a patch of

white scales; alula with narrow brown scales along margin; 1,2 short brown

remigial setae; upper calypter bare. Halier. Pedicel pale; capitellum dark

brown scaled. Abdomen. Terga dark brown scaled, II-VII with a small

laterobasal patch of white scales; sterna white scaled, III-VII with a narrow

apical band of brown scales. Genitalia (Fig. 8). Tergum IX strongly bilobed

with 2,3 short setae on each lobe; gonocoxite with an apical sternomesal lobe,

tergal surface with 3-5 long stout and 9-12 moderately long setae and one or 2

irregular rows of setae on basal 0.65 of tergomesal margin, setae short dis-

tally and increase in length proximally until they become moderately long at

base, lateral surface with a few long and moderately long setae, sternal sur-

face with a number of long stout and a few short setae on apical 0.3 and a sub-

apical elongate patch of approximately 20-25 moderately long and a few short

setae on mesal margin, lateral surface and lateral areas of tergal and sternal

surfaces with scales; gonostylus with pedicel narrow and distal portion bifid

and divided into a narrow club-shaped tergal arm bearing 12-15 short setae

and a file-like area of spicules, and a flattened rectangular shaped sternal

arm with small hair-like spicules on distal 0.55; basal mesal lobe divided at

base into 2 long arms, mesal arm with distal 0.5 narrower, curved tergally

and with 3 moderately long hook-like setae at apex, proximal 0.5 somewhat

flattened, expanded distally, and covered with short hair-like spicules, outer

arm long, middle of length somewhat swollen, distal 0.25 narrower and

covered with short to long simple spines, similar short spines near middle of

length, becoming more numerous distally, basal 0.3 cylindrical and with a

short finger-like projection near base which bears 2,3 short setae, one api-

cally and one or 2 subapically; phallosome with aedeagus divided into 2 heavily

pigmented lateral plates connected basally, each plate with one or 2 stout

teeth, apical 0.3 with a lightly pigmented sternal connection, distal portion of

Reinert: Aedes (Paraedes) in the Oriental Region 33

each plate bluntly rounded with apex pointed and projected mesally, paramere

approximately 0.88 length of lateral plate, parameral apodeme moderately

long and narrow; sternum IX large, with 2,3 long stout setae near center.

FEMALE. The female differs markedly from the male in the scale pat-

terns of the head, thorax and legs and the number of thoracic setae. The fol-

lowing features differ from the male. Head. Antenna 1.27-1.35 length of

proboscis; maxillary palpus 0.18-0.19 length of proboscis; proboscis 0.91-

0.99 length of femur I; vertex covered with broad decumbent dark blackish-

brown scales and a median triangular patch of narrow curved golden scales,

numerous erect forked golden scales scattered throughout golden scale patch

from occiput to interocular setae; interocular setae golden, well developed.

Thovax (Fig. 2). Scutum covered with narrow curved reddish-black scales,

narrow curved golden scales on following: anterior promontory area, extending

from anterior scutal fossal area over lateral scutal fossal area to scutal

angle, a few on scutal ridge, a broad stripe on acrostichal area extending from

anterior promontory area over posterior medial scutal area and along lateral

margins of prescutellar bare space, a more or less distinct narrow stripe on

anterior dorsocentral area to approximately scutal ridge, and a small patch

on supraalar area cephalad of wing base; scutellum with curved golden scales

on all lobes; setal differences are as follows: 3-5 anterior scutal fossal, 2-4

lateral scutal fossal, 6 posterior medial scutal, scutellum with 6-9 on median

lobe and 4-7 on lateral lobe, 8-11 antepronotal, 3-5 postpronotal, 6-8 pro-

pleural, 2-5 postspiracular, 1-3 upper and 7-14 posterior mesepisternal,

7-10 on prealar knob, and 10-13 mesepimeral; postspiracular area with a

patch of broad white scales. Legs. Femora I-III with a small dorsoapical

patch of white scales; posttarsi I-III (Fig. 28) with 2 ungues, I and II with

ungues equal in size and with a tooth, UI with ungues equal in size, both

simple. Wing. Upper calypter with dark setae onmargin. Halter. Capitel-

lum with dark scales at base and white scales apically. Abdomen. Tergum

VIII with a laterobasal pale scaled patch. Genitalia (Fig. 14). Tergum VIII

with scales absent, basal 0. 85-0.95 retracted into segment VII, base concave

mesally, apex straight, numerous short setae scattered over apical 0.68-0.77,

setae along apex short and thin, basolateral seta absent, VII-Te index 0. 68-

0.79, VIlI-Te/IX-Te index 2.02-2.30, length 0.19-0.22 mm, width 0. 25-0. 30

mm; sternum VIII with scales absent, base concave mesally, apex with a

moderately deep median indentation (0.13-0.16 of length) and a small lobe on

each side of midline, short to moderately long setae on apical 0. 82-0. 95,

seta 2-S appreximately 0.38 from 1-S, seta 3-S approximately 0.62 from 2-S,

apical intersegmental fold lightly pigmented, VIII-S index 0.59-0.65, length

0.20-0.22 mm, width 0.35-0.40 mm; tergum IX with apex with a small

median indentation and with 2-5 (usually 3, 4) short thin setae on each side of

midline, 5-9 total setae, IX-Te index 0.67-0.75, length 0.09-0.10 mm,

width 0.12-0.14 mm; insula with 4-6 small tuberculi; lower vaginal lip with

numerous minute to short spicules scattered over entire surface, lower vag-

inal sclerite absent; upper vaginal lip with upper vaginal sclerite moderately

to heavily pigmented, large size; postgenital lobe with a very small to moder-

ately deep median indentation (0.03-0.25 of length), 7-10 (usually 7, 8) setae

on each side of midline, 14-17 total setae, dorsal PGL index 0.94-1.03, ven-

tral PGL index 1.43-1.61, ventral length 0.10-0.12 mm; cercus with 2 long

stout setae at apex, dorsal surface with a number of short to moderately long

setae on apical 0.73-0.79, 0-4 broad scales (usually 1,2 scales), ventral sur-

face with a few short setae on lateral margin, cercus index 2.20-2.58, cercus/

dorsal PGL index 2.41-2.68, cercus length 0.17-0.19 mm; one large seminal

34 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

capsule, accessory gland duct moderately to heavily pigmented.

_ PUPA (Figs. 20, 22). Chaetotaxy as figured and recorded (Table 4).

Cephalothorax. Seta 1-CT with 2-7 branched; 5-CT with 3-8 branches.

Respiratory trumpet. Index 4.94-5.73, mean 5.28. Abdomen. Seta 4-I

usually 7-10(6-10) branched; 5-I usually 4, 5(3-6) branched; 1-II with 8-19

branches; 2-II inserted mesad of 3-II; 10-III usually single or 2(1-3) branched;

7-IV with 2-4 branches; 10-IV usually 2 (2-4) branched; 1-V with 2-5 branches;

6-VI single. Paddle. With minute spicules on apical 0.42-0. 54 of lateral and

apical 0.12-0.18 of mesal margins; index 1.42-1.76, mean 1.56.

LARVA (Fig. 26). Description based on 4 larval exuviae with associated

adults. Head. Moderately pigmented; setae 0,1,3,14,18, 20-C single; 4-C

with 8-12 (8) branches, short; 5-C with 6-8 (7) branches, very long; 6-C with

5,6 (5) branches, very long; 7-C with 10-14 (13) branches, long; 8-C with 2,3

(2) branches; 9-C with 3-5 (4) branches; 10-C with 2-4 (2) branches; 11-C

with 4-6 (5) branches; 12-C with 6-8 (8) branches; 13-C with 5-8 (6) branches;

15-C with 3-5 (4) branches; dorsomentum with 30-32 (30) teeth, heavily pig-

mented. Antenna. Scape long, index 11.75-12.50, mean 12.08; seta 1-A with

4,5 (4) branches, inserted 0.51-0.53 from base. Thovax. Seta 0-P with 9-16

(11) branches; 1,5,6,10, 12-P single; 2, 9-P single to 3 (2) branched; 3-P

with 3-5 (4) branches; 4-P with 2-4 (3) branches; 7-P with 3, 4 (3) branches;

8-P single or 2 (1) branched; 11-P with 2-5 (4) branches; 12-P with 2,3 (2)

branches; 1-M with 2-5 (4) branches; 2-M with 3-5 (4) branches; 3, 5, 7, 10,

12-M single; 4-M with 2-6 (5) branches; 6-M with 5,6 (6) branches; 8-M with

7-9 (7) branches; 9-M with 6-8 (8) branches; 11-M single or 2 (1) branched;

13-M with 19-26 (24) branches; 14-M with 11-16 (16) branches; 1-T single to

3 (2) branched; 2-T with 4-6 (4) branches; 3-T with 10-14 (12) branches;

4-T with 4-6 (5) branches; 5,10, 11-T single; 6-T with 2,3 (2) branches; 7-T

with 9-12 (10) branches; 8-T with 7-11 (11) branches; 9-T with 3, 4 (3) branches;

12-T single or 2 (1) branched; 13-T with 11-20 (15) branches. Abdomen.

Setae 0,2, 13-I single; 1-I with 4-9 (7) branches; 3-I with 3-7 (5) branches;

4-I with 17-20 (19) branches; 5-I with 4-8 (6) branches; 6, 9-I with 2,3 (2)

branches; 7, 11-I single or 2 (2) branched; 9, 10-I with 2,3 (2) branches;

0,2,9,10, 14-II single; 1-II with 2-4 (3) branches; 3-II with 5,6 (5) branches;

4-II with 10-15 (11) branches; 5-II single to 4 (2) branched; 6-II with 2,3 (2)

branches; 7-II with 3-7 (6) branches; 8-II with 2-4 (3) branches; 11-II with 3, 4

(4) branches; 12-II with 3,4 (3) branches; 13-II with 12-20 (16) branches; 0, 2,

6,9, 14-III single; 1-III with 6-8 (8) branches; 3-III with 3-5 (4) branches;

4-III with 2-4 (3) branches; 5,10, 11-III with 2,3 (2) branches; 7-III with 8-12

(9) branches; 8-III single or 2 (2) branched; 12-III with 2-4 (4) branches; 13-III

with 4-7 (5) branches; 0,2,6,9, 14-IV single; 1-IV with 5-7 (6) branches;

3-IV with 4,5 (4) branches; 4, 12-IV with 2,3 (2) branches; 5, 8-IV with 2,3

(2) branches; 7-IV with 8-10 (10) branches; 10-IV single to 3 (2) branched;

11-IV with 2,3 (3) branches; 13-IV with 4 branches; 0,2,6,9, 14-V single;

1-V with 4-6 (6) branches; 3-V with 3 branches; 4-V with 7-10 (10) branches;

5-V with 2-5 (3) branches; 7-V with 6-9 (6) branches; 8, 11-V with 2,3 (3)

branches; 10, 12-V with 2,3 (2) branches; 13-V with 4 branches; 0, 2, 6,9, 10,

14-VI single; 1-VI with 5-7 (6) branches; 3, 7-VI with 3-5 (3) branches; 4,

8-VI with 4, 5 (4) branches; 5-VI with 2-5 (5) branches; 11-VI with 2-4 (2)

branches; 12-VI with 2,3 (3) branches; 13-VI with 10-26 (18) branches; 0,

14-VII single; 1-VII with 6-8 branches; 2-VII single or 2 (1) branched; 3-VII

with 6,7 (6) branches; 4-VII with 3, 4 (3) branches; 5-VII with 4-8 branches;

6-VII with 15-17 (17) branches; 7-VII with 2,3 (2) branches; 8-VII with 10-13

branches; 9-VII with 2-4 (4) branches; 10-VII with 2,3 (3) branches; 11-VII

Reinert: Aedes (Pavaedes) in the Oriental Region 39

with 3 branches; 12-VII with 2,3 branches; 13-VII with 7-10 branches; 0, 4,

14-VIII single; 1-VIII with 4-6 (4) branches; 2-VIII with 2,3 (2) branches;

3-VIII with 7-9 (8) branches; 5-VIII with 7-10 (7) branches; comb on VIII with

9-12 (usually 9-11) scales; 1-X with 2,3 (2) branches; 2-X with 10-13 (13)

branches, moderately long; 3-X single, long; 4-X with 9,10 setae on grid, each

with 5-20 branches and one short precratal seta. Siphon. Heavily pigmented,

apical 0.4 very heavily pigmented, dark brownish-black colored; index 3.33-

3.56, mean 3.43; pecten on basal 0.40-0.45, with 13-16 (usually 13, 14) teeth,

each tooth with one or 2 small ventral denticles near base, apex long, narrow

and pointed, distal 1-3 teeth spine-like, without denticles and wider spaced

than remainder; 1-S with 2-4 (4) branches, inserted on basal 0.61-0.63 of

siphon and distad of last pecten tooth; 2-S single, short; 6,7, 9-S single;

8-S with 2,3 (2) branches.

TYPE-DATA. Leicester (1908: 194) stated that Aioretomyia Ostentatio

was described from a series of females hatched from larvae found in a small

jungle pool and some collected as adults. Mattingly (1958c: 106) reported that

Leicester's type-series is now represented by only 2 female cotypes which

are deposited in the BMNH. He designated one as lectotype and it contains the

following data: Aédes (Paraédes) ostentatio Leicester, HOLOLECTOTYPE,

P. F. Mattingly 1958; Patch of jungle, 4 1/2 miles, Pahang Rd., Kuala

Lumpur, 21/9/03; Cotype; Kuala Lumpur, Fed. Malay States, Dr. G. F.

Leicester, 1912-350; and Azovetomyia ostentatio Leic. The other female,

which is incomplete and in poor condition, was designated as a paratype by

Mattingly (1958c: 106) and bears the following data: Jungle, 5th mile, Gombak

Rd., 5/2/04; and other data as in lectotype.

DISTRIBUTION. Ninety-nine specimens examined: 2 “pl, 2%, 7 2pl, 5 2p,

592 and 1 L.

INDONESIA. Celebes, Paloe, 2 Aug. 1935, 35.825, 12. Java, Tasikmala-

ja, Padaherang, 14.339, 3°. Sumatra, vicinity of Bengkula, 2 Jan. 1958,

C. Soegiarto, 12; same except 30 June 1958, 12; same except 7 July 1958, 2°.

MALAYSIA. Pahang, Kg. Lamir, Pakan, 5 Apr. 1954, R. H. Wharton,

12. Sabah, Tenom, 9 Apr. 1970, S-440, S. bin Omar, C. Y. Wang and Ho,

22; North Borneo, Kalabakan, 1962, Bishop, 12. Savawak, Kuching, 3rd mile

Rook Rd., 27 Jul. 1914, J. C. Moulton, 12; Tarakan, 9 Jun. 1945, A. G.

Humes, 12. Selangor, Puchong Rd., May 1956, 0.27, 12; Puchong Rd., 14

miles, 3: Jun: 1959, 01230/1,°72, 7/8; °/9; 2:¢pl-and 2 Sol

REPUBLIC OF THE PHILIPPINES. Luzon Island, Mountain, Calaccad,

Stations 13 and 15, 25 Jun. and 9 Jul. 1963, L. E. Rozeboom and B. D.

Cabrera, 22. Palawan Island, Quezon, approximately 10 miles south of

Quezon, 1964, No. 39, L. E. Rozeboom and B. D. Cabrera, 12; same except

No. 64, 148.

THAILAND. Chon Buri, Amphoe Bang La Mung, Khao Mai Keo, 7-8 Oct.

1963, CL-1, CL 14-19, K. Mongkolpanya and team, 1 2pl and 22; same except

21, 22, 27 Aug. 1963, U-614, -616, -617, -618, -631, Udaya Sandhinand and

team, 21%; Khao Mai Ha Wa, 1 Jul. 1965, 00247-12, -100, -105, E. L. Peyton

and team, 1 fpl and 2 $p; same except 30 Jun. 1965, 00242-101, 1 2p, 00248-

10, 1 Spl. Khon Kaen, Amphoe Chumphae, Pha Dong Larn, 30 Nov. 1962,

T-1857, S. Esah, 12; same except Tham Pho Thi Yan, 16 Nov. 1962, T-1853,

-1854, 22. Nakhon Ratchasima, between Kabinburi and Pak Tong Chi, 24

May 1967, 02029, -100, K. Mongkolpanya, 1 fpland1L. Nakhon Sawan, Ban

Kaeng, 9 Nov. 1968, 03128, K. Mongkolpanya and team, 1°. Pyvachin Buvi,

Ban Tub Lan, 20 Jul. 1971, P3-103, K. Mongkolpanya and team, 1 2p; same

except 06031-100, 1%; same except 11 Aug. 1971, P 14, P 14-1, -100, 1¢,

36 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

1 Ppl and 1 Qp.

Distribution from literature.

INDONESIA. Sumatra, Moeara Tebo (Haga 1924: 831), Sumatra, Benkoelem,

Air Prioekan (Brug and Edwards 1931: 259), Dermajoe, Java, Tasikmalaja,

Padaherang (Mattingly 1958a: 26, questions the records from Ceram and

Indochina); Ceram, Java, Sumatra (Brug and Bonne-Wepster 1947: 185).

MALAYSIA (Edwards 1928: 53; Bohart 1945: 63; Brug and Bonne-Wepster

1947: 185); Sarawak (Moulton 1914: 47); Kuala Lumpur, Phang Rd. 4.5 miles

(Mattingly 1958c: 106); Pahang, near Kuantan (Macdonald 1957: 21); Tregganu,

Ulu Trengganu (Ramachandran et al. 1970: 511); Sarawak, Kampong Pangkalan

Kuap, Rajah Charles Brooke Memorial Hospital, 18 km south of Kuching on

Penrissen Rd. (Macdonald, Smith and Webb 1965: 338); Tarakan, Sarawak,

Rantau Panjang, Selangor (Mattingly 1958a: 26); Selangor, Rantau Panjang

(Ann. Rep. Inst. Med. Res. for 1956, 1968, Fed. Malaya, 1957: 107, 1968:

63); Telok Forest Reserve (Unif. Calif. Internat. Center Med. Res. Ann.

Prog. Rep. for 1973, 1973: 24); Sintok, Pacific Tin (Ann. Rep. Inst. Med.

Res. for 1964, Malaysia, 1968: 27, 31); Perak, Parit, Mukim Kampong Gajah

(Ann. Rep. Inst. Med. Res. for 1965, Malaysia, 1968: 32).

REPUBLIC OF THE PHILIPPINES (Edwards 1929: 3; Bohart 1945: 63;

Brug and Bonne-Wepster 1947: 185; Delfinado, Viado and Coronel 1962

(1963): 440); Palawan, Bacungan, Mindoro, San Jose (Knight and Hull 1953:

464); Palawan, Quezon Province, about 10 miles south of Quezon (Rozeboom

and Cabrera 1965: 208); Luzon, Mt. Makiling (Basio, White and Reisen 1970:

442); Laguna, Los Banos (Basio 1971: 26). |

THAILAND. Nakhon Phanom, Takhli, U-Tapao (Parrish 1968b: 7, 10, 13),

Ubon, Udorn (Reisen, Burns and Basio 1971, tables 4-8, 12,13; Biery and

Burns 1973: 20, 24, 27, 30, 33); Udorn (Biery and Burns 1972: 18); Prachinburi,

Bu Phram Valley (Wilkinson et al. 1978: 669).

VIETNAM (Brug and Bonne-Wepster 1947: 185); Pleiku, Phu Cat (Parrish

1968a: 18, 20; 1969: 554); Cam Ranh Bay, Phu Cat (Reisen, Burns and Basio

1971, tables 12,13; Biery and Basio 1973: 7,15).

DISCUSSION. Adults of osteniatio are similar in habitus to those of

chrysoscuta, therefore, for a comparison of the adults of these 2 species see

the discussion section of the latter species. The pale scaled patterns of the

head and scutum of the sexes are dimorphic for ostentatio. The patterns also

display variation between specimens from the same collection. Aedes ostenta-

ito adults are easily identified by the antepronotum which has both broad dark

and pale scales, the scutellum which has narrow curved golden scales on all

3 lobes, and the paratergite which is without scales.

Male genitalia of ostentatio are similar to those of chrysoscuta and are

discussed under the latter species. The development of the basal mesal lobe

is somewhat variable in the shape and number of spines on the one arm,

however, the spines are simple and the arm is swollen.

The female genitalia of ostentatio are most similar to those of chrysoscuta

but can be separated by several features (see discussion section for chryso-

scuta).

Aedes ostentatio pupae have the genital lobe shaped similarly to those of

chrysoscuta and thailandensis (see Figs. 18, 22). The length of seta 5-IV, V,

VI is less than 1.50 length of the attached segment which is similar to bonneae,

but differs from collessi in which it is less than 1.25 and from thailandensis in

which it is greater than 1.50. Aedes ostentatio pupae are very similar to

those of chrysoscuta, however, they can usually be separated by the characters

mentioned in the key.

Reinert: Aedes (Pavaedes) in the Oriental Region 37

Larvae of ostentatio are also similar to those of chrysoscuta and a com-

parison is given under the discussion section of the latter. The above descrip-

tion of the larva of osteniatio is taken from Malaysian specimens since it is

the type-locality. Specimens from Thailand share most characters with those

from Malaysia, however, several features differ between the 2 populations

and these follow (Malaysia listed first and Thailand in parentheses); seta 7-T

with 8-12 (7,8) branches; 3-VIII with 7-9 (3-5) branches; 2-X with 10-13

(6-8) branches; and siphon index 3.33-3.56 (2.30-2.91). Unfortunately, very

limited material was available from both these locations.

Larvae of osteniatio are easily separated from collessi and thailandensis

by the apical 0.4 of the siphon being very heavily pigmented. Aedes ostentatio

larvae are also distinguished from those of bDonneae by the siphon pigmentation

which is very heavy and brownish-black in ostentatio but paler and brown in

bonnedae.

BIONOMICS. In Malaysia females were taken once biting man in partial

shade of a secondary rain forest in hilly terrain. In Thailand adults have been

collected as follows: resting in a bamboo grove at the margin of a stream

(once); biting man in a plantation near a village in partial shade from 0600-

0900 h (once); biting man in sparse vegetation of a forest (once); biting man

outside from 1100-1200 h (twice) and 2200-2300 h (once); and during the day-

time (twice).

In Thailand larvae were taken twice from fresh water ina large footprint

of an elephant in partial shade of secondary scrub in a valley at 150 m eleva-

tion; from small crab holes in heavy shade of a secondary rain forest (twice)

and a tapioca plantation (once) in mountain terrain and at 150 m elevation; and

once from a small deep hole near a stream. Adults have been taken during

the months of May through August, October and November while immatures

were collected in May through August and October.

Leicester (1908: 194) stated the adults of osteniatio are sylvan and vicious

day biters in Malaysia. Also in Malaysia, Macdonald (1957: 21) reports the

adults occasionally were taken on human bait and Macdonald, Smith and Webb

(1965: 342) record this species taken biting in small numbers by day or in

the early evening. Ramachandran et al. (1970: 511) indicate ostentatio adults

made up 5 percent of all Aedes collected during 19 nights of human bait trap

and human bare leg catches in Tregganu, Malaysia, and when they were

examined for Brugia malayi they were negative. In Malaysia, ostentatio adults

were collected from Kera and human baited traps (Ann. Rep. Inst. Med. Res.

for 1963, Fed. Malaya, 1968: 53) and from traps baited with either monkeys

or humans, examined and found negative for filaria parasites (Ann. Rep.

Inst. Med. Res. for 1965, Malaysia, 1968: 23). In Selangor, Malaysia,

ostentatio was reported to be the sixth most prevalent mosquito species that

attacked man in nipah palm mangrove swamps (Ann. Rep. Inst. Med. Res. for

1956, Fed. Malaya, 1957: 107).

In the Philippine Islands adults were: netted ina densely shaded area

near a river (Knight and Hull 1953: 464); collected from undergrowth nearby

a larval breeding place and were a vicious daytime biter (Basio, White and

Reisen 1970: 443). Rozeboom and Cabrera (1965: 208) reported that of 21

ostentatio collected in the Philippine Islands from human baited traps in

a swamp-forest environment, 4 were dissected and found negative for Brugia

malayi parasites.

In Vietnam adults were collected in light traps from May through August

(Reisen, Burns and Basio 1971: tables 12, 13; Biery and Basio 1973: 7, 15).

Stojanovich and Scott (1965: 32; 1966: 136) report this species to be a vicious

38 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

daytime biter of man in densely shaded areas near rivers in this country.

This species has been taken in New Jersey light traps in Thailand during

the months of March, April, June, July and October (Reisen, Burns and

Basio: tables 4-8; Biery and Burns 1972: 18; Biery and Basio 1973: 7, 15).

Wilkinson et al. (1978: 669) in Thailand collected moist soil samples from

the bottoms and margins of former pools and flooded these in the laboratory;

larvae of Ae. jostentatio were produced from these samples.

AEDES (PARAEDES) PAGEI (LUDLOW)

(Figs. 2, 15, 29)

Danielsia paget Ludlow 1911: 128 (9); of Stone and Knight 1956: 233.

Aedes (?) ostentatio of Edwards 1922b: 468 (in pari).

Aédes (Aédes) ostentatio of Dyar and Shannon 1925: 78.

Aedes (Aedimorphus) ostentatio of Bohart 1945: 63. (in part); Knight and Hull

1953: 463 (in part).

Aédes (Paraédes) pagei of Mattingly 1958a: 21, 24, 31 (¢*, key).

Aedes (Paraedes) pagei of Stone, Knight and Starcke 1959: 200; Stone and

Delfinado 1973: 307; Knight and Stone 1977: 148.

MALE. Not known.

FEMALE. Head. Antenna dark brown, 1.18-1.34 length of proboscis,

pedicel golden-brown with a few small broad brown scales and several short

fine brown setae mesally, flagellomere 1 with a few small broad brown scales

on basal 0.5; maxillary palpus 0.17-0.20 length of proboscis; clypeus dark

brown; proboscis 0.97-1.05 length of femur I; vertex covered with broad de-

cumbent dark brown scales and a triangular patch of narrow curved golden

scales on median posterior area (size of patch small to moderate), erect

forked golden and a few brown scales in triangular patch; ocular line and

interocular space with narrow curved golden scales; lateral surface covered

with broad dark brown scales and with a small patch of broad white scales

anterior to antepronotum and posterior to eye margin; occiput with numerous

erect forked dark brown scales (Some specimens also with some golden

scales on median area); interocular and ocular setae dark brown and well

developed; eyes separated anteriorly. Thorax (Fig. 2). Scutal integument

dark reddish-brown; scutum covered with narrow curved dark reddish-brown

scales and narrow curved golden scales forming patterns (golden scaled pat-

terns variable) as follows: a patch on anterior promontory area, a narrow

stripe on acrostichal area from anterior promontory area to prescutellar

bare space at which it forks to form a narrow line along lateral margins,

stripe broader on posterior median scutal area, a small patch on anterior

scutal fossal area and similar scales extending laterally over lateral scutal

fossal area to scutal angle and mesad along scutal ridge, scales forming an

indistinct narrow stripe on dorsocentral area (usually a short patch on anteri-

or area and another connected to posterior portion of scutal ridge), small to

medium size patch on supraalar area; scutum with dark setae on following

areas: 2,3 anterior promontory, numerous dorsocentral (anterior and posteri-

or), scutal fossal (2-4 anterior, 1-3 lateral and 0-1 posterior), numerous

supraalar, 6 posterior medial scutal; scutellum with a patch of narrow curved

scales on each lobe, median lobe with golden scales on posterior and lateral

areas and reddish-brown scales (absent to numerous) on median basal area,

lateral lobe with golden scales (several specimens also with a few reddish-

Reinert: Aedes (Pavaedes) in the Oriental Region 39

brown scales); 5-7 setae on median lobe and 4, 5 setae on lateral lobe; pleural

integument dark reddish-brown; antepronotum with a patch of narrow curved

scales, dark reddish-brown scales on inner area and golden scales on outer

area, 7-12 dark setae; postpronotum with narrow curved reddish-brown scales

on dorsal and posterior areas, 3,4 dark posterior setae; propleuron with a

patch of broad white scales, 4-6 setae; paratergite with a patch of narrow

curved golden scales; postspiracular area with a small patch of moderately

broad or broad white scales posterior to setae, 2,3 dark setae; mesepisternum

with a large patch of broad white scales on upper area and a small patch of

Similar scales on lower area, 1-3 dark setae on upper area and 8-13 setae on

posterior area, lower setae short and pale; prealar knob with 4-7 dark setae;

mesepimeron with a large patch of broad white scales, 7-11 short setae dorsad

of scales. Legs. Coxae I-III with several setae, I with anterior surface

covered with broad scales, brown scales on lower 0.6 and white scales on dor-

sal area, II and III with a patch of white scales on anterior area; trochanters

I-III with broad brown and white scales; femora I and II with anterior surface

dark brown scaled, II with anterior surface brown scaled with a broad pale

scaled stripe on ventrobasal 0.65, I-III with posterior surface brown scaled

and with a broad white scaled stripe from base to near apex, stripe dorsal on

I and ventral on II and III, II and III also with a small apical white scaled patch

on anterior and posterior surfaces, I occasionally with similar pale scaled

patches; posttarsi I-III (Fig. 28) with 2 ungues equal in size and with a tooth.

Wing. Dorsal and ventral veins dark brown scaled; costa without a patch of

white scales; alula with narrow.dark brown scales on margin; one dark remigial

seta; upper calypter with a number of narrow dark setae onmargin. dHalter.

Pedicel pale; capitellum dark brown scaled. Abdomen. Terga I-VII dark

brown scaled and with a large patch of white scales on laterobasal area; sterna

II-VII with basal area white scaled and posterior area brown scaled. Genitalia

(Fig. 15). Tergum VIII with scales absent, basal 0.70-0.85 retracted into seg-

ment VII, base concave mesally, apex straight, numerous short setae scattered

over apical 0.70-0.81, setae along apex short and thin with the exception of 2,

3 moderately long thin ones, basolateral seta minute, VIII-Te index 0. 63-0. 73,

VIII-Te/IX-Te index 2.22-2.49, length 0.19-0.20 mm, width 0.26-0.29 mm;

sternum VIII with 0-4 broad scales (usually absent), base concave mesally,

apex with a moderately deep median indentation (0.11-0.14 of length) and with

a small lobe on each side of midline, short to moderately long setae on apical

0.82-0.92, seta 2-S approximately 0.48 from 1-S, seta 3-S approximately 0.52

from 2-S, apical intersegmental fold lightly pigmented, VIII-S index 0. 61-0. 71,

length 0.21-0.22 mm, width 0.30-0.34 mm; tergum IX with apex with a moder-

ately deep median indentation and with 3-5 (usually 4) thin setae on each side of

midline, 7-9 total setae, IX-Te index 0.67-0.82, length 0.08 mm, width 0. 10-

0.12 mm; insula with 4,5 small tuberculi; lower vaginal lip with numerous long

spicules scattered over entire surface, lower vaginal sclerite absent; upper

vaginal lip with upper vaginal sclerite moderately pigmented, medium to large

size; postgenital lobe with apex with a moderately deep median indentation

(0.11-0.25 of length), 7-10 (usually 7, 8) setae on each side of midline, 15-19

total setae, dorsal PGL index 0. 85-0.94, ventral PGL index 1.20-1.48, ven-

tral length 0.09 mm; cercus with apex sharply rounded with 2 long stout setae,

dorsal surface with a number of short to moderately long setae on apical 0.77-

0.91, scales absent, ventral surface with a few short setae on lateral margin,

cercus index 2.15-2.50, cercus/dorsal PGL index 2.5-2.8, cercus length 0. 15-

0.17 mm; one large seminal capsule, base of accessory gland duct heavily pig-

mented.

40 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

PUPA and LARVA. Not known.

TYPE-DATA. The lectotype female, deposited in the U. S. National

Museum of Natural History (USNM), possesses the following data on the adult

labels: Fort Pickit |[Malabang, Lanao], Mindanao, P. I. [Philippine Islands];

Danielsia pagei Ludl., Lectotype, Knight and Hull 1953; and Type No. 27799

U. S. N. M. [red rectangular label]. The lectotype is in good condition, but

it has the left and most of the right antennae missing, and tarsomeres 1-5 of

left midleg and tarsomeres 2-5 of left hindleg missing. Knight and Hull

(1953: 463) designated one of the 7 cotypes of pageias lectotype. Ludlow

(1911: 128) indicated that pagei was described from 7 specimens which were

collected in November at 0600 h by Major Henry Page, U. S. Army. Six

female paratypes of pagei are in the USNM and possess the same collection

data as the lectotype except one specimen which has a handwritten, folded

label with the following information: cotype, Mind. [Mindanao], Fort Pikit,

Danilsia Pagei n. sp. Lud. [Ludlow]. These paratypes are in fair to poor

condition.

DISTRIBUTION. Seventy-six specimens examined: 76%.

INDONESIA. Sulawesi (Central), Kulunera, 4-6 Dec. 1965, R. Straatman,

REPUBLIC OF THE PHILIPPINES. Leyte, Tacloban, Diit R., 23 Sept.

1945, H. R. Roberts, 1°. Mindanao, Malabang, Lanao, Fort Pikit (lectotype,

6 paratypes, 4 others), Henry Page, 11°; Agusan, S. Francisco 10 Km. SE,

12 Nov. 1959, L. W. Quate, 27%. Mindoro, San Jose, 21 Jan. 1945, E. S.

Ross, 1°; same except 30 Jan. 1945, 199; same except 28 Feb. 1945, 159;

same except 9 Mar. 1945, Set. Spry, 12.

Distribution from literature.

REPUBLIC OF THE PHILIPPINES (Delfinado, Viado and Coronel 1962

(1963): 440); Mindanao, Fort Pikit (Ludlow 1911: 128). .

DISCUSSION. Females of pagei are similar to those of collessi and

menoni and share somewhat similar markings of the scutum (see Fig. 2),

femora with white scaled apical patches, and eyes separated in front.

Aedes pagei adults are separated from these 2 species by possessing narrow

curved scales on the paratergite and wing with base of costa completely brown

scaled. The female genitalia of pagei have a single seminal capsule while those

of menoni have 3.

BIONOMICS. In the Philippine Islands females have been collected in a

forest and biting man ina jungle during the months of January through March,

September and November. In Indonesia an adult was collected in a Malaise trap

during December.

Ludlow (1911: 128) reported adults collected at 0600 h during November in

the Philippine Islands.

AEDES (PARAEDES) THAILANDENSIS REINERT

(Fiee,. 2,9, 16, 21623, 27)..28)

Aedes (Paraedes) thailandensis Reinert 1976b: 319 (¢*, o*, P*, L*); Knight

1978: 30.

MALE. Head. Antenna brown, 1.09-1.16 length of proboscis, pedicel

dark brown with a few short fine setae and small dark scales mesally; 0.11-

0.12 length of proboscis; clypeus dark brown; proboscis 1.07-1. 14 length of

femur I; several dark brown ocular setae, median 2 longer; vertex with broad

Reinert: Aedes (Paraedes) in the Oriental Region 41

decumbent dark brown scales and a few broad white ones intermixed, a few

narrow curved scales on median anterior area and on ocular line, also a few

long erect forked pale scales on median area, lateral surface covered with

broad dark brown scales except for a large patch of broad white scales on

area anterior to antepronotum, a few similar scales on postgena; occiput with

narrow curved decumbent creamy-white scales and a number of long erect

forked scales, median ones creamy-white and lateral ones dark brown; eyes

contiguous anteriorly. Thorax. Scutal integument dark reddish-brown; scutum

covered with narrow curved dark reddish-brown scales except for narrow

curved creamy-white scales on following areas: anterior promontory, scutal

fossal area (on anterior and along margin over lateral area to scutal angle),

along scutal ridge, a few on anterior dorsocentral area, several on supraalar

area, and a stripe of golden-brown narrow curved scales on acrostichal area

from anterior to posterior; scutum with dark brown setae as follows: 4,9

median anterior promontory, numerous dorsocentral (anterior and posterior),

scutal fossal (3-5 anterior and 2-4 lateral), numerous supraalar, 4-6 posteri-

or medial scutal; scutellum with narrow curved creamy-white scales on each

lobe, median lobe also with a few broader pale brown ones at base, 4,5 long and

3-6 short setae on median lobe, 2-4 long and 2,3 short setae on lateral lobe;

pleural integument dark brown; antepronotum covered with white scales, broad

ones on anterior area and narrow curved ones on posterior area, 8-14 dark

setae; postpronotum with narrow curved reddish-brown scales on dorsal area,

narrow curved white scales on median area and broad white scales on posteri-

or and lower areas, 4-7 dark posterior setae; propleuron with broad white

scales, 14-22 white and brown setae; postspiracular area with a patch of

broad white scales, 2-4 brown setae; paratergite with numerous narrow white

scales; mesepisternum with a large upper and small lower patch of broad

white scales, 3,4 upper and 10-15 posterior pale brown setae, lower setae

shorter; prealar knob with 6-12 golden-brown setae; mesepimeron with a

large patch of broad white scales on upper anterior area, 8-12 golden setae

dorsad of scales. Legs. Coxae I-III with golden to pale brown setae, I with

broad white and a small median patch of broad brown scales on anterior sur-

face, II and III with a small patch of broad white scales on lateral surface;

trochanters I-III with a small patch of broad white scales and a few short setae;

femora I-III with anterior surface dark brown scaled, I also with a narrow

dorsal pale scaled stripe on basal 0.6-0.7, II also with a broad ventral pale

scaled stripe on basal 0.75-0.85, III also with a broad ventral white scaled

stripe on basal 0.8-0.9, stripe narrower distally, I-III with posterior surface

dark brown scaled with a well developed white scaled stripe from base to apex,

stripe dorsal on I, ventral on II and III; posttarsi I-III (Fig. 27) with 2 ungues,

I and II with ungues unequal in size, larger one simple and smaller one with a

tooth, III with ungues equal in size, both simple. Wing. Dorsal and ventral

veins dark brown scaled except for 2-4 white scales at base of costa and 2-9

white scales dorsally and ventrally at base of radius next to remigium; alula

with several narrow brown scales on margin; one remigial seta; upper calypter

with a row of pale setae on margin. Halter. Pedicel pale; capitellum brown

scaled with a few pale scales dorsally. Abdomen. Terga II-VIII with dark

brown scales and a large laterobasal patch of white scales, white patches

longer on V-VII and extending slightly onto dorsal surface, VIII also witha

broad basal band of white scales which connects with laterobasal white scaled

patch, tergum VIII with apex broadly concave with numerous moderately long

thin setae forming a dense patch, a number of long stout setae basad of apical

patch, mesal ones stouter; sterna white scaled, V-VII also with a narrow pos-

42 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

terior band of dark brown scales, bands broader on lateral areas, VIII dark

brown scaled with a few white scales on laterobasal area, sternum VIII with

apex broadly concave with several moderately long and a few long thin setae

onmargin. Genitalia (Fig. 9). Tergum IX heavily pigmented, bilobed, with

3-6 short thin setae on each lobe, cephalic margin deeply emarginate, lateral

margins narrowly connected to sternum IX; gonocoxite heavily pigmented, ter-

gal surface with an elongated patch of short fine setae extending from near

base to near apex on mesal area and with a number of very long stout setae

along lateral margin and lateral surface, sternal surface with mesal margin

somewhat flared and with an elongated patch of very long stout setae from near

base to apex, a few basomesal setae short, long thin setae scattered over

remainder of sternal surface, numerous broad scales on sternal and lateral

surfaces, tergal, sternal and lateral surfaces covered with minute spicules;

gonostylus moderately long, approximately 0.77 length of gonocoxite, bifid,

inner arm thumb-like with 9-15 very short setae scattered over dorsal sur-

face, ventral surface with a row of 5-7 very short setae and apical area with

short spicules forming file-like ridges, outer arm with basal area narrow and

apical area expanded into a broad flat flap; basal mesal lobe divided at base

into 2 long caudally projecting arms, dorsomesal arm flattened, moderately

broad, apex with 5 moderately long flattened setae and a small mesal lobe, a

few short spicules at base of arm, outer area of arm connected by a delicate

membrane to mesal surface of gonocoxite, ventral arm long, narrow, basal

portion curved ventrad and apical portion curved dorsad and mesad of outer

arm, basal portion of arm covered with short spicules, apex with 3 long setae;

phallosome with aedeagus divided into 2 heavily pigmented lateral plates which

are connected basally, each plate curved tergomesad and with 2,3 long baso-

mesal teeth, paramere approximately 0.8 length of aedeagus, parameral

apodeme narrow, approximately 1.76 length of paramere; sternum IX heavily

pigmented, apical margin rounded and extended far cephalad, 2-4 long stout

setae on caudomesal area.

FEMALE. Essentially as in male but with the following differences. Head.

Antenna 1.14-1.20 length of proboscis, flagellomere 1 with basal 0.75 golden

and with several small brown scales, pedicel golden-brown; maxillary palpus

0.16-0.19 length of proboscis; proboscis 1.05-1.14 length of femurI. Thorax

(Fig. 2). Setal differences are as follows: 4-6 anterior promontory, scutal

fossal (4-6 anterior and 3-5 lateral), 2-4 anterior acrostichal, scutellar (5,6

long and 5-7 short ones on median lobe and 4-6 long and 4, 5 short setae on

lateral lobe), 10-15 antepronotal, 12-16 propleural, 5-7 postspiracular, 3,4

upper and 13-16 posterior mesepisternal, 9-13 on prealar knob, and 10-16

upper mesepimeral. Legs. White scaled stripes on posterior surfaces of

femora larger, white stripe on femur II covering most of posterior surface;

tarsomeres I-III (Fig. 28) with 2 ungues, I and II with ungues equal in size

and with a tooth, II with ungues equal in size, both simple. Wing. Remigium

with 1,2 setae. Genitalia (Fig. 16). Tergum VIII with scales absent, basal

0.9-1.0 retracted into segment VII, base concave mesally, apex straight or

slightly convex, numerous short setae scattered over apical 0.89-0.93, setae

along apex short and thin, basolateral seta absent, VIII-Te index 0. 92-0. 98,

VIll-Te/IX-Te index 2.23-2.60, length 0.22-0.26 mm, width 0.22-0.28 mm;

sternum VIII with a large lightly pigmented median area, scales absent, base

slightly concave mesally, apex with a moderately deep median indentation

(0.17-0.22 of length) and a medium size lobe on each side of midline, short to

moderately long setae on apical 0. 88-0.96, seta 2-S approximately 0.38 from

1-S, seta 3-S approximately 0.62 from 2-S, apical intersegmental fold lightly

Reinert: Aedes (Paraedes) in the Oriental Region 43

pigmented, VIII-S index 0. 86-0.96, length 0.26-0.29 mm, width 0. 30-0. 32

mm; tergum IX with mesal area lightly pigmented, apex with a small median

indentation and with 4-7 (usually 4,5) setae on each side of midline, 8-12

total setae, [X-Te index 0.90-0.94, length 0.09-0.10 mm, width 0.10-1.11

mm; insula with 3-5 small tuberculi; lower vaginal lip with numerous long

spicules scattered over surface, lower vaginal sclerite absent; upper vaginal

lip with upper vaginal sclerite moderately pigmented, medium size; postgeni-

tal lobe with apex with a small to moderately deep median indentation (0. 10-

0.22 of length), 4-7 setae on each side of midline, 9-14 total setae, covered

with short spicules, dorsal PGL index 0.98-1.09, ventral PGL index 1.68-

2.00, ventral length 0.10-0.13 mm; cercus with 2,3 long setae at apex, covered

with short spicules, dorsal surface with a number of short to moderately long

setae on apical 0.81-0.85, scales absent, ventral surface with a few short

setae on lateral margin, cercus index 2.85-2.97, cercus/dorsal PGL index

3.04-3.55, cercus length 0.20-0.24 mm; one large seminal capsule, base of

accessory gland duct heavily pigmented.

PUPA (Figs. 21, 22). Chaetotaxy as figured and recorded (Table 5).

Cephalothovax. Seta 1-CT with 3-6 branches; 6-CT short; 8-CT with 5-10

branches; 9-CT with 3-6 branches. Respiratory trumpet. Index 3.21-3. 63,

mean 3.35. Abdomen. Seta 1-II with 6-12 branches; 2-II inserted mesad of

3-IT; 1-III with 5-9 branches; 6-II with 4-8 branches; 5-IV-VI very long,

single; 1-V with 4-6 branches; 3-V with 3,4 branches; 6-V with 5,6 branches;

1-VI with 3-6 branches; 6-VI with 2-6 branches; 10-VI-VII very long, single;

1-VII with 2-4 branches; 6-VII with 4-7 branches; 9-VIII very long, single.

Paddle. Broad, with minute spicules on apical 0.58-0.'72 of lateral and apical

0.09-0.13 of mesal margins; index 1.01-1.19, mean 1.09.

LARVA (Fig. 23). Description based on 6 larval exuviae with associated

adults. Abdomen of each exuvium is twisted and damaged. Head. Moderately

pigmented; setae 0,1,3,14,18, 20-C single; 4-C with 5-7 (6) branches, inserted

mesad and only slightly caudad of 5-C; 5-C with 3 branches; 6-C with 2,3 (2)

branches; 7-C with 5-9 (7) branches; 8-C single to 3 (2) branched; 9-C with

2-4 (3) branches; 10-C with 2-5 (3) branches; 11-C with 4-8 (6) branches; 12-C

with 7,8 (8) branches; 13-C with 5-7 (6) branches; 15-C with 2,3 (3) branches;

dorsomentum with 26-28 (usually 28) teeth, heavily pigmented. Antenna.

Scape long, index 15.31-16.90, mean 15.98; seta 1-A with 3, 4 (3) branches,

inserted 0.45-0.50 from base. Thorax. Seta 0-P with 7-12 (9) branches; 1,5,

6,8,10, 12-P single; 2, 14-P with 2 branches; 3-P with 2-4 (2) branches; 4-P

single or 2 (1) branched; 7-P with 2,3 (2) branches; 9-P single or 2 (2) branched;

11-P with 2-4 (3) branches; 1-M with 2,3 (2) branches; 2,5,7, 10-12-M single;

3-M with 3,4 (3) branches; 4-M with 4-7 (4) branches; 6-M with 3, 4 (4)

branches; 8-M with 4, 5 (4) branches; 9-M with 4,5 (5) branches; 13-M with

10-16 (10) branches; 14-M with 5-8 (8) branches; 1-T with 2,3 (3) branches;

2-T single to 3 (2) branched; 3-T with 5-9 (9) branches; 4, 6-T with 2,3 (2)

branches; 5,10, 12-T single; 7-T with 4-6 (4) branches; 8-T with 8-13 (9)

branches; 9-T with 3 branches; 11-T single to 3 (1) branched; 13-T with 7, 8 (7)

branches. Abdomen. Setae 0,4, 14-VIII single; 1-VIII with 4-7 (4) branches;

2-VIII with 2,3 (2) branches; 3-VIII with 8-13 (10) branches; 5-VIII with 3-5

(4) branches; comb on VIII with 12-18 (usually 15-17) scales; 1-X single or

2 (2) branched; 2-X with 4,5 (5) branches; 3-X single; ventral brush with 9, 10

setae on grid, each with 3-9 branches, some setae always with 8 branches.

Siphon. Moderately pigmented throughout length; index 1.77-1.91, mean

1.87; pecten on basal 0.59-0.66, with 18-24 (usually 21,22) teeth, each tooth

with one large and 1,2 small ventral denticles near base, apex flattened or

44 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

with several long denticles, distal 1, 2 teeth spine-like, without denticles and

wider spaced than remainder; 1-S with 4-6 (4) branches, inserted on basal

0.63-0.67 of siphon and even with or slightly distad of last pecten tooth; 2-S

single, short; 6,7, 9-S single; 8-S with 2-4 (3) branches.

TYPE-DATA. The holotype with its associated pupal exuvium is deposited

in the USNM and possesses the following information on the adult labels and

collection data sheet: THAILAND, Pyvrachin Buri, Ban Bu Phram, 28 July

1971, 06045-102 [collection number], Kol Mongkolpanya and team [ collectors],

T75.18 | genitalia preparation number], collected as a pupa from clear, fresh,

temporary water ina small crab hole, in a heavily shaded area of secondary

scrub, ina valley, and at an elevation of 150 m. The holotype (USNM No.

76351) is in excellent condition. The genitalia are mounted in Canada balsam

on a microscope slide and are in excellent condition. The allotype with its

associated pupal exuvium possesses the following collection information:

THAILAND, Prachin Buri, Ban Tub Lan, 11 August 1971, 06061-101, Kol

Mongkolpanya and team, immature collection data as for holotype except for

elevation which is 80m. Paratypes are as follows: 06038-1 (2 pl), -2 (2 pl),

-100 (¢ pl), collection data as for allotype except date which is 27 July 1971;

06045-7 (2 pl), -9 (2 pl), -11 ( pl), -100 (“), -101 (% p), -104 (2 p), 06045

(2), collection data as for holotype; 06055-100 (“ p), -101 (% p), collection

data as for allotype except date which is 29 July 1971; 06061-1 (2 pl), -100

(S p), -102 (So p), collection data as for allotype; and 06062 (28 2), collected

biting man in a secondary deciduous forest with bamboo, in partial shade, in

mountainous terrain and at an elevation of 500 m, on 12 August 1971, at Khao

Chang Chalut, other data as for holotype. The type-specimens are deposited

in the USNM except for 2 female and one male paratypes (with associated

exuviae) which are deposited in the BMNH and 2 female paratypes (one with

associated exuvium deposited in the Bernice P. Bishop Museum.

DISTRIBUTION. Eighty-four specimens examined: 7 “p, 1°, 6 Spl,

3 2p and 458.

THAILAND. Khon Kaen, Amphoe Chum Phae, Pha Dong Larn, 25-28 Nov.

1962, T-1820, -1837, -1838, -1839, -1847, 82. Nakhon Ratchasima, Amphoe

Pak Chong, Nong Sa Rai, 20 May 1963, P. Boonyakanist, T-8404, 12. Nan,

Ban Sala, 6 Aug. 1966, S. Maneechai, 01364-104, 1%p. Pvachin Buri, Ban

Bu Phram, 12, 28 Jul. 1971, K. Mongkolpanya and team, 06025, 06045-7, -9,

-11, -100, -101, -102, -104, 1 “p (holotype), 1p, 1%, 3 2pl and 1 2p (para-

types), and 32; same except Ban Tub Lan, 27, 29 Jul. 1971, 06038-1, -2,

-100, 06055-100, -101, 06061-1, -100, -101, -102, 4p, 3 fpl, 1 2p (para-

types) and 1 £p (allotype); same except Khao Chang Chalut, 6 Jul. and 12 Aug.

1971, 06005, 06062, 28 2 (paratypes), and 39.

VIETNAM. Khanh Hoa, Duc My, 2 Oct. 1966, Lt. Erickson and VHL

Beardsley, 1505, 2°.

Distribution from literature.

THAILAND and VIETNAM. As reported above (Reinert 1976b: 325).

DISCUSSION. The adult habitus of both sexes of thailandensis are very

similar, a condition which differs from the marked sexual dimorphism of most

species of the subgenus. Adults of thailandensis are easily separated from

the other species of Paraedes (barraudi, bonneae, chrysoscuta, collessi,

menont, ostentatio and pagei) by the combination of antepronotum with broad

and a few narrow white scales, paratergite with narrow white scales, post-

pronotum with broad white scales, and the scale patterns of the vertex and

scutum. Reinert (1976b: 320) reported the posttarsi I and II of the male as

having the ungues simple; only the larger ungues are simple, the smaller ones

Reinert: Aedes (Paraedes) in the Oriental Region 45

each have a small tooth.

The male genitalia of thailandensis are very distinctive (see Fig. 9), as are

all species of the subgenus Pavaedes, in the development of the gonostylus and

basal mesal lobe. |

Female genitalia of thailandensis (see Fig. 16) can be separated from the

other species of the subgenus by the combination of the following features:

sternum VIII shape and pale pigmented median area; tergum VIII shape and

nearly covered with setae; and number of setae on tergum IX. From menoni

it is distinguished by having only a single seminal capsule while ™enonz has 3.

Pupae of thailandensis are easily separated from those of the other known

species by the following: paddle broad, index 1.01-1.19; 5-IV-VI length

greater than 1.5 length of attached terga; 10-VI, VII very long; 4-VIII very long;

and 9-VIII long, 0.60-1.18 length of paddle. Seta 2-II inserted mesad and

cephalad of 3-II which is similar to chrysoscuta and osteniatio, but differs

from bonneae and collessi which have 2-II inserted laterad and cephalad of 3-II.

Larvae of thailandensis possess a number of unique features which easily

distinguish them from the other known species of the subgenus. These are:

seta 5-C with 3 branches; 6-C with 2,3 branches; 2-M single; 3-M with 3, 4

branches; 2-X with 4,5 branches; pecten with 18-24 teeth; shape of pecten

teeth (see Fig. 23); siphon moderately pigmented throughout length; siphon

index 1.77-1.91; and antennal scape index 15.31-16. 90.

BIONOMICS. Adults have been taken in Thailand feeding on man in heavy

vegetation of a forest (1900-2000 h), a secondary deciduous forest, bamboo

groves (500 m elevation), and resting in forest vegetation. In Vietnam one

female was collected biting man (2000 h) near a salt marsh and an additional

female was taken in a CDC light trap.

The usual immature habitat is fresh water in crab holes located at eleva-

tions of 80-150 m. Immatures in Thailand were collected from clear, tem-

porary water in small crab holes (5 times), once from a small wheel track,

located in partially and heavily shaded areas of secondary scrub and ina bam-

boo grove, all located in valley terrain, and at elevations from 80-300 m.

ACKNOWLEDGMENTS

Grateful appreciation is expressed to Michael E. Faran and Ronald A.

Ward, Walter Reed Army Institute of Research, Washington, DC, E. L.

Peyton, Medical Entomology Project (MEP), USNM, Washington, DC, and

Peter F. Mattingly (retired), BMNH, London, England, for reading and com-

menting on the manuscript; to the following for the loan of types and other

specimens--Peter F. Mattingly, BMNH, Douglas J. Gould, E. L. Peyton and

Bruce A. Harrison, formerly of the South East Asia Treaty Organization Medi-

cal Research Laboratory, Bangkok, Thailand, Ronald A. Ward, Oliver S.

Flint, Jr. and E. L. Peyton, USNM, Wallace A. Steffan, Bernice P. Bishop

Museum, Honolulu, Hawaii, Shivaji Ramalingam, University of Malaya,

Kuala Lumpur, Malaysia, Lloyd E. Rozeboom (retired), Johns Hopkins Uni-

versity School of Hygiene and Public Health, Baltimore, Maryland, Nalini

Jayasekera, Medical Research Institute, Colombo, Sri Lanka, P. J. van

Helsdingen, Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands, and

Kenneth L. Knight (retired), North Carolina State University, Raleigh, North

Carolina; to Karl V. Krombein, for providing support for field collection of

specimens from the project, Biosystematic Studies of the Insects of Ceylon;

to Young T. Sohn and Vichai Malikul, MEP, for preparing the illustrations; to

46 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

Janet D. Rupp, MEP, for typing the manuscript for offset reproduction; and to

my wife, Mollie, for typing the manuscript drafts.

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The occurrence and known human-disease

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The occurrence and known human disease relationships

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Species composition and human disease relationships

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1970.

REISEN, W. K., J.

1971.

REINERT, J. F.

1974.

1976a.

1976b.

1979.

ROZEBOOM, L. E.

1965.

Filariasis in Ulu Trengganu, West Malaysia:

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Reinert: Aedes (Pavaedes) in the Oriental Region ol

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LIST OF FIGURES

Aedes (Paraedes) barvaudi--adult male

2. Aedes (Paraedes) bonneae, collessi, menoni, ostentatio, pagei and

thailandensis--female nota and antepronota

3. Aedes (Paraedes) barraudi--male genitalia

4. Aedes (Paraedes) bonneae--male genitalia

0. Aedes (Paraedes) chrysoscuta--male genitalia

6. Aedes (Paraedes) collessi--male genitalia

7. Aedes (Pavaedes) menoni--male genitalia

8. Aedes (Paraedes) ostentatio--male genitalia

9. Aedes (Paraedes) thailandensis--male genitalia

10. Aedes (Paraedes) bonneae--female genitalia

11. Aedes (Paraedes) chrysoscuta--female genitalia

12. Aedes (Paraedes) collessi--female genitalia

13. Aedes (Parvaedes) menoni--female genitalia

14. Aedes (Paraedes) ostentatio--female genitalia

15. Aedes (Paraedes) pagei--female genitalia

16. Aedes (Paraedes) thailandensis--female genitalia

17. Aedes (Paraedes) bonneae--pupa

18. Aedes (Paraedes) chrysoscuta--pupa

19. Aedes (Paraedes) collessi--pupa

20. Aedes (Paraedes) ostentatio--pupa

21. Aedes (Paraedes) thailandensis--pupa

22. Aedes (Paraedes) bonneae, collessi, ostentatio and thailandensis--male

genital lobes of pupae

23. Aedes (Paraedes) bonneae and thailandensis--larvae

24. Aedes (Paraedes) chrysoscuta--larva and female notum and

antepronotum

25. Aedes (Paraedes) collessi--larva

26. Aedes (Paraedes) ostenitatio--larva ,

27. Aedes (Pavraedes) barraudi, bonneae, chrysoscuta, collessi,

ostentatio and thailandensis--male tarsomeres 5 and posttarsi

28. Aedes (Paraedes) bonneae, chrysoscuta, collessi, menoni,

ostentatio, pagei and thailandensis--female tarsomeres 5 and

posttarsi

| Sees Boag +

ae: ae

ii a

Reinert: Aedes (Pavaedes) in the Oriental Region

FIGURE ABBREVIATIONS

Male Genitalia

Aedeagus

Basal mesal lobe

Gonocoxite

Gonostylus

Sternum 9

Tergum 9

Female Genitalia

Accessory gland duct base

Cercus

Line of attachment of

perianal membrane to

dorsal surface of PGL

Hinge

Insula

Tergum 9

Lower vaginal lip

Postgenital lobe

Pupa

Cephalothorax

Abdominal segments

1-9

Larva

Antenna

Head

Comb scale

Dorsomentum

Abdominal segments

1-8, 10

Mesothorax

Hed

eee |

Parameral apodeme

Paramere

Phallosome

Paraproct

Proctiger

Tergum 10

Seminal capsule

Seminal capsule pore

Tuberculus

Upper vaginal lip

Upper vaginal

sclerite

Sternum 8

Tergum 8

Setae 1, 2, 3-S of

VilI-s

Metanotal plate

Paddle

Respiratory trumpet

Prothorax

Pecten tooth

Siphon

Scape

Metathorax

Ventromedian

cervical sclerite

Aedes (Paraedes) barraudi

antepronotum

menoni/ 2 thailandensis 9°

co/llessi 2

ostentatio 2 | bonneae 2°

Aedes (Paraedes) barraudi

DAES Pee

IX, X-Te and Pr ;

BML

bor

lateral dorsal

Aedes (Paraedes) bonneae

IX, X-Te and Pr

Aedes (Paraedes) chrysoscuta

SRI LANKA

IX,X-Te and Pr

: ne ee an .

gl Vi RES

Aedes (Paraedes) col/lessi

Fig. 7

Cees y

0.1

Lee GA

EA er BRO) Lak eee OBIE

Aedes (Paraedes) menoni

IX,X-Te and Pr

Aedes (Paraedes) ostentatio

THAILAND

dorsal plate

X-Te

IX,X-Te and Pr membrane

Aedes (Paraedes) thailandensis

Fig.10

ventral

SCa

5 ;

ae AU ari i phone

ade feel yh [: lester] so) al ad | ar

age! Si Tal aiteen Bypee

Vill-Te

Aedes (Paraedes) bonneae

ventral

wil

revaverever ics}:

eae

Vill-Te

Aedes (Paraedes) chrysoscuta

Va,

Mpf2

SCa

ventral

0.1

he’. < yh ap

Daiie Sal ees 14

bey ily Vee ly

ae ae era!

Vill-Te

Aedes (Paraedes) co/lessi

Fig.13

ayes aa HR Al

Wheat att

ely ek: i

(VN yey [sf

\ \ i ¥ A

Akt As, " |

{ : | ;

sa) Abs el ee anaes

‘\ \ I | | [14 +

ree go

ventral

Vill-Te

dorsal

Aedes (Paraedes) menoni

ventral SCaP

nyt

nie

Vyrin| ty

(epee

ae’ yt)

| IM | 4

Aedes (Paraedes) ostentatio

THAILAND

Fig.15

ventral

dorsal

Vill-Te

Aedes (Paraedes) pagei

ventral

1X -Te SCa

dorsal

Aedes (Paraedes) thailandensis

Aedes (Paraedes) bonneae

Aedes(Paraedes) chrysoscuta

SRI LANKA

Aedes (Paraedes) collessi

Aedes (Paraedes) ostentatio

MALAYSIA

aN)

Aedes (Paraedes) thailandensis

Fig.22

Lobes of Paraedes Pupae

Male Genital

0-5

ae ame Te

toe

collessi

thailandensis

ostentatio

Hi ss

Aedes (Paraedes) bonneae

Aedes (Paraedes) thailandensis

Fig. 24

0:2

eae Nae

Antepronotum

—_

CE ae

—

A \

Aedes (Paraedes) chrysoscuta

SRI LANKA

ventral

Aedes (Paraedes) collessi

Aedes (Paraedes) ostentatio

MALAYSIA

Fige2y

Male Tarsomeres5 and Posttarsi

z= a aaa

a a ae

bonneae

chrysoscuta collessi

ostentatio

thailandensis

Fig.28

Female Tarsomeres5 and Posttarsi

col/less/ menoni

ostentatio q pagel

thailandensis lehai/llabehusl

vee

a BS

fats

Reinert: Aedes (Paraedes) in the Oriental Region

APPENDICES

Reinert: Aedes (Pavaedes) in the Oriental Region 85

TABLE 1. Record of the branching of the setae on the pupae of

Aedes (Pavaedes) bonneae (10 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Cephalothorax Abdomen III (Cont. ) Abdomen V (Cont. )

1 3-5 4 1 4-10 10 14 1 1

2 2-4 3 2 1 1

3 3-4 4 3 1 1 Abdomen VI

4 3-6 3 4 1-4 3

5 1-5 4 5 3-8 4 0 1 1

6 1 1 6 1-4 3 1 2-5 3

1 2-4 3 7 1-4 2 2 1 1

8 3-7 4 8 2-6 4 3 1-3 1

9 1-3 2 9 1 1 4 2-6 4

10 1-4 1 5 1 1

Metanotal Plate 11 1 1 6 1 1

14 1 1 7 1 1

10 3-5 5 8 2-5 3

11 1 1 Abdomen IV 9) 1 1

12 2-6 3 10 1 1

0 1 1 11 1 1

Abdomen I 1 4-9 6 14 1 1

2 1 1

1 21-48 28 3 1-7 5) Abdomen VII

2 1 1 4 1-2 1

3 1 1 5 1 1 0 1 1

4 4-7 6 6 1-2 1 1 1-3 p

5 2-5 4 7 1-3 1 2 1 1

6 1-2 1 8 1-4 3 3 1-4 2

7 1-4 2 9 1 1 4 2-4 2

9 1 1 10 1-4 Z 5 1 1

10 1 1 11 1 | 6 1-4 3

| 14 1 1 i 1 1

Abdomen II 8 2-5 3

Abdomen V 9 1 1

0 1 1 10 1-2 1

: 14-22 19 0 1 1 11 1 1

y 1-2 1 1 3-6 4 14 1 1

3 1 1 2 1 1

4 4-9 6 3 1-3 1 Abdomen VIII

5 2-5 3 4 2-7 4

6 1 i 5 1 1 0 1 1

7 1-4 1 6 1-3 1 4 1-3 1

9 1 1 7 2-8 4 9 1-2 1

8 2-4 i 14 1-2 1

Abdomen HI 9 1 1

10 1-2 1 Paddle

0 1 1 11 1 1

1 1 1

86 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

TABLE 2. Record of the branching of the setae on the pupae of

Aedes (Paraedes) chrysoscuta (10 specimens)

Seta Range Mode seta Range Mode Seta Range Mode

Cephalothorax Abdomen III (Cont. ) Abdomen V (Cont.)

1 2-5 3 1 4-8 4 14 1 1

2 2-4 3 2 1 1

3 2-5 3 3 1 1 Abdomen VI

4 2-7 6 4 2-6 3

5 3-6 5 5 2-4 3 0 1 1

6 1 1 6 1-4 2 1 2-5 3

7 2-3 2 7 2-5 3 2 1 1

8 4-6 6 8 2-5 3 3 1-3 1

9 2-4 3 9 1 1 4 3-6 4

10 3-5 4 5 1 1

Metanotal Plate 11 1 1 6 1 1

14 1 1 7 1 1

10 2-4 3 8 2-5 3

11 1 1 Abdomen IV 9 1 1

12 3-7 4 10 1-2 1

0 1 1 11 1 1

Abdomen I 1 3-8 9) 14 1 1

2 1 i

1 15-29 20 3 3-7 6 Abdomen VII

2 1 1 4 1-2 1

| 1 1 5 1 1 0 1 1

4 4-8 6 6 1-4 2 1 2-4 3

5 2-4 2 7 1-3 2 2 1 1

6 1 i 8 2-5 3 3 2-4 3

7 1-4 3 9 1 1 4 1-4 3

9 1 1 10 2-5 4 ) 1 1

10 1 1 11 1 1 6 2-5 3

14 1 1 7 1 1

Abdomen II 8 2-5 4

Abdomen V 9 1 1

0 1 it 10 1-4 3

1 10-23 13 0 il 1 11 1 1

2 1 1 1 2-5 3 14 1 1

3 1 1 2 1 1

4 4-9 5 3 1-3 2 Abdomen VIII

5 2-4 3 4 2-7 5

6 1-2 i! 5 1 1 0 1 1

7 1-4 1 6 1-3 a 4 2-4 3

9 1-2 1 7 3-7 6 9 1 1

8 2-4 3 14 1-3 1

Abdomen III 9 1 1

10 1-2 1 Paddle

0 1 1 11 1 1

1 1 1

Reinert: Aedes (Paraedes) in the Oriental Region 87

TABLE 3. Record of the branching of the setae on the pupae of

Aedes (Paraedes) collessi (7 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Cephalothorax Abdomen III (Cont. ) Abdomen V (Cont.)

1 2 2 1 2-3 2 14 i 1

2 2-3 2 2 1 1

3 2-4 2 3 1 1 Abdomen VI

4 1-3 2 4 1-4 1

5 2 2 5 2-3 2 0 1 it

6 1 1 6 1 1 1 1-2 1

7 2-3 2 7 2 2 2 1 1

8 3-4 3 8 2-4 2 3 1-3 1

9 1-2 2 9 1 1 4 1-3 3

10 1-3 2 5 1-2 1

Metanotal Plate 11 1 1 6 1 1

14 1 1 A 1 1

10 3-8 5 8 1-2 2

11 1 1 Abdomen IV 9 1 1

12 2-4 4 10 1 1

0 1 1 11 1 1

Abdomen I 1 1-2 2 14 1 1

2 1 1

1 17-35 28 3 2-5 4 Abdomen VII

2 1 1 4 1-2 1

3 | 1 5 1-2 1 0 1 1

4 6-9 6 6 1 1 1 1 1

9) 3-5 4 7 1-3 2 2 1 1

6 1 1 8 1-4 2 3 1-3 2

7 1-2 1 9 1 1 4 1-2 2

9 1 1 10 1-3 2 D 1-2 1

10 1 1 11 1 1 6 1-3 2

14 1 1 7 1 iL

Abdomen II 8 1-3 2

Abdomen V 9 1 1

0 1 1 10 1 1

1 5-14 6 0 1 1 11 1 1

y) 1 1 1 1-2 1 14 1 1

3 1 1 2 1 1

4 3-6 4 3 1 1 Abdomen VIII

5 2-4 5 4 2-4 4

6 1 1 3) 1 1 0 1 1

7 1 1 6 i 1 4 1-3 2

9 1 1 7 1-4 3 9 t 1

8 2-3 2 14 1 1

Abdomen III 9 Af 1

10 1 1 Paddle

0 1 1 11 1 1

1 1 1

88 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

TABLE 4. Record of the branching of the setae on the pupae of

Aedes (Paraedes) ostentatio (10 specimens)

Seta Range Mode Seta Range Mode seta Range Mode

Cephalothorax Abdomen III (Cont. ) Abdomen V (Cont.)

1 2-7 4 1 3-8 7 14 1 1

2 2-5 3 2 1 1

3 2-5 4 3 1 1 Abdomen VI

4 2-7 4 4 2-5 2

9) 3-8 a 5) 2-4 2 0 1 1

6 1 1 6 1-4 : 1 3-6 3

7 2-4 3 7 2-5 4 2 ead 1

8 2-7 5 8 2-6 3 3 1-3 |

9 2-5 3 9 1 1 4, 3-6 4

10 1-3 2 5) 1 1

Metanotal Plate 11 1 1 6 1 1

14 1 1 7 1 1

10 2-5 2 8 2-4 2

11 1 1 Abdomen IV 9 1 1

12 4-8 10 1 1

) 1 1 11 1 1

Abdomen I 1 3-8 9) 14 1 1

2 1 1

1 17-34 30 3 3-8 ‘i Abdomen VII

2 1-2 1 4 1-4 2

3 2 1 3) 1 1 0 1 i

4. 6-10 10 6 1-2 2 1 1-4 2

D 3-6 4 7 2-4 2 2 1 1

6 1 1 8 2-4 2 3 1-5 2

7 2-5 4 9 1 1 4 1-4 4

9 1 1 10 1-4 2 5 1 1

10 1 1 11 1 1 6 1-4 a

14 1 1 7 ek 1

Abdomen II 8 1-5 2

Abdomen V 9 1 1

0 1 1 10 1-2 1

1 8-19 12 0 1 1 11 1 1

2 1 1 1 2-5 3 14 1 1

3 1 1 2 1 1

4 ean oO 3 1-4 1 Abdomen VIII

5) 2-4 3 4 2-7 5

6 1 1 D 1 1 0 1 1

7 1-6 3 6 1-2 1 4 1-4 3

9 1 1 7 2-6 4 9 1-2 1

8 2-5 3 14 1 1

Abdomen III 9 1 1

10 1-2 1 Paddle

0 1 1 11 : 1

1 1 1

Reinert: Aedes (Pavaedes) in the Oriental Region 89

TABLE 5. Record of the branching of the setae on the pupae of

Aedes (Paraedes) thailandensis (10 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Cephalothorax Abdomen III Abdomen V (Cont.)

1 3-6 3 0 1 1 14 1 1

2 3-5 4 1 o-9 8

3 3-5 4 2 1 1 Abdomen VI

4 4-7 3) 3 1 1

5 4-8 4. 4, 2-3 2 0 1 1

6 1 1 5) 6-11 fi 1 3-6 6

r 2-5 4 6 4-8 4 2 1 1

8 9-10 9 7 1-2 1 3 2-3 3

9 3-6 4 8 2-5 3 4 3-6 a

9 1 1 5) 1 é

Metanotal Plate 10 3-5 3 6 2-6 3

3 1 1 7 1-2 2

10 4-6 D 14 1 1 8 3-5 a

11 1 i 9 1 1

12 3-6 Mi Abdomen IV 10 1 1

11 1 1

Abdomen I 0 1 1 14 1 1

1 4-8 7

1 32-53 48 2 1 1 Abdomen VII

2 1 1 3 Ged ictus

3 1-3 2 4 1 1 0 1 1

4 5-8 7 5 1 1 3 2-4 a

3) 2-3 2 6 2-6 3 2 1 1

6 1 1 7 1 1 3 2-5 4

7 2-4 2 8 2-4 2 4 2-4 5}

9 1 1 9 1 1 5) 1-2 2

10 1 1 10 2-4 3 6 4-7 4

i 1 1 11 1 1 7 1 1

14 1 1 8 3-6 4

Abdomen II 9 1-3 2

Abdomen V 10 1 1

0 1 1 11 1 :

1 6-12 8 0 1 1 14 1 1

2 1 1 1 4-6 3)

3 1-2 2 2 1 1 Abdomen VII

A 4-7 5) 3 3-4 3

5) 9-9 6 4, o-7 5) 0 1 1

6 1 1 5) 1 4 4 1-2 2

7 3-4 3 6 5-6 9) 9 1 1

9 1 1 4 4-7 5 14 1-2 1

8 2-8 3

9 1 1 Paddle

10 1-3 2

11 1 1 1 1 1

TABLE 6. Current taxonomic status of the subgenus Paraedes

Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1981

Species io SG = 2G P

barraudi > 2k

bonneae 2k * 2k * >

chrysoscuta > 2k x 2k x

collessi 2k 2k 2k 2 >k 2k

menont * * * x 2k 2k

ostentatio * > * x 2 >k ek kk

page > kk

thatlandensis > * 2 2k 2 2k ek

2K K

2K OK

* = Stage has been described.

Portion of stage or structure has been figured.

So = Male; “G = Male genitalia; 2 = Female; 2G = Female genitalia;

Pupa; L = Larva

Reinert: Aedes (Paraedes) in the Oriental Region 91

INDEX

Valid taxa are in roman type; invalid taxa are in italics.

numbers are those which begin the primary treatment of the taxon.

Italicized page

Numbers

in parentheses refer to figures illustrating the species and the suffix '"'k"'

indicates the page reference to a Key.

Aedes (genus) 1, 3, 7, 8, 10, 12, 13,

16, 17, 22, 26, 27, 28, G0, a1, 36,

36, 87, 38, 40, 52, 05, 86, 57, a8.

Aedes (subgenus) 1, 3, 7, 8, 31, 38

Aedimorphus 1, 3, 8, 17, 26, 31, 38

Aioretomyia 1, 3, 31, 35

argyrurus 1

aurotaeniatus 1, 2

barraudi 1, 3, 7, 9k, 10, 12, 44, 52,

), @), 2D, 20

bonneae 1, 3, 4, 5, 6, 7, 8k, 9k,

10k, 13, 16, 17, 36, 37, 44, 45,

52, (2), (4), (10), (17), (22), (23),

(27), (23), 33, OO

Bothaella 7

Cancraedes 7, 8

Christophersiomyia 7

chrysoscuta 1, 2, 4, 5, 6, 7, 9k,

10k, 16, 17, 21, 22,36, 37, 42,

45, 52, (5), (11), (18), (24), (27),

(28), 86, 90

collessi 1, 5, 6, 7, 8, 9k, 10k, 12,

16, 22, 26, 27, 36, 37, 40, 44,

45, 52, (2), (6), (12), (19), (22),

(25), (27), (28), 87, 90

curtipes 16

Danielsia 1, 3, 38, 40

Finlaya 8

Geoskusea 7

Huaedes 7

Indusius 7

Leptosomatomyia 7

lineatopennis 8

menoni 1, 4, %, 7, Gk, 26, 27, 2s,

30, 40, 44, 45, 52, (2), (7), (3),

(28), 90

Neomelaniconion 8

Nothoskusea 7

Ochlerotatus 1, 3, 17, 31

ostentatio 1, 2, 4, 5, 6, 9k, 10k, 16,

17, 21, 22, 26, 31, 20, 36, 3%,

38, 44, 45, 52, (2), (8), (14),

(20), (22), (26), (27), (28), 88,

pagei 1, 4, 7, 9k, 26, 27, 38, 40,

44, 52, (2), (15), (28), 90

Paraedes 1, 2, 3, 6, 1, 3, 40, 12,

13,16, 47, 22. 26,27, 20, 30,

31, 35, 38, 40, 44, 45, 52, 85,

86, 87, 88, 89, 90

Pseudohouardina 1, 3, 17, 21

Rhinoskusea 7, 8

Skusea 16

Stegomyia 2

thailandensis 1, 3, 4, 5, 6, 7, 8, 9k,

10k, 12, 16, 17, 36, 37, 40, 44,

45, 52, (2), (9), (16), (21), (22),

(23), (27), (28), 89, 90

Udaya 1, 7

Verrallina 7

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