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Ne, Contri butions

American Entomological Institute.

Volume 10, 1973-1974

CONTENTS \

No. 1. Klein, J. M. Contributions to the mosquito fauna of southeast

Asia. XVII. The Cambodian Aedes (Neomacleaya) species with

some new descriptions (Diptera: Culicidae). Pages 1-20.

Reinert, John F. Contributions to the mosquito fauna of south-

east Asia. XVIII. A reconsideration of Diceromyia Theobald with

inclusion of Aedes nummatus Edwards and Aedes pseudonummatus

new species (Diptera: Culicidae). Pages 21-40. November 23, 1973.

No. 2. Arnell, J. Hal. Mosquito studies (Diptera: Culicidae). XXXII.

A revision of the genus Haemagogus. 174 pages. November 25, 1973.

No. 3. Reinert, John F. Contributions to the mosquito fauna of southeast

Asia. XIX. Bothaella, a new subgenus of Aedes Meigen. 51 pages.

December 21, 1973.

No. 4. Kistner, David H. A taxonomic revision of the termitophilous

tribe Termitohospitini. Part III. The subtribe Termitusina with

notes on their behavior and the ultrastructure of certain setae

(Coleoptera: Staphylinidae). 63 pages. October 21, 1974.

No. 5. Wilder, D. Dee. A revision of the genus Syneches Walker

(Diptera: Empididae) for North America and the Antilles. 30 pages.

December 15, 1974.

No. 6. Phuoc, Duang Trang and Frederick W. Stehr. Morphology and

taxonomy of the known pupae of Coccinellidae (Coleoptera) of North

America, with a discussion of phylogenetic relationships. 125 pages.

December 15, 1974.

This is an irregular-appearing Series,

published by the

AMERICAN ENTOMOLOGICAL INSTITUTE

0950 Warren Road

Ann Arbor, Michigan 48105, USA

Volumes 1-7 available in individual numbers

Volumes 8 and following supplied only as

complete cloth-bound volumes

Contributions

of the

American Entomological Institute

Volume 10, Number 1, 1973

oy?

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA.

XVII. THE CAMBODIAN AEDES (NEOMACLEAYA) SPECIES

WITH SOME NEW DESCRIPTIONS (DIPTERA: CULICIDAE)

J. M. Klein

XVOI. A RECONSIDERATION OF DICEROMYIA THEOBALD

WITH THE INCLUSION OF AEDES NUMMATUS

EDWARDS AND AEDES PSEUDONUMMATUS NEW

SPECIES (DIPTERA: CULICIDAE)

John F. Reinert

Klein on Cambodian Aedes (Neomacleaya)

CONTENTS

Piso PRACT ee ee eee ee ee Sea ob ta Meee eine bee

PN TRODUCTION. oc se te aS tee oe FO ee ane eee Sicaptunraiie

NOTES AND DESCRIPTIONS

andamanensis Edwards........-. a are kw ae

ClQva is arr oue ea eee we ee

ChCIOINS. DCEO 2 os Gls so a ee ees ee ak we

cyrtolabis Edwards ....... Se a ae a a ae

GEVINGIOCN sys BUUR” so ks 4k ees oe sae 6 ke a ee

dux Dyar and Shannon, . 2... sss. pa ae vee aes Serene

es ee steY gs a lk wed ew Eee ole

indecorabilis (Leicester) . ae ee ee ee eS

WRHWEVUS Th. Book a es ea ee ee SoG Oa ee

ROWMIPONIUS UWSP. che cans 6 6 0 CU ea a mae eich ey. a vacuo ie

morose Denna es ee ek ee ele ce ce eee

DUROWIUS Te BO. oe ee ee ws De at eee fee Se ee

SHUMGUS Meeps oo 6 oe ee ts Ce 6 a ek Ge le eee we we

WICH VANE G RAI) os 6 eee Es ee oe a cw ee at alae es

CORSETS ae Paue ge ek ge ee eae ON eee ea

KEYS TO THE SPECIES

Bes te ea ee el ee a eae ee ee eS

Ee ke eae ee ean

NOMNOWLEMOEIIEINTS 0 0 ee 2s eg

Te re a ee ge ea ee ace ee eel aes

PC re a es eg eee eee ee Rw ale SO ec ea We

CO CO OC aI CO O1 O1 B® GC © DO DO bet eet

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. XVII

THE CAMBODIAN AEDES (NEOMACLEAYA) SPECIES

WITH SOME NEW DESCRIPTIONS (DIPTERA: CULICIDAE)!

iM. Rien

ABSTRACT

These notes concern new descriptions, distribution and biology of the

Aedes (Neomacleaya) species, which were collected by the author in Cambo-

die from 1966 to 1970. Fifteen species have been recorded, of which 4 are

described as new: khmerus n. sp., kompongus n. sp., phnomusn. sp. and

stungus n. sp. The unknown females of cyrtolabis Edwards and dermajoensis

Brug are also described and illustrated; finally, complementary descriptions

and figures on fragilis (Leicester) and indecorabilis (Leicester) are given.

A key to the adults of this Cambodian fauna is presented in conclusion.

INTRODUCTION

Up to the present a single species in this subgenus, namely, Neo-

macleaya vallistris has been reported from Cambodia (Delfinado 1968: 3).

The present addition of 10 known species and 4 new ones to the fauna is there-

fore quite a considerable advance in our knowledge of the fauna of Southeast

Asia. These new records show the presence either of species known to have

a wide distribution both east and west of Cambodia or to occur in neighbour-

ing countries as shown in the following summary:

andamanensis, Andaman Islands, Philippines, Thailand, India,

Malaya, Sumatra, Java, N. Borneo, Vietnam,

Cambodia.

clavatus , India, Thailand, Vietnam, Cambodia.

cretatus, Thailand, Cambodia.

cyrtolabis, Singapore, Thailand, Cambodia.

dermajoensis, Sumatra, Thailand, Cambodia.

aux, Philippines, Andaman Islands, Thailand, Java,

Hainan Island, Vietnam, Cambodia.

A ei nadunbed Tan |

Supported in part by Research Contract No. DA-49-193-MD- 2672 from the

U.S. Army Medical Research and Development Command, Office of the

Surgeon General, Washington, D.C.

Office de la Recherche scientifique et technique Outre-Mer, Paris.

2 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

fragilis, Malaya, Cambodia.

indecorabilis, Malaya, Thailand, N. Borneo, Cambodia.

notabilis, Thailand, Cambodia.

UNCUS , Philippines, Thailand, Java, N. Borneo, Malaya,

Sarawak, India, Cambodia.

Knowledge of the immatures of Neomacleaya unfortunately has not

been advanced. The summary given by Delfinado (1968), showed that of 42

species known at that time neither the larva nor the pupa of 23 had been de-

scribed. Now 4 more must be added to the latter.

The present contribution adds considerably to our knowledge of the

habits of adults. Previously 4 species, cautus, panayensis, uncus and

siamensis were known to bite man. At one place in Thailand the last mention-

ed (fide E. L. Peyton) occurred in large numbers during the day resting around

a ground pool. They fed freely on man and a number of progeny were subse-

quently reared from eggs laid by them. The present contribution now adds 9

more to the man-biting list, namely andamanensis, clavatus, cretatus, cyrto-

labis, dermajoensis, fragilis, indecorabilis, vallistris and stungus. One of

these (vallistris) is reported as occurring in large numbers near Phnom- Penh

during the second half of the rainy season with a maximum in October and bit-

ing man freely during the daytime. It does seem strange that there are so few

earlier records of biting and one wonders if, because of difficulty of identifica-

tion, specimens have not been misidentified or perhaps even discarded. The

present contribution plus the earlier work of Delfinado should now make it pos-

sible to identify the species and it is hoped that the subgenus will receive bet-

ter attention in the future.

Adults are often caught in vegetation, rockholes and crabholes but

only 6 species have been caught in light traps. If the species show a disincli-

nation to enter light traps then this may also account for the fact that there are

few records of attempts to isolate viruses from them.

AEDES (NEOMACLEAYA) ANDAMANENSIS EDWARDS

Aedes Cea andamanensis Edwards 1922, Indian J. med. Res. 10:

aia (So).

Aedes (Neomacleaya) andamanensis Edwards, Delfinado 1967, Contr. Amer.

ent. Inst. 1(8): 9 (o*, 9*, L*, P*).

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kandal:

Takmau 2c, Oct. 1967, resting on low vegetation in gardens; Kompong Speu:

Kirirom rainforest 3c, 292, Apr. 1968, resting in rockholes; Kompong Sela:

Stung Chral rainforest 12, March 1970, biting man at 10 pm; Battambang:

Pailin forest 12, Apr. 1969, biting man at 6 pm. Larvae. Kandal: Dey Eth,

July 1967, in shaded muddy road ruts; Kompong Speu: Kirirom hills, May

1968, in residual pools of irrigated fields drying up; Kompong Sela: Stung

Chral, June 1968 and June 1969, in shaded pools on edges of torrents.

AEDES (NEOMACLEAYA) CLAVATUS BARRAUD

Aedes (Aedes) clavatus Barraud 1931, Indian J. med. Res. 19: 614 (c*).

Aedes (Neomacleaya) clavatus Barraud, Delfinado 1967, Contr. Amer. ent.

Inst. 1(8): 13 (o*, 9%).

Klein: Aedes (Neomacleaya) in Cambodia 3

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Speu:

Kirirom rainforest 20°, March 1969, resting on low vegetation; Kompong Sela:

Stung Chral Febr. 1970, 2c resting on low vegetation, 9? biting man between

7 am and 5 pm; Kompong Som: Tuk Sap, sublittoral rainforest 19, March1970,

biting man at4 pm. Larvae. Kompong Sela: Stung Chral, June 1968, in rock

pools.

AEDES (NEOMACLEAYA) CRETATUS DELFINADO

Aedes (Neomacleaya) cretatus Delfinado 1967, Contr. Amer. ent. Inst. 1(8):

14 (ao *, 9*, L*, P*).

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Sela:

Stung Chral rainforest 19, March 1970, biting man at 8 am. ‘Males reared.

Larvae. Kompong Speu: Kirirom rainforest, Febr. 1968, in shaded pools

along edges of torrents; Kompong Seia: Stung Chral rainforest, March 1970,

in stream pools.

AEDES (NEOMACLEAYA) CYRTOLABIS EDWARDS

(Fig. 5)

Aedes (Aedes) cyrtolabis Edwards 1928, Bull. ent. Res. 18: 273 (*);

Edwards and Given 1928, Bull. ent. Res. 18: 344 (L*).

Aedes (Neomacleaya) cyrtolabis Edwards, Delfinado 1967, Contr. Amer. ent.

Inst. 1(8): 16 (c*, L*, P*).

Species characterized as follows: 1. Pale lateral patches on abdo-

minal terga. 2. No scales on anterior lobe of pronotum nor on postspiracular

area. 3. Fine hairs on anterior portion of sternopleuron and on lower mes-

epimeron. 4. Hind claws equal and simple in both sexes, fore and midclaws

unequal in male with only the larger claw toothed, equal in female, each claw

toothed. 5. Terminalia in male as figured by Delfinado (1967, Fig. 7) and in

female as here in Fig. 5.

FEMALE. Head. Vertex dark, with pale lateral patches and pale

scales along the eye margin; upright brown scales confined to the occiput;

torus with minute hairs and some little scales on inner side; palpus short,

about 1.5 / 10 the length of proboscis; proboscis slightly longer than fore fe-

mur (10/9), dark brown. Thorax. As described for male. In addition to

bristles and narrow curved scales on posterior part of pronotum, there are

fine hairs, mainly in lower and posterior part. Wing. Alula fringed withsome

broad scales. Legs. Hind claws equal and simple; fore and mid claws equal,

each claw toothed (Fig. 5). Abdomen. Terga dark with oblique pale lateral

patches not produced on to dorsum; sterna with basal half or 2/3 pale scaled.

Terminalia (Fig. 5). Cercus elongate, about two times as long as basal width;

postgenital plate broad with a wide and deep emargination; postatrial sclerite

rounded with 2 large convex lateral lobes; postatrial plate triangular with an

angulated apical structure and a cordate opening with many fine hairs in front

of it and basally; preatrial plate heart-shaped with a deep median emargina-

tion distally; 3 unequal ovate spermathecae, each with a short neck.

TYPE DATA. Female plesiotype (no. 507, with slide of terminalia,

in USNM), Phnom Praung, Kompong Sela, CAMBODIA, May 1969. Other

examined material, with slides of terminalia: 7<°, 69, same locality, July

1968 and May 1969; 1" Tuk Sap, Kompong Som, Dec. 1969 and 1c Sihanouk-

ville, Kompong Som, Jan. 1967 (3c, 39 with slides of terminalia in USNM,

others in Centre ORSTOM, Bondy, France).

4 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

TAXONOMIC DISCUSSION. On external characters cyrtolabis appears

to be indistinguishable from wncus, protuberans and torosus. Females of the

two last species are still unknown. Females of cyrtolabis and uncus are well

differentiated by their terminalia, in particular by their postgenital, postatrial

and preatrial plates.

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Sela:

Phnom Praung 5c, 39, July 1968 and May 1969, resting on low vegetation;

Kompong Som: Sihanoukville littoral rainforest 29, Dec. 1966 and July 1969,

biting man in daytime and 3c, Jan. 1967, resting on low vegetation; Tuk Sap

sublittoral rainforest 4c’, Dec. 1969, resting on low vegetation. Larvae.

Kompong Sela: Phnom Praung, May and July 1968, in jungle pools

AEDES (NEOMACLEAYA) DERMAJOENSIS BRUG

(Figs. 1, 2)

Aedes (Aedes) dermajoensis Brug 1931, Tijdschr. Ent. 74: 250 (<*).

Aédes (Neomacleaya) dermajoensis Brug, Delfinado 1967, Contr. Amer. ent.

Inst. 1(8): 17 (c*, P*).

Species characterized as follows: 1. Pale basal bands on abdominal

terga complete in male, mostly broken in female. 2. No scales on anterior

lobe of pronotum, nor on postspiracular area. 3. No fine hairs on anterior

portion of sternopleuron, nor on lower mesepimeron. 4. Hind claws equal

and simple in both sexes, fore and mid claws unequal in male, equal in female,

each claw toothed. 5. Terminalia as figured by Delfinado (1967, Fig. 8) and

here in Figs. 1 and 2.

FEMALE. Head. Vertex brown, with pale lateral patches, some

broad or narrow pale yellowish scales on the middle; upright brown scales

confined to the occiput; torus with a few small scales on inner side; palpus

short, about 1.5/10 the length of the proboscis; proboscis longer than fore fe-

mur (8/7), dark brown. Thorax. As described in Delfinado (1967: 18) inmale,

with following addenda: posterior part of pronotum with many narrow curved

brown scales; postspiracular area with 3 or 4 bristles and no scales; no fine

hairs on anterior portion of sternopleuron, or only with 1-3 near the upper

scale patch. Wing. Alula fringed with some broad elongated brown scales.

Legs. Hind claws equal and simple; fore and mid claws equal, each claw

toothed. Abdomen. Terga II-VI with pale mediobasal bands and oblique lat-

eral patches; sometimes basal bands and lateral patches are in contact or even

form a complete basal band as it is usual in male; sterna pale scaled in basal

half, more densely on the sides. Terminalia (Fig. 2). Cercus a little shorter

than two times the basal width; postgenital plate broad, with a very shallow

emargination; postatrial sclerite rounded; postatrial plate pocket-shaped, with

its outer wall covered with short hairs; preatrial plate represented by 2 small

hairy surfaces, poorly sclerotized; 3 unequal spermathecae, each with a long

narrow neck.

TYPE DATA. Female plesiotype (no. 628, with slide of terminalia,

in USNM), Sihanoukville, Kompong Som, CAMBODIA, March 1970. Other

examined material, with slides of terminalia: 8o, 72, May to July 1969 and

Febr. 1970, Stung Chral, Kompong Sela; 2°, July 1968, Angkor forest, Siem

Reap (3¢°,32 in USNM, others in Centre ORSTOM, Bondy). :

TAXONOMIC DISCUSSION. The females ascribed here to derma-

joensis were collected in association with males belonging to this species.

The terminalia of these males presentan apical process of basimere which is

gently rounded or with a smooth pointed apex, as in Fig. 1, never distinctly

pointed as figured by Delfinado (1967, Fig. 8). According to Dr. Botha de

Meillon (pers. com.), one specimen from Thailand in USNM has a rounded

Klein: Aedes (Neomacleaya) in Cambodia 5

apical process, and Brug's original illustration shows a somewhat intermediate

condition.

Female terminalia of dermajoensis are of the same type as those of

panayensis, incertus and the here below described khmerus n. sp. The pocket-

shaped postatrial plate is wrinkled in panayensis, smooth in incertus and

khmerus n. sp. and hairy in dermajoensis. Externally, panayensis is easy to

recognize by the pale scaling of the anterior lobe of pronotum, and khmerus

n. Sp. by its many hairs on anterior portion of sternopleuron and the pale

scaling of postspiracular area. It may be difficult to differentiate externally

_dermajoensis from cretatus and from incertus, which may have the same type

of abdominal scaling

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Siem Reap:

Angkor forest, 120’, 89, July 1968, resting on low vegetation; Kompong Sela:

Stung Chral forest, 160, 109, May to July 1969 and Febr. 1970, resting on

low vegetation; Kompong Som: Sihanoukville littoral rainforest, 3°, March

1970, biting man between 2 and 6 pm. Larvae. Not found.

AEDES (NEOMACLEAYA) DUX DYAR AND SHANNON

Aedes (Aedes) dux Dyar and Shannon 1925, Insect. Inscit. menst. 13: 81 (¢, 9);

Laffoon 1946, J. Wash. Acad. Sci. 36: 233 (o'*, 9*, L*).

Aedes (Neomacleaya) dux Dyar and Shannon, Delfinado 1967, Contr. Amer.

ent. Inst. 1(8): 18 (o*, 9*, L*, P*).

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kandal: Chrui

Changwar, in the suburb of Phnom-Penh, 1c, May 1967, in a light trap, near

cattle. Females not found. Larvae. Not found.

AEDES (NEOMACLEAYA) FRAGILIS LEICESTER

(Fig. 9)

Verrallina fragilis Leicester 1908, Cul. Malaya 3: 199 (cc).

Verrallina malayi Leicester 1908, Cul. Malaya 3: 198 (¢); Laffoon 1946, J.

Wash. Acad. Sci. 36: 238.

Aedes fragilis (Leicester), Edwards 1917, Bull. ent. Res. 7: 222 (c'*).

Aedes (Neomacleaya) fragilis (Leicester), Delfinado 1968, Contr. Amer. ent.

Inst. 2(4): 15 (o*, 9*),

As female terminalia of fragilis are only schematically represented

in Delfinado (1968, Fig. 8) and not at all in older descriptions, I draw them

here (Fig. 9) tentatively from 2 females assigned to this species on following

characters: 1. Pale lateral patches on abdominal terga. 2. Pale scales on

anterior lobe of pronotum, but no scales on postspiracular area. 3. No fine

hairs on anterior portion of sternopleuron, nor on lower mesepimeron. 4.

Proboscis as long as fore femur. 5. All female claws equal and toothed. 6.

Female terminalia as described and represented in Delfinado (1968). We note

that the horseshoe- shaped postatrial plate has broad flat extensions, one on

the top and one on each side at base; preatrial plate large, bulbous and hairy;

preatrial sclerite laterally widened; 3 unequal spermathecae without neck;

sternum VIII with 2 apical paramedian rounded lobes bearing strong bristles.

MATERIAL EXAMINED. 19 (no. 657, with slide of terminalia, in

USNM), March 1970, Tuk Sap, Kompong Som, CAMBODIA; 192 (no. 514, with

slide of terminalia, in Centre ORSTOM, Bondy), July 1968, Phnom Praung,

Kompong Sela, CAMBODIA.

TAXONOMIC DISCUSSION. As discussed under stungus n. sp..

6 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Sela:

Phnom Praung rainforest, 19, July 1968, biting man at 1 pm; Kompong Som:

Tuk Sap sublittoral rainforest, 12, March 1970, biting man at 4 pm.

AEDES (NEOMACLEAYA) INDECORABILIS (LEICESTER)

(Figs. 7, 8)

Verrallina indecorabilis Leicester 1908, Cul. Malaya 3: 200 (c, 9).

Verrallina imitator Leicester 1908, Cul. Malaya 3: 201 (2); Stone, Knight and

Starcke 1959, Thomas Say Found., Ent. Soc. Amer. 6: 206 (synon-

ymy @

Aedes (Neomacleaya) indecorabilis (Leicester), Delfinado 1967, Contr. Amer.

ent, Inst. 1(8): 25 (o*, ?*).

Species characterized as follows: 1. No definite pale lateral patches

on abdominal terga, but dull pale scales distributed laterally on the whole

length of terga. 2. Pale scales on anterior lobe of pronotum, but no scales

on postspiracular area. 3. No fine hairs on anterior portion of sternopleuron,

nor on lower mesepimeron. 4. Hind claws equal and toothed in both sexes;

fore and mid claws unequal in male, equal in female, each claw toothed. 5.

Terminalia as described and figured by Delfinado (1967, Figs. 13, 20) and

here, in Figs. 7, 8.

We note that in male terminalia, apex of basimere is darkened by a

sclerotization bearing a fine hair inserted in an apical concavity. In female

terminalia, postatrial plate is horseshoe-shaped, with a nearly rounded open-

ing; preatrial plate bulbous and hairy, widely open laterally; preatrial sclerite

M-shaped with widened lateral branches; 3 unequal spermathecae without neck;

sternum VIII with 2 apical paramedian rounded lobes bearing strong bristles.

MATERIAL EXAMINED. 4c, 59, with slides of terminalia, Nov.

1967, July and Oct. 1968, May 1969, Phnom Praung, Kompong Sela, CAMBO-

DIA; 12, Dec. 1969, Sihanoukville, Kompong Som (2c, 32 with slides of termi-

nalia in USNM, others in Centre ORSTOM, Bondy).

TAXONOMIC DISCUSSION. The apical sclerotized part of the basi-

mere in male indecorabilis is not accurately represented in Delfinado (1967,

Fig. 13) and it is confirmed by Dr. Botha de Meillon (pers. com. ) that the

structure with an apical seta, as shown in our Fig. 7, is observable in one of

the indecorabilis terminalia preparations in USNM from Thailand. Dr.

Delfinado did not dispose of any male specimen other than the type in BM and

it appears that fore and mid claws could not be re-examined thoroughly.

Leicester (1908: 201) described ''fore and mid ungues markedly unequal, the

larger unguis uniserrate, "' but the tooth of the small claw may have been over-

looked without a fine preparation. Our slides concerning 3c show distinctly

that the small claw of each pair is also toothed.

Externally indecorabilis in either sex is recognizable by the dull

palish scaling of the abdominal terga laterally; this differentiates it from

fragilis, which has definite lateral patches, and from uniformis, whose terga

are wholly dark:

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Sela:

Phnom Praung rainforest, 40°, 32, Nov. 1967, July and Oct. 1968 and May

1969, resting in low vegetation, and 29, Nov. 1968, biting man in daytime;

Kompong Som: Sihanoukville littoral rainforest, 22, Dec. 1969, biting man at

3pm. Larvae. Kompong Sela: Phnom Praung, June 1967, in jungle pools.

Klein: Aedes (Neomacleaya) in Cambodia 7

AEDES (NEOMACLEA TA) KHMERUS n. Sp.

(Figs. 3, 4)

Species characterized as follows: 1. Pale lateral patches on abdomi-

nal terga. 2. No scales on anterior lobe of pronotum, but some pale scales

on postspiracular area. 3. Many fine hairs on anterior portion of sternopleu-

ron, but no hairs on lower mesepimeron, nor on metameron. 4. Hind claws

equal and simple, fore and mid claws unequal in male, equal in female, each

claw toothed. 5. Terminalia as in Figs. 3 and 4.

FEMALE. Head. Vertex brown, with lateral pale scale patches;

some broad and narrow pale scales on the middle of vertex; upright brown

scales confined to the occiput; torus with some minute hairs and small scales

on inner side; palpus short, about 1/7 the length of proboscis; proboscis longer

than fore femur (8/7), dark brown. Thorax. Scutal and scutellar scales dark

or golden brown on dark or reddish brown integument; anterior lobe of prono-

tum with 8 to 10 bristles, without scales; posterior part of pronotum with 4 or

0 bristles, many narrow curved golden brown scales and sometimes 1 to 5 pale

broad or elongated curved scales; postspiracular area with 5 to 8 bristles and

1 to 5 pale scales, broad or narrow and curved; sternopleuron with loosely

arranged pale scale patches, lower patch mixed with fine hairs, some bristles

behind upper scale patch, and many fine hairs on anterior portion; lower mes-

epimeron bare; metameron bare; propleuron with pale scales and bristles.

Wing. Alula fringed with some broad brown scales. Legs. Fore coxa brown

scaled, with a pale spot above; mid and hind coxae with some pale scales; hind

claws equal and simple, fore and mid claws equal, each claw toothed. Abdo-

men. Terga dark brown with lateral oblique pale patches, not or slightly pro-

duced on to dorsum; sterna pale scaled in basal half, more densely on sides.

Terminalia. As in Fig. 4. Cercus short; postgenital plate broad with a shal-

low emargination; postatrial plate pocket-shaped with lateral arms and without

hairs; preatrial plate represented by 2 poorly sclerotized hairy surfaces; 3 un-

equal spermathecae, each with a long narrow neck.

MALE. General habitus as in female. There are no pale scales visi-

ble on postspiracular area, nor on posterior part of pronotum, but they may

be rubbed off, as many scales of thorax are lacking on our 2 specimens; fine

hairs on anterior part of sternopleuron not so numerous as in female, but up

to10 or 12. Legs. Hind claws equal and simple; fore and mid claws unequal,

each claw toothed. Abdomen. Terga with pale lateral patches, produced on

to dorsum, but not forming complete subbasal bands. Terminalia.(Fig. 3).

Basimere short, without apical projection, but with 2 large spines and a patch

of hairs inserted near base, and a flattened median projection with a hairy tip;

distimere expanded in the middle, narrowed and hook-shaped at apex; aedeagus

of andamanensis type; paraproct very small.

LARVA and PUPA. Unknown.

TYPE DATA. Holotype 2 (no. 380, with slide of terminalia and claws,

in USNM), Aug. 1967, Sre Klong, Kompong Speu, CAMBODIA; allotype ¢ (no.

497, with slide of terminalia, in USNM), June 1968, Kirirom hills, Kompong

Speu; 72, 1o paratypes, Aug. and Sept. 1967, Sre Klong, Kompong Speu (3 in

USNM, others in Centre ORSTOM, Bondy).

TAXONOMIC DISCUSSION. Female terminalia of kkmerus n. sp. are

nearest to those of incertus, having a similar smooth pocket- shaped postatrial

plate; kimerus n. sp. may be easily differentiated externally by its many hairs

on anterior portion of sternopleuron and pale scaling of postspiracular area.

Male terminalia of Rhmerus n. sp. are nearest to those of varus, but differ by

the median flattened projection of basimere, whose apex is hairy in the first

and denticulated in the second; other differences exist in basal spines and patch

of hairs on inner margin of basimere, others in distimere and structure of

phallosome; externally the two species may be indistinguisgable when the pale

scales of postspiracular area in khkmerus n. sp. are rubbed off.

8 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Speu;

Sre Klong forest, 1c, 79, Aug. and Sept. 1967, resting on low vegetation;

Kirirom hills rainforest, 1¢, 19, June 1968, resting on low vegetation. Lar-

vae. Not found.

AEDES (NEOMACLEAYA) KOMPONGUS n. sp.

(Fig. 11)

Based only on 2 male specimens, female, larva and pupa being un-

known. Species characterized as follows: 1. Lateral pale patches on abdom-

inal terga produced on to dorsum. 2. Pale broad scales on postspiracular

area and 1 or 2 narrow curved pale scales on anterior lobe of pronotum. 3.

No fine hairs on anterior portion of sternopleuron, nor on lower mesepimeron.

4. Hind claws equal and simple, fore and mid claws unequal, only the larger

claw toothed. 5. Terminalia as in Fig. 11.

MALE. Head, Vertex dark brown, with a few pale broad scales on

the sides and pale narrow scales on the middle; dark upright scales confined

to the occiput; torus bare or with one minute hair or scale on inner side; an-

tenna strongly plumose; palpus short, about 1/8 the length of proboscis; pro-

boscis dark brown, about as long as fore femur. Thovax. Scutal and scutellar

scales brown to golden, lighter on front margin of scutum; anterior lobe of

pronotum with 7 or 8 bristles and 1 or 2 narrow curved pale scales, inserted

near anterior border; posterior part of pronotum with 3 bristles and from 4 to

12 narrow golden brown scales; postspiracular area with 2 bristles and 2 or 3

pale broad scales; sternopleuron with anterior portion bare, upper scale patch

with a row of 6 or 7 bristles behind and lower patch mixed with some hairs and

some bristles behind; lower mesepimeron bare; metameron bare. Wing. Alu-

la fringed with some narrow scales. Legs. Dark scaled, the femora pale

ventrally; fore coxal scale patch brownish with pale scales above and below;

mid and hind coxae each with some pale scales; hind claws equal and simple;

fore and mid claws unequal, only the larger being toothed. Abdomen. Terga

dark brown, with pale lateral patches curved and produced on to dorsum, but

not forming complete subbasal bands; sterna pale scaled on basal half. Ter-

minalia.(Fig. 11). Basimere produced apically into a large process divided

into 2 or 3 finger-like projections, and an inner basal lobe with 3 or 4 blunt

spines and some bristles; distimere inserted subapically, slightly curved and

evenly tapered; paraproct developed into darkened triangular blades with a

denticulated apical margin and an inner apical blunt pointed apex; aedeagus of

the andamanensis type.

FEMALE, LARVA and PUPA. Unknown.

TYPE DATA. Holotype - and 1 paratype ° (with slides of terminalia,

in USNM), Febr. 1968, ex pupae, Kirirom hills, Kompong Speu, CAMBODIA.

TAXONOMIC DISCUSSION. Terminalia of Rompongus n. sp. are very

distinctive and externally the described males do not seem to belong to one of

the species known only by their females. Between these, the nearest may be

hispidus, which has, like kompongus n. sp. pale scaling on postspiracular

area; but its anterior lobe of pronotum is described as having narrow golden

scales and abdominal terga as being completely banded.

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. The new spe-

cies is known only from 2 males reared from pupae, collected in a little grassy

pool on the edge of a torrent in rainforest.

Klein: Aedes (Neomacleaya) in Cambodia g

AEDES (NEOMACLEAYA) NOTABILIS DELFINADO

Aedes (Neomacleaya) notabilis Delfinado 1967, Contr. Amer. ent. Inst. 1(8):

28 (2*).

The single female described by Delfinado has the fore legs missing.

I therefore mounted all legs of my specimen and noted: equal and simple hind

claws (not toothed as described), equal fore and mid claws, each claw toothed.

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Sela:

Stung Chral rainforest, 19, July 1969, resting on low vegetation. Males not

found. Larvae. Not found.

AEDES (NEOMACLEAYA) PHNOMUS n. sp.

(Fig. 10)

Based only on a unique male specimen, female, larva and pupa being

unknown. Species characterized as follows: 1. Lateral pale patches on ab-

dominal terga produced on to dorsum. 2. Pale broad scales on postspiracular

area and apparently also some on anterior lobe of pronotum. 3. No fine hairs

on anterior portion of sternopleuron, nor on lower mesepimeron. 4. Hind

claws equal and simple; fore and mid claws unequal, each claw toothed. 5.

Terminalia as in Fig. 10.

MALE. Head, Vertex dark brown with pale broad scales on the mid-

dle and on the sides; some dark upright scales confined to occiput; torus with

some minute hairs on inner side; palpus short, about 1/8 the length of probos-

cis; proboscis dark, longer than fore femur (8/7). Thorax. Scutal and scutel-

lar scales golden brown, lighter on front margin of scutum; anterior lobe of

pronotum with 3 or 4 broad pale scales on anterior border (but may be rubbed

from the vertex), 6 to 8 bristles and behind them 5 to 8 hairs; posterior pro-

notum with narrow golden brown scales and 3 or 4 bristles; postspiracular

area with 2-4 pale broad scales and 3 bristles; sternopleuron with anterior

portion bare, upper scale patch with a row of 5 bristles behind and a lower

patch mixed with some hairs; upper part of mesepimeron with a large patch of

scales and no hairs behind; lower mesepimeron bare; metameron bare. Wing.

Alula fringed with narrow or lanceolate scales. Legs. Dark brown, the fe-

mora pale ventrally; fore coxa brown scaled, with a pale spot above; mid and

hind coxae with some pale scales; hind claws equal and simple; fore and mid

claws unequal and each claw toothed. Abdomen. Terga dark, with pale later-

al patches extending on to dorsum, but not forming complete subbasal bands;

sterna not visible. Terminalia (Fig. 10). Basimere with a straight apical

projection, in side view curved and tapering, with a short rounded apex in lat-

eral position, and an inner basal lobe, bearing a group of strong bristles, 2

dark long spines and a longer pointed process, all nearly straight; distimere

slender, curved upwards and tapered to a pointed tip; aedeagus simple, of the

andamanensis type; paraproct enlarged at base, slightly curved inwards and

pointed distally.

FEMALE, LARVA and PUPA. Unknown.

TYPE DATA. Holotype < (no. 802, legs on one side missing, with

Slide of terminalia, in USNM), Nov. 1970, Chrui Changwar, Kandal, CAM-

BODIA.

TAXONOMIC DISCUSSION. Male terminalia of phnomus n. sp. are

near to those of indicus, but Barraud (1934: 283) described these as having on

basimere "a single slender elbowed process, arising from inner surface of

coxite; no spines at ventral root.'' Externally, phnomus n. sp. does not fit

the description of indicus, which has a postspiracular area with narrow curved

golden scales, according to Delfinado (1967: 24). A. phnomus n. sp. is near

10 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

to the species having pale scaling on postspiracular area and on anterior lobe

of pronotum, in particular to vallistris, from which it is well differentiated by

the male fore and mid claws being all toothed and by the terminalia.

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. The new spe-

cies is known only from a single male, collected in a light trap, near cattle.

AEDES (NEOMACLEAYA) STUNGUS n. sp.

(Fig. 6)

Based on only 3 females, male, larva and pupa being unknown. Spe-

cies characterized as follows: 1. Pale lateral patches on abdominal terga.

2. Proboscis distinctly shorter than fore femur. 3. Pale scales on anterior

lobe of pronotum, but no scales on postspiracular area. No fine hairs on ante-

rior portion of sternopleuron, nor on lower mesepimeron. 5. All claws equal,

each being toothed. 6. Terminalia as in Fig. 6.

FEMALE. Head. Vertex dark brown, with pale lateral patches and

pale scales along the eye margin; dark upright scales confined to occiput; torus

with some small scales on inner side; palpus short, about 1/6 the length of

proboscis; proboscis shorter than fore femur (6/7 or 5/6), dark brown. Tho-

vax. Scutal and scutellar scales dark to golden brown on light brown integu-

ment; anterior lobe of pronotum with 5 or 6 bristles and many broad pale

scales; posterior part of pronotum with 4 or 5 bristles near posterior border

and without scales; postspiracular area with 2 or 3 bristles and without scales;

sternopleuron with lower patch of scales mixed with fine hairs and without

hairs on anterior portion; upper mesepimeron with a patch of scales without

hairs behind; lower mesepimeron bare; propleuron with pale scales and bris-

tles. Wing. Alula fringed with narrow scales. Legs. Dark scaled, mid and

hind femora extensively pale scaled ventrally; hind claws equal and toothed;

fore and mid claws also equal and toothed. Abdomen. Terga dark brown, with

pale lateral patches, sometimes well rounded; sterna pale scaled on basal half

or 2/3. Terminalia (Fig. 6). Cercus long, tapered distally, a little more

than twice as long as basal width; postgenital plate rather narrow with a shal-

low emargination; postatrial plate horseshoe- shaped, dilated at top and with a

narrow opening; postatrial sclerite with an internal rib covered with fine hairs

in lower part, and an outer rib curved inwards in upper part with a short later-

al arm; preatrial plate large, bulbous and hairy; preatrial sclerite M- shaped

in the middle with broad lateral extensions; 3 unequal spermathecae without

neck; sternum VIII with 2 apical paramedian rounded lobes, bearing strong

bristles.

MALE, LARVA and PUPA. Unknown.

TYPE DATA. Holotype ? (no. 656, with slide of terminalia, in USNM),

July 1969, Stung Chral, Kompong Sela, CAMBODIA; 22 paratypes, May 1969

and Feb. 1970, same locality. (19 paratype in USNM, the other in Centre

ORSTOM, Bondy).

TAXONOMIC DISCUSSION. Female terminalia of stungus n. sp. with

their horseshoe- shaped postatrial plate, are near to those of indecorabilis,

fragilis, hamistylus, pahangi and robertsi, all species having broad pale scales

on anterior lobe of pronotum (to be confirmed for pahangi) and in females all

claws equal and toothed. The last three species have wholly dark abdominal

terga and robertsi is the only one having a proboscis shorter than fore femur

a. 6 or 8.3/10); this has not been mentioned by Laffoon (1946) nor by Delfinado

1968), but has been checked by Dr. Botha de Meillon (pers. com.) on female

specimens in USNM. Externally, robertsi may be differentiated, besides

wholly dark abdominal terga, by the female claws, which have a strongly con-

vex outline, according to the drawing in Delfinado (1968, Fig. 25). In female

terminalia, vobertsi differs in particular by the rib of postatrial sclerite,

Klein: Aedes (Neomacleaya) in Cambodia 11

which gives rise apically to a much longer lateral arm; there is no inner rib

with a hairy lower part and the preatrial M- shaped sclerite forms a much

sharper zigzag in the middle part. Female terminalia of hamistylus and

pahangi have strikingly different features of postatrial plate and ribs of post-

atrial sclerite; those of indecorabilis and fragilis, incompletely represented

in Delfinado (1967, Fig. 20; 1968, Fig. 8), have been figured here (Figs. 7, 8

and 9). We note also, that among the Aedes (Neomacleaya) species, whose fe-

males are still unknown, none seems to have the same combination of external

characters in their described males, as the female of stungus n. sp.

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. This new spe-

cies is known only from 3 females, collected in the rainforest of Stung Chral,

Kompong Sela, May and July 1969 and Febr. 1970, of which 2 were resting on

low vegetation and one was biting man at 2 pm.

AEDES (NEOMACLEAYA) UNCUS (THEOBALD)

Culex uncus Theobald 1901, Mon. Cul. 2: 53 (9*); Laffoon 1946, J. Wash.

Acad. Sci. 36: 237 oo Q*), 3

Aedes (Neomacleaya) uncus (Theobald), Delfinado 1967, Contr. Amer. ent.

Inst. 1(8): 30 (o*, 9*, P*).

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kompong Som:

Sihanoukville littoral rainforest, 1c, Dec. 1966, resting on low vegetation;

Kompong Speu: Sre Klong forest, 5c, 39, Aug. 1967 and June 1968, resting on

low vegetation; Kirirom rainforest, 3c’, March 1969, resting in rock holes;

Kompong Sela: Stung Chral rainforest, 12c", 69, June 1968, resting on low ve-

getation and 162, Febr. 1970, biting man between 5 am to 5 pm, mainly be-

tween 7am tollam. Larvae. Kompong Sela: Phnom Praung rainforest, May

1968 and Stung Chral rainforest, June 1968, in jungle pools.

AEDES (NEOMACLEAYA) VALLISTRIS BARRAUD

Aedes (Aedes) vallistris Barraud 1928, Indian J. med. Res. 16: 369 (c*, ?*);

Barraud 1934, Fauna Brit. India, Diptera 5: 290 (o*, 9*); Iyengar and

Menon 1956, Bull. ent. Res. 47: 791 (c).

Aedes (Neomacleaya) vallistris Barraud, Delfinado 1967, Contr. Amer. ent.

Inst. 1(8): 32 (o*, 9*).

DISTRIBUTION and BIOLOGY. CAMBODIA. Adults. Kandal: Phnom-

Penh Ville, 19, June 1967, in a light trap in gardens; Chrui Changwar, Arey

Ksatr, Svay Chrum, Dey Eth, Takmau, all localities near Phnom-Penh, in

small forests or nearby fields or gardens, hundreds of females, all through

the year from 1967 to 1970, mainly in the second half of the rainy season and

in the following months, i.e. from August to March, with a maximum in Oc-

tober, resting in the low vegetation, or biting man in daytime. Between 1965

and 1969, at the Institut Pasteur of Phnom-Penh, 1667 specimens of vallistris,

collected in the neighbourhood of the town, were pooted in 13 lots and inocu-

lated into suckling mice. No viruses were isolated (Chastel, C., Audebaud,

G., Salaun, J.J. and Klein, J.M., unpublished). Kompong Speu: Sre Klong

forest, 3?, Aug. 1967, resting on low vegetation; Kompong Chhnang: Trapeang

Chan forest, 39, Dec. 1969 and Jan. 1970, biting man at 4 pm and 11 pm.

Males not found. Larvae. Not found.

12.

Contrib. Amer. Ent. Inst.,

vol. 10, no. 1, 1973

KEYS TO THE ADULTS OF NEOMACLEAYA SPECIES IN CAMBODIA

2(1)

3(1)

4(3)

5(3)

6(5)

7(6)

8(7)

FEMALES!

Abdominal terga without definite pale markings ...

Abdominal terga with definite pale markings

Abdominal terga wholly dark or with light brown

lateral patches, sometimes with a small lateral

whitish spot on tergum I; hind claws simple.

Postatrial plate simple, with a pair of hairy

finger-like processes (Fig. 9 in Delfinado 1967)

Abdominal terga dark, with the sides dull pale

brown; hind claws toothed; postatrial plate

horseshoe- shaped (hig. 3) 6. ek es Be

Abdominal terga with median basal bands, mostly

separated on the sides from the lateral patches,

sometimes complete, rarely absent .......

Abdominal terga with only lateral pale patches ..

Postgenital plate with a very shallow median

emargination; postatrial plate pocket- shaped,

ee eee wall covered with short hairs

PP A a a ea a ee Ges

Postgenital plate with a deep median emargination;

postatrial plate with characteristic lateral

wing- like structures (Fig. 5 in Delfinado

1967) sos Be RO a

Postspiracular area with some broad pale

scales

Postspiracular area without pale scales ee ae

Metameron with fine hairs; postatrial plate with

bluntly pointed characteristic distal arms

(Fig. 1 in Delfinado 1967) . .

Metameron without fine hairs; postatrial plate

CITICTENE oo ee es Ty VEO ea ages ica er Gee

Proboscis distinctly shorter than fore femur;

hind claws toothed; postatrial plate hor seshoe-

shaped (Fig. 6) ..

Proboscis as long as or longer than fore femur;

ee eo e oe oe e® oe oe oe e @

hind claws simple; postatrial plate different...

Anterior lobe of pronotum with some pale scales;

anterior portion of sternopleuron bare; post-

atrial plate with numerous long hairs on basal

half (Fig. 17 in Delfinado 1967

Anterior lobe of pronotum without pale scales;

anterior portion of sternopleuron with numer-

ous fine hairs; aay nai plate pocket-shaped,

without hairs (Fig. AY NS oe

@ e @ @ e

Poe dax

indecorabilis

vallistris

khmerus Nn. sp.

I Females of kRompongus n. sp. and phnomus n. sp. are still unknown.

Klein: Aedes (Neomacleaya) in Cambodia 13

9(5) Anterior lobe of pronotum with some pale scales;

hind claws toothed; postatrial plate horseshoe-

shaped: (Pigs OV 5 vie a a ee fragilis

Anterior lobe of pronotum without scales; hind

claws simple; postatrial plate different .......... 10

10(9) Lower mesepimeron with numerous fine hairs ........ rE

Lower mesepimeron bare. .....668s es i ES As oe 12

11(10) Postgenital plate with a shallow median emargi-

triangular structure covering the cordate

opening and numerous hairs in front and bas-

ally (Pig 3) She gee errs Ss WR cyrtolabis

12(10) Postgenital plate with a shallow median emargi-

nation; postatrial plate characteristically

broadened basally and bearing long coarse

hairs around the opening (Fig. 17 in

Deliinade VOGT). I. ee See Sed an clavatus

Postgenital plate with a deep median emargina-

tion; postatrial plate with 2 characteristic

hairy lobed structures basally (Fig. 18 in

Delfinado 1967). 4.4% te 4 ees er re notabilis

MALES!

1 Abdominal terga without definite pale markings ........ 2

Abdominal terga with definite pale marking .......... 3

2(1) Abdominal terga wholly dark, or with light brown

lateral patches, sometimes with a small lateral

whitish spot on tergum I. Hind claws simple.

Basimere not produced distally. Distimere

Swollen at base, tapering to a curved tip (Fig.

10 in Delfinade VOGT) ar Oe ; dux

Abdominal terga dark, with the sides dull pale

brown; hind claws toothed; basimere bluntly

produced distally; distimere swollen in the

middle, abruptly narrowed and curved distally

(Chis Dee wn eae ee a eget ee ae gals OS age eu indecorabilis

3(1) Abdominal terga with median basal bands ........... 4

Abdominal terga with only lateral pale patches ........ 5)

4(3) Basimere with a short apical pointed or rounded

process and on inner margin 2 spines near

base and 1 subapical spine (Fig. 1)....... dermajoensis

Males of notabilis and stungus n. sp. are still unknown.

9(3)

6(5)

7(6)

8(7)

9(77)

10(5)

11(10)

Contrib. Amer. Ent. Inst.,

Basimere with 2 apical processes, the one

short and hairy, the other long and slender,

and on inner margin a peg-like process and

1-2 large spines near base (Fig. 6 in

Delinaco: LOG fecevetiie elela deus was. haves

Postspiracular area with some broad sie

scales

Metameron with fine hairs; basimere produced

apically into a large broad process with a

deep concave depression at apex and 2 other

slender processes (Fig. 2 in Delfinado 1967) .

Metameron without fine hairs; basimere dif-

TOPE 6 ae BG Oia ree BER. ace Teme Rie

Each claw of the unequal fore and mid claws

toothed .

Only the larger claw of the unequal fore and

mid claws toothed

Anterior portion of sternopleuron with numerous

ae Me ae basimere without apical projection

Fig m

Anterior portion of sternopleuron. without fine

hairs; basimere with an apical projection

(Fig. se @ eo 2. @.: 82 @) 1H 8. '® .@ e @ e e @ a

Basimere with a short apical and 2 subapical

projections and on inner margin a small

lobe bearing a group of strong spines; para-

proct slender, tapering at apex (Fig. 17 in

Deliinade L9G Daisies ahece cdig lbs. albu se

Basimere produced apically into a large process

divided into 2-3 finger-like projections; para-

proct ee broadened and indented at apex

CPig. EL) aivwineits mara ce. b ee ak rete SS? Geto

Anterior lobe of pronotum with some pale scales;

hind claws toothed; basimere joined subapi-

cally; distimere expanded at middle (Fig. 8

in Delfinado LOGS crs aod s

Anterior lobe of pronotum without pale scales;

hind claws simple; basimere not joined sub-

ApiCally: GiStMere CUO rene oie: ieee pel

Lower mesepimeron bare; paraproct divided;

basimere broadly produced distally, with ‘a

knob-like process at tip and a short pointed

subapical inner projection (Fig. 17 in

Delfinado 1967).

Lower mesepimeron with many fine hairs;

paraproct not divided; basimere different... .

VOL 10s now:

1973

cretatus

khmerus n. sp.

phnomus 0. sp.

vallistris

Rompongus N. sp.

fragilis

gig Mad we eres $1

clavatus

ey ae 12

Klein: Aedes (Neomacleaya) in Cambodia 15

12(11) Paraproct very long and wavy; basimere with

a short apical projection and a small subapi-

cal thumb- like process (Fig. 7 in Delfinado

1G) Se en See 3 eee Fe bese ae oS ie ee cyrtolabis

Paraproct very short; basimere with 2 long

slender apical projections, the one bearing

2-3 large spines, the other being split at tip

(Fig. 19 in Delfinado 1967)... . < 6'<-2. sete, pha Oe uNCUS

ACKNOWLEDGEMENTS

The author is indebted to Dr. Botha de Meillon, Principal Investigator

of SEAMP, for advice, helpful re-examination of material in the USNM, re-

viewing the manuscript and seeing it through the press. The author is also

especially appreciative of the assistance of MM. Kim Suor and Lim Thou,

from Institut Pasteur of Cambodia, Phnom-Penh, who accompanied hime very

helpfully on his field trips.

LITERATURE CITED

BARRAUD, P.J.

1934. The fauna of British India, including Ceylon and Burma.

Diptera 5, family Culicidae, tribes Megarhinini and Culicini.

Taylor & Francis, London, 463 pp., illus.

BRUG, S.L.

1931. New Culicidae from Sumatra. Tijdschr. Ent. 74: 245-250.

DELFINADO, M.D.

1967. Contributions to the mosquito fauna of Southeast Asia. I. The

genus Aedes, subgenus Neomacleaya Theobald in Thailand.

Contr. Amer. ent. Inst. 1(8): 1-35.

1968. Contributions to the mosquito fauna of Southeast Asia. III.

The genus Aedes, subgenus Neomacleaya Theobald in South-

east Asia. Contr. Amer. ent. Inst. 2(4): 1-74.

EDWARDS, F. W.

1917. Notes on Culicidae, with descriptions of new species. Bull.

ent. Res. 7: 201-229.

1928. Mosquito notes. VII. Bull. ent. Res. 18: 267-284.

LAFFOON, J.

1946. The Philippine mosquitoes of the genus Aedes, subgenus

Aedes. J. Wash. Acad. Sci. 36: 228-245.

LEICESTER, G. F.

1908. The Culicidae of Malaya. Stud. Inst. med. Res. F.M.S.

3(3): 18-261.

STONE, A., K.L. KNIGHT and H. STARCKE

1959. A synoptic catalog of the mosquitoes of the world (Diptera:

Culicidae). Ent. Soc. Amer. (Thomas Say Found. ), Wash. ,

D.C., 358 pp.

DERMAJOENSIS

: hike: igi

whee Me

elltageee AU

JUL ve

OAnam

KHMERUS

i) ES

ty [4g

iris

cat

~tRaty,

‘

—s.

mye

SSC

ST A

Kad f

oy ee

Sees = Sg

Rar

alps

’

ff Rinanlg

S «

wer)

La . ‘

° rf ah

cs Oar ay, or

Cay H

‘ a 4 :

4 ay

vs op

4 an “s

e . R a eh a

‘ ° ° fs z Sc

“ta ‘ 5 ° try

gyre ‘Ss ons

sane on es ing

¢ .

A i

SRO

‘

: .

> .

‘ as

* Oe 8g

/ At

CYRTOLABIS

II II]

FRAGILIS PHNOMUS

O,i am

wmemwawreaccaat?

KOMPONGUS

Klein: Aedes (Neomacleaya) in Cambodia ; 21

INDEX

Italicized pages are those which begin the primary treatment of the

taxon. Numbers in parenthesis refer to figures illustrating the species in

question.

andamanensis

cautus

clavatus

cretatus

cyrtolabis

dermajoensis

dux

fragilis

hamistylus

hispidus

incertus

indecorabilis

indicus

khmerus

kompongus

notabilis

pahangi

panayensis

phnomus

protuberans

rarus

robertsi

siamensis

stungus

torosus

uncus

uniformis

vallistris

» ty By Oe LZ, 14

Boxe, 14

ae ae ge

ae Oo ete A ee

46, 0% AALS fls6

LA py 2do

Hei el: a 5 Ps ee De

—_

OO, abe Bape Ee

Be» Sunt ode (6)

S, 12 14, (20)

9, 13

@e-~ ~ ~

~~.

)

Ral Deis ¢o deme We Wale e-em hi 2

KOm RE DOF ARONEN FEF ORUORHPHEHHHE pe

Sy LUG hy 1S

4, 2hOp ath. 12,4

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. - XVIII.

A RECONSIDERATION OF DICEROMYIA THEOBALD WITH THE

INCLUSION OF AEDES NUMMATUS EDWARDS AND AEDES

PSEUDONUMMATUS NEW SPECIES (DIPTERA: CULICIDAE)!

John F, Reinert2

ABSTRACT

Aedes nummatus Edwards is redescribed and the adult habitus, female

and male genitalia and larva are illustrated. Aedes pseudonummatus, new spe-

cies, is described and the female habitus and genitalia are illustrated. These

2 species are placed in the subgenus Diceromyia Theobald, nummatus being

transferred from the subgenus Aedimorphus Theobald. A map shows the

distribution of the above 2 species. Characters of Diceromyia are compared to

related subgenera of Aedes Meigen.

INTRODUCTION

Edwards (1923: 4) in his original description of Aedes nummatus

placed it in the subgenus Aedimorphus Theobald but stated that the reduced

male palpi and the thoracic adornment differed strikingly from all other

known species of the subgenus. He later (1932: 166) placed it in a monotypic

species group (Group B) of Aedimorphus. Barraud (1928: 667, 1934: 265) also

considered it as belonging to Aedimorphus but as an aberrant species differeing

in many respects from the other species in the subgenus. Reinert (1970b: 130)

indicated that nummatus possessed morphological and ecological resemblances

to the subgenus Diceromyia Theobald of Aedes.

Since the revision of the subgenus Diceromyia (Reinert 1970a) and

during the revision of the subgenus Aedimorphus (Reinert 1973a) it became

evident that Aedes nummatus shared many important features with both the

Oriental and African members of the subgenus Diceromyia and possessed

IThis work was supported in part by Research Contract No. DA-49-193-MD-

2672 from the U. S. Army Research and Development Command, Office of

the Surgeon General, and carried out at the Southeast Asia Mosquito Project,

Smithsonian Institution, Washington, D. C. 20560.

2Major, Medical Service Corps, U. S. Army, Department of Entomology,

Walter Reed Army Institute of Research, Walter Reed Army Medical Center,

Washington, D. C. 20012.

Reinert: Reconsideration of Aedes (Diceromyia) 23

fewer characters in common with the subgenus Aedimorphus. This species,

along with the following new species, appears morphologically to form a con-

necting link between the African and Oriental species of Diceromyia; therefore,

because of the similarities of these 2 species to each other and to this sub-

genus, I am placing pseudonummatus, new species, in the subgenus Diceromyia

and transferring nummatus to this subgenus. Notes on the subgenus Dicero-

myta and a comparison of characters with related subgenera are presented in

the taxonomic discussion section of zummatus.

Nomenclature and chaetotaxy used for females, males and male geni-

talia follow Knight (1970) and Knight and Laffoon (1970a, 1970b, 1971a, 1971b).

The terminology of the female genitalia follows Laffoon and Knight (1971) and

Reinert (1973a). Illustrations of nummatus (Figs. 1-4) and pseudonummatus

(Figs. 4-5) and a map (Fig. 6) showing their geographical distributions are

included.

AEDES (DICEROMYIA) NUMMATUS EDWARDS

(Fies, 1,.2, 3. 4)

Aédes (Aédimorphus) nummatus Edwards 1923, Bull. ent. Res. 14: 4 (ch 9);

Edwards 1932, Genera Insec., Fasc. 194: 167; Barraud 1934, Fauna Brit.

India, Diptera 5: 265 (o*, 9, L*).

Aedes (Aedimorphus) nummatus Edwards, Barraud 1928, Indian J. med. Res.

15: 667 (o%*, 9); Stone et al. 1959, Thomas Say Found. 6: 195.

FEMALE (Fig. 1). Head. Antenna dark brown, approximately 0. 87

length of proboscis, pedicel dark brown with several small, broad, brown

scales and a few short, fine, brown hairs mesally, flagellomere 1 with basal

0.40 pale and with a few small, broad, brown scales; clypeus dark brown,

bare; maxillary palpus dark brown scaled, approximately 0.19 length of pro-

boscis; proboscis dark brown scaled, approximately 1.10 length of femur I;

vertex covered with broad, decumbent, dark brown scales and with a small

posteromedian triangular patch of dusky-white ones; ocular line with a narrow

row of broad white scales; lateral surface covered with broad dusky-white scales

and connected to ocular line of white scales; occiput with a number of golden,

erect, forked scales; eyes contiguous in front; 1 pair of long brown interocular

setae; 8 long brown ocular setae. Thorax. Scutal integument dark brown;

scutum covered with narrow curved dark reddish-brown scales with a large

circular patch of narrow curved snowy-white scales covering most of anterior

0.50 of scutum but not reaching anterior and lateral margins which are reddish-

brown scaled; prescutellar space bare; scutellum with a large patch of broad

brown scales on median lobe and narrow curved brown ones on lateral lobe;

following setae brown and well developed: 2-3 long and 3-4 short anterior prom-

ontory, scutal fossal (4-6 anterior, 3-4 lateral and 1 posterior), dorsocentral

(0-1 in posterior of pale scaled area and several posterior ones), 34-49 supra-

alar, several posterior medial scutal, 1 postalar callar and scutellar (3-4 long

and 3-4 short on lateral lobe, 4-5 long and 5-6 short on median lobe), other

setae absent; pleural integument dark brown; antepronota normal size, widely

separated, each with narrow curved reddish-brown scales, 12-15 long brown

setae; postpronotum with narrow curved reddish-brown scales dorsally and 4-5

moderately broad white ones below, 3 golden posterior setae, upper one short

and other 2 long; propleuron covered with broad snowy-white scales, 7-9 golden

setae; prosternum bare; postspiracular area with 2-3 broad snowy-white scales;

24 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

mesepisternum with a large patch of broad snowy-white scales on upper area

and extending ventrally over posterior area, 4-5 upper and 16-19 posterior

golden-brown setae, lower ones shorter; prealar knob with 9-12 golden setae;

paratergite narrow, bare; mesepimeron with a patch of broad snowy -white

scales on upper area, 11-12 golden setae on upper area posterior to scale

patch; other pleural areas bare. Legs. Coxae [I-III each with several golden

and brown setae, I with broad light brown scales and a small dorsal patch of

snowy -white scales on anterior and lateral surfaces, II and III each witha

patch of broad snowy-white scales on anterior surface; trochanters I-III each

with several short pale setae and a patch of snowy-white scales; femora I and II

with dark brown scales on anterior surface, I also with a large number of pale

scales on basal 0.30-0.35, II also with a few pale scales at base and similar

ones forming an incomplete, anteroventral, longitudinal stripe on apical 0. 50-

0.60, III covered with snowy -white scales and a narrow longitudinal stripe of

brown scales on dorsal 0. 50-0. 60 which overlaps slightly onto anterior sur-

face and greatly onto posterior surface, I with posterior surface snowy-white

scaled with a narrow, longitudinal, brown scaled stripe on dorsoapical 0. 25-

0.30 and one on ventroapical 0. 50-0. 60, II with posterior surface snowy-white

scaled with a posterodorsal, narrow, longitudinal, brown scaled stripe on

apical 0. 40-0. 50; tibiae I-III. covered with dark brown scales, I also witha

posteroventral, longitudinal, snowy-white scaled stripe from base to apex, II

also with a posteromedian, longitudinal, snowy-white scaled stripe from base

to apex; tarsi I-II. dark brown scaled, II and III also with a ventral, longitudinal,

pale scaled stripe on tarsomere 1; posttarsi I-III each with 2 ungues, I and II

with ungues equal in size, each with a tooth, III with ungues equal in size and

simple. Wing. Dorsal and ventral veins with brown scales; alula with moder-

ately broad brown scales along margin and a short second row above; 1-2

remigial setae; upper calypter with numerous dark hairs. Halter. Pedicel pale;

capitellum covered with snowy -white scales and a few dusky ones at base. Ab-

domen. TergaI-VII each brown scaled with a large laterobasal triangular spot

of snowy -white scales, spot largest on II and smaller on others; sterna white

scaled with dusky scales along posterior margins; terga and sterna with numer-

ous golden setae, mostly along posterior margins. Genitalia (Fig. 2). Tergum

VIII with base and apex each slightly concave, 0.65 retracted into segment VI,

several long stout setae on apical margin and a number of short and a few

moderately long ones on apical 0.35, covered with minute spicules, numerous

broad scales covering most of surface, index 0.49; sternum VIII with base

slightly concave, apex with a small median indentation and a small lobe on each

side of midline, numerous setae and broad scales covering most of surface,

covered with minute spicules, index 0.72; tergum IX short, moderately pig-

mented, apex with a small median indentation and with 4-5 setae on each side

of midline, covered with minute spicules; insula long, tongue-like, covered

with minute spicules and with 2 small tuberculi on apical 0.20; lower and upper

vaginal lips narrow, moderately pigmented, covered with small spicules; upper

vaginal sclerite large and moderately pigmented; postgenital lobe moderately

long and broad, apex broadly rounded, several short and 5-6 long setae on

apical 0.45, covered with small spicules; cercus short, apex broadly rounded

with a few long stout setae, completely covered with minute spicules, dorsal

surface with a number of short and moderately long setae and broad scales,

index 2.15; 3 pigmented, spherical seminal capsules, 1 large and 2 slightly

smaller ones.

MALE (Fig. 1). Similar to female in general habitus but differs as

follows. Head. Maxillary palpus brown, slender and 5 segmented, segments 1,

Reinert: Reconsideration of Aedes (Diceromyia) 25

2 and 5 small, apical segment slightly downturned with 4-5 short and 2-3

longer dark brown setae at apex, 3-4 short dark brown setae ventrally at apex

of segment 3, overall length approximately 0.42 length of proboscis; vertex

and lateral surface of head covered with broad, decumbent, white scales with

light brown ones intermixed. Thorax. Antepronotum with narrow curved

brown scales and a few broad pale brown ones; propleuron with 5 setae;

mesepimeron with a patch of 9-11 upper setae and 1 lower one. Legs. Femur

II with posterior surface white scaled with a triangular dorsoapical brown

scaled spot which tapers to a point near mid femur; posttarsi I-III each with

2 ungues, I and II with ungues unequal in size, each with a tooth, III with ungues

equal in size and simple. Abdomen. Tergum VIII white scaled; sternum VII

brown scaled. Genitalia (Fig. 3). Tergum IX bilobed with 5-7 setae on each

lobe, entire surface covered with minute spicules; gonocoxite short and

moderately broad, dorsal surface with numerous moderately long setae,

lateral surface with a number of long stout setae from base to apex, ventral

surface with a number of short setae mesally on basal 0.50 and numerous

moderately long and long stout setae on distal 0.50, scattered scales on

lateral and ventral surfaces; gonostylus with basal 0.34 broad and apical 0, 66

forked into a slightly longer, narrow, lateral arm with a minute seta at the

apex and a mesal, narrow, slightly shorter arm with a short apical, pigmented

gonostylar claw; basal mesal lobe with an apical, moderately long, flattened

seta and 4-5 shorter setae, distal one of these somewhat stouter, entire sur-

face covered with short hair-like spicules; proctiger moderately long, para-

proct moderately to heavily pigmented and with 3 bluntly pointed teeth apically,

cercal setae absent; phallosome with aedeagus with 2 lateral plates connected

basally, each plate with 7-8 long, longitudinal, lateral teeth with tergally

curved apices and covered with a very lightly pigmented dorsal flap which has

scattered hair-like spicules on sternal surface, paramere moderately long,

approximately 0.65 length of lateral aedeagal plate, parameral apodeme

broad basally and tapering into a long, narrow, distal arm; sternum IX large,

entire surface covered with minute spicules, 4 setae near center (lateral 2

shorter).

PUPA and EGG. Not known.

LARVA (Fig. 4). The following description is from 2 badly damaged

skins, one of them from the holotype. Head. Seta 0-C minute and single; 1-C

long, stout and single; 3-C short and single; 4-C short and 12-14 branched;

8-C short and 4 branched; 9-C short and triple; 10-C short and 4 branched;

11-C moderately long, barbed and 5-6 branched; 12-C short and 9-10 branched;

13-C moderately long and 7 branched; 14-C short and single; 15-C short and

double to 5 branched; 6-MP short and single; mental plate with 24-25 teeth.

Antenna, Moderately long, apical 0.50 slightly incurved, moderately pig-

mented with a few small spicules on basal 0.50 of shaft; 1-A attached 0. 49

from base; 2, 4-A long, approximately equal in length; 3-A approximately

0.50 length of 2-A. Thorax. Seta 0-P with 18-20 branches; 1-P triple or 4

branched; 2, 6, 9, 10, 12-P single; 3-P with 5 branches; 4, 7, 8-P triple;

9-7-P long and barbed; 11-P single or double; 14-P double or triple; 1, 8-M

single; 2-M single or double; 3-M double; 4-M with 4 branches; 5, 7, 11-M

single; 6-M with 4-5 branches; 5, 6-M long and barbed; 13-M with 15-18 bran-

ches; 14-M with 22-26 branches; 1-T with 5-6 branches; 2-T with 4-5 branches;

3-T with 21-22 branches; 4-T with 5 branches; 5, 11, 12-T single; 6-T double;

7-T long, barbed with 8 branches; 8-T with 8-12 branches; 13-T with 4-5 bran-

ches. Abdomen. Seta 1-I with 7-8 branches; 2-I single; 3-I double or triple;

4-I with 23-25 branches; 5-I with 5-7 branches; 6-I long, barbed with 7

26 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

branches; 9-I triple; 1-II with 24-28 branches; 2, 9, 10-II single; 3-II double to

4 branched; 5-II with 6 branches; 6-II long, barbed with 6-7 branches; 7-II

moderately long, barbed with 8-10 branches; 8-II double or triple; 11-II double;

12-II triple; 13-II with 30-40 branches; 1, 3-[II with 4 branches; 2, 8, 9-II

single; 5-III with 5 branches; 6, 10, 11, 13-ID[ double; 7-IIl with 10-14 branches;

12-III triple or 4 branched; 1, 10, 11-IV triple; 2, 8, 9-IV single; 3-IV with 4

branches; 4, 12, 13-IV double; 5-IV with 6 branches; 7-IV with 6-9 branches;

2, 8, 9, 12-V single; 3-V double; 5-V triple; 6-V long and double; 7-V with

7-11 branches; 10, 11-V double or triple; 13-V with 4 branches; 6-VI long and

double; 1-VII long and single; 2, 11-VII single; 3-VII with 4-6 branches; 5-VII

with 10 branches; 6-VII with 18-27 branches; 7-VII long, double or triple;

8-VII with 28-30 branches; 9-VII with 4 branches; 10, 12-VII triple or 4 branched;

13-VII with 6-10 branches; 0, 14-VIII single; 1-VII with 5-6 branches; 2-VII

double or triple; 3-VIII long, barbed and triple or 4 branched; 4-VIII single or

double; 5-VIII barbed with 4-6 branches; comb with 21-24 scales arranged in 2

irregular rows, each scale with a long, bluntly rounded, median spine with

short denticles on lateral margins and apex; 1-X short and triple or 4 branched;

2-X long and single or double; 3-X long and single; ventral brush with 10 setae

on grid and no precratal ones, setae with 7-12 branches; 4 anal papillae, mod-

erately long and slender; saddle moderately pigmented, incompletely rings seg-

ment X, acus absent. Siphon. Moderately pigmented, index 3. 50-4.00; pecten

composed of 12-14 evenly spaced teeth, each tooth long, slender, pointed and

with a ventral row of tiny denticles from base to near apex; seta 1-S small,

triple or 4 branched, inserted 0.67-0. 73 from base; 2-S small and single.

TYPE-DATA. Aedes (Aedimorphus) nummatus Edwards, holotype

male with associated larval skin, Meenglas, Bengal, INDIA, VIII-1922,

M.O.T. Iyengar collector, adult reared from a larva collected from water ina

treehole, type deposited in British Museum (Natural History). Edwards (1923:

5) states the female allotype is in Mr. Iyengar's collection.

DISTRIBUTION. Specimens examined: INDIA, Assam, Haflong, 2

females, Aug. 1922, P. J. Barraud; Bengal, Meenglas, Jalpaiguri, 1 male

(holotype) with associated larval skin and 1 female with associated larval skin,

Aug. 1922, M. O. T. Iyengar; and Sukna, 1 male, 28 Aug. 1922, P. J. Barraud.

Other distribution. INDIA, Assam, Nongpoh, Khasi Hill Dist.; Haflong, Cachar

Hills (Barraud 1934: 267).

BIOLOGY. The type was reared from a larva which was collected from

water in a treehole (Edwards 1923: 5). Barraud (1928: 667) records larvae

collected from water in treeholes and similar situations and from a broken

bottle in the jungle. Barraud (1934: 267) also collected larvae from water in

bamboos and at an elevation of 500 feet in Sukna, India.

TAXONOMIC DISCUSSION. Aedes nummatus and pseudonummatus,

new species, posséss a greater number of morphological and ecological

similarities to the subgenera Diceromyia and Stegomyia than to the subgenus

Aedimorphus. A comparison of the adult females of these 2 species with the

other Diceromyia and related subgenera of Aedes follows: (1) head with only

decumbent broad scales on vertex and erect forked scales restricted to occiput;

Diceromyia with decumbent broad scales on vertex (a few African species with

moderately broad curved scales, e.g., flavicollis Edwards, furcifer Edwards,

taylovi Edwards) and most Oriental species have numerous erect forked scales

on occiput and none or only a very few near anterior portion of vertex, however,

periskelatus (Giles) and all African species have erect forked scales on vertex

as well as occiput; Stegomyia with decumbent broad scales on vertex and erect

forked scales restricted to occiput; Aedimorphus with decumbent, narrow, curved

Reinert: Reconsideration of Aedes (Diceromyia) 27

scales on vertex (broad scales present in punctifemoris (Ludlow) and a few

African species) and erect forked scales numerous on occiput and vertex ex-

tending to ocular line, (2) antennal pedicel with a few small broad scales and

short fine hairs mesally which is similar to both Diceromyia and Aedimorphus

but unlike Stegomyia which has a large patch of broad, overlapping, snowy -

white or silvery scales on mesal and ventral areas which usually extends onto

lower portion of lateral surface, (3) prosternum bare which is similar to both

Diceromyia and Aedimorphus except females of the vexans group which have a

few narrow scales on ventral area (scales absent in males of this group) but

unlike Stegomyia which has broad white scales on this structure, at least on

dorsolateral area, (4) propleuron of female with 7-9 setae; Diceromyia with 4-

10 setae; Aedimorphus with 10-38 setae (usually 12 or more) and Stegomyia

with 4-7 setae, (5) mesepisternum with 4-5 upper and 16-19 posterior setae

which is similar to Diceromyia (0-4 upper and 7-16 posterior) and Aedimor-

phus (2-6 upper, usually 3-4 and 10-24 posterior) while Stegomyia has 0-2

upper and 1-8 posterior setae (usually only 2-4 posterior setae), (6) acrosti-

chal setae absent which is similar to Stegomyia and most Oriental Diceromyia

(African species and the following Oriental species -- micropterus (Giles),

periskelatus and reginae Edwards with acrostichal setae) while all species of

Aedimorphus have these setae, (7) wing with 1-2 remigial setae which is

similar to Diceromyia and Aedimorphus but unlike Stegomyia in which these

setae are absent, (8) alula of wing with 2 rows of moderately broad scales (1

row on margin and a second shorter linear row above) which is similar to

Diceromyia and many Stegomyia but unlike all Aedimorphus which have a single

row of narrow scales along the margin and (9) female posttarsi I-III each with 2

ungues, I and II with ungues each with a tooth and III with ungues simple which

is similar to African species of Diceromyia (Oriental species with all ungues of

posttarsi simple), most species of Stegomyia (scutellaris group with all ungues

simple) and Aedimorphus except for a few species (e.¢., pampangensis (Ludlow)

which has ungues of III toothed, stenoetrus (Theobald), vexans vexans (Meigen)

and vexans nipponti (Theobald) which have ungues of III toothed or simple).

The female genitalia of nummatus and pseudonummatus are very similar

to both Diceromyia and Stegomyia; however, a greater number of features are

shared with the former subgenus. These 2 species resemble Oriental members

of Diceromyia in the development of tergum IX and African species of the sub-

genus in the development of tergum VIII, sternum VII and presence of numer-

ous broad scales on the cerci. Other features of the female genitalia are

similar to both the Oriental and African species of the subgenus. Female

genitalia of Stegomyia differ primarily in the development of ter gum VIII and

sternum VIII and may have broad scales present or absent on the cerci depend-

ing on the species. Both species differ sharply from Aedimorphus by the

short, blunt cerci with numerous scales, shape and presence of numerous

scales on sternum VIII and tergum VIII, and postgenital lobe which has a broad-

ly rounded apex. Aedimorphus species have long cerci with none to only a few

moderately broad scales, 0-4 scales on a heart-shaped sternum VIU, 0-4

scales on tergum VIII and postgenital lobe with a moderately deep to deep

median apical notch.

The male genitalia of nummatus resemble African members of Dicero-

myia in the development of the basal mesal lobe, proctiger and forked gonosty -

lus (e.g., fascipalpis (Edwards), furcifer, taylori) and Oriental species in the

development of the phallosome and gonocoxite. The aedeagus of nummatus is

composed of 2 lateral plates each with several longitudinal teeth and covered with

a lightly pigmented dorsal flap which is like other Oriental species of Dicero-

28 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

myia. The subgenus Ayuvakitia Thurman possesses a similar phallosome but

the adults are easily distinguished from Diceromyia by the absence of post-

spiracular setae. Aedimorphus genitalia have the aedeagus similarly developed

to Diceromyia except the dorsal flap is heavily pigmented, gonocoxite moder-

ately long, proctiger usually with a subapical, small, thumb-like process and

the gonostylus which is greatly expanded distally or blade-like (vexans group).

Genitalia of Stegomyia lack the aedeagal dorsal flap, usually have a long slender

gonostylus with an apical gonostylar claw (albolineatus group with a gonostylus

which resembles those of several African species of Diceromyia) and proctiger

differently developed.

The adult habitus of nummatus and pseudonummatus has a large white

scaled area on anterior of the scutum which superficially resembles members

of the Aedes (Finlaya) niveus group and Aedes (Chaetocruiomyia) spinosipes

Edwards. These 2 species are easily distinguished from the Finlaya by the

aedeagus of the male genitalia which has 2 lateral toothed plates and the female

genitalia which have a tongue-like insula without setae while the Finlaya have

the male aedeagus simple, trough-like and without teeth and the female insula

is lip-like with a few long well developed setae. From spinosipes these 2 spe-

cies differ by the absence of acrostichal setae and the absence of a patch of

long, narrow, forked scales arising from base of the remigium.

The male maxillary palpi of nummatus are slender, approximately

0.42 length of the proboscis, with only a few short setae and have the apical

2 segments short which are similar to Aedes (Finlaya) okinawanus Bohart,

several species of the albolineatus group of Stegomyia (group as outlined by

Knight and Rozeboom, 1946) and the subgenus Diceromyia (except the overall

length). Aedes nummatus can be separated from okinawanus by the characters

of the Finlaya mentioned above and from the Stegomyia by a combination of

characters given by Reinert (1973b). Some of these characters of the Stegomyia

that differ from nummatus and pseudonummatus not mentioned above are:

mesepimeron without lower setae, tarsus III always with a large white dorso-

basal patch or band of white scales on tarsomeres 1 and 2 and female maxillary

palpi with a large dorsoapical spot or apical band of snowy-white scales (palpi

dark scaled in albolineatus group).

The larva of nummatus has head seta 4-C medium size, multiple

branched and mesad to 6-C, 5-C is caudad to 6-C, abdominal setae 6-I-VII

moderately long to long, 1-VII long, comb scales thorn-like, abdominal seg-

ment X without precratal setae in ventral brush, saddle and siphon without an

acus, and larval habitat in water in bamboos and treeholes. This combination of

larval features differs from Aedimorphus but fits Diceromyia and is somewhat

similar to Stegomyia; however, the presence of 10 setae in the ventral brush

on segment X, each of which has 7-12 branches, resembles Aedimorphus ex-

cept for the absence of precratal setae.

From the above comparison of nummatus and pseudunummatus with the

subgenera of Aedes it becomes evident they share many important features

with Diceromyia and fewer with other subgenera and are therefore placed in

Diceromyia but in a separate species group. Also from a comparison of fea-

tures it appears the subgenus Diceromyia contains species that share a number

of characters with both Aedimorphus and Stegomyia but with an apparent closer

relationship with the latter subgenus.

Aedes nummatus and pseudonummatus adults are easily identified from

the other species of Diceromyia by the large white scaled patch on the anterior

0.50 of the scutum. The larva of nummatus is distinguished from other mem-

bers of the subgenus by the pecten teeth which are long, slender and with hair-

Reinert: Reconsideration of Aedes (Diceromyia) 29

like denticles along the ventral margins.

AEDES (DICEROMYIA) PSEUDONUMMATUS, NEW SPECIES

(Figs. 4, 5)

FEMALE (Fig. 4). Head. Antenna dark brown, approximately 0.90

length of proboscis, pedicel dark brown with a few small, broad, brown scales

and a few short, fine, brown hairs mesally, flagellomere 1 with basal 0. 50

pale and with several small, broad, brown scales; clypeus dark brown, bare;

maxillary palpus dark brown scaled, approximately 0.19 length of proboscis;

proboscis dark brown scaled, approximately 1.08 length of femur I; vertex

covered with broad, decumbent, dark brown scales with a posteromedian

triangular patch of dusky-white ones, patch with posterior margin moderately

broad and tapered to a long narrow apex before reaching ocular line which has

a narrow row of similar scales; lateral surface covered with broad dusky -white

scales and connected to ocular line of similar scales; occiput with a number of

golden, erect, forked scales; eyes contiguous in front; 1 pair of long brown

interocular setae; 8 long brown ocular setae. Thorax. Scutal integument dark

brown; scutum covered with narrow curved scales with prescutellar space bare,

posterior 0.50 of scutum with dark reddish-brown scales and small patches of

similar scales on anterior promontory and anterior scutal fossal areas, anterior

0.50 of scutum covered with snowy-white scales extending to lateral margins

and a few similar scales along lateral margins of prescutellar bare space;

scutellum with a large patch of broad snowy-white scales on median lobe and

narrow curved snowy-white scales on lateral lobe; following setae brown and

well developed: 4 long and 3 short median anterior promontory, scutal fossal (8-9

anterior, 2-3 lateral and 2 posterior), dorsocentral (2-3 in pale scaled area

and several posterior ones), 52-57 supra-alar, several posterior medial scutal,

1 postalar callar and scutellar (4-5 long and 3-4 short on lateral lobe, 6 long

and 6 short on median lobe), other setae absent; pleural integument dark brown;

antepronota normal size, widely separated, each with narrow curved snowy -

white scales, 12 long golden and brown setae; postpronotum with narrow curved

scales, reddish-brown ones dorsally and a small lower posterior patch of snowy -

white ones, 4 posterior golden setae, upper and lower ones short and middle 2

long; propleuron covered with broad snowy-white scales, 9-10 golden setae;

prosternum bare; postspiracular area with a posterior patch of broad snowy -

white scales, 5 long golden bristles; subspiracular area with a small patch of

5 broad snowy-white scales; mesepisternum with a large patch of broad snowy-

white scales on upper area and extending ventrally over posterior area, 4 upper

and 19 posterior golden setae, lower ones shorter; prealar knob with 15-16

golden setae; paratergite narrow, bare; mesepimeron with a large patch of broad

snowy -white scales on upper area, 17-18 short golden setae on upper area

posterior to scale patch; other pleural areas bare. Legs. Coxae I-III each with

several golden and brown setae, I with broad snowy-white scales on anterior and

lateral surfaces, a small patch of broad pale brown scales near middle of pale

area, II and III each with a patch of broad snowy-white scales on anterior surface;

trochanters I-III each with several short pale setae and a patch of snowy-white

scales; femora I and II with broad dark brown scales on anterior surface, I also

with a large number of pale scales on basal 0.33, II also with a narrow, antero-

ventral, longitudinal, snowy-white scaled stripe from base to near apex, II with

anterior surface covered with snowy-white scales and a narrow, dorsal, longi-

tudinal brown scaled stripe on apical 0.40, I with posterior surface snowy-white

30 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

scaled with a narrow, posteroventral, longitudinal, dark brown scaled stripe

on apical 0.75, II and III each with posterior surface snowy-white scaled with

a posterodorsal dark brown scaled area on apical 0. 40; tibiae I-III each dark

brown scaled, I also with a posteroventral, longitudinal white scaled stripe

from base to apex, II also with a posterior, longitudinal, wide white scaled

stripe from base to apex, III also with a ventral, longitudinal, indistinct, pale,

scaled stripe from base to near apex; tarsi I-III each dark brown scaled, II

also with a posterior, longitudinal, narrow pale scaled stripe on tarsomeres

1-3, II also with a ventral, longitudinal, narrow, pale scaled stripe on tarso-.

meres 1 and 2; posttarsi I-III each with 2 ungues, I and II with ungues equal in

size, each with a tooth, III with ungues equal in size and simple. Wing. Dorsal

and ventral veins covered with dark brown scales except for a small patch of

snowy-white scales at base of costa; alula with moderately broad brown scales

along margin and a short second, longitudinal row above; 2 remigial setae,

upper calypter with numberous dark hairs. Halter. Pedicel pale; capitellum

covered with snowy-white scales and with brown ones dorsally and mesally.

Abdomen. TergaI-VII each dark brown scaled with a large laterobasal patch

of snowy-white scales; sterna snowy-white scaled, V-VIII each also with a

posterior, narrow dark brown scaled band; terga and sterna with numerous

short golden setae, mostly along posterior margins. Genitalia (Fig. 5). Ter-

gum VIII with base and apex slightly concave, 0.50 retracted into segment VII,

several long stout setae on apical margin and a number of short and a few

moderately long ones on apical 0.42, covered with minute spicules, numerous

broad scales covering most of surface, index 0.50; sternum VIII with base

slightly concave mesally, apex with a small median indentation and a small lobe

on each side of midline, numerous setae and broad scales covering most of

surface, covered with minute spicules, index 0.70; tergum IX short, moder-

ately pigmented, apex with a small median. indentation and with 6-7 setae on

each side of midline, covered with minute spicules, index 0.59; insula long,

tongue-like, covered with minute spicules and with 2 small tuberculi on apical

0.20; lower and upper vaginal lips narrow, moderately pigmented, covered

with small spicules; upper vaginal sclerite large and moderately pigmented;

postgenital lobe moderately long and broad, apex broadly rounded, 23 short

and 4 long setae on apical 0.50, covered with small spicules, dorsal PGL

index 0. 80, ventral PGL index 1.30; cercus short, apex broadly rounded with

a few long stout setae, completely covered with minute spicules, dorsal sur-

face with a number of short and moderately long setae and broad scales, index

2.02; 3 pigmented, spherical seminal capsules, 1 large and 2 slightly smaller

ones.

MALE, PUPA, LARVA and EGG. Not known.

TYPE-DATA. Holotype female (genitalia removed and mounted ona

microscope slide, preparation number T72.610), Doi Jom Jang, near Wat Phra

Bat Mon Kaeo, Amphoe Sanpatong, Chiang Mai, THAILAND, 25 September 1963,

Mr. Seham Esah collector, collection number CM 119-1-33, female reared from

a larva collected from water in a treehole, at an elevation of 382 meters, type

deposited in U. S. National Museum (Natural History), Washington, D.C.

Unfortunately the immature skins were lost and I did not see them.

DISTRIBUTION. Known only from the type-locality.

BIOLOGY. The holotype female was reared from a larva which was

collected from water in a treehole.

TAXONOMIC DISCUSSION. Aedes pseudonummatus is very similar in

adult habitus and female genitalia to nummatus but can be separated by the

following features: pseudonummatus possesses: scutum with a large patch of

Reinert: Reconsideration of Aedes (Diceromyia) 31

snowy -white scales on anterior 0.50 which extends to lateral margins of scu-

tum; scutellum with snowy-white scales on each lobe; 52-57 supra-alar setae;

antepronotum with snowy-white scales; mesepimeron with 17-18 setae on upper

posterior area; and female genitalia with 6-7 setae on each lobe of tergum IX

while nummatus possesses: scutum with a large patch of snowy-white scales

on anterior 0.50 which does not reach lateral margins of scutum; scutellum

with dark brown scales on each lobe; 34-49 supra-alar setae; antepronotum with

reddish-brown scales; mesepimeron with 11-12 setae on upper posterior area;

female genitalia with 4-5 setae on each lobe of tergum IX. Additional informa-

tion is included in the taxonomic discussion section of nummatus.

ACKNOWLEDGEMENTS

I am grateful to Dr. Botha de Meillon, Principal Investigator, Southeast

Asia Mosquito Project (SEAMP) and Lieutenant Colonel Bruce F. Eldridge,

Chief of the Department of Entomology, Walter Reed Army Institute of Research

for reviewing the manuscript. Special thanks are given to Dr. Peter F. Mattingly,

Department of Entomology, British Museum (Natural History), who kindly per-

mitted me to examine the type of nmummatus and other specimens in the British

Museum. Gratitude is expressed to Lieutenant Colonel Alexander A. Hubert,

U. S. Army 406th Medical Laboratory, Tokyo, Japan, and his staff of artists,

Mr. Young T. Sohn (SEAMP) and Mr. Vichai Malikul (SEAMP) for preparing

the illustrations, and Mrs. Janet Rupp for typing the manuscript for offset

reproduction. I am especially appreciative to my wife, Mollie, who typed the

drafts.

LITERATURE CITED

BARRAUD, P. J.

1928. A revision of the culicine mosquitoes of India, Part XXIII. The

genus Aedes (sens. lat.) and the classification of the subgenera.

Descriptions of the Indian species of Aedes (Aedimorphus),

Aedes (Ochlerotatus), and Aedes (Banksinella), with notes on

Aedes (Stegomyia) variegatus. Indian J. med. Res. 15 (3): 653-

669.

BARRAUD, P. J.

1934. The fauna of British India including Ceylon and Burma. Diptera

V, family Culicidae, tribes Megarhinini and Culicini. Taylor

and Francis, London. 463 p.

EDWARDS, F. W.

1923. Mosquito notes. --IV. Bull. ent. Res. 14 (1): 1-9.

EDWARDS, F. W.

1932. Genera insectorum. Diptera, Fam. Culicidae. Fasicle 194.

Desmet-Verteneuil, Brussels. 258 p.

KNIGHT, K. L.

1970. A mosquito taxonomic glossary I. Adult head (external).

Mosq. Syst. Newsletter 2 (1): 23-33.

32 Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

KNIGHT, K. L. and J. L. LAFFOON.

1970a. A mosquito taxonomic glossary III. Adult thorax. Mosq. Syst.

Newsletter 2 (3): 132-146.

KNIGHT, K. L. and J. L. LAFFOON.

19 70b. A mosquito taxonomic glossary IV. Adult thoracic appendages.

Mosq. Syst. Newsletter 2 (4): 165-177.

KNIGHT, K. L. and J. L. LAFFOON.

1971a. A mosquito taxonomic glossary V. Abdomen (except female

genitalia). Mosq. Syst. Newsletter 3 (1): 8-24.

KNIGHT, K. L. and J. L. LAFFOON.

1971b. A mosquito taxonomic glossary VII. The larval chaetotaxy.

Mosq. Syst. Newsletter 3 (4): 160-194.

KNIGHT, K. L. and L. E. ROZEBOOM.

1946. The Aedes (Stegomyia) albolineatus group (Diptera, Culicidae).

Proc. Biol. Soc. Wash. 59: 83-98.

LAFFOON, J. L. and K. L. KNIGHT.

1971. A mosquito taxonomic glossary VI. Female genitalia. Mosq.

Syst. Newsletter 3 (2): 32-41.

REINERT, J. F.

1970a. Contributions to the mosquito fauna of Southeast Asia. -- V.

Genus Aedes, subgenus Diceromyia Theobald in Southeast Asia.

Contr. Am. ent. Inst. 5 (4): 1-43.

REINERT, J. F.

1970b. The zoogeography of Aedes (Diceromyia) Theobald (Diptera:

Culicidae). J. ent. Soc. sth. Afr. 33 (1): 129-141.

REINERT, J. F.

1973a. Contributions to the mosquito fauna of Southeast Asia. -- XVI.

Genus Aedes Meigen, subgenus Aedimorphus Theobald in South-

east Asia. Contr. Am. ent. Inst. 9 (5): 1-218.

REINERT, J. F. |

1973b. Aedes wainwrighti Baisas, a synonym of Aedes (SStegomyia)

meronephada (Dyar and Shannon), with notes on the subgenus

Stegomyia Theobald (Diptera: Culicidae). Mosq. Syst. 5 (1):

27-30.

STONE, A., K. L. KNIGHT and H. STARCKE.

1959. A synoptic catalog of the mosquitoes of the world (Diptera,

Culicidae). Ent. Soc. Am., Thomas Say Found. 6: 1-358.

Dok © he

Reinert: Reconsideration of Aedes (Diceromyia) 33

LIST OF FIGURES

Aedes (Diceromyia) nummatus - adult morphology

Aedes (Diceromyia) nummatus - female genitalia

Aedes (Diceromyia) nummatus - male genitalia

Aedes (Diceromyia) nummatus - larva; pseudonummatus - adult morphology

Aedes (Diceromyia) pseudonummatus - female genitalia

Distribution of Aedes (Diceromyia) nummatus and pseudonummatus

pneseetrtretrir

perenne sii,

PT aro ealaa

jpeispcststrtrttPt trees

nummatus

* fi +

pee Ori caod

. R ry tine

WAN Ne att -

nummatus

a Ol |

EAU

ts Lesa

‘ ANS Ny

VENTRAL

nummatus

Figs)

—K—

——

SEZ

—

sign 4

‘

By erat

a) ew ie

ae , Voy ae a

Fev \ van x

Ry 5 ) aL

Fly ‘

Cercus a

\ oa

‘ /

tuts, o ee

eS pane”

ia ze Aatet

| WA. S yas

anes LM ier

\ HS y

{ \ ; ;

R DIMGIRGHEE

Lane §

<==

z LZ =

VMZZZE

_—_ BE

— =

——

Postgenita

Lobe

— DSRS = ws

OOO =

Seats LS ae Sh

——s 2

= ==

: fe

Bee eg ce

ace

Fae.

Peed ee

- ee

cad

oe eae, ic ¢ eo

Say inp: CL —— -

poner (7 See,

—=

Upper

Vaginal

Lip

Upper

Vaginal

Sclerite

Lower __

Vaginal Lip

Insula ee

Tak

Tuberculus ——

Seminal

Capsules ae

VENTRAL

i t A if

ae y é Nh

th m q w ‘A i vy Ap

Lon ce

| | Pia

a ee ih ve mt

ag! Wee

\ y et nee 4

\ ny nw.

\ \ ) ie |

‘ \ nae

SAIN SS a Ao nae

Serge : tN Mii Ky WET eh fee aang oe

f Ar va SMP ACR NO NAL Le

rea A y

~ cn \ cua, (ay

i a hs. SS ° BAN \ . :

, x Bt a

Vill-Tergum BX Sy

m “ Mi i it

——=

——

=————

—S——

ers

i alah

—

SS SS

——<——

SSS

—s

S53

——

Kk :

e ae 2

b NS EN

‘ e OA:

Ne \it

“0 \\

| AWAY:

-S—

Cercus

if ie y)

\ Vi Vy} lay

——

\ jy.

pseudonummatus

IX-Tergum

BANGLADESH

CAMBODIA

e-nummatus

w- pseudonummatus

Contrib. Amer. Ent. Inst., vol. 10, no. 1, 1973

INDEX

Valid names are set in roman type. The italicized pages are those

which begin the primary treatment of the taxon. Numbers in parenthesis

refer to the figures illustrating the species in question.

Aedes 22, 26, 28

Aedimorphus 22, 23, 26, 27, 28

albolineatus group 28

Ayurakitia 28

Chaetocruiomyia 28

Diceromyia 22, 23, 26, 27, 28

fascipalpis 27

Finlaya 28

flavicollis 26

furcifer 26, 27

micropterus 27

niveus group 28

nummatus 22, 23, 26, 27, 28, 30,

a1 (4, 2,°S,,. 4)

okinawanus 28

pampangensis 27

periskelatus 26, 27

pseudonummatus 22, 23, 26, 28, 29,

30 (4, 5)

punctifemoris 27

reginae 27

scutellaris group 27

spinosipes 28

Stegomyia 26, 27, 28

stenoetrus 27

taylori 26, 27

vexans group 27, 28

vexans nipponii 27

vexans vexans 27

hala

5) =

Re Pe :

Des

ere

Contributions

of the

American Entomological Institute

Volume 10, Number 2, 1973

MOSQUITO STUDIES (Diptera, Culicidae)

XXXII. A revision of the genus Haemagogus

By J. Hal Arnell

CONTENTS

INTRODUCTION .. .

MATERIAL AND METHODS

SYSTEMATICS

Taxonomic History .

Taxonomic Characters

Proposed Classification

Distribution

Affinities. . .

BIONOMICS AND MEDICAL IMPORTANCE

TAXONOMIC TREATMENT . hee

Genus Haemagogus .

Keys to Groups and Species.

Subgenus Conopostegus .

Subgenus Haemagogus

Albomaculatus Section ,

1. Haemagogus (H.) eo.

2. Haemagogus (H.) chrysochlorus .

3. Haemagogus (H.) spegazzinii

4. Haemagogus (H.) baresi

5. Haemagogus (H.) andinus

6. Haemagogus (H.) nebulosus

7. Haemagogus (H.) janthinomys

8. Haemagogus (H.) capricornii

9. Haemagogus (H.) mesodentatus .

10. Haemagogus (H.) albomaculatus .

11. Haemagogus (H.) panarchys

12. Haemagogus (H.) soperi

13. Haemagogus (H.) acutisentis

14. Haemagogus (H.) equinus

Tropicalis Section : :

15. Haemagogus (H.) tropicalis ;

Splendens Section .. .

16. Haemagogus (H.) splendens .

17. Haemagogus (H.) celeste .

18. Haemagogus (H.) iridicolor .

19. Haemagogus (H.) aeritinctus . .

20. Haemagogus (H.) regalis .

21. Haemagogus (H.) lucifer .

22. Haemagogus (H.) argyromeris .

23. Haemagogus (H.) chalcospilans

24. Haemagogus (H.) boshelli

REFERENCES CITED

FIGURES =...

TABLE OF DISTRIBUTIONS Aoi

CONSPECTUS OF TAXONOMIC CHANGES»

INDEX TO SCIENTIFIC NAMES

COTDNNDNANAMNWNNN

MOSQUITO STUDIES (Diptera, Culicidae)

XXXII. A REVISION OF THE GENUS HAEMAGOGUS’

J. Hal Arnell’

INTRODUCTION

The Neotropical genus Haemagogus contains several important, if not the prin-

cipal, vectors of jungle yellow fever. It has been the subject of several relatively

recent regional studies (Kumm, Osorno-Mesa and Boshell-Manrique, 1946; Kumm

and Cerqueira, 195la; Levi-Castillo, 1951b) and numerous restricted studies on

the taxonomy, ecology and/or disease relations of 1 or more species. However,

since Dyar’s Mosquitoes of the Americas (1928) the only study encompassing

the entire genus, in Lane’s Neotropical Culicidae (1953), has a very unsatisfactory

coverage. Recently Zavortink (1972:116) enlarged the genus by the transfer of the

subgenus Conopostegus from the subgenus Finlaya of Aedes, recognizing 4 species

and 4 unnamed forms.

In the present revision I am recognizing only 2 subgenera, Conopostegus and

Haemagogus, the latter including all the generic group taxa listed under the genus

Haemagogus by Stone, Knight and Starcke (1959). As nothing can be added to the

knowledge of the subgenus Conopostegus at this time, its treatment is restricted

here to its inclusion in the generic description and discussion and in the keys but

without consideration of the species. In the subgenus Haemagogus, treated in

detail, I am recognizing 24 species in 3 sections, the Albomaculatus Section with

14 species, the Tropicalis Section with only the nominate species and the Splendens

Section with 9 species. The taxonomy of Haemagogus has been based almost

entirely on characters of the male genitalia with little regard to the external

morphology of the adults or the immature stages. In this study the internal

classification of the subgenus Haemagogus is based on a consideration of immature

stages associated with adults and a critical comparison of usually large series of

adults of most species.

I wish to especially thank Pedro Galindo of the Gorgas Memorial Laboratory

for discussions allowing me to draw on his considerable experience with and

knowledge of the biology, habits and distribution of the Central American Haema-

gogus fauna, and for the loan of specimens. I also wish to thank John N. Belkin

for suggesting this study and for reading and criticizing the manuscript; Thomas J.

Zavortink for suggestions, advice and many hours of stimulative discussion of the

study; Willis W. Wirth for arranging the loan of specimens from the U.S. National

‘Contribution from project “Mosquitoes of Middle America’ supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

* Department of Biology, University of California, Los Angeles, California 90024.

Z Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Museum of Natural History; Peter F. Mattingly for the loan of specimens from the

British Museum (Natural History); Erwin Lindner of the Staatliches Museum fur

Naturkunde in Stuttgart for the loan of the holotype of lindneri; Oswaldo P.

Forattini for the loan of specimens of baresi from the Faculdade de Higiene e

Saude Publica; Sebastiao H. Xavier for the loan of specimens from the Centro de

Pesquisas Rene Rachou; L.T. Nielsen for the loan of specimens from the University

of Utah; William A. Powder and Sandra J. Heinemann for the preparation of

material; L. Margaret Kowalczyk and Nobuko Kitamura for preparation of the

preliminary and final drawings; Caryle A. Stallard for typing a part of the rough

draft; and Angeliki Demos for typing the remainder of the rough draft and pre-

paring the text copy for lithoprinting.

MATERIAL AND METHODS

The material for this study came primarily from collections made for the project

*““Mosquitoes of Middle America” (Belkin, Schick et al. 1965) and is deposited

at the University of California, Los Angeles [UCLA]. Additional specimens were

borrowed from the U.S. National Museum of Natural History [USNM]; the British

Museum (Natural History) [BM], including the holotypes of albomaculatus and

equinus and the lectotypes of obscurescens and splendens, the Staatliches Museum

fur Naturkunde in Stuttgart [SMNS], the holotype of lindneri; the Faculdade de

Higiene e Saude Publica, Sao Paulo [FH], including the holotype of baresi; the

Centro de Pesquisas Rene Rachou, Belo Horizonte, Brazil [BH]; Gorgas Memorial

Laboratory, Panama [GML]; the Department of Biology, University of Utah [Utah] ;

and the Natural History Museum of Los Angeles County [LACM]. A total of 17,479

specimens was examined, 3088 males, 4296 females, 3830 pupae and 6265 larvae.

Included in this total were 2419 individual rearings of which 1326 were larval

rearings, 616 were pupal rearings and 477 were incomplete.

The procedures used in this study were primarily those of comparative morpho-

logical taxonomy. The form of presentation and terminology and abbreviations

used in the descriptions follow Belkin (1962) with later modifications (Belkin,

1968a:72). Descriptions are composites of all available specimens; in 4 instances

only a single specimen was available. Illustrations were prepared in general from

a single specimen but were corrected to show the modal condition by comparison

with most available specimens. On the maps, localities from which I examined

specimens are indicated by closed symbols and records from the literature by open

symbols. Political subdivisions and locality names in the distribution lists conform

to the recommended usage in the Official Standard Names Gazeteers of the United

States Board on Geographic Names.

SYSTEMATICS

TAXONOMIC HISTORY. Williston erected the genus Haemagogus in 1896 for

his new species splendens from the island of St. Vincent. Theobald (1901:239)

recognized Haemagogus as a distinct genus but synonymized splendens with Culex

cyaneus Fabricius, 1805 and later (1903a:308; 1903b:282) added 2 new species

described from females only, albomaculatus from British Guiana and equinus from

Jamaica. The genus Cacomyia was erected by Coquillett (1906) for the latter 2

species on the basis of wing venation and long palpus in the male, although the

Arnell: Genus Haemagogus 3

male of albomaculatus was unknown and actually has a short palpus. Lutz (1904)

described Haemagogus capricornii from Brazil and later (1905) placed it in a new

genus, Stegoconops. In volume 2 (1913) of their monograph, Howard, Dyar and

Knab, illustrating male genitalia and larvae, placed 4 species, equinus, regalis Dyar

and Knab, 1906 (=albomaculatus, in part), capricornii and lucifer n. sp. in Lutz’

Stegoconops because of the presence of bristles on the postnotum of the type of

Haemagogus splendens which they referred to the Sabethini. However, in volume 4

of the monograph (1917), they recognized 4 species in the genus Haemagogus,

dividing it into 2 groups based on wing venation, female claws, male genitalia and

larval comb scales: splendens (=lucifer, in part) and albomaculatus (=regalis, in part)

in one group and equinus and capricornii in the second group. Dyar (1921), in

attempting to correct the confusion in the 2 Howard, Dyar and Knab volumes

divided the genus into 2 subgenera, Haemagogus, with simple female claws and

short male palpus and Stegoconops, with toothed female claws and long male

palpus. This interpretation was followed in Dyar’s Mosquitoes of the Americas

(1928). Edwards, in his catalog (1932) recognized that the 2 subgeneric characters

were not always correlated and redefined the subgenera strictly on the basis of the

male palpus, Stegoconops with a long palpus and Haemagogus with a short palpus.

Antunes (1939) included Stegoconops, whose type species, capricornii, has a short

male palpus in the synonymy of Haemagogus and proposed a new subgeneric name

for the species with long male palpus. This name was unavailable since no type

species was designated. Levi-Castillo (1951b) recognized the same subgenera and

proposed the name Longipalpifer for the species with long male palpus with

panarchys Dyar, 1921 as type species. Lane, in his Neotropical Culicidae (1953)

recognized 3 “groups”? in Haemagogus, corresponding, with 1 exception, to the

3 subgenera recognized later by Stone, Knight and Starcke (1959): Longipalpifer,

with long male palpus; Stegoconops, with toothed female claws, short male palpus

and larvae with a single row of comb scales; and Haemagogus, with simple female

claws, short male palpus and larvae with comb scales in a patch.

Haemagogus leucomelas, described in 1904 by Lutz in Bourroul’s thesis, was

placed in the genus Aedes by Dyar and Shannon (1924:484) and since the name

leucomelas was preoccupied they proposed leucocelaenus as a replacement name.

Dyar (1925:141) erected the subgenus Conopostegus for leucocelaenus and sub-

sequently, Edwards, in his catalog (1932) placed Conopostegus in the synonymy of

Finlaya. Zavortink (1972:116), in removing all New World aedines from the sub-

genus Finlaya resurrected the name Conopostegus for leucocelaenus and related

forms and placed Conopostegus in the genus Haemagogus.

TAXONOMIC CHARACTERS. The genus Haemagogus is one of the most highly

derived of the Aedini, yet it exhibits a number of primitive characters. It is often

difficult to determine if the expression of some character states is primitive or

derived, but evaluations can be made by determining the tendency within the taxon

and comparing the character states in other aedine groups. Derived characters in

the adults include the increase in broad, flat metallic scales, accompanied by a

reduction in thoracic chaetotaxy, enlarged pronotal lobes, the enlargement of the

hindcoxae and the development of an apical sternomesal scale patch on the side-

piece of the male genitalia. The derived condition is also expressed in the tendency

toward short male palpus, setae on the postnotum, simple claws in both the male

and female, the absence of setae on tergite IX in the male and female, specialized

scales and setae on tergite VIII in the male, and reduced number and length of

bristles on the male antenna. In the larvae, the derived condition is the reduction

of the ventral brush, fewer comb scales and a reduction in sclerotization.

4 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Adult characters. Studies of the taxonomy of Haemagogus have for the most part

neglected adult characters, other than the male genitalia. Group characters such as

the female claws, wing venation (the ratio of R,43 to R,; see fig. 18), length of

the male palpus and postnotal setae have been recognized but good species char-

acters have often been overlooked. Good specific and sometimes group characters,

in addition to the above, used in the present study are: the length of the proboscis

as compared to the length of the forefemur, the number of bristles on the male

antennal segments, the color of the integument of the coxae and trochanters, the

width of the interocular space as measured by ommatidial diameters, the presence

or absence of bristles on the lower stp, the male claws, and the color and distribu-

tion of scales on the head, thorax, legs and abdomen.

The iridescent metallic colors of the scales and their tremendous range of hue

presented a special problem. Because definitions of colors are usually a matter of

subjective interpretation and rather subtle differences in color are often reliable

specific characters in this group, it was necessary to define as accurately as possible

some of the hues that appear in Haemagogus scales. Comparisons were made with

color samples in Ridgway (1912) and, in the interest of simplicity, the terminology

was taken from Kelly and Judd (1955), except for the colors which are better

described by comparison with the metals silver, gold, copper and bronze, these

colors and terms not appearing as such in the above cited references. Gold and

silver can be defined simply as having the color of the respective metals. Copper

is similar to gold but containing hues of red. Bronze, likewise, is similar to gold

but with varying green hues. The colors violet and purple predominate on the

proboscis, palpus, wing, legs and often abdomen of most species. Both colors are

combinations of dark red and dark blue, in violet the blue predominating and in

purple the red being stronger.

Male and female genitalic characters. The species of Haemagogus have been

separated almost exclusively on the basis of the male genitalia, as this structure

is highly derived and, for the most part, exhibits reliable species and group char-

acters. Tergite VIII, often overlooked, is one of the most important male genitalic

structures in this group, offering good characters in its shape and in the setae and/or

scales on the distal margin. Tergite IX is reduced but exhibits subgeneric and group

characters. The distribution of setae, the development of the basal and apical tergo-

mesal lobes and the apical sternomesal scales of the sidepiece are important, and

the development of the clasper shows group characters and species differences in

the Splendens Section. The aedeagus has a characteristic development in most

Species, especially in those of the Albomaculatus Section, as does the claspette

in nearly all species.

In the subgenus Conopostegus and 2 species of the subgenus Haemagogus,

anastasionis and chrysochlorus, the claspette filament is inserted in a distinct alve-

olus on the apex of the claspette stem, however, in the remaining species the alve-

olus is either incomplete or entirely absent. The claspette filament is a specialized

seta, arising from an alveolus, supplied with a nerve ending and functioning as a

sensory organ. In those Haemagogus in which no alveolus is present or the alveolus

is incomplete there is then no true claspette filament. The membranous structure

at the apex of the claspette is only an elaborate development of the claspette stem,

possibly without a sensory function, and formed possibly from the failure of the

apical portion of the claspette to differentiate completely during ontogenetic devel-

opment. Although not a true claspette filament in most species, the structure is

sufficiently differentiated from the stem to be readily recognizable and will con-

tinue to be referred to as the filament.

Arnell: Genus Haemagogus 5

The female genitalia of species of Haemagogus exhibit good group characters

in the presence or absence of setae on tergite IX.

Pupal characters. The pupae of the subgenus Haemagogus are remarkably uniform

in structure and for the most part are difficult or impossible to recognize to species.

I have not found any species or group characters sufficiently reliable to prepare a

key. However there are several characters that differ somewhat from species to

species and since they are of some value they are included in the species descrip-

tions. A few species are recognizable with reasonable certainty and several distinc-

tions can be made within species complexes. These are discussed in the species

accounts. The most useful characters are the development of hair 7-C, the relative

lengths of the large dorsal hairs 3-II,III,5-IV,V, the position of hair 2-VII and the

branching of the paddle hair, 1-P. Hair 1 on segments II to VII varies in development

from small and single to very large and dendritic depending on the hairiness of the

corresponding larva and is not a taxonomically reliable character.

Larval characters. Species of Haemagogus often exhibit a wide range of intra-

specific variation in larval hairiness, so much so that the larvae have the appearance

of different species. The hairs most often affected are 7,11,14,15-C; 0,4,8,14-P;

1,14-M; 1,3,5,13-T; 1,2,4,5,9,11,13-I; 1,2,5,7,13-II-VI; 1,2,5,6,9,13-VU; 1,3,5-VIll

and 1-X, the hairs varying in length, thickness and number of branches. Rosen and

Rozeboom (1954) and Colless (1956), discussing hairiness in the Aedes (Stegomyia)

scutellaris (Walker, 1859) group and Aedes (S.) albopictus (Skuse, 1894) respec-

tively, conclude that hairiness is environmentally induced by some physical factor

in the treehole or treehole water. This generally would seem to be true in Haema-

gogus, however there are several instances in which both hairy and nonhairy larvae

have been taken in the same collection. Hairiness is accompanied, in most cases,

by increased sclerotization of the tubercles of several hairs and of the boss of the

ventral brush, and in the species with spiculose integument, an increase in the

density of spiculation. Hairiness is carried over into the, pupa but usually affects

only hair 1-II-VII, the hair being small and simple in nonhairy forms and extremely

large and dendritic in hairy forms. As in other groups with hairy and nonhairy

immatures, all forms give rise to identical adults. Several species examined in this

study, in both subgenera, consisted of entirely hairy or nonhairy forms, however

in those instances few immatures were available. All species in which a large series

of immatures was examined exhibited some variation in hairiness. Because of this

variability in larval hairiness, the role of larval chaetotaxy as a taxonomic tool is

reduced. However, some of the hairs other than those noted above are not generally

affected by variation in hairiness and their relative, rather than absolute, develop-

ment shows specific differences.

It should be emphasized that the larval illustrations cannot accurately represent

the entire range of variability of a species because of the absence of intraspecific

constancy in larval chaetotaxy. They should not be used without the keys and

species diagnoses in species determinations.

PROPOSED CLASSIFICATION. I am in full agreement with Zavortink’s inclu-

sion (1972:111-131) of the subgenus Conopostegus in Haemagogus and his treat-

ment of the included species, 4 named and 4 unnamed. As noted in the discussion

of Conopostegus below, this subgenus has retained many primitive characters and

is probably closer to the ancestral stock of the genus.

I am including in the subgenus Haemagogus all the species formerly placed in

the subgenera Haemagogus, Stegoconops and Longipalpifer. | do not consider the

subdivision into the above 3 subgenera to be justified, especially with the inclu-

6 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

sion of Conopostegus in Haemagogus, as the criteria for separating the present

subgenera Conopostegus and Haemagogus are of a different order of magnitude

than those for subdividing the subgenus Haemagogus. The 24 species included in

the subgenus Haemagogus form a relatively homogeneous assemblage of species,

all having the mesonotum completely covered with broad, flat metallic scales, a

single broad band of silver scales on the pleuron, a sclerotized dorsal process on

the aedeagus and specialized scales or setae on tergite VIII in the male genitalia,

while the subgenus Conopostegus has brown and silver mesonotal scales, 3 arcs

of silver scales on the pleuron, an aedeagus without a sclerotized dorsal process

and no specialized scales or setae on tergite VIII in the male genitalia.

I prefer to recognize 3 sections within the subgenus Haemagogus. The Albo-

maculatus Section of 14 species, with 1 exception, contains the species formerly

placed in the subgenera Stegoconops and Longipalpifer. This section is character-

ized in the adults by the absence of setae on the postnotum, toothed female

claws, a relatively short vein R, and a densely plumose male antenna; in the

female genitalia by setae on tergite IX; in the male genitalia by setae on tergite

IX and a relatively simple clasper and sidepiece; and in the larvae by 5 pair of

hairs in the ventral brush and a single row of few comb scales. The Splendens

Section of 9 species includes the nominal species formerly placed in the sub-

genus Haemagogus. This section is characterized in the adults by the presence of

setae on the postnotum, simple female claws, long vein R, and a moderately

plumose male antenna; in the female genitalia by the absence of setae on tergite

IX; in the male genitalia by the absence of setae on tergite [X and a more complex

clasper and sidepiece; and in the larvae by 6 pair of hairs in the ventral brush and

numerous comb scales in a double row or triangular patch. One species, tropicalis,

being annectent between the above 2 sections, with characters of the Albomaculatus

Section in the adult and of the Splendens Section in the larva, is placed in the

monotypic Tropicalis Section. The interrelations of these sections and the included

species are discussed below under the respective sections.

DISTRIBUTION. The distribution of the genus Haemagogus is centered in Central

America and northern South America and adjacent islands where 19 of the 28

species occur. The genus has a wide distribution in South America to northern

Argentina except for the Pacific coast south of the Gulf of Guayaquil and the

higher elevations of the Andes, and is found as far north as Sonora, Mexico on

the Pacific and Texas on the Atlantic of North America. It is known from the

island of Jamaica in the Greater Antilles and several islands of the Lesser Antilles

south of Martinique (see fig. 1).

AFFINITIES. Haemagogus is one of the more derived genera of the tribe Aedini

(in the sense of Belkin, 1962), arising in northern South America or the adjacent

Caribbean area where the largest number of species occur today, and undoubtedly

derived from the genus Aedes, although there is no element of the present New

World aedine fauna that shows any obvious affinities to Haemagogus.

Members of the Kochi Group of Aedes (Finlaya) of Southeast Asia and the

South Pacific show remarkable similarities to Haemagogus in the male genitalia,

especially in the aedeagus and proctiger, the sidepiece including the specialized

scales on the distal sternomesal margin, the clasper and the reduction of tergite IX.

However there is little in the adults or immature stages to indicate relationship to

Haemagogus.

A close affinity between Haemagogus and the Oriental aedine genus Heizmannia

has been proposed by Edwards (1922:445) and Mattingly (1957:4), among others.

Arnell: Genus Haemagogus a

The statement by Edwards (1922) of relationship was made to clarify the earlier

suggestion of Theobald (1910:588) and Howard, Dyar and Knab (1915:50) that

Heizmannia scintillans Ludlow, 1905 was a sabethine mosquito. Obviously Heizman-

nia is closer to Haemagogus than to the Sabethini. There is a striking resemblance

in many aspects of the adults of the 2 genera, however the immature stages and the

female genitalia show little evidence of close affinity and the male genitalia are

remarkably different. The most reliable group characters in the male genitalia of

the Aedini are the structure of the phallosome and proctiger according to Belkin

(1962:326), with the clasper and basal mesal derivatives of the sidepiece also im-

portant. Haemagogus has a simple aedeagus and a paraproct with cercal setae while

in Heizmannia the aedeagus is divided and has toothed lateral plates and the para-

proct lacks cercal setae. This divergence in structure indicates that the affinities

of the 2 genera lie in 2 separate branches of the Aedini. This interpretation was

also that of Edwards (1932:180). The similarities in the structure and ornamenta-

tion of the adults may be attributed to convergence, the 2 genera apparently occu-

pying much the same ecological situation, treeholes or bamboo usually in tropical

forest with peak biting activity near midday. Many of the same adult characters

are found in day flying members of the tribes Sabethini and Toxorhynchitini, such

as reduced chaetotaxy, metallic scales and often enlarged apn lobes and hindcoxae.

BIONOMICS AND MEDICAL IMPORTANCE

Females of most species of Haemagogus attack man readily and with few excep-

tions usually bite about the lower body. Several species tend to be arboreal in

habit, janthinomys being strongly arboreal and seldom descending to ground level,

and then only when the forest canopy has been disturbed. Mesodentatus, lucifer

and equinus are arboreal but not as strongly as janthinomys. Males of several species

have been observed near the hosts where they mate with females approaching for

a blood meal.

Species of Haemagogus breed primarily in treeholes and cut or broken bamboo

internodes but are often found in other plant containers such as bromeliads and

fallen fruits and occasionally in ground pools and rockholes. Species of this genus

are common in tropical rain forest, open deciduous forest and second growth and

in coastal mangrove associations, with many species being quite specific in their

habitat preferences. For example, janthinomys is found almost exclusively in trop-

ical rain forest, aeritinctus, regalis, chalcospilans and boshelli are restricted to man-

grove, and anastasionis and argyromeris are found primarily in scrub forest and

second growth and are somewhat peridomestic. Equinus appears to be the most

adaptable in terms of habitat utilization, being found commonly in rain forest,

scrub and thorn forests, mangrove, and in peridomestic situations.

The genus Haemagogus plays a primary role in the transmission of sylvan or

jungle yellow fever in Central and South America. In laboratory transmission

experiments, all species of Haemagogus tested have been found to be capable of

harboring the virus or transmitting it by bite. Janthinomys is undoubtedly the most

important vector and has been repeatedly incriminated in the jungle yellow fever

cycle. Other Haemagogus species found to be capable laboratory vectors, found

naturally infected with yellow fever virus or at least suspected to be involved in

the transmission cycle are leucocelaenus, capricornii, mesodentatus, albomaculatus,

soperi, equinus, celeste and lucifer. In addition, janthinomys has been found infect-

8 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

ed with the Ilheus virus in Panama and leucocelaenus has been found infected with

the Una virus at Belem, Brazil. The disease relations of all species except leuco-

celaenus are discussed further in the bionomics paragraphs of the species accounts.

The role of leucocelaenus in yellow fever transmission is discussed by Kumm and

Cerqueira (1951b:195-196), and in the transmission of Una virus by Causey, Casals

et al. (1961:778).

TAXONOMIC TREATMENT

Genus HAEMAGOGUS Williston

1896. Haemagogus Williston, 1896:271. TYPE SPECIES: Haemagogus splendens Williston,

1896, St. Vincent; monobasic.

For synonymy see under subgenus.

FEMALES. Small to medium-sized species; head, abdomen and legs elongate, pre-

dominantly dark scaled; head, thorax and abdomen densely covered with moder-

ately broad to broad flat scales, usually with brilliant metallic reflections; scales

of proboscis, palpus, wing and legs with metallic reflections; thoracic chaetotaxy

reduced.

Head: Integument dark brown to black. Eyes narrowly to broadly separated

above antennae, interocular space usually with broad, flat, silver scales. Frontal

bristles absent. Orbital bristles moderately to well developed, numerous, 6-12

pairs. Postgenal bristles moderately developed, numerous. Decumbent scales broad,

flat, dark to light dorsally, silver laterally, ventrally and in narrow orbital line.

Erect scales confined to single row on occiput, usually dark. Clypeus large, bare.

Proboscis slender, about 0.80-1.50 of forefemur, usually entirely dark scaled,

with basal bristles. Palpus short, about 0.12-0.21 of proboscis; 3-or 4-segmented,

segment 4 small when present; entirely dark scaled; segments 1-3 with bristles.

Antenna about 0.50-0.75 of proboscis, torus with fine setae and occasionally

small, dark scales mesally; flagellar segment 1 with a few small dark scales, seg-

ments 2-13 with basal whorl of usually 6 rather short bristles.

Thorax: Integument dark brown to black. Mesonotum strongly arched. Acros-

tichal and dorsocentral bristles well developed only on anterior promontory; supra-

alar bristles numerous and well developed; elsewhere acrostichal and dorsocentral

bristles as well as humeral and posterior fossal bristles present but minute and

visible only in slide preparations; prescutellar bristles well developed only in Cono-

postegus. Scutellum with 2-4 strong bristles on midlobe and 3-6 on lateral lobes,

several weak additional bristles on all lobes. Mesonotum with broad, flat, ap-

pressed scales over entire surface or interrupted by rather inconspicuous anterior

inner dorsocentral and median and lateral prescutellar bare spaces (Conopostegus).

Scutellum covered with broad, flat scales. Paratergite with broad, flat silver scales.

Apn lobes considerably to greatly enlarged, often nearly meeting at midline. Meron

moderately large. Bristles reduced in number on all pleural sclerites; present and

well developed on apn, moderately to well developed on ppn, ppl, pra and upper

mep, minute to well developed on psp and stp near base of midcoxa; Conopostegus

in addition with 2 or 3 well developed setae near middle of stp; sp, ssp and lower

mep bristles absent. Pleuron with broad, flat silver scales forming single broad,

vertical band from midcoxa to antealar area with scales on pra, psp, ssp, mep

Arnell: Genus Haemagogus 9

and stp; pcx, ppl and pst with silver scales, apn with silver or dark scales, ppn

bare or with silver or dark scales; or, in Conopostegus, with silver scales forming

3 nearly vertical arcs, the anterior extending from apn through pst to forecoxa,

the middle extending from ppn through ssp and lower stp to midcoxa and the

posterior extending from antealar area through paratergite, psp, upper stp, lower

mep to hindcoxa. Hypostigial area, anterior stp and metameron bare.

Legs: Hindcoxa larger than midcoxa, its base at level of or slightly below upper

margin of meron. Legs moderately long to long, forefemur 1.25-1.80 of distance

from top of thorax to apex of midcoxa. Integument of coxae uniformly dark

brown or dark brown basally grading to yellow apically; all with silver scales,

occasionally with additional small patch of dark scales in middle of forecoxa

and midcoxa. Trochanters dark brown or yellow, all with silver scales. Femora

dark scaled with silver to silvery yellow scales basally on anterior, ventral and

posterior surfaces, usually extending apically in short to long streaks; small knee

spots of silver scales on apex of midfemur and hindfemur present or absent.

Tibiae and tarsi entirely dark scaled or with white to dingy scales on outer surface

of basal midtarsal segments. Claws of foreleg and midleg simple or with subbasal

tooth, claws of hindleg simple.

Wing: Entirely dark scaled; R43 about 0.31-1.60 of R,.

Haltere: Stem usually pale, knob dark scaled with silver scales at tip.

Postnotum: Integument dark brown to black, bristles present or absent.

Abdomen: Tergite I dark scaled dorsally, laterotergite with silver scales. Tergites

II-VII with basolateral silver patches, usually extending to distal margins of I-IV

or V but constricted distally; restricted to base on V-VII; dorsal silver scales

present or absent; VII silver or dark scaled, strongly produced caudad. Sternites

II-VII dark scaled with basal silver band or basolateral silver patches; sternite VII

with few dark scales laterally.

FEMALE GENITALIA. Segment VIII: Tergite narrowed apically; on midline

about 0.52-0.71 of tergite VII; distal 0.90 with scales and bristles, or densely

scaled with bristles only on apical 0.25-0.33, apical bristles usually well developed.

Sternite about 1.30-1.62 of tergite VIII, broader distally; distal margin straight,

slightly rounded or with rounded lobe immediately laterad of midline; all but

extreme base and basolateral area with bristles, bristles more numerous distally

and along midline, predominantly weaker distally, some bristles moderately devel-

oped proximally along midline; lateral margin with scales. Tergite IX: Well devel-

oped, divided distally, V-shaped; about 0.50-0.69 of tergite VIII; moderately to

strongly sclerotized; with or without setae. Insula: Weakly sclerotized, connected

to sigma, with 1 or 2 pairs of weakly developed setae. Cercus: Relatively short,

about 0.61-0.83 of tergite VIII; apex obliquely truncate in lateral aspect; bristles

numerous, several apical ones strongly developed. Postgenital Plate: Moderately

long and broad, about 0.49-0.67 of tergite VIII; index 1.4-2.2; apex straight,

rounded or slightly emarginate in ventral aspect; distal portion with numerous

weakly developed bristles and usually 2 stronger ones on distal margin; basal

median longitudinal apodeme developed or undeveloped. Cowl: Strongly sclerotized.

Atrial plates absent. Sigma: Connected to cowl; weakly to moderately sclerotized.

Basal portion of spermathecal duct strongly sclerotized. Spermathecae 3, strongly

sclerotized, spherical, 1 slightly enlarged.

MALES. Generally similar to females except for sexual differences. Head: Clypeus

smaller than in females. Proboscis slender, about 0.95-1.70 of forefemur. Palpus

short, 0.14-0.21 of proboscis, 4-segmented with segment 4 minute to small; or

10 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

long, 0.56-0.79 of proboscis, 5-segmented, with segments 2 and 3 ankylosed and

long, making up 0.50-0.65 of palpus; entirely dark scaled. Antenna 0.44-0.65 of

proboscis; torus moderately to greatly enlarged, without scales or with inconspicu-

ous small setae; flagellum sparsely to strongly plumose, segments 1-12 with 6 to

many moderate to long bristles; flagellar segment 1 slightly elongate, with small

dark scales; segments 12 and 13 elongate, the 2 combined slightly shorter than

total length of first 11 segments. Legs: Claws of foreleg and usually midleg enlarged,

unequal; larger claw simple or with acute or blunt submedian tooth; smaller claw

simple or with acute submedian, subbasal or basal tooth; claws of hindleg small,

simple.

MALE GENITALIA. Segment VIII: Sternite with apical 0.75 scaled; apex round-

ed, with several to many well developed setae. Tergite moderately long, about

0.52-0.98 of sternite; apical 0.75 scaled; variously shaped, distal margin rounded,

straight or emarginate, with specialized scales or setae apically. Segment IX: Poorly

developed dorsally; middorsal portion of tergite straight or emarginate, with sclero-

tization weak or absent; lobes poorly developed, setae present or absent; sternite

moderately large, 2-9 small setae distally. Sidepiece: Well developed, elongate,

mesal surface membranous from base to apex; basal tergomesal area usually not

enlarged but sometimes moderately well developed, with long setae or cluster

of long flattened attenuate setae proximally; apical lobe present or absent; distal

sternomesal area with specialized scales of varied size and shape; lateral half of

dorsal surface, ventral surface and lateral surface with scales and setae, setae

usually well developed especially laterally and distally on ventral surface. Claspette:

Well developed; stem long, narrow, usually curved or angled dorsad, spiculose

basally, 1 to 3 short to moderately long setae near base; filament simple to

highly modified. Clasper: Short, length (including spiniform) usually about 0.55-

0.67 of sidepiece, usually simple but occasionally hypertrophied; apical spini-

form short or long, 0.20-0.61 of clasper. Phallosome: Aedeagus moderate to large,

tip excavated or with variously developed dorsal process. Proctiger: Strongly

developed, broad at base, narrowed distally, the basolateral sclerotization nearly

horizontal; paraproct well sclerotized, apical knob with fine serrations; cercal setae

2 10 8:

PUPAE. Cephalothorax: Weakly to moderately pigmented, usually darker dor-

sally. Hair 1-C strongly developed, considerably stronger than 2,3-C. Hair 5-C

weakly to strongly developed, weaker or stronger than hair 7-C. Hair 8-C usually

single, rarely double. Trumpet: Moderately to strongly pigmented, light to dark

brown, usually lighter apically; usually strongly broadening from base to apex,

occasionally fusiform or cylindrical; tracheoid sculpturing weakly developed in

basal 0.10; reticulate sculpturing weak to moderate. Abdomen: Weakly to moder-

ately pigmented, stronger at base of anterior tergites. Float hair (1-I) displaced

mesad. Hair 1-II-VII weakly to very strongly developed, single, branched or strong-

ly dendritic, usually more or less subequal on all segments. Hair 2-VII close to or

considerably cephalad of hair 1-VII. Hair 3-II, III moderately to strongly developed,

0.6-1.3 of corresponding tergite; 3-[V usually weak, forked; 3-V-VII weak, single.

Hair 5-II,NI usually weak, single to triple, rarely well developed; 5-IV-VII well

developed, 0.6-3.0 of corresponding tergite, usually single, rarely multiple. Hair

6-II-VI fine, usually subequal on all segments, 6-VII fine to moderately developed,

single or double. Hair 9-II-VI short, fine, subequal on all segments, dorsal or

ventral; 9-VII at or cephalad of caudolateral margin of tergite, well developed, 1-6b;

9-VIII at caudolateral margin of tergite, with 2-12 long primary branches. Hair

Arnell: Genus Haemagogus 11

10-VI considerably mesad of hair 11-VI. Terminal Segments: Male genital lobe

moderately long, 0.95-1.8 of tergite VIII. Paddle: Pigmentation varied, often con-

spicuously darker than remainder of abdomen; shape varied, longer than broad,

apex rounded or pointed, rarely emarginate; midrib conspicuous to apex; marginal

spicules few or absent. Hair 1-P weakly to strongly developed, single to multiple.

LARVAE. Head: Rounded, uniformly moderately pigmented, yellowish to brown,

with darker collar and slightly lighter ocular area. Labial plate subquadrate. Hair

1-C stout, blunt, occasionally slightly to conspicuously serrate laterally. Hair 4-C

moderately to strongly developed, 8-20b, mesad of hair 1-C and cephalad of or at

level of hair 6-C. Hair 5-C single, rarely double, usually laterad of hair 6-C. Hair 6-C

single or double, usually distinctly flattened in middle part when single; mesad or

laterad of hair 1-C. Hair 7-C 2-11b. Hair 14-C well developed, short, stout, 4-1 0b.

Hair 15-C well developed, long, 2-8b. Mental plate with 8-10(7-12) teeth on each

side of median tooth.

Antenna: Relatively short, slender; moderately to strongly pigmented, uniform

or lighter distally; with few spicules, more numerous basally. Hair 1-A long, usually

single (single or double).

Thorax: Epidermis and fat body without conspicuous pigmentation. Integument

with or without spicules. Hairs 1-3-P on common tubercle. Tubercles of hairs 5-7-P

free or joined. Hair 1-M, T moderately long, branched. Hair 3-T subequal to 1-T.

Hair 5-P 1-5b, hair 5-M single or double. Hair 11-P,M,T moderately developed,

2-5b, subequal to 9-P. Hairs 13-T and 14-M moderately to strongly developed,

2-12b.

Abdomen: Hair 1-I-VII moderately to strongly developed, becoming stronger

on more distal segments but somewhat weaker on segments VI and VII. Hair 2-I-VII

moderately developed, 2-10b, usually stout and often stellate; usually in line with

or laterad of hair 1 of corresponding segment. Hair 3-VII very strongly developed,

long, single (very rarely double). Hair 5-I-VI moderately developed, 4-10b. Hair 6

usually triple on I,II, double on III-VI. Hair 7 long, single (1-2b) on I, branched

and usually subequal to hair 5 on III-VI. Hair 9-I-VII 2-6b. Hair 12-I present or

absent. Hair 13-I-VI moderately to strongly developed, 2-10b, usually subequal

to or larger than hair 1 of corresponding segment.

Segment VIII: Hairs 1,5 moderately to strongly developed, 3-8b. Hair 3 strongly

developed, 4-10b. Comb scales 4-75, with single sharply pointed to broadly rounded,

minutely fringed spine; in single row, irregular double to triple row or large tri-

angular patch.

Siphon: Moderately to strongly pigmented, light to dark brown, with darker

basal band. Index about 2.0-3.0(1.7-3.9). Acus distinct, free or attached, or indis-

tinct, fused with base of siphon. Pecten teeth 10-18(6-24), with main spine and

1 to 3 basal denticles; in rather even, straight row on basal half; proximal teeth

often not developed. Hair 1-S inserted distad of pecten, on basal 0.50-0.65 of

siphon; moderately to strongly developed, 2-4b.

Anal Segment: Saddle extending about halfway around segment; moderately

to strongly pigmented, light to dark brown with darker basal band; caudal margin,

with spines, short and with multiple teeth laterally, stronger and with fewer teeth

dorsally. Hair 1-X strongly developed, longer than saddle, 2-6b(1-10b). Hair 2-X

2-6b. Hair 3-X single. Ventral brush (4-X) moderately to well developed, with 5

or 6 pair of hairs, all hairs but proximal 1 or 2 arising from weakly to strongly

sclerotized boss; hair 4a-X 2-8b, long or short. Anal gills 0.45-5.0 of dorsal saddle

length; usually tapered distally, occasionally short and bluntly rounded; dorsal

gill usually slightly longer than ventral gill.

i2 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

DISCUSSION. The members of the genus Haemagogus can be distinguished from

all other New World aedines by the following characters: in the adults by the broad,

flat, appressed scales, usually with metallic reflections, covering the mesonotum

and scutellum, and the pleuron with silver scales in a single broad vertical band or

three vertical arcs extending from the mesonotum to the coxae; in the female

genitalia by the cerci obliquely truncate in lateral aspect and tergite [X well devel-

oped, divided distally and V-shaped; in the male genitalia by the distal sternomesal

area of the sidepiece with a cluster of specialized scales of varied sizes and shapes;

in the pupae by the alveoli of the float hairs (1-I) conspicuously mesad of the

alveoli of hairs 1-II so that the distance between the float hairs is about 0.5 of

the distance between hairs 1-II; and in the larvae by the combination of (1) hair

3-VII very strongly developed, long and usually single, (2) hair 12-I present and/or

hair 7-III-V conspicuously stronger than hair 9 of the corresponding segment and

(3) the pecten of the siphon more or less straight and never distinctly arcuate

dorsally.

Discussions of the taxonomic history, classification and affinities of the genus

as a whole are presented in the chapter on Systematics.

KEYS TO GROUPS AND SPECIES

ADULTS

hs Scales of mesonotum dark brown with silver scales in conspicuous patches

or lines in acrostichal, antealar and prescutellar areas; pleuron with 3

vertical arcs of silver scales . . . .. . . (subgenus Conopostegus)

Scales of mesonotum metallic green, blue, copper or bronze with silver

scales only in antealar area; pleuron with single broad vertical band

of silver scales (figs. 18,27,38) (subgenus Haemagogus) . .... .2

2(1). Female fore and midclaws with subbasal tooth; postnotum bare; R24;

usually greater than 0.55 of R, (figs. 18,27) a oe opal

Female claws simple; postnotum with 2 small setae posteriorly: R, 43

usually less than 0.50 of R, (fig. 38) (Splendens Section) . . . . 15

Tropicalis Section

3(2). Ppn apparently without scales; midclaws of male small, subequal (fig. 36)

(Tropicalis Section). . . puberty. abil J ineptealis

Ppn densely scaled; midclaws ‘of male eiiareed) unequal (Albomaculatus

ee os hs Gre ay Mi cna as Uitte od

Albomaculatus Section

4(3). Midtarsus with white or gray scales on outer surface of proximal 2 or 3

SeOMeNts? vo sil lita ya A ieltah <8 0 Wig papel er tarts

Midtarsus entirely dark Lanied ¢ sot RS RINRKD ecu ni Mila once Maa. hia A par a HCG

5(4). Proboscis relatively long, 1.50 of forefemur in females, 1.50-1.70 of fore-

femur in males; eyes widely separated above antennae (4 ommatidial

diameters); male palpus long, about 0.75 of proboscis; abdomen with

silver scales dorsally on tergites II-VII (fig. 27) . . . .J1J1. panarchys

6(5).

7(6).

8(7).

97),

10(6).

11(10).

12(10).

Arnell: Genus Haemagogus 13

Proboscis shorter, less than 1.40 of forefemur in females, 1.50 of fore-

femur in males; eyes narrowly separated above antennae (1 or 2 omma-

tidial diameters); male palpus short or long, but less than 0.70 of

proboscis; abdomen with dorsal silver scales restricted to tergites IV-

Vibor absent. bi bibiion. eaileie Bie Pi aaa

Conspicuous knee spots of silver scales present anteriorly on apex of

mid and hindfemora . . sll nanan Were al iia oak p! GS bee

Knee spots on mid and hindfemors absent kee Mati nlel site mention

Proboscis short, about 1.10 of forefemur in females, 1.30 of forefemur

in males; scales of mesonotum copper; abdominal tergites without

dorsal silver scales; male palpus short, less than 0.20 of proboscis . .8

Proboscis longer, 1.25-1.40 of forefemur in females, 1.35-1.50 of fore-

femur in males; scales of mesonotum hues of blue or green; abdo-

men with dorsal silver scales on tergites IV or VI-VII; male palpus

lone, about .O0-0. 70 OL DIODOSCIS 13. tie WOOP i tae ee Oe Se

Abdominal tergites green or light bronze; decumbent scales of vertex

COPpPer- cies -4 . . . J. andinus

Abdominal fergites Bias. decumbent scales of Sea bluish green

6. nebulosus

R43 0.70-0.95 of R,; abdomen with dorsal silver scales usually re-

stricted to tergites VI,VII. . . . . . 12. soperi; 13. acutisentis

R43 1.05-1.25 of R,; abdomen with dorsal silver scales usually on ter-

wites VV a oe we eer ae OO Oe ae 2 4 eaaeenens

Lower stp with well developed bristle; larger claw of foretarsus of male

with acute submedian or subbasal tooth . lop ea abd was

Lower stp without well developed bristle; larger claw of foretarsus of

UEC AACR gl oem entai Ce Mur MT ERG hen My tg Sorinentios ge welt: ARE gy Poste Sedey os ~ cane BF

Hindfemur with silver scales extending nearly to apex anteriorly; dark

scales of abdomen usually predominantly purple but with bluish green

scales on distal margins of tergites V-VIII; female proboscis 1.10-1.15

of forefemur; large and small claws of foreleg and midleg of male

toothed (fig. 18) . ee eae hae . . . 4%, janthinomys; 8. capricornii

Hindfemur with silver scales not extending beyond basal 0.75 anteriorly;

dark scales of abdominal tergites I-VII purple; female proboscis 1.25

of forefemur; larger claw of foreleg of male toothed, smaller claw of

foreleg and claws of midleg simple (fig. 25). . . . 10. albomaculatus

Female proboscis short, 1.10 of forefemur; scales of mesonotum primarily

dark blue; coxae with conspicuous patches of dark scales; R,43 0.55

of R,; abdominal dark scales purple. . .. Siew deel, aol ae enaaNeaE

Female proboscis at least 1.20 of forefemur; scales of mesonotii pri-

marily copper to light bronze; coxae all silver or rarely with dark scales

on midcoxa; R243 1.20-1.50 of R,; abdominal dark scales hues of

Die aren oF ROI Mtiny Losey Moi Wa wel one wer eo ee ES

13(12).

14(13).

1S@).

16(15).

17(15).

1817).

19(17).

2019):

Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Apn lobes almost entirely silver scaled; abdominal tergites without silver

scales dorsally; ppn scales gold to copper, decumbent scales of vertex

green togold . . one ; 3. spegazzinii

Apn lobes with few silver scales: abdomen with silver scales dorsally on

tergites IV-VII; ppn scales blue or light bronze; decumbent scales of

vertex copper, blue Or ish ietTeen bh ee ee eS

Decumbent scales of vertex, scales of ppn and dark scales of abdomen

blue to bluish green; apn scales blue; smaller claw of foreleg and midleg

of male with submedian tooth (fig. 8). . . . . . . .Jl. anastasionis

Decumbent scales of vertex copper, scales of apn, and darker scales of

abdomen light bronze; all claws of foreleg and midleg of male simple

(Hie, SO) or er a eee lee: 2 2 ehrysochiorus

Splendens Section

Porn witht larce patch of scales posteriorly. =. ee oe ee DO

Pon Sare or with very few silver scales posteriorly ©... 2%... 17

Scales of vertex and occiput copper to light green; male antenna sparsely

plumose, flagellar segment 4 with about 8-10 bristles (fig. 38) . .

y 26. splendens

Scales blue to violet on vertex, green on occiput: male antenna moder-

ately plumose, flagellar segment 4 with about 18-22 bristles

Dr ee Nee a a ee _ 17. celeste

Proboscis shorter than forefemur, 0.80-0.95 of forefemur in females,

about 0.90-0.95 in males. .°. . Beg les

Proboscis equal to or longer than forefemur, about 0. 95- 1 20 of fore-

heimur ti females aie sidies 3. 0) es ere, a. OED

Integument of coxae and trochanters yellow; scales of mesonotum usu-

ally bronze to copper with distinct purple or occasionally dark blue

patch in fossa; abdominal scales usually purple; large claw of foreleg

of male with blunt submedian tooth, large claw of midleg simple

(gS). Goo ; . . .23. chalcospilans

Integument of coxae ad frocnaniters ae coe of mesonotum rather

uniformly greenish blue; abdominal ccales blue to violet; larger claw of

foreleg and midleg of ‘male with acute subbasal tooth dis® 51)

Se A Ee pe ee en Oo 2 areyroments

Proboscis about equal in length (0.95-1.05) to forefemur; eyes narrowly

separated above antennae (1 or 2 ommatidial diameters). . 24. boshelli

Proboscis longer than forefemur; eyes widely separated above antennae

(3 or Fommatidial diameters) so sO ee ee ee 0

Integument of apex of coxae, trochanters and base of femora yellow;

scales purple on vertex, light yellowish brown on occiput; scales of

mesonotum copper .... . . 19. aeritinctus

Integument of coxae, ochre and peadors danke scales of vertex blue

to violet; scales of mesonotum bronze to green or blue but rarely

pee

21(20):

pace |).

01).

3(2).

4(3).

5(4).

6(4).

Arnell: Genus Haemagogus 15

Scales of mesonotum usually greenish blue or blue; apn lobes blue; abdo-

minal scales blue with median basal silver patches on III or IV-VII

NL) 20. regalis

ese os ecu eealty bree ‘to. Se reer apn lobes violet to

purple; abdominal scales usually purple or violet with median basal

silver patches only on VILVUl (at. oot wi Ca emg ee ganna

Ppn bare or with small patch of silver scales posteriorly . . 18. iridicolor

Pon bare never Witt Giver SCaleS be es ie a a ae, ee

MALE GENITALIA

(6. nebulosus unknown)

Distal margin of tergite VIII without long, lanceolate scales or large

spiniform setae mesally; tip of aedeagus excavated, without sclerotized

dorsal process 2. . . . . (subgenus Conopostegus)

Distal margin of tergite VIII with fone! lanceolate scales or large spiniform

setae mesally; tip of aedeagus with variously developed sclerotized

dorsal process (subgenus Haemagogus) ............2

Sidepiece with apical lobe poorly developed; tergite IX with 1-3 setae .3

Sidepiece with apical lobe well developed and prominent or represented

by dense cluster of well developed setae; tergite IX without setae

CSPIeMGeis GOCEION) i iwir ue wi teas palin a ae im ee Rea Cad an A

Tropicalis Section

Distal margin of tergite VIII straight and with large spiniform setae

mesally; ventral half of claspette filament large, erect, rugose and

bilobed Gig: 36):(Troptealis Section) |. °° hak ye oe SY Prepiedhs

Distal margin of tergite VIII usually emarginate, if not emarginate then

with elongate, lanceolate scales mesally; claspette filament without

large, bilobed, erect ventral half (Albomaculatus Section) . ... .4

Albomaculatus Section

Tergite VIII with large spiniform setae mesally; basal tergomesal lobe of

sidepiece extending distad to near distal third of sidepiece as slightly

raised area with numerous short setae. . . . dies 2

Tergite VIII with elongate lanceolate scales cea didepiese witout

raised ared distad of Dasal tergomesal lobe: hw er eee OG

Claspette filament narrowly sickle-shaped; claspette stem broader beyond

middle; distal margin of tergite VIII deeply emarginate (figs. 8,36)

1. anastasionis; 2. chrysochlorus

Claspette filament broadly sickle-shaped; claspette stem uniform in thick-

ness; distal margin of tergite VIII ails shallowly emarginate (fig.

ao A Ge a ie aA iaip Maal avait was SLE ROL

Tergite VIII short, about 0.50 of sternite VIII, distal margin convex;

claspette stem densely pilose from distal fourth to apex (fig. 28)

hia hea ben ee AS ART RA AOU Bi se UA Sule RE ‘ . 11. panarchys

7(6).

8(7).

9(8).

10(9).

11(10).

12(10).

13(12).

14(12).

Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Tergite VIII at least 0.55 of sternite VIII, distal margin shallowly to

deeply emarginate; claspette stem without pilosity . . ..... .7

Distal half of claspette stem with membrane projecting anteriorly and

posteriorly and enclosing basal portion of filament; filament with

basal supporting ribs (fig. 34) . . . . . ee . . 14. equinus

Distal half of claspette stem without membrane: filament without support-

ing ribs

Aedeagus with apex deeply cleft mesally with an acute sclerotized pro-

jection on each slide of cleft (fig. 25) . . . . . . 10. albomaculatus

Aedeagus with apex pointed or produced distad mesally ..... .9

Apex of aedeagus produced into long, thin, coarsely serrated carina

making up about 0.3 of aedeagus length, dorsum with elliptical open-

ing into which base of carina projects (fig. 23). . . . 9. mesodentatus

Apex of aedeagus not produced into long serrated carina, dorsum of

Cee ne ar eM pi ee rene ie ag

Tergite VIII relatively short, about 0.55-0.60 of sternite VIII, distal

margin very shallowly emarginate . . pet Bd

Tergite VIII longer, about 0.70-0.85 of sternite VILL, distal margin shal-

Wey PO eel y CMM etite ee es ea Or ry ED

Aedeagus with large spicules on venter of apical, beaklike keel; apical

knob of paraproct with hooklike process mesally (fig. 19) . :

7. janthmomr

Aedeagus with: apex slightly incised just taterad of short, heavily sclero-

tized median keel; apical knob of paraproct without hooklike process

ne Me re gl nek ee eke ite. pa Oy, PP ECOP TEL

Tergite VIII broadly emarginate distally; aedeagus with apex pointed. 13

Tergite VIII with distal margin declivous mesally; aedeagus with apical

PIOCess DIDAC ANG IN CRtea ida ea ere pe

Aedeagus with distal fourth broadly triangular, tip with a strongly sclero-

tized dorsal carina proximad to a small beak; ne stem sharply

angled dorsad at apical third (fig. 14) . .... . . 6. andinus

Distal fourth of aedeagus broadly rounded, with | a small, weakly

sclerotized beak at apex; claspette stem uniformly curved dorsad (fig.

OS a CoO: ADC ACN GR NC OU AG Mahe ats LO ea SN Ce ROOM AE) 7 i

Mesal half of sidepiece between basal sternomesal area and distal fourth

with moderately developed setae laterally and bare space mesally (fig.

OO os . « « dd SOperi

Mesal half of sidepiece between basal sternomesal area and distal fourth

with uniformly distributed, pepsliands short setae ie a2)!

By at Caceres HAYS : rant : 23. acutisentis

PS(2)-

16(15).

PACTS),

18(17).

19(18).

20(19).

21(20).

Arnell: Genus Haemagogus 17

Splendens Section

Clasper cylindrical; spiniform cylindrical, inserted apically. . . . . 16

Clasper flattened; spiniform spatulate, inserted subapically. . . . . 17

Distal sternomesal scales of sidepiece lanceolate to oblanceolate; apical

lobe represented by cluster of well developed setae; tergite VIII with

distal margin rounded, lateral setae shorter than mesal setae (figs. 39,

oe Ae oa, 2 B6.splendens; 17. celeste

Distal sterorieeat scales of tideprers circular and lanceolate dorsally

and sinuous ventrally; apical lobe prominent; tergite VIII emarginate

distally, lateral setae much longer than mesal setae (fig.53) . .

nee & chalcospilans

Clasper angled sharply inward near middle with elbowlike flap on outer

surface distad of angle; sidepiece with large oval area of blunt, peglike

setae near middle (fig. 55) .. . . DD ew eke. DORE

Clasper broadly curved inward over entire length: sidepiece without peg-

Hike setae Hear Tie 6 sk 3 Pe aie ee ee ee a

Clasper gradually broadened to bulbous apex; claspette stem expanded

distally into apieel knob with seta on posterior margin (fig. 51).

. 22. argyromeris

Clasper Breuate or pointed distally: claspette stem not expanded into

SORELLE SOB iets cena abet eagles ae cor RRM AO 3 SO UPAR 2 MON aL Se ELePRm

Claspette stem angled dorsad at middle, narrowed beyond angle; filament

a broad, cuneiform leaf inserted distally on stem (fig. 43) heads

EES 2. 9 jnidioptor

Claspette stem ‘angled dorsad Ys ‘apical third; expanded beyond angle to

broad apex or expanded slightly then tapered to acute apex .. . 20

Claspette stem with apex expanded; filament a flat, expanded leaf in-

serted distally on stem and terminating in a slightly recurved point

CLP 2G 1) er . . 19. aeritinctus

Claspette stem expanded slightly beyond aiele Hien tapered to acute

apex; filament inserted distally on stem and extending posteriorly

over distal third of stem as attached, folded membrane . .... 21

Clasper arcuate and terminating in rounded point; claspette stem uni-

formly narrow proximad of angle (fig.47) . . . . . . 20. regalis

Clasper expanded on distal fourth then tapered to sharn point; claspette

stem stout, expanded at basal third then constricted beyond middle

CT I RAD LER St a Rah SOU POR eR SN utp Anes aig SECO a7 yaar er rr

LARVAE

(2. chrysochlorus unknown)

Comb scales 5-12 (4-20) in single row, ventral brush with 5 pairs (very

rarely 6) of hairs .

2(1).

3(1).

4(2).

5(4).

6(4).

7(6).

8(7).

9(7).

10(9).

11(6).

Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Comb scales 10-50 (6-75) in irregular double row or triangular patch;

ventral brush with 6 pairs of hairs (subgenus Haemagogus, in part). .3

With the following combination of characters: integument without spic-

ules, hair 12-I present, boss strongly sclerotized, hair 12-II subequal

to or weaker than JO-[]. 2... . . (subgenus Conopostegus)

Not as above; differing in at least 1 of the above characters (subgenus

Haemagogus, Albomaculatus Section) .

Tropicalis Section

Boss weakly sclerotized or absent; hair 6-C single, rarely double (fig. 37)

(Tropicalis Section). . . . . . IS. tropicalis

Boss strongly sclerotized; fale 6-C double (very rarely veer oe

DECOM) fF Boe ee eee : te 5

Albomaculatus Section

HPeeUMicnt WIUMSOICUICS © ee ea ee ee 5

INTeetenT WAMOUL SPictiles of eee oe ee ee

Comb scales attached basally to sclerotized plate; hair 12-I absent (figs.

ZOG2 2 oes . . . 7. janthinomys; 8. capricornii

Comb scales not attached basally to sclerotized plate; hair 12-I present

(9S 2A ey he oii On mesodentatus

Pie ease ee

di TE PrCseme or ee ee ee oe Sk

BibmOM iigex lessen Sse Oe oe er eee og

Sion tnde< renter Mats. ee Oy ate Oe oe

Siphon index about 2.8; hair 6-C double; hair 7-IJ-IV larger than hair 5

of corresponding sepmient Ce Sy ce . . 4. baresi

Siphon index about 2.0; hair 6-C single; bait ale IV subequal to hair 5

Of Corresponding segment (fig.9)... ... ..°. .° 40d. anastasionis

Siphon index about 3.2-3.4; pecten with 20-24 teeth (fig. 11).

oe: spegazzinii

Siphon index ushelly 3. Sc or greater: pecten with about 16-22 teeta, + 10

Siphon index about 3.5 (3.3-3.8); hair 5-C single (1-2b); hair 4a-X 2b

(Fie 15) ey . . 6. andinus

— index ee 3. 8: an 5-C EAE (2 3b) Pan 4a-X 4-5b (fig. 17)

: Pa : : 7. nebulosus

Boss strongly sclerotized; tubercles of hairs 5-7-P broadly connected;

comb scale spine oe double length of basal, attached portion (fig.

0) ea eee . . . L0. albomaculatus

Boss oe erotics: Or useae. hilpes ies of hairs 6-7-P very rarely

connected though tubercles of hairs 5-6-P often connected; comb

scale spine short, subequal in length to basal, attached portion . . 12

Arnell: Genus Haemagogus 19

12(11). Hair 1-C conspicuously serrate laterally; hair 6-C double. . ... . 13

Hair 1-C smooth to apex; hair 6-C usually single, rarely double . . . 14

13(12). Comb scales with broadly rounded to spatulate spine (fig. 31). 12. soperi

Comb scales with short, sharply pointed spine (fig. 33) . .13. acutisentis

14(12). Comb scales usually 10-12 (8-20) in irregular single row; spines on caudal

margin of saddle reduced, most very small (fig. 29). . . JJ. panarchys

Comb scales usually 7-10 (5-13) in single, regular row; spines on caudal

margin of saddle well developed, prominent (fig. 35) . . .14. equinus

Splendens Section

15(3). Comb scales more than 35; hair 4a-X with at least 6 branches; hair

7-III-VI smaller than hair 5 of corresponding segment. . ... . 16

Comb scales fewer than 30; hair 4a-X with fewer than 6 branches; hair

7-III-VI subequal to hair 5 of corresponding segment. . . ... 17

16(15). Comb scales usually 45-55 (40-75); hair 5-M single; hair 4a-X 6-9b (fig.

: ot ee . . .23. chalcospilans

Comb ees 35-42 9 49), jee “5-M aoa hair 4a-X 6b (fig. 56)

Bis ! ' cone . 24. boshelli

Lit S).: Combscales ustiaty tower tian 22 go a es ik, eee ee

Comb. scales usudlly more Tain 230 a oe a ora eae

18(17). Comb scales 10-13 (6-20); pecten extending to about basal 0.40-0.45

of siphon; hair 4a-X 3-4b (fig. 40) . . . . . . 16. splendens

Comb scales 17-22 (12-27); pecten extending’ to about basal 0.50 of

Siphon: ha 4a-y +5 (ie oa 2) a ee oo ee 2 eereste

19(17). Spines on caudal margin of saddle reduced, very short, each with multi-

ple teeth; pecten extending to about basal 0.45 of siphon (fig. 46)

. 19. aeritinctus

Spines « on - eaudal margin of saddle stronger, several spines long, with

single tooth; pecten extending to about basal 0.50 of siphon (figs. 44,

48,50,52). . . .18. iridicolor; 20. regalis; 21. lucifer; 22. argyromeris

Subgenus CONOPOSTEGUS Dyar

1925. Conopostegus Dyar, 1925b:141. TYPE SPECIES: Aedes leucocelaenus Dyar and Shannon,

1924; monobasic.

Zavortink (1972:111-131) removed leucocelaenus and related species from Aedes

and placed them in Haemagogus, resurrecting the subgenus Conopostegus for them.

This group of species had remained in Aedes since 1924 primarily on the basis of

the presence of postspiracular and prescutellar bristles and the absence of these

bristles in Haemagogus. However, these bristles are present in the metallic Haema-

gogus, though small and poorly developed. Considering the many similarities be-

tween Conopostegus and the metallic Haemagogus in all stages, the main difference

being the thoracic scaling of the adults, and their bionomics and distribution, I

am confident that these 2 groups constitute a single phylad.

20 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

The subgenus Conopostegus can be separated from the subgenus Haemagogus:

as adults by the deep brown color of the mesonotal scales, conspicuous patches

or lines of silver scales in acrostichal, antealar and prescutellar areas and 3 vertical

areas of silver scales on the pleuron; in the male genitalia by the absence of large,

spiniform setae or long, lanceolate scales mesally on the distal margin of tergite VIII

and the aedeagus without a sclerotized dorsal process; in the pupae by the strongly

developed hairs 5-C and 1-P; and in the larvae by the combination of (1) few comb

scales in a single row, (2) integument without spicules, (3) hair 12-I present, (4) boss

strongly sclerotized, and (5) hair 12-II subequal to or weaker than hair 10-II.

Conopostegus retains several of the more primitive characters not present in the

more highly derived subgenus Haemagogus, and is undoubtedly closer to the original

stock which gave rise to the genus. Among the more primitive characters of Cono-

postegus are the absence of metallic scales on the scutum and scutellum and the

well developed prescutellar and postspiracular bristles in the adults, and, in the

male genitalia, the absence of specialized scales or setae on the distal margin of

tergite VIII and the absence of dorsal sclerotizations on the aedeagus. In the pupae,

Conopostegus exhibits several unusual specializations not present in the subgenus

Haemagogus such as the development of hair 1-I (float hair), hair 1-P and the shape

of the paddle in 1 species. The larvae of Conopostegus and Haemagogus are so

similar that they are not easily separable.

The subgenus Conopostegus and the included species are discussed thoroughly

by Zavortink (1972:111-131).

Subgenus HAEMAGOGUS Williston

1896. Haemagogus Williston, 1896:271. TYPE SPECIES: Haemagogus splendens Williston,

1896, St. Vincent; monobasic.

1905. Stegoconops Lutz, 1905:83-84. TYPE SPECIES: Haemagogus capricornii Lutz, 1904,

in Bourroul, 1904:66, Brazil; monobasic. Synonymy with Haemagogus by Howard,

Dyar and Knab (1917:863). As subgenus of Haemagogus by Dyar (1921:102-104).

NEW SYNONYMY.

1906. Cacomyia Coquillett, 1906:25. TYPE SPECIES: Haemagogus albomaculatus Theobald,

1903, Guyana; the first of 2 included species, designation by Coquillett (1910:516).

Synonymy with Haemagogus by Howard, Dyar and Knab (1917:863).

1939. Cyanoconops Antunes, 1939:106. NOMEN NUDUM.~—As subgenus of Haemagogus.

1951. Longipalpifer Levi-Castillo, 1951b:11. TYPE SPECIES: Haemagogus panarchys Dyar,

1921, Ecuador; original designation.—As subgenus of Haemagogus. NEW SYNONYMY.

1955. Osornomyia Levi-Castillo, 1955:360. NOMEN NUDUM.—As subgenus of Haemagogus.

Haemagogus of Williston (1896:271); Theobald (1901:239; 1903a:308; 1905b:37; 1907:550,

in part; 1910:493, in part); Coquillett (1906:25, in part); Howard, Dyar and Knab (1917:

863-877); Kumm, Osorno-Mesa and Boshell-Manrique (1946:13); Vargas (1950:63, in part);

Kumm and Cerqueira (1951:169); Komp (1954d:264); Zavortink (1972:111).

Haemagogus (Haemagogus) of Dyar (1921a:105-114; 1928:136-141); Dyar and Shannon (1924:

483); Edwards (1932:179); Antunes (1939:106); Lane (1939:118-120; 1953:779-789); Levi-

Castillo (1951b:11,16-31); Stone, Knight and Starcke (1959:217-218).

Haemagogus (Stegoconops) of Dyar (1921a:102-104; 1928:134-135); Dyar and Shannon (1924:

483); Edwards (1932:179); Lane (1953:789-802); Stone, Knight and Starcke (1959:216-

ory). |

ee, (Longipalpifer) of Levi-Castillo (1951b:31-37); Stone, Knight and Starcke (1959:

215-216).

Haemagogus (Osornomyia) of Levi-Castillo (1955:361).

Arnell: Genus Haemagogus 21

Haemagogus (Cyanoconops) of Antunes (1939:106); Lane (1939:121; 1953:802-810).

Cacomyia of Coquillett (1906:25); Theobald (1907:554; 1910:493-494).

Stegoconops of Lutz (1905:83-84); Howard, Dyar and Knab (1913:figs. 162-165 ,438,439, pl. 77).

Aedes in part of Dyar and Knab (1906a:190,191,195,202, fig. 18).

FEMALES. Scales of head, thorax and abdomen broad, flat, with brilliant

metallic reflections. Light scales silver or silvery white. Head: Eyes narrowly to

broadly separated above antennae (1 to 5 ommatidial diameters). Proboscis 0.80-

1.52 of forefemur. Palpus 4-segmented, segment 4 small. Thorax: Mesonotum

and scutellum densely covered with broad, flat scales. Prescutellar bristles not

developed. Apn lobes considerably to greatly enlarged, often nearly meeting at

midline; ppn with or without scales; setae near middle of stp not developed;

lower stp bristles weakly or strongly developed. Pleuron with broad, flat, silver

or silvery white scales forming single broad, vertical band from antealar area to

midcoxa with scales on paratergite, pra, psp, ssp, mep and stp; pcx, ppl and pst

with silver scales. Legs: Hindcoxa with base at level of or slightly below upper

margin of meron; midcoxa with single well developed bristle or row of 2-4 well

developed bristles laterally; forefemur and midfemur usually with silver scales at

base and often extending to basal third or half and occasionally to apex ventrally;

hindfemur with silver scales on anterior, ventral and posterior surfaces usually

extending to about basal 0.75 anteriorly but not beyond basal half posteriorly;

knee spots of silver scales at apex of midfemur and hindfemur present or absent.

Claws of foreleg and midleg simple or with subbasal tooth. Wing: R, 43 about

0.30-1.56 of R,. Postnotum: Bristles present or absent.

FEMALE GENITALIA. Tergite IX: Setae present or absent.

MALES. Generally similar to females except for sexual differences. Head: Pro-

boscis about 0.95-1.64 of forefemur. Palpus either short or long. Antenna 0.44-

0.65 of proboscis with torus moderately to greatly enlarged, flagellum sparsely

to densely plumose, segments 1-12 with 6 to many moderate to long bristles.

Legs: Claws of foreleg enlarged, unequal, tooth present or absent; claws of mid-

leg usually enlarged, unequal, rarely small, subequal, tooth present or absent.

MALE GENITALIA. Segment VIII: Tergite with long, lanceolate scales or large

spiniform setae mesally. Segment IX: Tergite with or without setae. Sidepiece:

Apical lobe present or absent. Claspette: Filament simple or highly modified.

Clasper: Simple to highly modified; spiniform apical or subapical, short or long,

0.18-0.65 of clasper. Phallosome: Aedeagus tip with variously developed dorsal

process.

PUPAE. Cephalothorax: Hair 1-C considerably stronger than 2,3-C. Hair 5-C

weaker than or rarely subequal to 7-C. Trumpet: Strongly broadening from base

to near apex. Abdomen: Hair 1 weakly to very strongly developed, usually

weaker on posterior segments. Hair 5-IV-VII 0.4-1.2 of corresponding tergite.

Paddle: Apex rounded or pointed. Midrib usually considerably darker than re-

mainder of paddle; marginal spicules few. Hair 1-P weakly developed, usually

single (1-4b).

LARVAE. Head: Hair 5-C single (rarely double). Hair 6-C single or double.

Thorax: Integument with or without spicules. Tubercles of hairs 5-7-P free or

joined. Abdomen: Hair 12-I present or absent. Segment VIII: Comb scales 6-50

(4-75) with single, minutely fringed spine; in single row, irregular double row or

large triangular patch. Anal Segment: Spines on caudal margin of saddle strongly

or weakly developed. Ventral brush (4-X) with 5 or 6 pair of hairs arising from

strongly to weakly sclerotized boss. Hair 4a-X 2-8b, long or short.

22 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

DISCUSSION. The subgenus Haemagogus can be distinguished from the subgenus

Conopostegus: in the adults by the dense metallic green, blue, copper or bronze

mesonotal scales and the single broad vertical band of silver scales on the pleuron;

in the male genitalia by long lanceolate scales or large spiniform setae on the

distal margin of tergite VIII and a variously developed sclerotized dorsal process

on the aedeagus; in the pupa by relatively weakly developed hairs 5-C and 1-P;

and in the larva by at least 1 of the following characters: (1) comb scales numerous

and in a triangular patch, (2) integument with spicules, (3) a weakly sclerotized

boss, (4) hair 12-I absent or (5) hair 12-II stronger than hair 10-II.

As discussed above in the chapter on Systematics, the subgenus Haemagogus

includes the species formerly included in the subgenera Haemagogus, Stegoconops

and Longipalpifer (Stone, Knight and Starcke, 1959:215-218). The subgenus consti-

tutes a rather homogeneous assemblage of species and I do not consider the sub-

division into the above 3 subgenera to be warranted, especially with the inclusion

of the subgenus Conopostegus in the genus Haemagogus, as the criteria for sepa-

rating the present subgenera Conopostegus and Haemagogus are of a different

magnitude than those for subdividing the subgenus Haemagogus. In addition, the

species formerly placed in Longipalpifer, those with long male palpus, are not a

monophyletic group, as this character is found in 3 separate phyletic lines. I

prefer, instead, to recognize 3 sections in the subgenus Haemagogus: the Albo-

maculatus Section containing the species formerly placed in the subgenera Stego-

conops and Longipalpifer with the exception of tropicalis; the annectent mono-

typic Tropicalis Section; and the Splendens Section containing the species formerly

placed in the subgenus Haemagogus.

Although several species complexes are apparent within the subgenus Haema-

gogus, formal species groups are not recognized, primarily because more than half

of the species groups would be monotypic, and such an arrangement would prob-

ably be unduly cumbersome. The species complexes are discussed under the Section

discussion paragraphs and in the systematics paragraphs of the species accounts.

ALBOMACULATUS SECTION

FEMALES. Head: Eyes narrowly to broadly separated above antennae (1-4

ommatidial diameters). Proboscis 1.10-1.52 of forefemur. Thorax: Ppn with scales;

single lower stp bristle usually well developed. Legs: Midcoxa usually with row of

2-4 well developed bristles laterally; forefemur about 1.25-1.60 of distance from

top of thorax to apex of midcoxa. Knee spots present or absent. Claws of foreleg

and midleg with subbasal tooth. Wing: R,,3 about 0.55-1.56 of R,. Postnotum:

Bristles absent.

FEMALE GENITALIA. Tergite IX: Setae (1-3) present.

MALE. Head: Proboscis about 1.31-1.64 of forefemur. Palpus either short or

long. Antenna 0.55-0.65 of proboscis, with torus much enlarged, flagellum densely

plumose, segments 1-12 with very numerous moderate to long bristles. Legs: Claws

of foreleg and midleg enlarged, unequal; larger claw simple or with submedian

tooth; smaller claw simple or with submedian or basal tooth.

MALE GENITALIA. Segment IX: Tergite with 1-3 poorly to moderately devel-

oped setae. Sidepiece: Apical lobe absent. Claspette: Filament simple or modified.

Clasper: Simple, cylindrical, curved inward at apex; seta present near apex of inner

surface; spiniform apical, long, about 0.40-0.65 of clasper, slightly curved inward.

Arnell: Genus Haemagogus 23

LARVAE. Head: Hair 5-C usually single (single to triple). Hair 6-C usually single,

occasionally double. Thorax: Integument with or without spicules. Tubercles of

hairs 5-6-P usually joined and hair 7-P free but often with tubercles of hairs 5-7-P

either all connected or all free. Abdomen: Hair 12-I present or absent. Segment

VIII: Comb scales 6-12(4-20), with single sharply pointed, minutely fringed spine,

in single row. Anal Segment: Spines on caudal margin of saddle usually relatively

strongly developed and with few teeth dorsally. Ventral brush (4-X) with 5 pairs

of hairs, all hairs arising from weakly to strongly sclerotized boss. Hair 4a-X 2-5b,

long or short.

DISCUSSION. The Albomaculatus Section can be distinguished from the remain-

der of the subgenus Haemagogus: in the adults by the combination of (1) toothed

claws on the foreleg and midleg of the female, (2) postnotum without setae, (3)

vein R,,3 at least 0.55 of R, and (4) a dense patch of scales on the ppn, in the

female genitalia by the presence of setae on tergite IX; in the male genitalia by

(1) 1 to 3 setae on tergite IX and (2) the distal margin of tergite VIII emarginate,

or if not emarginate then with lanceolate scales mesally; and in the larvae by a

single row of comb scales and 5 pairs of hairs in the ventral brush.

The Albomaculatus Section contains, with the exception of tropicalis, the species

formerly placed in the subgenera Stegoconops and Longipalpifer. In general this

section appears to be the least derived of the subgenus Haemagogus in the adults

and in terms of numbers of individuals and extent of distribution contains the

dominant elements of the Haemagogus fauna, janthinomys, mesodentatus and

equinus.

The relationships within the Albomaculatus Section are difficult to determine

with the exception of a few obvious species complexes.

The species of the anastasionis complex, anastasionis, chrysochlorus and spe-

gazzinii, share the absence of a well developed lower stp bristle in the adults, a

simple large claw on the foreleg and midleg of the males, the presence of large

spiniform setae in place of elongate scales on the distal margin of tergite VIII in

the male genitalia, and, in the larvae, the absence of hair 12-I, single hairs 5,6-C

and a strongly sclerotized boss.

The species of the andinus complex, andinus and nebulosus, differ in the adults

only in the color of the dark scales of the head, mesonotum and abdomen and in

the larvae only by the shorter siphon and fewer branches on hair 4a-X on andinus.

The relationships of the andinus complex are obscure, but it may be related to

the anastasionis complex through baresi. The latter has the male genitalia much

like those of andinus and lacks a lower stp bristle and has a simple large claw on

the male foreleg and midleg, characters found in the anastasionis complex.

The capricornii complex consists of janthinomys and capricornii, which are

apparently indistinguishable except for details of the aedeagus. Possibly closely

related to the capricornii complex is mesodentatus, which also has a spiculose

larval integument and an extreme development of the aedeagus, although it differs

in details of adult ornamentation and has larval hair 12-I developed, which janthi-

nomys and capricornii lack. Janthinomys and mesodentatus have a similar biology

and have a basically complementary distribution.

The species of the soperi complex, soperi and acutisentis, differ only in the

distribution of setae on the sidepiece of the male genitalia and the shape of the

larval comb scales. The soperi complex is probably allied to equinus with which

it shares the long male palpus and several important larval characters, however

the male genitalia differ considerably.

24 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

The relationships of albomaculatus are not clear, and though it is unusual only

in the unique development of the aedeagus, it seems not to be particularly closely

allied to any other members of the Albomaculatus Section.

Panarchys is, in many respects, the most unusual member of the subgenus and

since it retains several primitive characters, it may be annectent to Conopostegus.

It has no close affinities to any other member of the subgenus Haemagogus.

1. Haemagogus (H.) anastasionis Dyar

Figs. 2,8,9

1921. Haemagogus anastasionis Dyar, 1921c:155. TYPE. Lectotype male (1529) with geni-

talia slide, Puntarenas, Puntarenas, Costa Rica, 15 July 1921, A. Alfaro [USNM, 24864;

selection of Stone and Knight, 1955:287].

1971. Haemagogus (Stegoconops) dominguezi Duret, 1971:83-86. TYPE. Holotype male (7258)

with genitalia slide, Department of Zelaya, Nicaragua, 1961, F. Dominguez [A]. NEW

SYNONYMY.

Haemagogus (Haemagogus) anastasionis of Edwards (1932:179); Anduze (1947:353).

Haemagogus (Stegoconops) anastasionis of Stone, Knight and Starcke (1959:216); Cova Garcia,

Sutil and Rausseo (1966a:61-62, fig. 120; 1966b:114, fig. 202); Diaz Najera (1966:61).

Haemagogus anastasionis of Komp and Kumm (1938:259); Kumm, Komp and Ruiz (1940:

398); Kumm and Zuniga (1942:404); Hovanitz (1946:40); Kumm, Osorno-Mesa and Boshell-

Manrique (1946:17); Woke (1947:365); Trapido and Galindo (1956a:306); Galindo and Trapido

(1957:148); Vargas and Diaz Najera (1959:362); Diaz Najera (1960:185).

Haemagogus (Haemagogus) anastationis of Dyar (1928:137); Lane (1939:118); Levi-Castillo

(1951b:11,23-24).

Haemagogus (Stegoconops) anastationis of Lane (1953:798-800).

Haemagogus anastationis of Vargas and Martinez Palacios (1953:37); Barreto Reyes (1955:79).

Haemagogus uriartei of Komp (1949:73).

FEMALE. Wing: 3.23 mm. Proboscis: 2.68 mm. Forefemur: 2.05 mm. Abdo-

men: 2.90 mm. Dark scales of proboscis, palpus, wing and legs primarily violet,

with some purple reflections. Head: Eyes narrowly separated above antennae

(1 ommatidial diameter). Decumbent scales of vertex and occiput blue anteriorly

and laterally, bluish green posteriorly. Proboscis relatively long, about 1.30 of fore-

femur; palpus about 0.15 of proboscis; antenna relatively long, about 0.71 of

proboscis. Thorax: Scales of mesonotum copper dorsally, more blue anteriorly,

laterally and posteriorly, usually blue over supraalar bristles, silver in antealar

area; scales on scutellum copper, blue on lobes. Apn lobes moderately enlarged,

scales blue, often with silver scales anteriorly; ppn scales blue to bluish green;

lower stp without well developed bristle. Legs: Midcoxa with patch of violet

scales near middle; legs relatively short, length of forefemur about 1.30 of distance

from top of thorax to apex of midcoxa. Wing: R43 about 1.25-1.50 of R,.

Abdomen: Dark scales blue to bluish green, occasionally with purple reflections;

silver scales dorsally on tergites IV-VII, in small basal patches on IV and V and

rather broad basal bands on VI and VII.

MALE (fig. 8). Wing: 2.60 mm. Proboscis: 2.45 mm. Forefemur: 1.70 mm.

Abdomen: 2.55 mm. Head: Proboscis about 1.45 of forefemur; palpus short,

about 0.14 of proboscis; antenna about 0.65 of proboscis. Legs: Larger claw of

foreleg and midleg simple; smaller claw of foreleg with acute submedian tooth,

smaller claw of midleg with acute subbasal tooth.

Arnell: Genus Haemagogus 25

MALE GENITALIA (fig. 8). Segment VIII: Tergite about 0.80 of sternite;

distal margin rather deeply emarginate, with enlarged spiniform setae. Segment IX:

Tergite emarginate and weakly sclerotized mesally, lobes usually with 1 or 2 setae.

Sidepiece: Conical, length about 3.0 times median width; basal tergomesal lobe

not enlarged, but extending distad to near distal third of sidepiece as slightly

raised area with numerous short setae, basally with numerous setae, the more

proximal ones considerably enlarged; apical sternomesal scales lanceolate to broadly

lanceolate. Claspette: Stem slightly broader beyond middle, slightly angled dorsad

at distal third; filament striate, narrowly sickle-shaped. Clasper: Broadest at. base,

spiniform about 0.50 of clasper. Phallosome: Aedeagus large, broadly obovate,

tip with a dorsal carina proximad to a small beak; venter open except on basal

fourth, with heavily sclerotized ridges lateroventrally which are expanded basally

but not joined at midline. Proctiger: Cercal setae 6; apical knob of paraproct with

about 10-15 serrations.

PUPA (fig. 8). Abdomen: about 3.50 mm. Trumpet: 0.45 mm. Paddle: 0.70

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs

4,5-C more or less subequal, 1-4b, weaker than 7-C which is single. Trumpet:

Medium brown, lighter distally, reticulate sculpturing moderate. Abdomen: Weakly

to moderately pigmented, genital lobe and anterior segments darker. Float hair

(1-I) with about 10-12 primary branches and about 2-5 secondary branches. Hair

1-II-VII weakly to rather strongly developed, single, branched or sometimes den-

dritic. Hair 2-VII short, considerably cephalad of 1-VII. Hairs 3-II,ITI,5-IV,V more

or less subequal. Hair 6-VII slightly stronger than 6-III-VI which are subequal.

Hair 9-VII slightly cephalad of caudolateral margin of tergite, single or double,

often brush tipped; 9-VIII about 4-7b. Terminal Segments: Male genital lobe

relatively small, about 0.95 of tergite VIII. Paddle: Weakly to moderately pig-

mented, apex broadly rounded or slightly acuminate. Hair 1-P single.

LARVA (fig. 9). Head: 0.80 mm. Siphon: 0.68 mm. Anal Saddle: 0.33 mm.

Head: Hairs 5,6-C single. Thorax: Integument without spicules. Tubercles of hairs

5-6-P connected, tubercle of hair 7-P narrowly connected to 6-P. Abdomen: Hair

12-I absent. Segment VIII: Comb scales 6-10, with single sharply pointed, minutely

fringed spine, in single row. Siphon: Index about 2.0. Pecten teeth about 18(12-

22), extending to basal 0.50 of siphon. Hair 1-S double or triple (1-4b), inserted at

basal 0.55-0.60 of siphon. Anal Segment: Spines on caudal margin of saddle well

developed. Hair 4a-X 4b(3-5b), long. Boss strongly sclerotized.

SYSTEMATICS. Haemagogus anastasionis is separated from the other members

of the Albomaculatus Section: in the adults by the combination of (1) absence of

well developed lower stp bristle, (2) scales of vertex and occiput blue to bluish

green, (3) scales of mesonotum copper with blue scales laterally, (4) dark scales

of abdomen blue to bluish green with silver scales dorsally at base of tergites

IV-VII and (5) simple large claw of foreleg and midleg of male; in the male

genitalia from all species except chrysochlorus by (1) presence of large spiniform

setae and absence of large lanceolate scales on distal margin of tergite VIII and

(2) narrowly sickle-shaped claspette filament; and in the larva by the combination

of (1) single hairs 5,6-C, (2) integument without spicules, (3) absence of hair 12-I,

(4) hair 12-II weaker than 10-II, (5) short siphon (index about 2.0) and (6) strongly

sclerotized boss.

Haemagogus anastasionis is closely allied to chrysochlorus and spegazzinii. Char-

acters common to this anastasionis complex include the absence of a well developed

lower stp bristle, the simple large claw of the male foreleg and midleg, the distal

26 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

margin of tergite VIII of the males with enlarged setae but no elongate scales, and,

in the larvae, single hairs 5,6-C, the absence of hair 12-I and a strongly sclerotized

boss.

As interpreted here, anastasionis is composed of several apparently disjunct

populations. I have seen specimens from the Pacific versant of Central America,

the upper Magdalena River valley of Colombia, the Zulia River valley of Colombia

and Venezuela and the northern coast of Venezuela. In view of the discontinuous

distribution of this species. the literature records of anastasionis from the Orinoco

drainage of Venezuela and the Atlantic coast of Mexico can reasonably be assumed

to be valid. The single female specimen from Puerto La Cruz, Venezuela is unusual

in the amount of silver scaling on the apn and the more greenish hue of the

scales of the head and abdomen. I am placing it here until more specimens of

all stages can be secured and examined.

Haemagogus dominguezi was described by Duret on the basis of a single male

captured in the department of Zelaya, Nicaragua. This specimen appears to be

teratological, differing from anastasionis in the loss of the claspette filament and

in the abnormal coloration of some of the leg scales. It seems to agree in all other

aspects with typical males of anastasionis.

BIONOMICS. On the Pacific versant of Central America anastasionis inhabits

primarily scrub forest, second growth and open deciduous forest. The immature

stages are most commonly found in treeholes, but have also been taken from

bamboo internodes and fruits on the ground. It is apparently peridomestic in some

areas. It has been found at elevations up to approximately 500 meters in Nicaragua

and Colombia.

DISTRIBUTION (fig. 2). Haemagogus anastasionis is known from the Atlantic

coast of Veracruz and Campeche, Mexico and the Pacific versant of Central America

from Guatemala to San Jose, Costa Rica with one record from the Atlantic slope of

Nicaragua. Apparently absent from southern Costa Rica and Panama, it is present

in the Caribbean drainage of Colombia and Venezuela, with | record from the

Orinoco drainage of Venezuela. The records in Stone, Knight and Starcke (1959:

216) reporting anastasionis from Curacao and Peru refer to chrysochlorus and

obscurescens (=janthinomys) respectively. Material examined: 338 specimens; 36

males, 54 females, 33 pupae, 215 larvae; 33 individual rearings (15 larval, 13 pupal,

5 incomplete).

COLOMBIA. Cundinamarca: Girardot (10 km NW), 500 m, 11 Nov 1965, C. Marinkelle

(COM 16), 1 P, 19 L [UCLA]. Villeta, 1000 m, 1 M [USNM]; same data (GML), 1 M [UCLA];

same data, H. Kumm, 5 M, 4 F, 41 [USNM, UCLA]; same data, 1943, G. Boshell-Manrique and

H. Kumm, 3 1M gen, 3 M, 1 F, 61 [USNM]. Norte de Santander: Bachaquoro, Zulia, Mar 1947

P. Anduze, 1 M, 1 1 [UCLA, USNM].

COSTA RICA. Guanacaste: Las Canas, 9 June 1943, H. Kumm, 5 F [UCLA]. Palo Verde,

OTS Station, 10 m, 18 Aug 1971, D. Heinemann (CR 445), 1 lpF (445-10) [UCLA]. Santa Cruz,

12 Oct 1937, H. Kumm, 5 M, 5 F, 21, 4 L [UCLA, USNM, BM]. Puntarenas: Esparta (5 km E),

250 m, 13 Aug 1971, D. Schroeder (CR 355), 1 IP (355-13); same data (CR 358), 1 IpM (358-22)

[UCLA]. Las Loras, nr. Puntarenas, F. Knab, 1 F [USNM]. Puntarenas, 15 July 1924, A. Alfaro,

1 M, 3 F [USNM]. San Jose: San Jose, F. Knab, 1 F [USNM].

EL SALVADOR. San Miguel: San Miguel, 1 Aug 1941, H. Kumm, 1 M gen, 1 F [USNM].

San Salvador: Hacienda San Ramon, 17 June 1953 (GML), 1 IpF (01669), 1 lp (01668), 1 1

[UCLA].

GUATEMALA. Suchitepequez: Patulul, 20 July 1964, P. Cowsill (GUA 57), 1 pF (57-100)

[UCLA].

HONDURAS. Choluteca: Choluteca, Dec 1945, B. Avila, 1 M, 2 F [UCLA].

NICARAGUA. Chinandega: Castanones Peninsula, nr. Corinto, 31 Aug 1944, H. Crowell, 1 F

Arnell: Genus Haemagogus 27

[UCLA]. Corinto, 24 Oct 1944 (GML), 1 M gen; same data, 18 Sept 1945, H. Crowell, 1 M

[UCLA]. Leon: Puerto Somoza, 12 June 1964, A. Quinonez (NI 2), 4 lpM (2-101-103,105),

2 lpF (2-104,106), 25 L; same data, 13 June 1964 (NI 4), 3 IpF (4-107,108,111), 1 pM (4-105),

6 pF (4-101-104,110,112), 23 L; same data, 18 June 1964 (NI 20), 1 F [UCLA]. Simonillo

(nr. Nagarote), 14 June 1964, A. Quinonez (NI 8), 6 F; same data, 15 June 1964 (NI 10),

1 IpF (10-103), 1 pM (10-101), 1 pF (10-102), 1 P, 40 L; same data, 16 June 1964 (NI 12),

3 IpF (12-105,106,108), 3 pM (12-102,103,107), 1 P, 70 L [UCLA]. Rivas: San Juan del Sur,

24 Nov 1943, 1 M, 1 M gen [USNM]. Department unknown: Bozbollow, 28 Oct 1953 (GML),

2 F [UCLA].

VENEZUELA. Anzoatequi: Puerto La Cruz, 26 Sept 1944, W. Komp, 1 F [UCLA].

LOCALITY UNSPECIFIED. 1 F [USNM].

Additional Records From the Literature

MEXICO. Campeche: Campeche (Diaz Najera, 1960:185). Vera Cruz: Paso de San Juan (Vargas

and Martinez Palacios, 1953:37).

NICARAGUA. Zelaya (Duret, 1971:83).

VENEZUELA. Cojedes: Galeras del Pao (Cova Garcia, Sutil and Rausseo, 1966b:114). Zulia:

Sucre district (Anduze, 1947:353).

2. Haemagogus (H.) chrysochlorus Arnell, n. sp.

Figs. 2,36

TYPES: Holotype male with genitalia slide (721117-1), Curacao, E. van der Kuyp [USNM].

Allotype female, Curacao, E. van der Kuyp [USNM] . Paratype: 1 M with genitalia slide (46.V.13°),

Curacao, E. van der Kuyp [UCLA].

Haemagogus (Haemagogus) anastasionis of van der Kuyp (1954:54).

Haemagogus (Stegoconops) anastasionis in part of Forattini (1965:32-34).

Haemagogus anastasionis of van der Kuyp (1949a:69; 1949b:142; 1953a:146 1953b:38).

FEMALE. Wing: 2.70 mm. Proboscis: 2.30 mm. Forefemur: 1.95 mm. Abdo-

men: 2.45 mm. Dark scales of proboscis, palpus, wing and legs primarily violet

with some purple and green reflections. Head: Eyes narrowly separated above

antennae (1 ommatidial diameter). Decumbent scales of vertex bronze. Proboscis

about 1.20 of forefemur; palpus about 0.12 of proboscis; antenna about 0.70

of proboscis. Thorax: Scales of mesonotum copper, more green posteriorly, silver

in antealar area; scales of scutellum bronze. Apn lobes moderately enlarged, scales

light bronze, silver anteriorly and laterally; ppn scales light bronze; lower stp with-

out well developed bristle. Legs: Relatively short, length of forefemur about 1.30

of distance from top of thorax to apex of midcoxa. Wing: R, 43 about 1.50 of R,.

Abdomen: Darker scales light bronze with some copper reflections; silver scales

dorsally on tergites IV-VII, in basal patches on IV and V and basal bands continuous

with lateral silver patches on VI and VII.

MALE (fig. 36). Wing: 2.38 mm. Proboscis: 2.30 mm. Forefemur: 1.70 mm.

Abdomen: about 2.25 mm. Head: Proboscis about 1.35 of forefemur; palpus

short, about 0.13 of proboscis; antenna about 0.65 of proboscis. Legs: Larger

and smaller claws of foreleg and midleg simple.

MALE GENITALIA (fig. 36). Generally similar to anastasionis. Distal margin

of tergite VIII with very large spiniform setae.

PUPA. Unknown.

LARVA. Unknown.

28 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

SYSTEMATICS. Haemagogus chrysochlorus can be distinguished: in the adults

by (1) conspicuous light bronze color of the darker abdominal scales and basal

silver scales dorsally on tergites IV-VII, (2) copper scales on mesonotum and

head, (3) absence of well developed lower stp bristle, and (4) simple large and

small claws of male foreleg and midleg; and in the male genitalia from all but

anastasionis by (1) presence of large spiniform setae and absence of large lanceo-

late scales on distal margin of tergite VIII and (2) narrowly sickle-shaped claspette

filament.

A member of the anastasionis complex, chrysochlorus is closely allied to anasta-

sionis and less closely to spegazzinii with both of which it shares the absence of a

well developed lower stp bristle, the simple large claw of the foreleg and midleg

of the male and the absence of elongate scales on the distal margin of tergite VIII

of the male. The male genitalia are apparently indistinguishable from anastasionis,

although the spiniform setae on the distal margin of tergite VIII may possibly be

larger and fewer in chrysochlorus. It is readily separated from both anastasionis

and spegazzinii by the adult ornamentation and the male claws as indicated above.

Chrysochlorus undoubtedly arose as an isolate of the anastasionis stock.

BIONOMICS. Nothing is known of the bionomics of chrysochlorus other than

the report by van der Kuyp (1949b:143) of larvae found primarily in treeholes

following heavy rains and adults attacking man readily in shade.

DISTRIBUTION (fig. 2). Haemagogus chrysochlorus is known only from the

islands of Aruba and Curacao in the Netherlands Antilles. Material examined: 3

specimens; 2 males, 1 female.

NETHERLANDS ANTILLES. Curacao: type series, see above.

Additional Record From the Literature

NETHERLANDS ANTILLES. Aruba (van der Kuyp, 1953:146).

3. Haemagogus (H. ) spegazzinii Brethes

Figs. 2,10,11

1912. Haemagogus spegazzinii Brethes, 1912:39. TYPE: Holotype female (7273) and 2 slides

(3028), one with a wing and the other with genitalia, an antenna and a palpus, Jujuy

Province, Argentina, adult taken in dense woodland, 1907, D. Carlos Spegazzini [BA].

Synonymized with capricornii by Dyar (1921b:150); considered a synonym of equinus

by Dyar (1923:183); resurrected from synonymy by Cerqueira (1943:10-12) but mis-

interpreted as being conspecific with janthinomys Dyar, 1921b.

1928. Haemagogus uriartei Shannon and Del Ponte, 1928:68-69. TYPE: Holotype male (128-3)

with genitalia slide (2353) and pupal slide (V3), Vipos, Tucaman, Argentina, larva

collected in a treehole, 22 Mar 1927, R. Shannon and E. Del Ponte [USNM; see Stone

and Knight, 1955:289]. Synonymized with spegazzinii by Martinez, Carcavallo and

Prosen (1960:72).

1931. Haemagogus lindneri Martini, 1931a:118. TYPE: Holotype female, (?San Jose de)

Chiquitos, Santa Cruz, Bolivia, Oct 1926 [SMNS]. Synonymized with uriartei by Lane

(1939:121).

Haemagogus (Stegoconops) spegazzinii of Martinez, Carcavallo and Prosen (1961:72); Stone

(1963:132); Forattini (1965:34-35); Martinez and Carcavallo (1965:40); Morales-Ayala (1971:

142).

Haemagogus (Haemagogus) uriartei of Edwards (1932:179); Lane (1939:120-121); Levi-Castillo

(1951b:11,26-28).

Arnell: Genus Haemagogus 29

Haemagogus (Stegoconops) uriartei of Dyar (1928:135); Martini (1931a:117, in part; 1931b:211,

in part); Lane (1953:800-802); Stone, Knight and Starcke (1959:217); Galindo and Trapido

(1967:108).

Haemagogus uriartei of Antunes and Whitman (1937:827); Cerqueira (1943:12-13); Martinez

(1950:61); Kumm and Cerqueira (1951:174); Levi-Castillo (1951a:15); Bevier, Torrez-Munoz

and Doria-Medina (1953:474); Manso Soto, Martinez and Prosen (1953:33).

Haemagogus (Stegoconops) lindneri of Martini (1931b:212).

Haemagogus lindneri of Mattingly (1955:28).

FEMALE. Wing: 2.63 mm. Proboscis: 2.20 mm. Forefemur: 1.74 mm. Abdo-

men: 2.38 mm. Dark scales of proboscis, palpus, wing and legs predominantly

purple to violet. Head: Eyes narrowly separated above antennae (1 ommatidial

diameter). Decumbent scales of vertex and occiput light green, bluish green or

gold. Proboscis relatively long, about 1.25 of forefemur; palpus relatively short,

about 0.12 of proboscis; antenna about 0.70 of proboscis. Thorax: Scales of meso-

notum and scutellum light bronze to copper with red reflections, bluish green to

gold over supraalar bristles and on scutellar lobes, silver in antealar area. Apn lobes

moderately enlarged, scales almost entirely silver, often a few copper scales posterio-

dorsally; ppn scales gold to copper; lower stp without well developed bristle.

Legs: Midcoxa usually with patch of blue to violet scales near middle; legs moder-

ately long, length of forefemur about 1.45 of distance from top of thorax to apex

of midcoxa. Wing: R,,3; about 1.20-1.50 of R,. Abdomen: Dark scales of tergites

light bronze to bluish green with more purple reflections laterally; sternites purple

to blue.

MALE (fig. 10). Wing: 2.45 mm. Proboscis: 2.40 mm. Forefemur: 1.80 mm.

Abdomen: 2.40 mm. Head: Proboscis about 1.35 of forefemur; palpus short, about

0.14 of proboscis; antenna about 0.68 of proboscis. Legs: Larger claw of foreleg

and midleg simple; smaller claw with acute subbasal tooth. Abdomen: Distal seg-

ments and genitalia often with purple scales.

MALE GENITALIA (fig. 10). Segment VIII: Tergite about 0.70 of sternite;

distal margin broadly emarginate, with enlarged setae laterally and dense cluster

of about 15-20 large spiniform setae mesally. Segment IX: Tergite emarginate

mesally, weakly sclerotized at midline, strongly sclerotized immediately laterad,

lobes with 1 or 2 setae. Sidepiece: Conical, length about 3.5 times median width;

basal tergomesal lobe not enlarged but extending distad as slightly raised area

with numerous short setae to near distal third of sidepiece, basally with numerous

setae, the more proximal ones enlarged; apical sternomesal scales mostly broadly

lanceolate. Claspette: Slightly bowed outward before middle in dorsal aspect:

uniform in thickness, slightly curved dorsad, usually 2 large setae near middle of

inner surface; filament striate, broadly sickle-shaped. Clasper: Broadest at base;

spiniform about 0.50 of clasper. Phallosome: Aedeagus large, broadly obovate, tip

with a strongly sclerotized dorsal carina proximad to a small beak; venter broadly

open except on basal fourth, with heavily sclerotized ridges lateroventrally, expanded

basally to meet on midline and forming 2 small distally projecting spines. Proctiger:

Cercal setae 6-8; apical knob of paraproct with about 15-20 serrations.

PUPA (fig. 10). Abdomen: about 3.50 mm. Trumpet: 0.50 mm. Paddle: 1.00

mm. Cephalothorax: Moderately pigmented, slightly darker dorsally. Hairs 4,5-C

moderately developed, subequal, triple, weaker than 7-C which is double. Trumpet:

Medium brown, lighter distally; reticulate sculpturing moderate. Abdomen: Weakly

pigmented, genital lobe and anterior segments slightly darker. Float hair (1-I) with

about 14-16 primary branches and 2-6 secondary branches. Hair 1-II-VII weakly

30 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

to strongly developed, branched or dendritic. Hair 2-VII short, considerably cepha-

lad of 1-VII. Hairs 3-IL,III,5-[IV,V more or less subequal or with 3-III weaker. Hair

6-VII stronger than 6-IIJ-VI which are subequal. Hair 9-VII slightly cephalad of

dorsolateral margin of tergite, double; 9-VIII 8-12b. Paddle: Weakly pigmented,

apex slightly acuminate. Hair 1-P single.

LARVA (fig. 11). Head: 0.90 mm. Siphon: 1.00 mm. Anal Saddle: 0.41 mm.

Head: Hairs 5,6-C single. Thorax: Integument without spicules. Tubercles of hairs

5,6-P connected, tubercle of hair 7-P free. Abdomen: Hair 12-I absent. Segment

VIII: Comb scales 8-10, with single sharply pointed, minutely fringed spine, in

single row. Siphon: Index about 3.2-3.4. Pecten teeth 20-24, extending to about

basal 0.50 of siphon. Hair 1-S 3-4b, inserted at basal 0.60 of siphon. Anal Segment:

Spines on caudal margin of saddle very well developed, long, numerous. Hair 4a-X

4b, long. Boss strongly sclerotized.

SYSTEMATICS. Haemagogus spegazzinii can be distinguished from other species

of the Albomaculatus Section: in the adults by the combination of (1) almost

entirely silver apn lobes, (2) darker abdominal scales light bronze or occasionally

bluish green and absence of silver scales dorsally on abdominal tergites, (3) light

bronze to copper mesonotal scales, (4) absence of a well developed lower stp

bristle and (5) simple large claw of male foreleg and midleg; in the male genitalia

by spiniform setae mesally on the distal margin of tergite VIII and a broadly

sickle-shaped claspette filament; and in the larva by the combination of (1) single

hairs 5,6-C (2) integument without spicules, (3) absence of hair 12-I, (4) hair 12-II

weaker than 10-II, (5) relatively long siphon (index about 3.2-3.4) and (6) strongly

sclerotized boss.

Haemagogus spegazzinii is related to anastasionis and chryochlorus and is in-

cluded with these species in the anastasionis complex. It shares with these species

the absence of a well developed lower stp bristle, the simple large claw of the foreleg

and midleg of the male, the absence of elongate scales on the distal margin of tergite

VIII of the male and, in the larva, the absence of hair 12-I, single hairs 5,6-C and a

well-sclerotized boss. It can be separated from them on the basis of the adult

Ornamentation as discussed above, the broader claspette filament and the longer

siphon.

There appears to be considerable variation in the color of the dark scales of the

head, mesonotum and abdomen of spegazzinii, the abdomen, most conspicuously,

being usually light bronze but occasionally dark bluish green. However this variation

apparently has no geographical pattern.

Two female specimens in the USNM collection labeled only ““Ecuador/R. Levi-

Castillo” are typical spegazzinii, and, assuming the specimens are correctly labeled,

they significantly extend the range of this species.

It is evident from Brethes’ original description that spegazzinii is a species distinct

from janthinomys. Although I was not able to see the holotype of spegazzinii lam

confident that the material on which the above descriptions are based is conspecific

with it. I also find that the synonymy of uriartei with spegazzinii by Martinez,

Carcavallo and Prosen (1960:72) is correct. A review of the confusion of spegazzinii,

capricornii and janthinomys is given below in the discussion of the systematics of

the latter.

BIONOMICS. The larvae of this species are commonly found in treeholes and

bamboo internodes. Antunes and Whitman (1937) showed spegazzinii (as uriartei)

to be capable of harboring yellow fever virus but they were unable to effect trans-

mission. Bevier, Torres-Munoz and Doria-Medina (1953) found spegazzinii, as well

Arnell: Genus Haemagogus 31

as janthinomys, to be present in areas of yellow fever outbreaks near Santa Cruz,

Bolivia, where it may possibly play a role in transmission.

DISTRIBUTION (fig. 2). Haemagogus spegazzinii extends from eastern and south-

eastern Brazil to Paraguay, northern Argentina and eastern Bolivia, with 1 record

from the upper Amazon in Ecuador. The record in Stone, Knight and Starcke

(1959:217) of spegazzinii (as uriartei) from Venezuela undoubtedly refers to anasta-

sionis. Material examined: 47 specimens; 13 males, 18 females, 1 pupa, 15 larvae;

1 individual rearing (incomplete).

ARGENTINA. Cordoba: Alta Gracia, 12 Feb 1918, 1 F [USNM]. Formosa: El Coati, Nov

1949, 1 F [USNM]. Jujuy: Ledesma (Libertador General San Martin), G. Boshell-Manrique and

H. Kumm, 3 F, 1 M, 1 L [UCLA, USNM]; same data, 1943, H. Kumm, 1 1 [USNM]. Locality

unspecified, Dec 1968, W. Patterson, 3 M, 2 F [BM]. Santiago del Estero: Frias, 24 Feb 1960,

11 L [USNM]. Tucuman: Raco, 13 Feb 1927, R. Shannon, 1 F [USNM].

BOLIVIA. Santa Cruz: Cercado, Jan 1939, 1 F [USNM]. Pozo Redondo, Cordillera, 7 Mar

1944, Carr, 2 F [UCLA]. Santa Cruz, 30 Dec 1943, Torrez-Munoz, 1 M [UCLA]. Locality unspeci-

fied, 9 Dec 1943, 3 M, 7 F [UCLA]. Locality unspecified: Carr, 2 M [USNM].

BRAZIL. Bahia: Esplanada, 18 Jan 1936, 1 M [BH]. Sacco do Soares, 1931, H. Kumm, 1 F

[BM]. Ceara: Fortaleza, Mar-Apr 1938, 3 M, 4 F [USNM, BH]. Fortaleza, J. Aragao, 1 L [BH];

same data, 8 Mar 1947, V. Iracema, 1 lp [USNM]. Locality unspecified, 2M [BH] . Para: ?Belem,

Apr 1930, N. Davis, 4 M, 1 F [BM]. Pernambuco: Sao Goncalo, May 1933, 1 F [USNM]. Guana-

bara: Rio de Janeiro, 1 M, 1 F [GML]. Sergipe: Nossa Senhora das Dores, 22 Nov 1946,

F. Pedrinhas, 1 M gen [USNM].

ECUADOR. Locality unspecified: R. Levi-Castillo, 2 F [USNM].

LOCALITY UNSPECIFIED. 1 M [USNM].

Additional Records From the Literature

BRAZIL. Goias: Sao Pedro on Bananal Island (Kumm and Cerqueira, 1951:174). Mato Grosso:

Aquidauana (Kumm and Cerqueira, 1951:174).

PARAGUAY. Boqueron: Fortin M. Estigarriba and Puerto Casado (Manso Soto, Martinez and

Prosen, 1953:39).

4. Haemagogus (H.) baresi Cerqueira

“Pigs. 2512513

1960. Haemagogus (Stegoconops) baresi Cerqueira, 1960:1-5, 2 plates. TYPE: Holotype male

(1607.3) with associated pupal skin (slide 316; 2961) but genitalia (slide 332) missing,

Igarape do Taruma, Manaus, Amazonas, Brazil, larva from treehole in dark forest,

6 Dec 1956, C. Elias [FH, 15105].

Haemagogus (Stegoconops) baresi of Stone (1961:44).

FEMALE. Wing: 2.40 mm. Proboscis: 2.05 mm. Forefemur: 1.85 mm. Abdo-

men: 2.50 mm. Dark scales of proboscis, palpus, wing and legs violet with some

purple reflections. Head: Eyes narrowly separated above antennae (1 ommatidial

diameter). Decumbent scales of vertex and occiput dark blue with some bluish green

reflections. Proboscis short, about 1.10 of forefemur; palpus about 0.12 of pro-

boscis. Antenna long, about 0.80 of proboscis. Thorax: Scales of mesonotum and

scutellum dark blue with some green reflections. Apn lobes considerably enlarged,

scales dark blue; ppn scales dark blue with some purple reflections; lower stp with-

out well.developed bristle. Legs: Coxae with patches of violet to purple scales

near middle; legs moderately long, forefemur length about 1.50 of distance from

32 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

top of thorax to apex of midcoxa; forefemur and midfemur entirely dark scaled;

hindfemur with few silver scales in streak on anterior surface extending to basal

0.75. Wing: R43 about 0.55 of R,. Abdomen: Dark scales purple with some

violet reflections; bluish green scales present laterally on tergite VII.

MALE (fig. 12). Wing: 2.75 mm. Proboscis: 2.50 mm. Forefemur: 1.85 mm.

Abdomen: about 2.60 mm. Head: Proboscis about 1.35 of forefemur; palpus

very short, about 0.09 of proboscis; antenna long, about 0.80 of proboscis. Legs:

Larger claw of foreleg and midleg simple; smaller claw of foreleg and midleg with

small subbasal tooth.

MALE GENITALIA (fig. 12). Segment VIII: Tergite about 0.85 of sternite;

distal margin broadly emarginate, with about 15 lanceolate scales mesally. Segment

IX: Tergite broadly and deeply emarginate, weakly sclerotized mesally; lobes with

1 or 2 setae. Sidepiece: Conical, length about 3.5 times median width; basal tergo-

mesal area not enlarged, with numerous setae, the more proximal ones much

elongate, somewhat flattened and attenuate; mesal half distad of basal sterno-

mesal area with setae rather evenly distributed; apical sternomesal scales narrowly

lanceolate with acuminate tips. Claspette: Stem slightly curved dorsad; filament

striate, broadly sickle-shaped, tip slightly recurved. Clasper: Broadest at base;

spiniform about 0.60 of clasper. Phallosome: Aedeagus large, broadly obovate,

tip with a small beak; venter broadly open except on basal-third, with heavily

sclerotized ridges lateroventrally which are expanded basally, ovoid sclerotized

lobes present basally at midline. Proctiger: Cercal setae 4; apical knob of para-

proct with about 10 striations.

PUPA (fig. 12). Abdomen: about 3.55 mm. Trumpet: 0.45 mm. Paddle: 0.75

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C sub-

equal, single, weaker than 7-C which is single. Trumpet: Dark brown, lighter

distally, reticulate sculpturing moderate. Abdomen: Float hair (1-I) with 10-12

primary branches and 2-6 secondary branches. Hair 1-I]-VII rather weakly devel-

oped, single or double. Hair 2-VII short, considerably cephalad of 1-VII. Hair

3-II single, considerably longer than hairs 3-III,5-IV,V, which are single. Hair 6-VII

rather weakly developed, subequal to 6-VI. Hair 9-VII near caudolateral margin of

tergite, single; 9-VIII 2-6b. Terminal Segments: Male genital lobe about 1.0 of

tergite VIII. Paddle: Rather broad, apex acuminate, midrib dark. Hair 1-P single.

LARVA (fig. 13). Head: 0.95 mm. Siphon: 0.90 mm. Anal Saddle: 0.35 mm.

Head: Hair 6-C double. Thorax: Integument without spicules. Tubercles of hairs

5,6-P broadly connected, tubercle of hair 7-P narrowly connected to 6-P. Abdo-

men: Hair 12-I absent. Hair 7-IJ-[V well developed, double, much larger than

hair 5 of corresponding segment. Hairs 1,13-III-V very well developed. Segment

VIII: Comb scales 8-10, with single, pointed, minutely fringed spine, fringe at

apex of spine larger and more conspicuous, in single row. Siphon: Index about

2.8. Pecten teeth about 11-13, extending to about basal 0.45 of siphon. Hair 1-S

double, inserted at basal 0.55 of siphon. Anal Segment: Spines on caudal margin

of saddle enlarged, conspicuous dorsolaterally. Hair 1-X double, long. Hair 4a-X

double or triple, long. Boss strongly sclerotized.

SYSTEMATICS. Haemagogus baresi can be separated from the remaining species

of the Albomaculatus Section: in the adults by the combination of (1) absence of

well developed lower stp bristle, (2) short female proboscis (1.10 of forefemur),

(3) dark blue scales on head, mesonotum, apn and ppn, (4) dark scales on coxae,

(5) entirely dark scaled forefemur and midfemur, and (7) simple large claw of male

foreleg and midleg; in the male genitalia by the combination of (1) elongate,

Arnell: Genus Haemagogus 33

lanceolate scales mesally on distal margin of tergite VIII, (2) broadly sickle-shaped

claspette filament and (3) aedeagus broadly rounded on distal fourth with very

small, weakly sclerotized beak at apex; and in the larva by the combination of

(1) double hair 6-C, (2) integument without spicules, (3) absence of hair 12-I,

(4) hair 12-II weaker than 10-II and (5) hair 7-II-[V double and much larger than

hair 5 of corresponding segment.

Haemagogus baresi shares with anastasionis, chrysochlorus and spegazzinii the

absence of a well developed lower stp bristle and the absence of a tooth on the

large claw of the foreleg and midleg of the male. In the male genitalia baresi

closely resembles andinus. Baresi may be an annectent from between the anastasi-

onis complex and the andinus complex.

BIONOMICS. Nothing is known of the bionomics of baresi.

DISTRIBUTION (fig. 2). Haemagogus baresi is known only from 2 localities

on the middle Amazon, Iquitos, Peru and Manaus, Brazil. Material examined:

8 specimens; 2 males, 1 female, 2 pupae, 3 larvae; 3 individual rearings (1 larval,

1 pupal, 1 incomplete).

BRAZIL. Amazonas: Igarape do Taruma, Manaus, 6 Dec 1956, C. Elias, 1 pM (holotype),

1 lpF (allotype) [FH].

PERU. Loreto: Iquitos, 28 May 1931, R. Shannon, 1 IM, 11 [USNM].

5. Haemagogus (H. ) andinus Osorno-Mesa

Figs. 4,14,15

1944. Haemagogus andinus Osorno-Mesa, 1944b:170-175. TYPE: Holotype male with genitalia

slide, coffee plantation near Fusagasuga, Cundinamarca, Colombia, elev. 1746 m, larva

collected in “guamos” rothole (Inga sp.), May 1942, E. Osorno-Mesa [USNM] .

Haemagogus (Haemagogus) andinus of Levi-Castillo (1951b:11,28-30).

Haemagogus (Stegoconops) andinus of Lane (1953:794-796); Stone, Knight and Starcke (1959:

216, :

Haemagogus andinus of Hovanitz (1946:35); Kumm, Osorno-Mesa and Boshell-Manrique (1946:

17); Barreto Reyes (1955:78).

FEMALE. Wing: 3.35 mm. Proboscis: 2.72 mm. Forefemur: 2.45 mm. Abdo-

men: 3.20 mm. Dark scales of proboscis, palpus, wing and legs mixed purple and

violet. Head: Eyes narrowly separated above antennae (2 ommatidial diameters).

Decumbent scales of vertex and occiput copper. Proboscis relatively short, about

1.10 of forefemur; palpus relatively short, about 0.13 of proboscis; antenna about

0.62 of proboscis. Thorax: Scales of mesonotum deep copper, green over supraalar

bristles, a few silver scales in antealar area; scales on scutellum copper, green on

lobes. Apn lobes considerably enlarged, scales green, with silver scales anteriorly; ppn

scales deep copper; psp scales with light green or copper reflection. Legs: Fore-

coxa with small patch of purple or violet scales near middle; legs moderately long,

forefemur length about 1.50 of distance from top of thorax to apex of midcoxa;

knee spots present on midfemur and hindfemur. Wing: R,,3; about 0.95 of Ro.

Abdomen: Dark scales light green to light bronze except for blue scales on tergite

VIII.

MALE (fig. 14). Wing: 2.95 mm. Proboscis: 2.85 mm. Forefemur: 2.20 mm.

Abdomen: 3.25 mm. Head: Proboscis about 1.30 of forefemur. Legs: Larger claw

of foreleg and midleg with acute submedian tooth, smaller claw of foreleg and

midleg with acute subbasal tooth.

34 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

MALE GENITALIA (fig. 14). Segment VIII: Tergite about 0.70-0.75 of sternite;

distal margin broadly emarginate, with enlarged setae laterally and cluster of 25-35

lanceolate scales mesally. Segment IX: Tergite deeply emarginate and weakly scle-

rotized mesally; lobes with 1 or 2 setae. Sidepiece: Conical; length 3.5 times median

width; basal tergomesal lobe slightly enlarged, with numerous setae, the more

proximal ones much elongate and flattened and attenuate; mesal half distad of

basal tergomesal area with setae rather evenly distributed, short, longer on distal

fourth; apical sternomesal scales oblanceolate, usually with acuminate tip. Claspette:

Stem broader near base, constricted at apical third and sharply angled dorsad

beyond constriction; filament striate, broadly sickle shaped. Clasper: Broadest just

before middle; spiniform about 0.55 of clasper. Phallosome: Aedeagus large, broadly

obovate, tip produced distally into a strongly sclerotized dorsal carina proximad

to a small beak; venter broadly open except on basal fourth, with heavily sclerotized

‘ ridges lateroventrally which are expanded basally into ovoid sclerotized lobes at

midline. Proctiger: Cercal setae 6-8; apical knob of paraproct with about 10 stria-

tions.

PUPA (fig. 14). Abdomen: about 3.45 mm. Trumpet: 0.40 mm. Paddle: 0.85

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C sub-

equal, 1-4b, weaker than 7-C which is usually single. Trumpet: Dark brown, lighter

distally, reticulate sculpturing moderate. Abdomen: Moderately pigmented, genital

lobe and anterior segments darker. Float hair (1-I) with 10-14 primary branches

and 4-10 secondary branches. Hair 1-II-VII strongly developed, dendritic. Hair

2-VII short, considerably cephalad of 1-VII. Hairs 3-IL,III,5-IV,V subequal or with

5-IV,V slightly weaker than 3-II, III; 5-IV,V less than 0.75 of corresponding tergite.

Hair 6-VII moderately well developed, single or double, stronger than 6-VI, which

is stronger than 6-III-V. Hair 9-VII near caudolateral margin of tergite, usually

triple; 9-VIII usually 5-7b. Terminal Segments: Male genital lobe 1.2 of tergite

VIII. Paddle: Relatively narrow, apex acuminate; with darker pigmentation laterally

and along midrib. Hair 1-P single.

LARVA (fig. 15). Head: 0.85 mm. Siphon: 0.85 mm. Anal Saddle: 0.33 mm.

Head: Hair 5-C usually single, occasionally double. Hair 6-C single. Thorax: Integu-

ment without spicules. Tubercles of hairs 5-6-P usually connected, tubercle of hair

7-P free. Abdomen: Hair 12-I absent. Segment VIII: Comb scales about 10(8-14),

with single sharply pointed, minutely fringed spine, in single row. Siphon: Index

about 3.5(3.3-3.8). Pecten teeth 16-20(14-23), extending to about basal 0.50 of

siphon. Hair 1-S 4b(3-5b), inserted at basal 0.55 of siphon. Anal Segment: Spines

on caudal margin of saddle very well developed, extremely elongate and conspicuous

dorsolaterally. Hair 4a-X double, considerably shorter than hair 4b-X. Boss strongly

sclerotized.

SYSTEMATICS. Haemagogus andinus can be distinguished from the remaining

members of the Albomaculatus Section: in the adults by the combination of (1)

light green to light bronze abdominal scales with no silver scales dorsally at bases

of abdominal tergites, (2) knee spots of silver scales on midfemur and hindfemur,

(3) copper scales on vertex, mesonotum and ppn (4) relatively short female pro-

boscis (1.10 of forefemur) and (5) single well developed lower stp bristle; in the

male genitalia by the combination of (1) long, lanceolate scales mesally on distal

margin of tergite VIII, (2) apical fourth of aedeagus broadly triangular with strongly

sclerotized dorsal carina and (3) claspette stem angled sharply dorsad at apical

third; and in the larva by the combination of (1) integument without spicules,

(2) hair 12-I absent, (3) long siphon (index usually about 3.5), (4) hair 5-C usually

Arnell: Genus Haemagogus a5

single and (6) hair 4a-X usually double.

Andinus is closely related to nebulosus, with which it forms the andinus complex.

It differs in the female in the color of the dark scales of the head, thorax and

abdomen and in the larva by the slightly shorter siphon and fewer branches on

hair 4a-X. Its relationship to other members of the Albomaculatus Section is

obscure, although it may be related to the anastasionis complex through baresi,

as the male genitalia are much like those of baresi.

BIONOMICS. Virtually nothing is known of the bionomics of andinus. It is

uncommon and appears to be restricted to high elevations along the Eastern

Cordillera of the Colombian Andes. The few available larval collections have been

from treeholes.

DISTRIBUTION (fig. 4). Haemagogus andinus is known only from elevations

near 2000 meters on the western slopes of the Eastern Cordillera of the Colombian

Andes. Material examined: 75 specimens; 16 males, 11 females, 11 pupae, 37

larvae; 11 individual rearings (9 larval, 2 incomplete).

COLOMBIA. Cundinamarca: Fusagasuga, 1746 m, 1 F [BM]; same data, 2 IM gen, 1 lp, 4M,

5 M gen, 6 F, 141 [USNM]; same data, H. Kumm, 1 F, 1 1 [UCLA]. Hacienda Normandia, nr.

Fusagasuga, 1740 m, 23 Sept 1966, E. Osorno-Mesa and Morales (COB 107), 6 lpM (107-10,12-

16), 3 IpF (107-11,17,18), 1 P, 9 L [UCLA]. Santander: Locality unspecified, 2 L [USNM].

Additional Record From the Literature

COLOMBIA. Santander: Jesus Maria (Kumm, Osorno-Mesa and Boshell-Manrique, 1946:17).

6. Haemagogus (H.) nebulosus Arnell, n. sp.

Fig. 4,16,17

TYPES: Holotype female with associated larval and pupal skins (VZ 151-10), Rancho Grande,

Aragua, Venezuela, elev. 1100 m, larva taken from small treehole, 5 July 1969, T. and J. Zavortink

[USNM]. Paratypes: 2 \pF (VZ 151-11,12), 2 pF (VZ 151-100,101), same data as holotype

[UCLA, BM, MDM].

FEMALE. Wing: 3.25 mm. Proboscis: 2.65 mm. Forefemur: 2.40 mm. Abdo-

men: 3.00 mm. Dark scales of proboscis, palpus, wing and legs violet with purple

reflections. Head: Eyes narrowly separated above antennae (1 ommatidial dia-

meter). Decumbent scales of vertex and occiput light bluish green. Proboscis short,

about 1.10 of forefemur, with small patch of silver scales ventrally at base; palpus

about 0.15 of proboscis; antenna about 0.68 of proboscis. Thorax: Scales of

mesonotum predominantly copper, green mixed with copper laterally and posteri-

orly, greenish blue over supraalar bristles and silver in antealar area; scales green and

copper on scutellum. Apn lobes slightly enlarged, scales blue with silver scales

anteriorly; ppn scales green, a few silver scales posteriorly; psp scales with light

green reflection. Legs: Legs moderately long, length of forefemur about 1.5 of

distance from top of thorax to apex of midcoxa; knee spots present on midfemur

and hindfemur. Wing: R243 about 0.70-0.95 of R,. Abdomen: Dark scales dark

blue with purple reflections; basolateral silver patches usually extending to distal

margin of tergites II-VII forming nearly complete lateral silver line.

MALE. Unknown.

PUPA (fig. 16). Abdomen: about 4.00 mm. Trumpet: 0.55 mm. Paddle: 1.05

mm. Cephalothorax: Moderately pigmented, darker dorsally. Hair 5-C slightly strong-

er than 4-C and weaker than 7-C. Trumpet: Dark brown, lighter distally, reticulate

36 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

sculpturing moderate. Abdomen: Moderately pigmented, genital lobe and anterior

segments considerably darker. Float hair (1-I) with about 12-16 primary branches

and 2-6 secondary branches. Hair 1-II-VII very strongly developed, dendritic. Hair

2-VII short, considerably cephalad of 1-VII. Hair 3-II subequal to or slightly strong-

er than 3-III, and weaker than 5-IV,V which are subequal and as long as correspond-

ing tergite. Hair 6-VII moderately well developed, single or double, stronger than

6-VI which is stronger than 6-III-V. Hair 9-VII slightly cephalad of caudolateral

margin of tergite, 4-6b; 9-VIII usually 5-7b. Paddle: Relatively narrow, apex acumi-

nate, brown pigmented, midrib darker. Hair 1-P single.

LARVA (fig. 17). Head: 0.95 mm. Siphon: 1.05 mm. Anal Saddle: 0.38 mm.

Head: Hair 5-C double (double or triple). Hair 6-C single. Thorax: Integument with-

out spicules. Tubercles of hairs 5-7-P free or narrowly connected. Abdomen: Hair

12-I absent. Segment VIII: Comb scales 9-10(9-12), with single sharply pointed,

minutely fringed spine, in single row. Siphon: Index about 3.8. Pecten teeth 18-22

(16-28), extending to about basal 0.55 of siphon. Hair 1-S 4-5b(3-6b), inserted at

basal 0.60 of siphon. Anal Segment: Spines on caudal margin of saddle very well

developed, extremely elongate and conspicuous dorsolaterally. Hair 4a-X 4-5b. Boss

strongly sclerotized. |

SYSTEMATICS. Haemagogus nebulosus can be separated from the remainder.

of the Albomaculatus Section: in the adults by the combination of (1) enlarged

lower stp bristle, (2) dark blue abdominal scales with no silver scales dorsally at

bases of abdominal tergites, (3) knee spots of silver scales at apex of midfemur

and hindfemur, (4) scales of vertex bluish green and (5) relatively short proboscis

(1.10 of forefemur) with small patch of silver scales ventrally at base; and in the

larva by the combination of (1) integument without spicules, (2) hair 12-I absent,

(3) long siphon (index about 3.8), (4) hairs 5-C usually double and (6) hair 4a-X

usually 4-or 5-branched.

Nebulosus is closely allied to andinus but can be distinguished by the color

of the dark scales of the head, thorax and abdomen of the female and the longer

siphon and more branched hair 4a-X of the larva. Both species are apparently

restricted to high elevations in the northern ranges of the Andes.

BIONOMICS. Nothing is known of the bionomics of nebulosus. The only collec-

tion of this species was made in a very small treehole. The general habitat was

a dense cloud forest which is described in a thorough study by Beebe and Crane

(1947).

DISTRIBUTION (fig. 4). Haemagogus nebulosus is known only from a single

locality in the Cordillera de la Costa near Maracay, Venezuela. Material examined:

13 specimens; 5 females, 5 pupae, 3 larvae; 5 individual rearings (3 larval, 2 pupal).

VENEZUELA. Aragua: Rancho Grande, type series, see above.

7. Haemagogus (H.) janthinomys Dyar

Figs. 3,18 ,19,520

1921. Haemagogus (Haemagogus) janthinomys Dyar, 1921a:112-113. TYPE: Lectotype male

(17-1) with associated larval skin and genitalia slide (219), St. Ann’s, Port of Spain,

St. George, Trinidad, Trinidad and Tobago, larva collected in a treehole, June 1905,

F.W. Urich [USNM, 24335; selection of Stone and Knight, 1955:288]. Synonymized

with capricornii by Antunes (1939:106); synonymized with spegazzinii by Cerqueira

(1943:10); considered a subspecies of capricornii by Martinez, Carcavallo and Prosen

(1960:76).

Arnell: Genus Haemagogus 37

1931. Haemagogus uriartei var. obscurescens Martini, 1931b:212. TYPE: Lectotype female

(8863 ;3/30), Ucayali River, Loreto, Peru, 24 Oct 1903 [BM; selection of Mattingly,

1955:28]. Synonymized with anastasionis by Stone (1957:339-340). NEW SYNONYMY.

1946. Haemagogus spegazzinii falco Kumm, Osorno-Mesa and Boshell-Manrique, 1946:14-28,

pl. 1-5. TYPE: Syntypes male, female, larvae, Volcanes Forest in Pitas River Valley nr.

Caparrapi, Cundinamarca, Colombia, May and June 1943, H.W. Kumm, E. Osorno-Mesa

and J. Boshell-Manrique [USNM; lectotype not selected, Stone and Knight, 1955:288].

Synonymized with janthinomys by Levi-Castillo (1956:346); resurrected from synonymy

with janthinomys by Galindo (1957:121-122); considered a subspecies of capricornii by

Martinez, Carcavallo and Prosen (1960:76). NEW SYNONYMY.

1961. Haemagogus (Stegoconops) capricornii petrocchiae Martinez, Carcavallo and Prosen,

1961:79-82. TYPE: Holotype male, Salvador Mazza (Pocitos), Departmento General

Jose de San Martin, Salta, Argentina, Jan 1960, A. Martinez and R. Carcavallo [BA].

NEW SYNONYMY.

Haemagogus (Haemagogus) janthinomys of Bonne and Bonne-Wepster (1925:435); Dyar (1928:

140); Edwards (1932:179).

Haemagogus (Stegoconops) capricornii janthinomys of Martinez, Carcavallo and Prosen (1961:

76); Stone (1963:132); Forattini (1965:19); Martinez and Carcavallo (1965:42).

Haemagogus spegazzinii janthinomys of Levi-Castillo (1956:346).

Haemagogus janthinomys of Shannon (1931:8); Kumm and Frobisher (1932:352); Komp (1936:

61); Antunes (1937:75); Antunes and Whitman (1937:827); Boshell-Manrique (1938:415);

Kumm and Novis (1938:501); Bugher (1941 :303).

Haemagogus uriartei var. obscurescens of Lane (1939:121); Mattingly (1955:28).

Haemagogus (Haemagogus) spegazzinii falco of Levi-Castillo (1951b:11,21-22).

Haemagogus (Stegoconops) spegazzinii falco of Lane (1953:794); Stone, Knight and Starcke

(1959:217); Fauran (1961:31); Cova Garcia, Sutil and Rausseo (1966a:61-62, fig. 117; 1966b:

114-115, fig. 200).

Haemagogus (Stegoconops) capricornii falco of Martinez, Carcavallo and Prosen (1961:76); Stone

(1963:132); Forattini (1965:19); Martinez and Carcavallo (1965:42); Morales-Ayala (1971:

142).

Haemagogus spegazzinii falco of Anderson and Osorno-Mesa (1946:613); Hovanitz (1946:35);

Bates (1947:1); Osorno-Mesa (1947:455); Waddell and Kumm (1948:247); Galindo, Carpenter

and Trapido (1949:277; 1951a:114-116; 1951b:104-106; 1955:158); Galindo, Trapido and

Carpenter (1950:546); Kumm and Cerqueira (1951:172-173); Levi-Castillo (1951a:14; 1954:

84); Carpenter, Galindo and Trapido (1952:162); Elton (1952a:157; 1952b:428); Komp

(1952:330; 1956:37); Bevier, Torres-Munoz and Doria-Medina (1953:474); Vargas-Mendez

and Elton (1953:857); Mackie, Hunter and Worth (1954:19); Barreto Reyes (1955:78); Galindo

and Trapido (1955:545; 1957:147); Trapido and Galindo (1955:669; 1956a:303; 1957:122);

Trapido, Galindo and Carpenter (1955:525); Galindo, de Rodaniche and Trapido (1956:1022);

Galindo, Trapido, Carpenter and Blanton (1956:544); de Rodaniche (1956:480); Galindo

(1957:121-124); de Rodaniche and Galindo (1957:236; 1961:393); de Rodaniche, Galindo

and Johnson (1957:682); Foote and Cook (1959:141); Galindo, de Rodaniche and Johnson

(1959:557); Groot, Morales and Vidales (1961:399); Galindo and de Rodaniche (1964:846).

Haemagogus (Stegoconops) capricornii petrocchiae of Stone (1963:132); Forattini (1965:19);

Martinez and Carcavallo (1965:42).

Haemagogus (Haemagogus) capricornii in part of Lane (1939:118).

flaemagogus (Stegoconops) capricornii of Bonne-Wepster and Bonne (1923:127); Bonne and

Bonne-Wepster (1925:431); Stone, Knight and Starcke (1959:216, in part); Morales-Ayala

(1971:142).

Haemagogus capricornii of Howard, Dyar and Knab (1917:877, in part); Dyar (1921b:150);

Lutz and Nunez-Tovar (1928:25); Antunes (1939:106); Shannon, Whitman and Franca (1938:

111, in part); Shannon (1939:137, in part); Bates (1943:23; 1944a:84; 1944b:160; 1945:17;

1946:47); Bugher, Boshell-Manrique, Roca-Garcia and Osorno-Mesa (1944:31); Boshell-Manrique

and Osorno-Mesa (1944:172); Osorno-Mesa (1944a:45); Bates and Roca-Garcia (1945:204);

Floch and Abonnenc (1947:11); Bevier, Torres-Munoz and Doria-Medina (1953:474).

38 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Stegoconops capricornii of Howard, Dyar and Knab (1913:figs. 165,438).

Aedes capricornii in part of Dyar and Knab (1906b:163-164).

Haemagogus (Haemagogus) spegazzinii of Cerqueira and Boshell-Manrique (1946:193-198); Anduze

(1947:353); Levi-Castillo (1951b:11,19-21, in part).

Haemagogus (Stegoconops) spegazzinii of Martinez (1950:61); Stone, Knight and Starcke (1959:

216); Fauran (1961:31); Aitken, Worth and Tikasingh (1968:257).

Haemagogus (Stegoconops) spegazzinii spegazzinii in part of Lane (1953:792-793).

Haemagogus spegazzinii of Shannon and Del Ponte (1928:67); Martini (1931b:212, in part);

Cerqueira (1943:10-12); Waddell (1945:329; 1949:568); Cerqueira and Lane (1945:286);

Waddell and Taylor (1945:229); Bates and Roca-Garcia (1946:586); Hovanitz (1946:35);

Kumm, Osorno-Mesa and Boshell-Manrique (1946:17-19); Laemmert, Ferreira and Taylor

(1946: 43); Causey and Kumm (1948:470); Waddell and Kumm (1948:247, in part); Causey

and Dos Santos (1949:477); Causey, Kumm and Laemmert (1950:302); Kumm (1950:677);

Kumm and Laemmert (1950:750); Laemmert and Kumm (1950:724); Kumm and Cerqueira

(1951a:171-172; 1951b:196); Levi-Castillo (1951a:13, in part); Komp (1952:330; 1955b:

137-138; 1955e:277-280); Bevier, Torres-Munoz and Doria-Medina (1953:474); Manso Soto,

Martinez and Prosen (1953:33); Mackie, Hunter and Worth (1954:19); Horsfall (1955:536);

Trapido (1955:619); de Rodaniche and Galindo (1957:235); Stone (1957:340); Boshell-

Manrique and Bevier (1958:27); Kerr, Roca-Garcia and Bugher (1960:26).

Haemagogus spegazzinii spegazzinii of Aitken (1955:575); Downs, Aitken and Anderson (1955:

840); Levi-Castillo (1956:345); Foote and Cook (1959:142).

_ Haemagogus (Stegoconops) anastasionis in part of Morales-Ayala (1971:142).

Haemagogus anastasionis in part of Stone (1957:340); Forattini (1965:32-34).

Haemagogus (Stegoconops) equinus in part of Dyar (1923:183; 1928:134); Edwards (1932:179).

Haemagogus equinus of Dyar (1925b:138, in part); Shannon (1931:8, in part); Soper (1935:172).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:867).

Haemagogus splendens in part of Howard, Dyar and Knab (1917:867).

Haemagogus regalis in part of Theobald (1910:493-494).

Haemagogus cyaneus in part of Theobald (1903a:308; 1907:550); Lutz (1904:13).

FEMALE (fig. 18). Wing: 3.25 mm. Proboscis: 2.55 mm. Forefemur: 2.25 mm.

Abdomen: 3.20 mm. Dark scales of proboscis, palpus, wing and legs predominately

purple, with some violet reflections. Head: Eyes narrowly separated above antennae

(1 ommatidial diameter). Decumbent scales of vertex and occiput violet to bluish

green with some purple reflections, dark scales often continuing laterally to post-

gena. Proboscis short, about 1.10-1.15 of forefemur; palpus about 0.15 of proboscis;

antenna about 0.74-0.77 of proboscis. Thorax: Scales of mesonotum usually dark

green, occasionally copper, usually blue or bluish green in fossa and posteriorly,

bright blue over supraalar bristles, silver in antealar area; scales dark green to

bright blue on scutellar lobes. Apn lobes moderately enlarged, scales blue to bluish

green, occasionally with a few silver scales laterally; ppn scales bronze to dark

green. Legs: Forecoxa and midcoxa with small to extensive patch of purple scales

near middle; legs moderately long, forefemur length about 1.44-1.50 of distance

from top of thorax to apex of midcoxa; hindfemur with silver scales on anterior

surface in conspicuous streak to near apex. Wing: R,,3; about 0.90-1.60 of R,.

Abdomen: Dark scales purple with some violet reflections; distal margins of tergites

V-VIII with dark bluish green scales, more numerous laterally, a few bluish green

scales on distal margins of more proximal segments; silver scales dorsally in basal

band on tergites VII and VIII; dark scales of sternites purple with bluish green

scales distally.

MALE (fig. 18). Wing: 2.90 mm. Proboscis: 2.65 mm. Forefemur: 2.05 mm.

Abdomen: 2.80 mm. Head: Proboscis about 1.25-1.40 of forefemur; palpus short,

Arnell: Genus Haemagogus 39

about 0.15 of proboscis; antenna about 0.62-0.68 of proboscis. Legs: Larger claw

of foreleg and midleg with acute subbasal tooth, smaller claw of foreleg and mid-

leg with acute basal tooth.

MALE GENITALIA (fig. 19). Segment VIII: Tergite about 0.60-0.65 of sternite;

distal margin broadly emarginate, with enlarged setae laterally and cluster of 20-30

lanceolate scales mesally. Segment IX: Tergite emarginate and weakly sclerotized

mesally; lobes with usually 2 setae. Sidepiece: Conical, length about 3.5 times

median width; basal tergomesal lobe slightly enlarged, with numerous setae, the

more proximal ones elongate, flattened and attenuate; mesal half distad of basal

tergomesal lobe with setae rather evenly distributed, short, longer on distal third;

apical sternomesal scales mostly oblanceolate with acuminate tip, some lanceolate.

Claspette: Stem bowed inward near middle in dorsal aspect; uniformly curved

dorsad; slightly constricted near middle, filament striate, broadly sickle-shaped,

expanded somewhat near middle, slightly expanded and rounded posteriorly,

slightly recurved at apex. Clasper: Slightly narrowed at base, spiniform about

0.55 of clasper. Phallosome: Aedeagus large, broadly obovate;: tip produced dis-

tally into sclerotized beaklike keel below a second beaklike process or with keel

expanded distally into single, enlarged beak; numerous spicules present on venter

of beak; venter broadly open except on basal fourth, with sclerotized ridges ventro-

laterally which are expanded basally and terminate in sclerotized lobes mesally.

Proctiger: Cercal setae about 6; apical knob of paraproct with about 15-20 ser-

rations and a small hooklike process mesally near apex.

PUPA (fig. 19). Abdomen: about 3.30 mm. Trumpet: 0.45 mm. Paddle: 0.65

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

subequal, weak, usually single or double, weaker than 7-C which is single. Trumpet:

Medium to dark brown, lighter distally; reticulate sculpturing moderate. Abdomen:

Weakly to moderately pigmented, genital lobe and anterior segments darker. Float

hair (1-1) with about 12-14 primary branches and 2-6 secondary branches. Hair

1-II-VII weakly to strongly developed, single, multiple or strongly dendritic. Hair

2-VII short, considerably cephalad of 1-VII. Hairs 3-II,IJ1,5-IV,V variable in develop-

ment, subequal, with 3-III weaker, or with 3-II,III stronger or weaker than 5-IV,V.

Hair 6-VII weak, subequal to or weaker than 6-III-VI which are subequal. Hair

9-VII usually considerably cephalad of caudolateral margin of tergite, single or

double; 9-VII usually 4-7b. Terminal Segments: Male genital lobe about 1.2 of

tergite VIII. Paddle: Weakly to moderately pigmented; relatively narrow to broad,

apex broadly rounded or acuminate. Hair 1-P single.

LARVA (fig. 20). Head: 0.80 mm. Siphon: 0.78 mm. Anal Saddle: 0.33 mm.

Head: Hair 6-C single. Thorax: Integument with spicules. Tubercles of hairs 5-7-P

connected. Abdomen: Hair 12-I absent. Segment VIII: Comb scales 6-8(4-11),

with single pointed, minutely fringed spine, in single row, attached basally to

sclerotized plate; 1 or more occasionally detached from sclerotized plate. Siphon:

Index about 2.8(2.5-3.1). Pecten teeth about 10-14(6-15), extending to about

basal 0.50 of siphon. Hair 1-S double or triple, inserted at basal 0.55 of siphon.

Anal Segment: Spines on caudal margin of saddle well developed, much elongate

and conspicuous dorsally. Hair 4a-X 3-5b, short, less than length of saddle, or long,

nearly the length of 4b-X. Boss strongly sclerotized.

SYSTEMATICS. Haemagogus janthinomys can be separated from the remaining

species of the Albomaculatus Section by the following characters: in the adults

from all except capricornii by (1) silver scales extending to apex anteriorly on

hindfemur, (2) patches of dark scales on forecoxa and midcoxa, and (3) dark

40 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

bluish green scales on distal margins of abdominal tergites V-VIII which are more

numerous laterally and contrast with predominantly purple abdominal scales; in

the male genitalia by (1) conspicuous spicules on venter of aedeagus proximad

to beaklike apical process and (2) small hooklike process mesally near apex of

paraproct; and in the larva from all species except capricornii by the combination

of (1) spiculose integument, (2) absence of hair 12-I and (3) comb scales attached

basally to sclerotized plate.

Haemagogus janthinomys is apparently indistinguishable from capricornii except

in the male genitalia where it is characterized by the spicules and apical process of

the aedeagus and a hooklike process on the paraproct.

The apical process of the aedeagus of janthinomys varies in development, the

extremes being an elongate, dorsally curving beaklike process and a short, beaklike

process distad to a heavily sclerotized beaklike keel (see fig. 19). On the basis of

this variation, janthinomys (as spegazzinii) was divided into 2 subspecies (s. spega-

zzinii and s. falco) by Kumm, Osorno-Mesa and Boshell-Manrique (1946:14). Kumm

and Cerqueira (1951a:172) recognized the 2 subspecies in Brazil, including a num-

ber of intermediate forms and Martinez, Carcavallo and Prosen (1961) interpreted

the species as a complex of 3 subspecies of capricornii, c. falco, c. janthinomys

and c. petrocchiae, based on a long, intermediate and short aedeagus tip. Having

examined a large number of male genitalia from throughout the range, for the most

part, I can recognize no geographical pattern in this particular character, although

specimens from Central America tend to have a longer apical process and those

from Argentina a shorter one. Specimens from the remainder of South America

including the islands of Trinidad and Tobago show a completely random pattern

of short, long and intermediate development. In fact, a single collection from near

Villavicencio, Colombia contains both extremes. All possible degrees of develop-

ment are evident in the numerous specimens examined from Trinidad, this being

true also of specimens from Peru, Brazil and Venezuela. I can see no justification

for subdividing janthinomys on the basis of this 1 variable character.

As noted in the preceding paragraph and in the long and varied list of synonyms

and taxonomic references, there has been considerable confusion and differences

of opinion regarding the taxonomy and nomenclature of this most important spe-

cies. Janthinomys was first placed in synonymy with capricornii by Antunes (1939:

106) and subsequently with spegazzinii by Cerqueira (1943:10) when he resurrected

the latter from synonymy with equinus. Most authors followed Cerqueira’s synon-

ymy and used the name spegazzinii for this species until Martinez, Carcavallo and

Prosen (1961:63-86) determined that spegazzinii was a different species and the

same as uriartei. These authors considered janthinomys as a subspecies of capri-

corniit, which in their opinion included 3 other subspecies, the nominate form,

falco and petrocchiae. 1 find that capricornii is specifically distinct from janthino-

mys and as noted in the preceding paragraph see no justification for recognizing

falco or petrocchiae as subspecies distinct from typical janthinomys. Having exam-

ined the lectotype of obscurescens Martini, 1931, described as a variety of uriartei,

I find it to be conspecific with janthinomys. | am also convinced that spegazzinii

is a distinct species as shown by Martinez, Carcavallo and Prosen.

BIONOMICS. Haemagogus janthinomys is found almost exclusively in primary

tropical rain forest throughout most of its range and is decidedly arboreal in habit,

seldom descending to ground level to bite and then only when the forest canopy

has been disturbed. It breeds primarily in treeholes, though it is often taken in

bamboo oviposition traps. The treeholes are probably high and inaccessable for

Arnell: Genus Haemagogus 41

sampling as the number of larvae found in an area is seldom commensurate with

the number of adults present. In the rain forests of Central America, janthinomys

reaches its maximum density at elevations between 100 and 1000 meters. Numer-

ous studies have shown janthinomys to be a very efficient vector of yellow fever

virus. Because of its ability to harbor and transmit the virus and its arboreal habits,

janthinomys is the primary vector in the cycle of sylvan yellow fever which is

endemic in several areas in South America.

There is an extensive literature on the biology, ecology and disease relations

of janthinomys. Informative papers on the ecology of janthinomys in Central

America include Galindo, Trapido and Carpenter (1950), Galindo, Carpenter and

Trapido (1951la, 1955) and Trapido and Galindo (1956). Papers discussing the role

of janthinomys as a vector of yellow fever include Antunes and Whitman (1937),

Bates and Roca Garcia (1945, 1946b), Laemmert, Ferreria and Taylor (1946) and

Trapido and Galindo (1956). Ilheus virus has been isolated from janthinomys

captured in Panama (de Rodaniche and Galindo, 1961).

DISTRIBUTION (fig. 3). Haemagogus janthinomys extends from the Atlantic

versant of Honduras and Nicaragua through Central America and the entire Atlantic

and Caribbean drainages of South America to northern Argentina and southeastern

Brazil including the islands of Trinidad and Tobago. Material examined: 1407

specimens; 231 males, 463 females, 303 pupae, 410 larvae; 294 individual rear-

ings (179 larval, 20 pupal, 95 incomplete).

ARGENTINA. Jujuy: Ledesma (Libertador General San Martin), 2 IM, 2 1F [USNM] ; same data,

21 Mar 1926, N. Davis and R. Shannon, 2 F [USNM]; same data, 1943, H. Kumm, 1 M,5 F, 81

[UCLA, USNM]; same data, Mar 1944, G. Boshell-Manrique, 1 IM [USNM]. Salta: Acambuco,

4 F [USNM].

BOLIVIA. Beni: Rurrenabaque, Oct 1921, W. Mann, 1 F [USNM]. La Paz: Mapiri, 800 m,

8 Jan 1905, S. Carlos, 1 F [BM]. Santa Cruz: Cercado, Feb 1936, 1 F [USNM]. Lagunillas,

Cordillera, 20 Mar 1944, 5 F [UCLA]. Pozo Redondo, Cordillera, 7 Mar 1944, 1 F [UCLA].

Locality unspecified: Carr, 1 F [UCLA].

BRAZIL. Acre: Xapuri, 20 Aug-1 Sept 1947, H. Kumm, 1 lp, 4M [USNM]; same data, 26 Aug

1947, 1 lpF [BM]; same data, 10 Oct 1947, 1 lpM gen [UCLA]. Amapa: Locality unspecified,

20 Sept 1948, H. Kumm, 2 M [BM]. Amazonas: Borba, 23 Mar 1948, H. Kumm, 2 lpM gen

[UCLA]. Coari, 10 Aug 1948, H. Kumm, 3 M [UCLA, USNM]. Codajaz, 13-16 Aug 1948, H.

Kumm, 2 F [BM]. Manaus, 13 Oct 1947, 1 lpF [BM]. Manicore, 22 Mar 1948, H. Kumm, | lpM

gen [UCLA]. Tefe, Meta Patrimonio, 11 Jan 1949, 2 F [UCLA]. Bahia: Ilheus, 1 M [UCLA];

same data, 12 Jan 1944, 5 lp [USNM]; same data, 20 June 1944, 1 lp, 1 M gen [USNM]; same

data, 8 Mar 1945, 1 lp, 1 M gen [USNM]. Ilheus, Faz. Pirataguisse, 19 Jan 1944, 1 lpF [BH];

same data, 27 Feb 1944, H. Kumm, 1 lp [USNM]. Ilheus, Ribeirao da Fortuna, 8 Mar 1944, 1 lpM

[BH] ; same data, 23-26 Feb 1944, 8 lp, 1 M gen, 3 F [USNM]. Rio Cururipe, H. Kumm, | M, 2 F

[BM]; same data, 22 Aug 1930, N. Davis and R. Shannon, 1 F [USNM]; same data, 22 Apr 1936,

E. Silva, 1 M gen, 1 F, 1 p [USNM]. Salvador, P. Antunes, 1 F [USNM]; Salvador, 4 Apr 1944,

9 lp, 9M, 1 p [USNM]. Espirito Santo: Vale do Canaa, 3 Nov 1947, H. Kumm, 1 lp [USNM];

same data, 29 June 1948, 1 lpM [BM]. Goias: Agua Fria, Jan 1938, 1 F [USNM]. Anapolis, 1935-

1936, 13 F [USNM]. Cachoeira, May 1935, 1 F [USNM]. Caldas Novas, 25 Aug 1948, H. Kumm,

1 IpF, 1 lp [USNM]; same data, 15 Sept 1948, 1 lpM [BM]. Morrinhos, May 1935, 1 F [USNM];

same data, 16 Sept 1948, H. Kumm, 1| Ip [USNM]. Maranhao: China, 25 Aug 1948, H. Kumm,

1 F [USNM]. Colinas, 26 Aug-17 Sept 1948, H. Kumm, 1 IpF, 1 lp [USNM, BM]. Mato Grosso:

Arado, Matta das Palmeiras, 12 May 1944 (SFA Ent.), 3 IpM (6423,6438,6439), 4 IpF (6436,6444,

6445,6449), 2 M, 3 F [USNM]. Florida, 16 Oct 1947, H. Kumm, 1 lpF [BM]; same data, 3 Nov

1947, 1 lpF, 1 F [USNM]. Frei Ambrosis, 16-23 Sept 1947, H. Kumm, 1 IpM, 1 lp, 2M [USNM,

BM]. Sitio Encantadinho, 14 Aug 1947, 1 lp [USNM]; same data, 24 Oct 1947, 2 lpM gen, 1 M

gen [UCLA]. Minas Gerais: Cabui, 18 Nov 1946, H. Kumm, 1 lp [USNM]. Campina Verde, Jan

1949, 5 M, 4 F [UCLA]. Lavras, 2 M [USNM]. Leopoldina, 16 Sept 1948, 1 lpF, H. Kumm,

42 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

1 lpF [USNM]. Para: Abaetetuba, 25 Aug 1947, 1 lp, 1 M gen [USNM]. Belem, 24 June 1970,

T. Aitken (BRA 46A), 5 F; same data, 1 July 1970 (BRA 46B), 8 F; same data, 8 July 1970

(BRA 46C), 6 F [UCLA]. Belem, Bosque Rodriques Alves, 1 M [USNM]. Castenhal, 14 Feb

1947, 1 lp [USNM]. Curralinho, Apr 1936, 5 F [USNM]. Curralinho, H. Kumm, 1 F [UCLA].

Juruti, 17 Dec 1947, H. Kumm, 1 IpF, 2 F; same data, 19 Mar 1948, 1 lpF [USNM]; Nova

Timboteua, 18 Dec 1947, H. Kumm, 1 lp [USNM]; same data, 23-24 Jan 1948, 2 M [BM];

same data, 4 May 1918, 1 lpM [BM]. Locality unspecified (?Belem), Apr 1930, N. Davis, 1 M,

2 M gen, 5 F [USNM, BM]. Parana: I. de Maio, 24 Dec 1946, H. Kumm, 1 Ip [USNM]; same data,

27 Dec 1946, 1 lpM [BM]; same data, 27 May 1947, 1 lpM [USNM]. Rio de Janeiro: Duque de

Caxias, 28 Feb 1946, H. Kumm, 1 Ip [USNM]. Rondonia: Guajara-Mirim, 15 July-9 Sept 1947,

H. Kumm, 1 lpF, 2 lp, 2 M, 7 F [USNM, BM]; same data, 24 Mar 1948, 1 pM gen [UCLA].

Roriama: Caracarai, 13 Aug 1948, H. Kumm, 1 F [BM]. Rio Branco, 16-18 Sept 1948, H. Kumm,

1 lpF, 1 lp, 1 F [USNM]. Sao Paulo: Fernandopolis, 9 Apr 1947, H. Kumm, 1 IpM [BM]; same

data, 11 Apr 1947, 2 lp, 1 F [USNM]. Locality unspecified: J. Lane, 2 M gen; Feb 1944, 2 M, 3 F;

Apr 1944, 1M, 1 F; 21 Oct 1947, H. Kumm, 3 lpM gen, 2 M gen; same data, 19-21 Sept, 1 IpM,21

[USNM].

COLOMBIA. Boyaca: Nunchia, 11 1 [USNM]. Caldas: Rio La Miel, 1 M [USNM]. Caqueta:

Florencia, 1 M, 15 M gen [USNM]. Cundinamarca: Caparrapi, Volcanes Forest, 1000-1500 m,

1 M, 1 M gen, 2 F, 3 lp, 21 1 [USNM] . Guayabetal, 10 May 1946, E. Chapin, 1 F [USNM]. Malta,

280 m, 18 Feb 1943, H. Kumm, 1 1 [USNM]. Magdalena: El Retiro, 1 M [USNM]. Meta: El

Danubio, nr. Villavicencio, 490 m, 19 June 1965, E. Osorno-Mesa et al. (COB 40), 8 lpM (40-

10,20,21,40,42,50-52), 10 IpF (40-11-13,23,24,30,41,43,54,55), 7 L, E; same data (COB 41),

5 IpM (41-10,13-16), 2 lpF (41-11,12) [UCLA]. La Florista, nr. Restrepo, 380 m, 23 June 1965,

E. Osorno-Mesa et al. (COB 49), 4 IpF (49-20-23), 1 P [UCLA]. Restrepo, 8 M [USNM]; same

data, P. Antunes, 1 M, 3 F [UCLA, USNM]; same data, 20 July-23 Aug 1935, W. Komp, 6 M, 2M

gen, 2 F, 2 p, 291 [UCLA, USNM]. Retiro, 19 Aug 1935, 1 M;same data, July 1935, W. Komp, 21

[USNM] . Salinas de Upin, nr. Villavicencio, 440 m, 18 June 1965, E. Osorno-Mesa et al. (COB 38),

1 p, 1 P, 31 [UCLA]. Vanguardia, nr. Villavicencio, 350 m, 23 June 1965, E. Osorno-Mesa et al.

(COB 47), 3 IpM (47-10,12,16), 5 IpF (47-11,13-15,18), 3 L [UCLA]. Villavicencio, 28 June

1941, W. Komp, 1 M gen [USNM]; same data, Sept 1942, 1 M, 1 1 [USNM]; same data, Jan 1943,

1 F [USNM]; same data, 8 June 1944 [USNM] ; same data, 28 Sept 1943, M. Bates, 1 M, 1 M gen,

[USNM]; same data, 1944, 1 lp [USNM]; same data, 15 Oct 1945, 1 M [USNM]; same data,

5 Nov 1947, L. Rozeboom (CV 376A), 2 F [UCLA]. Locality unspecified, 1 M gen [UCLA].

Norte de Santander: Tibu, H. Kumm, 3 M, 3 1, [UCLA, USNM]. Vaupes: Teresita, 1 IM [USNM].

Locality unspecified: Kerr, 1 F [UCLA]; 19,23 June 1941 (MB 278), 2 M, 1 F [UCLA].

ECUADOR. Napo-Pastaza: La Coca (Coca and Napo Rivers), 250 m, 23 Apr-12 May 1965,

L. Pena (ECU 8), 3 F [UCLA]. ?Pambay, R. Levi-Castillo, 1 lpM gen, 4 M, 6 M gen, 1 F [USNM].

Locality unspecified: R. Levi-Castillo, 1 F [USNM].

FRENCH GUIANA. Guyane: Cabassou, 31 Jan 1965, T. Aitken (FG 13), 18 F; same data,

1 Feb 1965 (FG 16), 1 F; same data (FG 17), 1 F; same data, 5 Apr 1968, J. Clastrier (FGC 3348),

1 lpM (3348-10); same data (FGC 3359), 1 1; same data (FGC 3361), 1 1 [UCLA]. Galion, 30 Jan

1965, T. Aitken et al. (FG 4), 1 F [UCLA]. Chemin Vidal, nr. Remire, 13 Mar 1967, J. Carmen

(FG 124A), 1 F [UCLA]. La Chaumiere, Cayenne, 11 June 1967, J. Clastrier (FGC 3271), 1 F;

same data, 4 Jan 1968 (FGC 3333), 1 lpM (3333-10); same data (FGC 3334), 1 IpF (8334-11),

1 pF (8334-10) [UCLA]. Matoury, J. Clastrier (FGC 3833), 1 IpM (3833-44), 1 IpF (3833-45)

[UCLA]. Raban (5 km SE Cayenne), 3 Feb 1965, T. Aitken et al. (FG 45), 3 F [UCLA]. Rorota

(12 km E Cayenne), 6 Feb 1965, T. Aitken et al. (FG 66), 1 F; same data (FG 67), 1 F [UCLA].

Locality unspecified: 4 F [GML] ; J. Clastrier (FGC 3492), 5 F [UCLA].

GUYANA. Demerara: Hyde Park, 1 Aug 1941, L. Rozeboom (BGR 5), 3 F [UCLA] . Essequibo:

23 June 1936, 1 F [GML].

HONDURAS. Atlantida: Tela, 17-25 May, 4 Sept 1954 (GML), 2 lpM (01580,01583), 4 IpF

01581,01582,01585,01586) [UCLA].

NICARAGUA. Chontales: Villa Somoza, July 1953 (GML), 1 lpM (00989) [UCLA]. Zelaya:

El Recreo, 5 July 1954, K. Neiland (NI 3), 1 L [UCLA].

PANAMA AND CANAL ZONE. Canal Zone: Barro Colorado Island, 7 May 1943, W. Komp,

Arnell: Genus Haemagogus 43

1 M gen; same data, 1 May 1945, 1 IpF (5-106) [USNM]. Ft. Sherman, 23 Sept 1949, 4 F

[UCLA]. Chiriqui: Chorcha, 5 Jan 1950, P. Galindo (GML), 2 IpM (00402,00410), 1 IpF (00411)

[UCLA]. Darien: Tacarcuna, 9 Sept 1958 (GG 1), 1 IpF (1-118); same data (GG 123), 12 IpM

(123-102,107-109,111,115-117,119,121-123), 11 IpF (123-101,103-106,110,112-114,118,120)

[UCLA]. Tacarcuna River Valley, 600 m, 20 June 1963 (PA 406), 1 F; same data, 8 July 1963

(PA 444), 1 F; same data, 12 July 1963 (PA 457), 2 F [UCLA]. Panama: Cerro Azul, 1954

(GML), 4 1 [UCLA]. Cerro La Victoria, 1949, H. Trapido, 15 F [UCLA]. Pacora, June-Aug

1950, 33 F [UCLA].

PERU. Huanuco: Cochicoto, 12 Nov 1965, J. Hitchcock, 1 M gen [USNM]. Loreto: Iquitos,

Mar-Apr 1931, R. Shannon, 12 F, 1 1 [USNM]. Ucayali River, 24 Oct 1903, 1 F (obscurescens

lectotype) [BM].

SURINAM. Saramacca: Matta, 15 Jan 1960, T. Aitken, 2 F [UCLA]. Locality unspecified:

J. Bonne-Wepster, 1 F [USNM].

TRINIDAD AND TOBAGO. TOBAGO. St. Paul: Roxborough, Main Ridge, 31 July 1957, T.

Aitken, 2 F [UCLA]. TRINIDAD. Caroni: Brasso Venado, 120 m, 21 June 1964, R. Manuel and

R. Martinez (TR 513), 2 F [UCLA]. Mayaro, 30 m, 23 July 1966, T. Aitken (TR 1561), 2 pM

(1561-10,11), 1 pF (1561-12) [UCLA]. Nariva: Big Bush Bush, Nariva Swamp, 29 Aug 1961,

T. Aitken (1-29/VIII/61), 2 lpF (2,4), 1 M, 2 F; same data, 7 Sept 1961, 1 F [UCLA]. Bush Bush

Forest, Nariva Swamp, 3 Jan 1964, TRVL (TR 13), 1 lpM (13-106), 1 IpF (13-108), 3 IP (13-

101,104,105); same data (TR 20), 1 IP (20-102); same data, 17 Feb 1964 (TR 72), 1 lpM (72-

136), 1 pM (72-131), 3 pF (72-132-134); same data, 26 Feb 1964 (TR 101), 3 IpF (101-149-151);

same data, 4 Mar 1964 (TR 153), 2 IpF (153-197,198); same data, 11 Mar 1964 (TR 172), 2 IpF

(172-120,180); same data, 15 Apr 1964(TR 197), 2 lpM (197-101,114), 3 IpF (297-136,199,200);

same data (TR 298), 2 lpM (298-111,141), 1 lpF (298-140); same data (TR 299), 7 IpM (299-104,

105,110,112,113,123,132); same data (TR 300), 3 IpM (300-102,124,159), 2 lpF (300-139,151),

6 L; same data, 27 May 1964 (TR 427), 1 lpM (427-157); same data (TR 429), 2 lpM (429-130,

131), 1 lpF (429-148); same data (TR 431), 4 lpM (431-126-128,129) 1 lpF (431-147); same data

(TR 432), 1 IpF (432-149), 1 IP (432-159); same data, 10 June 1964 (TR 485), 1 pM (485-

181); same data, 5 Dec 1965, T. Aitken (TR 1427), 1 F; same data, 22 Dec 1960 (5-22/XII/60),

2 p; same data, 2 Mar 1961 (4-2/III/61), 5 Ip (1-4,6), 1 p; same data, 27 Apr 1961 (S-27/IV/61),

12 Ip (2-13), 1 p; same data, 31 May 1961 (3-31/V/61), 5 lp (1-5); same data (4-31/V/61), 3 Ip

(1-3); same data, 4 Sept 1961 (3-4/IX/61), 1 pM(1), 1 M, 1 F; same data, 7 Sept 1961 (2-7/IX/61),

1 F; same data, 30 Jan 1963 (5-30/I/63), 1 IpM (1); same data, 19 Feb 1963 (6-19/II/63), 1 lpF

(1); same data, 11 Mar 1963 (4-11/III/63), 2 lpM (2,3), 1 IP (1); same data (5-11/TII/63), 2 IlpM

(1,5), 3 IpF (2-4); same data, 12 Mar 1963 (13-12/III/63), 1 lpM (1), 1 lp (2) [UCLA]. Charuma

Forest, 150 m, 27 Aug 1964, A. Guerra (TR 644), 2 F; same data, 8 Oct 1964 (TR 761), 4 F

[UCLA]. Rio Claro, 45 m, 5 Nov 1964, A. Guerra (TR 815), 2 lpF (815-108,110), 1 L [UCLA].

St. Andrew: Caratal Road, 75 m, 3 Dec 1964, F. Powdhar (TR 866), 1 F [UCLA]. Coryal, 75 m,

18 June 1964, A. Guerra (TR 498), 7 F [UCLA]. El Quemado Road, 75 m, 3 July 1964, F.

Powdhar (TR 547), 5 F [UCLA]. Guaico-Tamana Road, 75 m, 10 Apr 1965, F. Powdhar (TR

1216), 3 pF (1216-100-102) [UCLA]. Mt. Harris, 150 m, 23-31 July 1924, C. Withycombe, 13 F

[BM]; same data, 16 July 1964, F. Powdhar (TR 573), 11 F [UCLA]. Mt. Tamana, 100 m, 19

June 1964, A. Guerra (TR 512), 1 F [UCLA]. Nestor Village, 30 m, 12 June 1964, A. Guerra

(TR 476), 2 L; same data (TR 482), 6 L; same data (TR 483), 2 IpM (483-116,117), 1 pM (483-

103); same data (TR 484), 1 M, 2 F; same data, 24 Apr 1965, F. Powdhar (TR 1232), 1 F

[UCLA] . Sangre Grande, 75 m, 17 Dec 1964, F. Powdhar (TR 894), 1 F [UCLA]. Sangre Grande,

Biche Road, 15 Oct 1957, T. Aitken, 1 M [UCLA]. Turure Forest, 60 m, Oct 1966, A. Guerra

(TR 1618), 1 F [UCLA]. Vega de Oropouche Road, St. John’s Estate, 12 Dec 1960, T. Aitken

(12/XII/60), 5 lp (2-6), 1 p [UCLA]. St. David: Cumana, 90 m, 19 Nov 1964, A. Guerra (TR

840), 1 F [UCLA]. St. George: Agua Santa, 75 m, 29 July 1965, A. Guerra (TR 1295), 2 F

[UCLA]. Aripo Valley, L’Orange Road, 150 m, 17 Sept 1964, A. Guerra (TR 711), 1 F [UCLA].

Guanapo Valley, 20 m, 17 Apr 1964, A. Guerra (TR 332), 1 IpF (332-129), 1 pF (332-133)

[UCLA]. Heights of Guanapo (8 km N Arima), 250 m, 26 Mar 1964, A. Guerra (TR 259), 1 F

[UCLA]. Las Lapas Trace, 600 m, 3 Apr 1964, A. Guerra (TR 281), 1 F [UCLA]. Lopinot,

300 m, 30 Apr 1964, A. Guerra (TR 374), 2 F [UCLA]. Monos Island, Grand Fond Bay Valley,

4a. Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Om, 1 Aug 1964, R. Manuel (TR 584), 1 pF (584-103) [UCLA] . Mt. Beck, 300 m, 29 Apr 1965,

A. Guerra (TR 1137), 1 F; same data (TR 1143), 1 F [UCLA]. St. Ann’s, 26 Aug 1945, 1 F;

same data, 1906, F. Urich, 1 lp; same data, 26 Aug 1945, W. Komp, 3 Ip, 1 P, 1 1 [USNM].

St. Augustine, Oct 1923, C. Withycombe, 1 F [BM]. St. Pats, 12 Nov 1953, 1 F; same data,

15 Dec 1954 (GML), 1 lp (02031) [UCLA]. Talparo, 75 m, 26 June 1964, A. Guerra (TR 531),

6 F [UCLA]. Tunapuna, Caura Road, 240 m, 14 May 1964, A. Guerra (TR 397), 1 F [UCLA].

Verdant Vale, 300 m, 10 Sept 1964, A. Guerra (TR 679), 2 pF (679-114,115), 1 L; same data

(TR 686), 7 F; same data, 150 m, 12 Nov 1964 (TR 821), 5 L; same data (TR 823), 1 L [UCLA].

St. Patrick: Cedros, 27 Mar 1921, G. Pawan, 1 F [BM]. Victoria: Mayo, 25 June 1914, J. Dickson,

1 M [BM]. County unknown: (?)Franca, Rio Grande, 23 Sept 1903, 1 F [BM] . Locality unspeci-

fied: S. Vister, 1 F [BM]; F. Urich, 2M [USNM]; June 1905, A. Busk, 3 F [USNM].

VENEZUELA. Aragua: Maracay, Apr 1935, P. Anduze, 1 F [USNM]. Maracay (20 km N on

road to Choroni), 800 m, 6 Aug 1969, J. Valencia (VZ 312), 4 IpF (312-10-13) [UCLA]. Rancho

Grande, 1095 m, 9 July 1969, T. Zavortink (VZ 168), 1 lpF (168-10), 1 L [UCLA]. Rancho

Grande (8 km S), 800 m, 15 July 1969, T. Zavortink (VZ 203), 1 pM (203-101); same data,

11 Aug 1969, J. Valencia (VZ 325), 4 F; same data, 18 Aug 1969, Pulido and Clarijo (VZ 371),

1 1; same data (VZ 375), 1 pM (375-100); same data, 25 Aug 1969, J. Valencia and Clarijo

(VZ 409), 1 lpM (409-10) [UCLA]. Rancho Grande (park entrance), 1 Dec 1967, Hansell (VZ

82), 1 1 [UCLA]. Turiamo, 16 Sept 1944, 3 M, 5 F [UCLA]; same data, Sept 1944, W. Komp,

1 p, 81 [USNM]. Carabobo: San Jean, 24 Dec 1937, P. Anduze, 1 M gen [USNM]. Delta Amacuro:

Manoa Woods, Orinoco River, 10 Jan, 1 F [USNM]. Monagas: Guamito, 30 July 1927, M. Nunez-

Tovar, 1 F [USNM].

Additional Records From the Literature

ARGENTINA. Catamarca: Alrededores La Vina and Cuesta del Totoral (Manso Soto, Martinez

and Prosen, 1953:37).

BRAZIL. Ceara: Pacoti (Kumm and Cerqueira, 1951:172). Pernambuco: Recife (Kumm and

Cerqueira, 1951:172).

COSTA RICA. Alajuela: San Gerardo (Galindo and Trapido, 1955:546). Limon: Wauchope

(Galindo and Trapido, 1955:545). Puntarenas: Cordillera Brunquena, 40 km NW Golfito (Galindo,

Carpenter and Trapido, 1951b:105).

&. Haemagogus (H.) capricornii Lutz

Figs. 3,21,22

1904. Haemagogus capricornii Lutz, in Bourroul, 1904:13. TYPE: Neotype female reared from

egg, Horto Florestal, Serra da Cantareira, Sao Paulo, Brazil, Apr 1944, L. Gomes [LU;

designation by Cerqueira and Lane, 1945:186]. Synonymized with equinus by Dyar

(1923:183); revalidated by Antunes (1939:106).

Haemagogus (Haemagogus) capricornii of Lane (1939:118, in part); Levi Castillo (1951b:11,

18-19).

Haemagogus (Stegoconops) capricornii of Dyar (1921a:103); Lane (1953:790-792); Stone, Knight,

and Starcke (1959:216, in part); Cova Garcia, Sutil and Rausseo (1966a:61-62, fig. 118; 1966b:

46, fig. 201).

Haemagogus (Stegoconops) capricornii capricornii of Martinez, Carcavallo and Prosen (1961:

77); Stone (1963:132); Forattini (1965:19); Martinez and Carcavallo (1965:42).

Haemagogus capricornii of Blanchard (1905:633); Theobald (1907:551; 1910:493); Howard,

Dyar and Knab (1917:877, in part); Shannon and Del Ponte (1928:68); Shannon, Whitman

and Franca (1938:111, in part); Shannon (1939:137, in part); Cerqueira (1943:8-9); Cerqueira

and Lane (1945:279-288); Cerqueira and Boshell-Manrique (1946:193); Waddell and Kumm

(1948:247); Waddell (1949:568); Kumm and Laemmert (1950:750); Laemmert and Kumm

(1950:724); Kumm and Cerqueira (1951a:173-174; 1951b:196); Levi-Castillo (1951a:14);

Arnell: Genus Haemagogus 45

Mackie, Hunter and Worth (1954:19); Horsfall (1955:535); Foote and Cook (1959:141).

Stegoconops capricornii of Lutz (1905 :83-84).

Aedes capricornii in part of Dyar and Knab (1906b:163-164).

_ Haemagogus (Stegoconops) equinus in part of Dyar (1923:183; 1928:134); Edwards (1932:179).

Haemagogus equinus in part of Dyar (1925b:138).

FEMALE. Wing: 3.30 mm. Proboscis: 2.60 mm. Forefemur: 2.30 mm. Abdo-

men: 3.00 mm. Apparently indistinguishable from janthinomys.

MALE (fig. 21). Wing: 3.30 mm. Proboscis: 2.90 mm. Forefemur: 2.20 mm.

Abdomen: 3.40 mm. Apparently indistinguishable from janthinomys.

MALE GENITALIA (fig. 21). Segment VIII: Tergite about 0.55-0.60 of sternite;

distal margin straight or slightly broadly emarginate, with enlarged setae laterally

and cluster of about 20 lanceolate scales mesally. Segment IX: Tergite deeply

emarginate and weakly sclerotized mesally; lobes with 1 to 3 setae. Sidepiece:

Conical; length 3.5 times median width; basal tergomesal lobe slightly enlarged,

with numerous setae, the more proximal ones enlarged, flattened and attenuate;

mesal half distad of basal tergomesal area with setae rather evenly distributed,

short, longer on distal fourth; apical sternomesal scales oblanceolate with acuminate

tips. Claspette: Stem bowed inward near middle in dorsal aspect, slightly con-

stricted near middle, sharply curved dorsad beyond middle; filament striate, broadly

sickle-shaped, slightly expanded and rounded posteriorly, apex slightly recurved.

Clasper: Broadest at base, spiniform short, about 0.40 of clasper. Phallosome:

Aedeagus large, oval, apex slightly emarginate in dorsal view laterad of median dorsal

sclerotized keel, which is produced forward to form a short, blunt tip; venter broadly

open except on ventral fourth; sclerotized ridges present lateroventrally which are

expanded basally and terminate in sclerotized lobes mesally. Proctiger: Cercal setae

6-8; apical knob of paraproct with about 20 serrations. 7

PUPA (fig. 21). Abdomen: about 3.30 mm. Trumpet: 0.45 mm. Paddle: 0.70

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

subequal, weak, single, weaker than 7-C which is also single. Trumpet: Medium to

dark brown, lighter distally; reticulate sculpturing moderate. Abdomen: Weakly

to moderately pigmented, genital lobe and anterior segments often darker. Float

hair (1-I) with about 12-16 primary branches and about 2-6 secondary branches.

Hair 1-II-VII weakly developed, 1-4b. Hair 2-VII short, considerably cephalad of

I-VII. Hairs 3-I1,I1,5-IV,V subequal or 3-III slightly weaker. Hair 6-VII weak,

fine, subequal to 6-III-VI. Hair 9-VII usually Slightly cephalad of caudolateral

margin of tergite, single or double; 9-VIII usually 4-8b. Terminal Segments: Male

genital lobe about 1.3 of tergite VIII. Paddle: Weakly to moderately pigmented;

relatively broad, apex broadly rounded. Hair 1-P single.

LARVA (fig. 22). Head: 0.95 mm. Siphon: 0.80 mm. Anal Saddle: 0.40 mm.

Head: Hair 6-C single. Thorax: Integument with spicules. Tubercles of hairs 5-7-P

connected. Abdomen: Hair 12-I absent. Segment VIII: Comb scales 8-10(5-13),

with single sharply pointed, minutely fringed spine, in single row, attached basally

to sclerotized plate. Siphon: Index about 2.4-2.7. Pecten teeth about 10-14(9-18),

extending to about basal 0.45 of siphon. Hair 1-S double, inserted at basal 0.55

of siphon. Anal Segment: Spines on caudal margin well developed, much elongate

and conspicuous dorsally. Hair 4a-X double or triple, long, nearly the length of

4b-X. Boss strongly sclerotized.

SYSTEMATICS. Although capricornii is apparently indistinguishable from janthi-

nomys as adults and larva I consider it as a distinct species. It can be distinguished

46 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

from janthinomys and the remainder of the Albomaculatus Section in the male

genitalia by the combination of (1) absence of hooklike process on apex of

proctiger, (2) aedeagus slightly emarginate just laterad of median dorsal sclerotized

keel and (3) relatively short tergite VIII which is about 0.55-0.60 of sternite VIII.

Described by Lutz in 1904, capricornii was synonymized with equinus by Dyar

(1923) because the males of capricornii were unknown to him. Antunes (1939)

resurrected capricornii and Cerqueira and Lane (1945) later reared specimens from

the type locality and confirmed its validity. For a review of the confusion of

capricornii, spegazzinii and janthinomys see the section on systematics under the

latter.

BIONOMICS. Little is known of the bionomics of capricornii as the majority

of the papers containing information on the biology and habits of this species

actually refer to janthinomys. Capricornii is found only in the drier, more temperate

region of southeastern Brazil and apparently breeds almost exclusively in treeholes.

Capricornii is an efficient vector of yellow fever virus as demonstrated in labora-

tory transmission studies (Waddell and Kumm, 1948; Waddell, 1949). Yellow fever

has been transmitted by bite from wild caught females of presumably capricornii

from Affonso Arinhos, Brazil (Shannon, Whitman and Franca, 1938), although

there is some question as to whether these authors were dealing with capricornii

or janthinomys in their transmission studies. I am unaware of any instances of

outbreaks of sylvan yellow fever in areas of southeastern Brazil where capricornii

would be the only possible vector species.

DISTRIBUTION (fig. 3). Haemagogus capricornii is found only in southeastern

Brazil, from southern Bahia to northern Rio Grande do Sul. The record of capri-

cornii in Stone, Knight and Starcke (1959:216) from Bolivia undoubtedly refers

to janthinomys. That of Cova Garcia, Sutil and Rausseo from Venezuela is also

unquestionably erroneous; their drawings of male genitalia which identify this

species are apparently copied directly from Levi-Castillo (1951b:fig. 3, p. 40).

Material examined: 87 specimens; 30 males, 10 females, 24 pupae, 23 larvae;

24 individual rearings (13 larval, 1 pupal, 10 incomplete). |

BRAZIL. Espirito Santo: Santa Teresa, 13 Oct 1947, H. Kumm, 1 lpM gen [UCLA]. Minas

Gerais: Cabiri, 12 Nov 1946,H. Kumm, 1| lpM [BM]. Lavras, 3 M [UCLA]. Parana: Jandaia do Sul,

23 Dec 1946, 1 M gen [USNM]. Rio de Janeiro: Duque de Caxias, Mar-Apr 1950, 7 M, 6 F

[UCLA]; same data, 4 Mar 1946, H. Kumm, 4 lp [USNM]. ?Neocacios, 23 July 1946, H. Kumm,

1 lpM [BM]. Paraiba do Sul, Mar 1938, 1 M [USNM]. Sao Sebastiao, 15 Oct 1947, H. Kumm,

3 lpM gen [UCLA]. Santa Clara, 30 Apr 1947, 5 lpM gen, 1 pM gen, 2 M gen [USNM]. Sao Paulo:

?Gapira, A. Lane, 2 F [UCLA]. Mirasol, L. e Gui, 1 F [UCLA]. Serra da Cantareira, 24 Apr 1944,

L. Gomes, 1 lpM, 1 lpF, 6 lp [UCLA, USNM, BH]. Locality unspecified: 3 M gen [UCLA].

Additional Records From the Literature

BRAZIL. Bahia: Malhada (Kumm and Cerqueira, 1951:174). Rio Grande do Sul: Santo Augusto

and Sao Luiz Gonzaga (Kumm and Cerqueira, 1951:174). Santa Catarina: Ibirama (Kumm and

Cerqueira, 1951:174).

9. Haemagogus (H.) mesodentatus Komp & Kumm

Figs. 3,23,24

1938. Haemagogus mesodentatus Komp and Kumm, 1938:253-259. TYPE: Holotype male,

Parque Bolivar, San Jose, San Jose, Costa Rica, 20 Dec 1937, H.W. Kumm [LU; see

Stone and Knight, 1955:288].

1956. Haemagogus mesodentatus gorgasi Galindo and Trapido, 1956:228-231. TYPE: Holo-

type female (01591) with associated larval and pupal skins, Tapachula, Chiapas, Mexico,

Arnell: Genus Haemagogus 47

reared from eggs from female taken biting man, 4 Aug 1953 [USNM]. NEW SYN-

ONYMY.

- 1956. Haemagogus mesodentatus alticola Galindo, Trapido and Boshell-Manrique in Galindo

and Trapido 1956:228-231. TYPE: Holotype female (01920) with associated larval and

pupal skins, summit of Sumidero Canyon of the Rio Grijalva, 24 km N of Tuxtla

Gutierrez, Chiapas, Mexico, elev. 4000 ft, reared from eggs from female taken biting

man, 29 June 1953 [USNM]. NEW SYNONYMY.

Haemagogus (Haemagogus) mesodentatus of Lane (1939:120).

Haemagogus (Stegoconops) mesodentatus of Lane (1953:796-798) Stone, Knight and Starcke

(1959:216); Bertram (1971:475).

Haemagogus mesodentatus of Kumm, Komp and Ruiz (1940:395,399); Kumm and Zuniga

(1942:404); Vargas and Martinez Palacios (1953:38); Trapido and Galindo (1956a:304-305;

195 126%

Haemagogus (Stegoconops) mesodentatus mesodentatus of Diaz Najera (1963:191; 1966:61);

Forattini (1965:26-29).

Haemagogus mesodentatus mesodentatus of Galindo and Trapido (1956:228,231); Galindo, de

Rodaniche and Trapido (1956:1022); Galindo and Trapido (1957:147); de Rodaniche and

Galindo (1957:235); Boshell-Manrique and Bevier (1958:27); Foote and Cook (1959:141);

Vargas and Diaz Najera (1959:362); Diaz Najera (1960:187).

Haemagogus (Stegoconops) mesodentatus gorgasi of Stone, Knight and Starcke (1959:216); Diaz

Najera (1963:191; 1966:61); Stone (1963:132); Forattini (1965:26-29).

Haemagogus mesodentatus gorgasi of Galindo, de Rodaniche and Trapido (1956:1022); de

Rodaniche and Galindo (1957:236); Foote and Cook (1959:141); Vargas and Diaz Najera

_ (1959:362); Diaz Najera (1960:187).

Haemagogus (Stegoconops) mesodentatus alticola of Stone, Knight and Starcke (1959:216);

Diaz Najera (1963:191); Stone (1963:132); Forattini (1965:26-29).

Haemagogus mesodentatus alticola of Boshell-Manrique and Bevier (1958:33); Vargas and Diaz

Najera (1959:362); Diaz Najera (1960:187).

FEMALE. Wing: 3.05 mm. Proboscis: 2.55 mm. Forefemur: 2.10 mm. Abdo-

men: 3.20 mm. Dark scales of proboscis, palpus, wing and legs dark blue to violet

with some purple reflections. Head: Eyes narrowly separated (1 ommatidial dia-

meter) or contiguous above antennae. Decumbent scales of occiput light green to

light bluish green. Proboscis relatively long, about 1.20-1.35 of forefemur; palpus

about 0.16 of proboscis; antenna about 0.66-0.71 of proboscis. Thorax: Scales

of mesonotum and scutellum variable, from hues of green and blue to copper

and gold, usually bluish green over supraalar bristles and on scutellar lobes, silver

in antealar area. Apn lobes moderately enlarged, scales blue to green, with or

without silver scales; ppn scales with same range of color as mesonotum, often

with silver scales ventrally. Legs: Relatively short, length of forefemur about 1.25-

1.40 of distance from top of thorax to apex of midcoxa; forecoxa occasionally

with small patch of violet to purple scale near middle; knee spots present or

absent on midfemur and hindfemur; midtarsus with white or gray scales on outer

surface of proximal 2 or 3 segments. Wing: R, 43 usually about 0.75-0.90 of R,.

Abdomen: Dark scales blue to violet with some purple reflections; dorsal silver

patches of scales basally on tergites III-VIII, often expanded distally along mid-

line and larger on distal segments; lateral silver patches extending to distal margins

of tergites II-VII forming broad lateral silver stripe, or with patches only on basal

half of more distal segments.

MALE (fig. 23). Wing: 2.90 mm. Proboscis: 2.90 mm. Forefemur: 2.05 mm.

Abdomen: 3.20 mm. Head: Proboscis about 1.40-1.55 of forefemur; palpus short,

about 0.15 of proboscis; antenna about 0.60 of proboscis. Legs: Larger claw of

foreleg and midleg with acute submedian tooth, smaller claw of foreleg and mid-

leg with acute subbasal tooth.

48 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

MALE GENITALIA (fig. 23). Segment VIII: Tergite about 0.70-0.75 of sternite;

distal margin broadly emarginate, with enlarged setae laterally and cluster of 20-30

lanceolate scales mesally. Segment IX: Tergite emarginate and weakly sclerotized

mesally; lobes with 1 or 2 setae. Sidepiece: Conical, length 3.5 times median width;

basal tergomesal lobe slightly enlarged, with numerous setae, the more proximal

ones much elongate, flattened and attenuate; mesal half distad of basal tergomesal

area with setae rather evenly distributed, short, longer on distal fourth; apical

sternomesal scales lanceolate to oblanceolate with acuminate tips. Claspette: Stem

bowed inward near middle in dorsal aspect, relatively thin, slightly narrowed and

curved dorsad beyond middle; filament striate, broadly sickle-shaped, broader near

middle. Clasper: Broadest at base; spiniform about 0.50 of clasper. Phallosome:

Aedeagus large, broadly obovate, tip produced distally into long thin beak, which

makes up about 0.30 of aedeagus length, the dorsum of tip with a series of proxi-

mally projecting coarse serrations forming a median carina; dorsum with broad,

elliptical opening into which base of carina projects; venter broadly open except

at basal third, with sclerotized ridges lateroventrally which are expanded basally

and nearly joined near midline by sclerotized lobes. Proctiger: Cercal setae 6-10;

apical knob of paraproct with about 20 serrations.

PUPA (fig. 23). Abdomen: about 3.45 mm. Trumpet: 0.45 mm. Paddle: 0.75

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

subequal, single to triple, weaker than 7-C which is single. Trumpet: Dark brown,

lighter distally; reticulate sculpturing moderate. Abdomen: Weakly to moderately

pigmented, genital lobe and anterior segments darker. Float hair (1-1) with 8-16

primary branches and 2-6 secondary branches. Hair 1-II-VII weakly to very strongly

developed, single to dendritic. Hair 2-VII considerably cephalad of 1-VII. Hairs

3-I1,I11,5-IV,V somewhat variable in development but more or less subequal. Hair

6-VII weakly to moderately developed, subequal to or stronger than 6-III-VI,

usually single. Hair 9-VII slightly cephalad of caudolateral margin of tergite, 1-4b;

hair 9-VIII 6-10b. Terminal Segments: Male genital lobe about 1.2 of tergite VIII.

Paddle: Weakly to moderately pigmented; apex broadly rounded. Hair 1-P single.

LARVA (fig. 24). Head: 0.90 mm. Siphon: 0.75 mm. Anal Saddle: 0.32 mm.

Head: Hairs 5,6-C single (single or double). Thorax: Integument with spicules.

Tubercles of hairs 5-7-P connected. Abdomen: Hair 12-I present. Segment VIII:

Comb scales 8-11(6-12), with single sharply pointed, minutely fringed spine, in

single row. Siphon: Index about 2.2-2.5(1.9-2.9). Pecten teeth about 12-16(8-17),

extending to about basal 0.50 of siphon. Hair 1-S double (2-4b), inserted at basal

0.55-0.60 of siphon. Anal Segment: Spines on caudal margin of saddle well devel-

oped. Hair 4a-X 3-5b, short or long. Boss strongly sclerotized.

SYSTEMATICS. Haemagogus mesodentatus can be distinguished readily: in the

adults by white or gray scales on outer surface of proximal 2 or 3 midtarsal

segments; in the male genitalia by aedeagus tip expanded into long, thin, coarsely

serrated carina that projects posteriorly into elliptical opening in dorsum of aede-

agus; and in the larva by the combination of (1) spiculose integument, (2) hair

12-I present and (3) comb scales not attached basally to sclerotized plate.

This species was divided into 3 subspecies by Trapido and Galindo (1956) on

the basis of the presence or absence of light knee spots and the amount of silver

scales on the apn and ppn in the female. Mesodentatus is unstable in several

characters of the adults and larvae. The color of the scales of the mesonotum

varies from copper through gold to green and blue, there is a great deal of variation

in the number of silver scales on some of the thoracic pleurites and femora and

Arnell: Genus Haemagogus 49

considerable differences in the development of spiculation and of some hairs in

the larva. The characters proposed by Galindo and Trapido for their subspecies

are among the more variable and do not hold in the specimens examined in this

study. I consider mesodentatus to be a somewhat inconstant but monotypic spe-

cies.

Mesodentatus may be closely related to janthinomys, both having a spiculose

larval integument and an extreme development of the apex of the aedeagus. They

differ in details of adult ornamentation by which they are easily distinguished.

The 2 species occupy much the same ecological situations, the canopy, in the

rain forests of Central America. The 2 species are sympatric in the forests on the

Atlantic side north of the Isthmus of Panama with mesodentatus gradually replacing

janthinomys in Nicaragua and Honduras and becoming abundant beyond the north-

ern limits of janthinomys. Both species are capable of transmitting yellow fever

virus.

BIONOMICS. Haemagogus mesodentatus breeds primarily in treeholes and bam-

boo internodes although it is found in limestone rock holes at higher elevations

in southern and south-central Mexico. As mentioned above, mesodentatus is sym-

patric with janthinomys in the rain forests from Panama, where it is rare, to

Honduras, where it outnumbers janthinomys. Mesodentatus reaches its maximum

density in the forests of the Department of Peten in Guatemala and in southern

Mexico. Apparently absent from the Pacific slope of Nicaragua and Honduras,

it is common in El Salvador, Guatemala and southern Mexico and extends north-

ward to Sonora in the tropical deciduous forest of the Pacific coast. It has been

taken at elevations of over 1200 meters in the highlands of southern Mexico and

Costa Rica. Although not as efficient a vector as janthinomys, mesodentatus is

capable of transmitting yellow fever by bite (Galindo, de Rodaniche and Trapido,

1956) and this virus has been isolated from wild caught females in Guatemala

(de Rodaniche and Galindo, 1957).

DISTRIBUTION (fig. 3). Haemagogus mesodentatus extends from southern So-

nora and southern San Luis Potosi, Mexico through both the Atlantic and Pacific

versants of Central America to the Isthmus of Panama. Material examined: 437 speci-

mens; 66 males, 136 females, 104 pupae, 131 larvae; 80 individual rearings (57

larval, 16 pupal, 7 incomplete).

COSTA RICA. Alajuela: Esparta (11.8 km E), 19 June 1963, C. Hogue (CR 97), 2 F [UCLA].

San Jose: Lourdes (nr. San Jose), 1150 m, 29 Oct 1971, D. Schroeder (CR 494), 3 F [UCLA].

San Jose, H. Kumm, | M, 1 F, 121 [USNM, BM]; same data, 14 Aug 1953 (GML), 1 lpF (01661),

1 Ip (01660) [UCLA]; same data, 8 June 1963, C. Hogue (CR 88), 1 lpM (88-108), 1 lpF (88-

wl ie 2 1, 2 L [UCLA]; same data (CR 89), 2 IpM (89-102,105), 2 lp (89-101,201), 2 L

EL SALVADOR. Liberdad: La Liberdad, 15 June 1953 (GML), 1 lpF (01838), 3 pM (01929,

01933,01446) [UCLA]. San Salvador: Los Planes, H. Kumm, 4 M, 2 M gen, 4 F, 8 1 [USNM].

Locality unspecified: 1 L [USNM].

GUATEMALA. Escuintla: El Salto, 5 Aug 1953 (GML), 1 lpF (01732), 5 1 [UCLA]. Santa

Lucia Cotzumalguapa, 3 July 1964, T. Zavortink and P. Cowsill (GUA 31), 1 F [UCLA]. Izabal:

Rio Chiquito, nr. Bananera, 28 Dec 1954 (GML), 1 lpF (01960) [UCLA]. Jutiapa: Jutiapa,

9 July 1943, H. Hargis, 4 F [UCLA]. Peten: Locality unspecified, 9 Dec 1955 (GML), 6 1

[UCLA]. Retalhuleu: San Felipe, 780 m, 2 July 1964, T. Zavortink and P. Cowsill (GUA 22),

1 IpM (22-10); same data (GUA 27), 1 lpF (27-10), 1 pF (27-100); same data (GUA 28), 1 lpM

(28-14), 10 IpF (28-10,13,15-22), 1 IF (28-11), 1 pM (28-100), 1 IP (28-12), 7 M, 5 F, 19 P,

6 L; same data, 8 Sept 1964, P. Cowsill and Almengor (GUA 130), 1 IpM (130-10), 2 pM (130-

101,102), 1 pF (130-100) [UCLA]. San Sebastian, 300 m, 2 July 1964, T. Zavortink and

P. Cowsill (GUA 29), 3 IpM (29-12,14,16), 2 IpF (29-18,20), 2 pM (29-102,103), 2 IP (29-15,

50 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

22), 2M, 1 F,3 p,2P,31,2 L [UCLA]. Suchitepequez: Patulul, 250 m, 20 July 1964, P. Cowsill

(GUA 56), 1 IpF (56-11), 1 P, 9 L [UCLA]. Rio Bravo, 175 m, 3 July 1964, P. Cowsill (GUA 24),

1 pF (24-100) [UCLA]. Department unknown: El Penon, 24 Aug 1953 (GML), 2 IpF (01639,

01640) [UCLA]. Rio Blanco, P. Woke, 41 [USNM].

MEXICO. Chiapas: Arriaga, 21 May 1963, 1 F [USNM]. La Esperanza, Salto de Agua, 12 Sept

1958, 1 F [USNM]. Sumidero (24 km N Tuxtla Guteirrez), 1030 m, 26-29 June 1953, P. Galindo,

H. Trapido, G. Boshell-Manrique (GML), 5 IpM (01528,01631,01715,01718,01923), 4 IlpF (01567,

01568,01572,01922), 1 pM (01711), 5 M, 9 F, 2 L [UCLA, USNM, GML]; same data, 23 July

1963, E. Fisher (MF 8), 6 F [UCLA]; same data, 17 Aug 1964 (MEX 120), 13 F [UCLA]; same

data (MEX 121), 3 F [UCLA]; same data (MEX 128), 2 F [UCLA]. Tapachula, 4 Aug 1953,

P. Galindo and H. Trapido (GML), 2 lpM (01588,01593), 1 IpF (01589), 1 M [UCLA, USNM].

Guerrero: Chilpancingo (30 km S), 670-880 m, 31 Aug 1964, E. Fisher (MEX 166), 3 F; same

data, 7 Aug 1966, D. Schroeder (MEX 411), 1 F; same data, 8 Aug 1966 (MEX 413), 2 F; same

data (MEX 414), 16 F [UCLA]. Puerto Marquez (El Marques), nr. Acapulco, 30 Aug 1964, E.

Fisher (MEX 141), 1 F [UCLA]. Jalisco: El Tuito, 500 m, 29 Aug 1972, J. Belkin, (MEX 720),

1 F [UCLA]. Morelos: Canyon de Lobos, 8 km E Cuernavaca, 1330 m, 30 Sept 1954 (GML), 3 F

[GML]. Cuernavaca, 17 May 1945, N. Krauss, 1 F [USNM]. Nayarit: Tepic (ca 30 km NW),

1000 m, 7 June 1971, T. Zavortink and L. Nielsen (MEX 658), 1 lpM (658-26); same data

(MEX 662), 1 lpM (662-12) [UCLA]. Oaxaca: Matias Romero (42 km N), 26 July 1963, E. Fisher

(MF 11), 1 IlpM (11-10) [UCLA]. San Luis Potosi: Taman (1 km S), 14 Sept 1955 (GML), 2 lpM

(02319,02320) [UCLA]. Sinaloa: Hwy 15, 40 km NE Hwy 40, 450 m, 6 June 1971, T. Zavortink

and L. Nielsen (N-14-71, same as MEX 651), 2 IpF (N-14-71-8,10) [Utah]. Hwy 15, 55 km NE

Hwy 40, 970 m, 24 June 1970, K. and D. Schroeder (MEX 501), 1 lpM (501-71), 1 IpF (501-70),

1 M, 1 L [UCLA]. Sonora: Alamos (12 km SE), 5 June 1971, T. Zavortink and L. Nielsen

(MEX 643), 1 lpM (643-16) [UCLA]. Veracruz: Amatlan de los Reyes, 800 m, 28 July 1965,

D. Schroeder (MEX 243), 1 lpF (243-12), 1 L [UCLA]. Cordoba, 900 m, 13 July 1964, E. Fisher

and D. Verity (MEX 26), 3 F; same data, 14 July 1964 (MEX 32), 1 F; same data, 16 July 1964

(MEX 37), 2 F; same data, 29 June 1964 (MEX 55), 1 F; same data, 30 July 1965, D. Schroeder

(MEX 248), 1 F; same data, 7 Aug 1965, L. Ramirez (MEX 279), 1 pF (279-100); same data

(MEX 280), 1 pF (280-10), 1 1; same data, 11 July 1965, C. Hogue (MEX 377), 1 F [UCLA].

Cueva del Nacemiento del Agua, Rio Atoyac (10 km NE Cordoba), 700 m, 13 July 1965, C. Hogue

(MEX 381), 1 M, 3 F [UCLA]. Fortin, 950 m, 5 Oct 1954 (GML), 1 lpM (01895), 2 lpF (01894,

01897), 1 pM (01899), 1 M [UCLA]. Presidio (ca 25 km SE Cordoba), 335 m, 14 July 1965,

C. Hogue (MEX 386), 1 F [UCLA].

PANAMA. Panama: Cerro Azul, 1957 (GML), 1 IpM (03241) [UCLA].

Additional Record From the Literature

BRITISH HONDURAS. Cayo: Chiquibul Road (Bertram, 1971:748).

10. Haemagogus (H.) albomaculatus Theobald

Figs. 4,25 ,26

1903. Haemagogus albomaculatus Theobald, 1903:308-310. TYPE: Holotype female, Cara Cara

[Kara Kara] Creek, Demerara River, Demerara, British Guiana [Guyana], G.C. Low

(123) [BM].

Haemagogus (Haemagogus) albomaculatus of Bonne and Bonne-Wepster (1925:435); Dyar (1928:

141); Edwards (1932:179); Lane (1939:118); Anduze (1947:353); Levi-Castillo (1951b:11,

17-18).

Haemagogus (Stegoconops) albomaculatus of Lane (1953:800); Stone, Knight and Starcke (1959:

216); Fauran (1961:30); Forattini (1965:30-32); Cova Garcia, Sutil and Rausseo (1966a:

61-62, fig. 116).

Haemagogus albomaculatus of Theobald (1903b:283; 1905:37); Aiken and Rowland (1906:

Arnell: Genus Haemagogus 51

22); Aiken (1907:77; 1909:24); Howard, Dyar and Knab (1917:867, in part), Dyar (1921a:

113); Floch and Abonnenc (1947:11); Komp (1954a:50; 1954b:148-153); Foote and Cook

(1959:141).

Cacomyia albomaculatus of Coquillett (1906:25).

Cacomyia albomaculata of Theobald (1907:554; 1910:494).

Haemagogus splendens in part of Howard, Dyar and Knab (1917:867).

Haemagogus cyaneus in part of Theobald (1903a:308; 1910:493).

FEMALE. Wing: 3.00 mm. Proboscis: 2.70 mm. Forefemur: 2.15 mm. Abdo-

men: 3.05 mm. Dark scales of proboscis, palpus, wing and legs purple to violet.

Head: Eyes narrowly separated above antennae (1 ommatidial diameter). Decum-

bent scales of vertex and occiput primarily blue to greenish blue with some light

green or purple reflections. Proboscis relatively long, about 1.25 of forefemur;

palpus short, about 0.12 of proboscis; antenna about 0.65 of proboscis. Thorax:

Scales of mesonotum copper, with blue reflections on lateral and posterior margins,

deep blue to purple over supraalar bristles, silver in antealar area; scales on scutellar

lobes deep blue to purple. Apn lobes moderately enlarged, scales greenish blue,

often silver on anterior margin; ppn scales copper with some blue reflections. Legs:

Forecoxa with small to large patch of purple scales near middle; legs moderately

long, length of forefemur about 1.50 of distance from top of thorax to apex of

midcoxa; forefemur and midfemur with silver scales restricted to base of ventral

surface. Wing: R43; about 0.70-0.80 of R,. Abdomen: Dark scales of tergites

I-VII purple with some violet reflections, tergite VIII greenish blue; silver scales

in small basal median patch on tergites VI,VII; dark scales of sternites purple.

MALE (fig. 25). Wing: 2.90 mm. Proboscis: 2.90 mm. Forefemur: 2.05 mm.

Abdomen: 3.30 mm. Head: Proboscis about 1.40 of forefemur; palpus short, about

0.12 of proboscis; antenna about 0.60 of proboscis. Legs: Larger claw of foreleg

with acute submedian tooth, smaller claw of foreleg and claws of midleg simple.

Abdomen: Basal median silver scale patches on tergites VI,VII absent; lateral silver

scale patches somewhat reduced.

MALE GENITALIA (fig. 25). Segment VIII: Tergite about 0.80 of sternite,

distal margin rounded but slightly emarginate mesally, with enlarged setae laterally

and about 30 long, lanceolate scales mesally. Segment IX. Tergite deeply declivous

and weakly sclerotized mesally; lobes relatively conspicuous, with 3 or 4 setae.

Sidepiece: Conical, length about 2.5-3.0 times median width; basal tergomesal lobe

not enlarged, basally with numerous setae, the more proximal ones much elongate,

flattened, attenuate; mesal half distad of basal tergomesal lobe with setae rather

evenly distributed, short, longer on distal fourth; apical sternomesal scales lanceo-

late. Claspette: Stem bowed inward near middle in dorsal aspect, slightly narrowed

and sharply curved dorsad’ near middle, expanded slightly at apex; filament striate,

more or less narrowly triangular, expanded posteriorly and forming a long, narrow

point anteriorly. Clasper: Broadest near base, spiniform about 0.40 of clasper.

Phallosome: Aedeagus large, obovate, tip greatly produced forward, with deep

cleft mesally forming an acute sclerotized projection on each side of cleft, a

sclerotized carina dorsad at base of cleft; venter broadly open except on basal

third, with heavy sclerotized ridges lateroventrally which are expanded basally and

terminate in sclerotized lobes mesally. Proctiger: Cercal setae about 6; apical knob

of paraproct with about 20 serrations.

PUPA (fig. 25). Abdomen: about 3.65 mm. Trumpet: 0.50 mm. Paddle: 0.80

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C sub-

equal, single to triple, weaker than 7-C which is single. Trumpet: Dark brown,

52 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

lighter distally, reticulate sculpturing moderate. Abdomen: Moderately pigmented,

genital lobe and anterior segments darker. Float hair (1-1) with about 10 primary

branches and 2-5 secondary branches. Hair 1-II-VII moderately to strongly devel-

oped, dendritic, strongest on segments V,VI. Hair 2-VII moderate, considerably

cephalad of hair 1-VII. Hairs 3-II,III,5-IV,V single, strongly developed, subequal. Hair

6-VII moderately well developed, double or triple. Hair 9-VII near caudolateral

margin of tergite, 5-6b; 9-VIII 8-11b. Terminal Segments: Male genital lobe about

1.2 of tergite VIII. Paddle: Relatively narrow, apex acuminate, with darker pigmen-

tation along midrib. Hair 1-P double.

LARVA (fig. 26). Head: 0.90 mm. Siphon: 0.95 mm. Anal Saddle: 0.35 mm.

Head: Hair 5-C single or double; hair 6-C double (single or double). Thorax:

Integument without spicules. Tubercles of hairs 5-7-P broadly joined. Abdomen:

Hair 3-II-V well developed, single, as long as corresponding segment. Hair 12-l

present. Hair 12-II well developed, subequal to hair 3-II and considerably stronger

than hair 10-II. Segment VIII: Comb scales 8(7-10), with single sharply pointed,

very minutely fringed spine, spine long, usually double length of basal, attached

part of comb scale; in curved row. Siphon: Index about 3.0(2.8-3.1). Pecten teeth

about 14-16(12-17), extending to about basal 0.50-0.55 of siphon. Hair 1-S double,

inserted at basal 0.60 of siphon. Anal Segment: Spines on caudal margin of saddle

well developed, long. Hair 4a-X triple (double or triple), long. Boss strongly sclero-

tized.

SYSTEMATICS. Haemagogus albomaculatus can be distinguished from the re-

maining members of the Albomaculatus Section: in the adults by (1) well developed

lower stp bristle, (2) relatively long female proboscis (1.25 of forefemur), (3) dark

scales of female abdomen purple except for greenish blue tergite VIII, (4) ab-

sence of knee spots on midfemur and hindfemur, (5) scales of mesonotum and

ppn copper and (6) simple smaller claws of foreleg and midleg and larger claw of

midleg of male; in the male genitalia by aedeagus tip deeply cleft mesally with

sclerotized carina dorsally at base of cleft and acute sclerotized projections on each

side of cleft; and in the larva by the combination of (1) integument without spic-

ules, (2) hair 12-I present, (3) strongly sclerotized boss and (4) hair 12-II consider-

ably stronger than hair 10-II.

The larva of albomaculatus has previously been unknown and the larva described

and illustrated here is presumed to be that species. The association is not certain as

no larval rearings were available, although 5 of the 7 larvae examined were from a

collection which produced only a single male pupal rearing of albomaculatus in

addition to the larvae.

The relationships of albomaculatus are not clear, as it seems not to be particularly

closely allied to any of the other members of the Albomaculatus Section. It may be

a rather early offshoot of the main evolutionary line considering the unique develop-

ment of the aedeagus and its rather restricted distribution.

BIONOMICS. Little is known of the bionomics of albomaculatus. It breeds pri-

marily in treeholes and appears to be fairly common along the coast of the Guianas.

Its abundance in the more heavily forested interior is not known as relatively

little collecting has been done in this area.

The status of albomaculatus as a disease vector is uncertain. Sneath (1939; 1940)

reports yellow fever immunity among the inhabitants of the forested interior of

Guiana, undoubtedly as a result of exposure to jungle yellow fever. Albomaculatus

is probably present in these areas and may be involved in the transmission cycle,

although janthinomys is present also, and being a proven vector of jungle yellow

Arnell: Genus Haemagogus 53

fever, would be more highly suspect. There has been, to my knowledge, no research

on the ability of albomaculatus to transmit yellow fever, though it is probably a

capable vector as nearly all species of Haemagogus have been shown to be able to

harbor and transmit the virus.

DISTRIBUTION (fig. 4). Haemagogus albomaculatus extends from the mouth

of the Orinoco River to French Guiana. Material examined: 34 specimens; 6 males,

20 females, 1 pupa, 7 larvae, 1 individual rearing (pupal).

FRENCH GUIANA. Guyane: Kaw, 7 Mar 1969, J. Clastrier (FGC 3947), 1 lpM, 5 L; same

data (FGC 3948), 1 L; same data (FGC 3950), 1 L [UCLA].

GUYANA. Demerara: Cara Cara and Demerara River, Low, 1 F (holotype) [BM]. Georgetown,

H. Moore, 1 M, 1 F [USNM]. Georgetown, Hiawakilly, Demerara River, Dec 1908, H. Moore, 1 F

[USNM]. Essequibo: Issaroro, 17 Sept 1921, G. Bodkin and G. Gevers, 3 F [BM]. Mazaruni,

23 June 1936, 6 F [GML]; same data, 25 June 1936, W. Komp, 1 M [USNM]. Pickersgill, Low,

1 F [BM]. Potaro (?Landing), May 1909, L. Cleare, 2M, 1 F [BM]; same data, 21 Feb 1939,

A. Donovan, 1 F [UCLA]. Rupununi, H. Wise, 1 F [USNM]. Tumatumari, Potaro River, Dec

1915, G. Bodkin, 1 F [BM]. Turu Camp, Barima River rt. bank, 11 Aug 1901, Low, 1 F [BM].

Warapoka Mission, 1 F [USNM].

SURINAM. Suriname: Paramaribo, J. Bonne-Wepster, 1 F [USNM].

VENEZUELA. Delta Amacuro: Jotakui Island, Orinoco Delta, 20 Dec 1952, I. Ortiz, 1 M

[USNM] . State unknown: Guanaco River, 2 Jan 1929, F. Urich, 1 F [BM].

11. Haemagogus (H.) panarchys Dyar

Figs. 4,27,28,29

1921. Haemagogus (Stegoconops) panarchys Dyar, 1921a:104-105. TYPE: Holotype male (70)

with most of abdomen mounted on slide (1466), El Salado, Guayaquil, Guayas, Ecuador,

F. Campos Ribadeneira [USNM, 24331].

Haemagogus (Stegoconops) panarchys of Bonne and Bonne-Wepster (1925:430); Dyar (1928:

135); Edwards (1932:179).

Haemagogus (Longipalpifer) panarchys of Levi-Castillo (1951b:12,33-35); Stone, Knight and

Starcke (1959:216); Forattini (1965:53).

Haemagogus (Cyanoconops) panarchys of Lane (1939:121; 1953:806-807); Levi-Castillo (1949:

165-171).

Haemagogus panarchys of Dyar (1925a:30); Levi-Castillo (1954:84); Komp (1955:237-239).

FEMALE (fig. 27). Wing: 2.90 mm. Proboscis: 3.05 mm. Forefemur: 2.00 mm.

Abdomen: 2.95 mm. Dark scales of proboscis, palpus, wing and legs mostly blue

to violet with some purple reflections. Head: Eyes widely separated above antennae

(4 ommatidial diameters). Decumbent scales of vertex and occiput silvery blue to

bluish green. Proboscis long, about 1.50 of forefemur; palpus about 0.16 of pro-

boscis; antenna about 0.72 of proboscis. Thorax: Scales of mesonotum primarily

bluish green, blue over supraalar bristles, silver in antealar area; scales on scutellum

bluish green, blue on lobes. Well developed bristle present in antealar area immedi-

ately caudad of scutal angle. Apn lobes slightly enlarged, scales silver; ppn scales

silver; lower stp usually with 1 very well developed bristle and 1 to 3 moderately

developed bristles. Legs: Hindcoxa relatively small, base considerably below upper

margin of meron; legs relatively short, length of forefemur about 1.25 of distance

from top of thorax to apex of midcoxa; knee spots present on midfemur and hind-

femur. Wing: R,,3-about 0.65-0.75 of R,. Abdomen: Dark scales blue to greenish

54 Conte. deen Botclesee val Osu. 2 1973

blue, often with purple reflections; silver scales in broad basal band on tergites

II-VII, often expanded laterally, but not connected to lateral silver patches.

MALE (fig. 27). Wing: 2.40 mm. Proboscis: 2.80 mm. Forefemur: 1.65 mm.

Abdomen: 2.80 mm. Head: Proboscis relatively long, about 1.50-1.70 of forefemur;

palpus long, slender, straight throughout, about 0.73-0.79 of proboscis; 5-segmen-

ted; segments 2 and 3 ankylosed and long, making up about 0.50-0.55 of palpus;

segment 4 making up about 0.18-0.22 of palpus, segment 5 making up about

0.16-0.18 of palpus; segments 3-5 with several moderately to well developed

bristles near apex of each segment; antenna about 0.65 of proboscis. Legs: Knee

spots absent on midfemur and hindfemur; larger claw of foreleg and midleg with

acute submedian tooth; smaller claws with acute basal tooth.

MALE GENITALIA (fig. 28). Segment VIII: Tergite about 0.50 of sternite,

crescent shaped, distal margin convex with setae and scales, scales dense mesally,

elongate, lanceolate with acuminate or truncate tips. Segment IX: Tergite shallow-

ly emarginate and weakly sclerotized mesally; lobes prominent, strongly sclerot-

ized, with 1 or 2 moderately developed setae. Sidepiece: Conical, length 3.0-3.5

times median width; basal tergomesal lobe slightly enlarged, with numerous setae,

the more proximal ones much enlarged, flattened and attenuate; short, flattened,

apically curved setae extending distad from basal tergomesal area on extreme mesal

margin to near middle of sidepiece, shorter setae laterad of flattened setae, followed

by bare space to lateral half of sidepiece; distal half with longer setae rather evenly

distributed; a row of heavy, short setae dorsad of apical sternomesal scales extending

from middle to near apex, usually in row of 3 followed by row of 4; apical sterno-

mesal scales lanceolate to broadly lanceolate. Claspette: Stem bowed inward near

middle in dorsal aspect; relatively stout, curving sharply dorsad in basal third, with

large, dorsal setiferous tubercle near apical third, and a pilosity ventrally on apical

fourth, the hairs long and thick at apex; filament striate, leaflike, arising slightly

proximad of stem apex. Clasper: Broadest at base, spiniform long, about 0.60-0.65

of clasper. Phallosome: Aedeagus large, more or less pandurate, apex sharply incised

with a pointed, spiculose protuberance mesally; venter broadly open except on

basal third, with heavily sclerotized ridges lateroventrally which are expanded basally

but not meeting at midline and terminating in two heavily sclerotized lobes basally.

Proctiger: Cercal setae about 6; apical knob of paraproct with about 10 serrations.

PUPA (fig. 28). Abdomen: about 3.20 mm. Trumpet: 0.45 mm. Paddle: 0.65

mm. Cephalothorax: Very weakly pigmented, slightly darker dorsally. Hair 4-C

usually single. Hairs 5,7-C moderately well developed, subequal, single. Trumpet:

Light brown; reticulate sculpturing moderate. Abdomen: Weakly pigmented, slightly

darker mesally at base of anterior segments. Float hair (1-I) with about 10 primary

branches, secondary branches few, usually about 2-4. Hair 1-II-VII weakly to

moderately developed, single or multiple. Hair 2-VII short, within own length of

base of 1-VII. Hairs 3-II,]HI usually subequal, shorter than 5-IV,V which are usually

subequal. Hair 6-VII weak, subequal to 6-IIJ-VI. Hair 9-VII near caudolateral margin

of tergite, usually double; 9-VIII usually 4-5b. Terminal Segments: Male genital lobe

about 1.6 of tergite VIII. Paddle: Weakly pigmented; apex broadly rounded. Hair

1-P single, rarely double.

LARVA (fig. 29). Head: 0.80 mm. Siphon: 0.80 mm. Anal Saddle: 0.35 mm.

Head: Hair 6-C single. Thorax: Integument without spicules. Tubercles of hairs

5-7-P free. Abdomen: Hair 12-I present. Segment VIII: Comb scales 10-12(8-20),

with single pointed, minutely fringed spine, in irregular single row. Siphon: Index

about 2.7(2.1-3.2). Pecten teeth about 11-14(8-15), extending to about basal 0.45

Arnell: Genus Haemagogus 55

of siphon. Hair 1-S 4b, inserted at basal 0.50 of siphon. Anal Segment: Spines on

caudal margin of saddle reduced, most very small, very few elongate. Hair 4a-X

2-4b, elongate. Boss weakly sclerotized or absent.

SYSTEMATICS. Haemagogus panarchys is among the most unusual members

of the genus in the adult stage and one of the few species in the subgenus Haema-

gogus that can be distinguished with reasonable certainty in the pupal stage. It can

be distinguished: in the adults by (1) eyes widely separated above antennae (4 om-

matidial diameters), (2) proboscis long (1.50 of forefemur in the female and 1.50-

1.70 of forefemur in the male), (3) apn and ppn scales silver, (4) abdominal tergites

II-VII silver scaled dorsally at base and (5) male palpus long, about 0.75 of pro-

boscis; in the male genitalia by (1) tergite VIII short, crescent-shaped, (2) claspette

stem with dense pilosity distally and (3) aedeagus sharply incised at apex with

pointed, spiculose mesal protuberance; in the pupa by (1) hairs 5,7-C well devel-

oped, subequal and (2) hair 2-VII short and within its own length of base of hair

1-VII; and in the larva by the combination of (1) integument without spicules,

(2) hair 12-I present, (3) weakly sclerotized boss, (4) hair 1-C without serrations

and (5) comb scales usually 10-12 and in an irregular single row.

Panarchys does not appear to be particularly closely related to any other species

of Haemagogus. It may be a relict species close to the main evolutionary stem

which gave rise to the subgenus Haemagogus as it appears to share more characters

with Conopostegus than do any of the other species of the subgenus Haemagogus.

Among the characters which may indicate this relationship are the long male palpus

and male claws, somewhat better developed thoracic chaetotaxy and widely spaced

eyes in the adults and some details of the pupal chaetotaxy.

BIONOMICS. Larvae of panarchys are found most often in bamboo, both cut

or broken and uncut internodes, and treeholes. It has never been implicated in

any disease transmission.

DISTRIBUTION (fig. 4). Haemagogus panarchys is known from the lowlands

of Canar, Guayas and Los Rios provinces, Ecuador. Material examined: 719 speci-

mens; 131 males, 136 females, 120 pupae, 332 larvae; 69 individual rearings (32

larval, 33 pupal, 4 incomplete).

ECUADOR. Canar: Cochancay (Hwy 8, km 86), 280 m, 13 Feb 1966, J. Belkin and E. Gerberg

(ECU 164), 1 P [UCLA] . Guayas: Chongon (Hwy 3, km 18), 9 Feb 1966, J. Belkin and E. Gerberg

(ECU 134), 8 L [UCLA]. Cordillera de Chongon, R. Levi-Castillo, 1 M gen [USNM]. El Salado,

F.Campos R., 1 F; same data, R. Levi-Castillo, 8 M, 8 F [USNM]. Guayaquil, 14 M, 23 F [UCLA];

same data, 5 Apr 1943, 6 M, 21 [UCLA, USNM] ; same data, Apr 1955 (GML), 2 p [UCLA] ; same

data, Dec 1955 (GML), 12 L [UCLA]. Guayaquil, F. Campos R., 3 F, 4 M [USNM, BM]; same

data, 1944, R. Levi-Castillo, 6 M, 3 M gen, 12 F, 1 p 11 [USNM]. Guayaquil (Hwy 3, km 10),

12 Feb 1966, J. Belkin and E. Gerberg (ECU 160), 10 L; same data (ECU 161), 4 IpM (161-10,32,

34,37), 7 IpF (161-30,31,33,35,36,36A,39), 5 pM (161-100,103,104,108,112),6M,4 F,7p,45 L

[UCLA]. Guayaquil (Hwy 3, km 12), 11 Feb 1966, Hjort and Schroeder (ECU 140), 1 lpM (140-

10), 1 M, 1 p, 3 L; same data(ECU 141), 2 IpM-(141-10,14), 2 IpF (141-11,13), 8 pM (141-15,

100,101,104-107,110), 3 pF (141-103,108,109), 1 lp (141-18), 1 p, 1 P, 25 L; same data (ECU

147), 3 IpM (147-10-12), 1 IpF (147-13), 1 F, 2 P, 29 L; same data (ECU 148), 1 M,1 P, 2 L;

same data (ECU 149), 2 L; same data (ECU 151), 1 pM (151-100), 1 L; same data (ECU 152),

1 IpM (152-11), 2 IpF (152-10,12), 4 L; same data (ECU 153), 1 pM (153-101) [UCLA].

Guayaquil (Hwy 3, km 14), 9 Feb 1966, J. Belkin and E. Gerberg (ECU 155), 2 lpM (135-10,

12), 1 IpF (135-13), 3 pM (135-100,100,101), 1 IP (135-14), 1 P, 40 L; same data (ECU 136),

4 IpF (136-10,20,21,30), 4 pM (136-102,104-106), 1 pF (136-101), 1 IP (136-11), 4 P, 12 L

[UCLA]. Guayaquil Country Club, J. Murdock, 2 F; same data, Apr 1941, H. Hansons, 2 M

[UCLA]. Hermanitos Experimental Farm (km 3, Guayaquil-Salinas Rd.), 12 M, 36 F, 2 P,1L

[UCLA] ; same data, R. Levi-Castillo, 9 M, 7 F, 30 P, 32 L [USNM, BM]. Los Ciebos, Guayaquil,

56 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

15 Feb 1966, J. Belkin (ECU 171), 24 L [UCLA]. Pascuales (Hwy 2, km 9.5), 5 Feb 1966, Hjort

(ECU 105), 1 lpM (105-11), 1 lpF (105-10), 1 pF (105-102), 1 IP (105-12), 6 1; same data

(ECU 106), 5 L; same data (ECU 107), 1 pF (107-102), 29 L [UCLA]. Los Rios: Valencia

(4 km W), 6 Feb 1966, J. Belkin and E. Gerberg (ECU 114), 1 M [UCLA]. Locality unspecified:

F. Campos R., 4 M, 4 F [USNM] ; same data, R. Levi-Castillo, 2 M, 8 M gen, 2 F [UCLA, USNM];

same data, 1938, R. Levi-Castillo, 4 M, 4 F [USNM]; same data, 1946, 3 M gen [USNM]; same

data, 1950, 41 [USNM].

12. Haemagogus (H.) soperi Levi-Castillo

Figs. 5,30,31

1955. Haemagogus (Longipalpifer)- soperi Levi-Castillo, 1955b:480-484, 2 plates. TYPE: Holo-

type male with associated larval and pupal skins, Juan Montalvo, Los Rios, Ecuador

[USNM].

Haemagogus (Longipalpifer) soperi in part of Stone, Knight and Starcke (1959:216); Forattini

(1965:53).

Haemagogus spegazzinii falco in part of Levi-Castillo (1952:76-81).

FEMALE. Wing: 2.90 mm. Proboscis: 2.85 mm. Forefemur: 2.05 mm. Abdo-

men: 2.80 mm. Dark scales of proboscis, palpus, wing and legs purple and violet.

Head: Eyes narrowly separated above antennae (1 or 2 ommatidial diameters). De-

cumbent scales of vertex and occiput bluish green to light green. Proboscis relatively ©

long, about 1.30-1.40 of forefemur; palpus about 0.16 of proboscis; antenna about

0.65-0.70 of proboscis. Thorax: Scales of mesonotum copper to dark green, often

dark bluish green to blue in fossa, light bluish green posteriorly, greenish blue over

supraalar bristles, silver in antealar area; scales on scutellum bluish green to greenish

blue. Apn lobes considerably enlarged, scales blue to violet; ppn scales usually green

to bluish green, occasionally copper, often a few silver scales ventrally. Legs: Moder-

ately long, forefemur length about 1.50-1.60 of distance from top of thorax to

apex of midcoxa; conspicuous silvery white knee spots anteriorly on mid and hind-

femur. Wing: R,,3 about 0.70-0.95 of R,. Abdomen: Dark scales of tergites

purple to violet; silver scales dorsally on tergites VI and VII forming basal band,

silver scales occasionally forming a small patch dorsally on IV and V.

MALE (fig. 30). Wing: 2.55 mm. Proboscis: 2.90 mm. Forefemur: 1.95 mm.

Abdomen: 2.75 mm. Head: Proboscis about 1.45-1.55 of forefemur; palpus long,

slender, straight, about 0.65-0.70 of proboscis, 5-segmented, segments 2 and 3

ankylosed and long, making up about 0.60-0.65 of palpus, segment 4 making

up about 0.15-0.18 of palpus, segment 5 making up about 0.12-0.14 of palpus;

segments 3-5 with several moderately to well developed bristles near apex of each

segment; antenna about 0.62 of proboscis. Legs: Larger claw of foreleg and smaller

claw of foreleg and midleg with acute subbasal tooth; larger claw of midleg simple.

MALE GENITALIA (fig. 30). Segment VIII: Tergite about 0.75 of sternite,

distal margin declivous mesally, with enlarged setae laterally and about 20-30

lanceolate scales mesally. Segment IX: Tergite deeply emarginate and weakly scle-

rotized mesally; lobes with 1 or 2 setae. Sidepiece: Conical, length about 3 times

median width; basal tergomesal lobe slightly enlarged, with numerous setae, the

more proximal elongate, flattened and attenuate; mesal half of sidepiece between

basal sternomesal area and distal fourth with moderately developed setae laterally

Arnell: Genus Haemagogus 57

and bare space mesally; distal fourth with well developed setae; apical sternomesal

scales lanceolate to oblanceolate with acuminate tips. Claspette: Stem bowed in-

ward near middle in dorsal aspect; narrowed at base and slightly expanded to apex,

angled dorsad near middle; filament striate, broadly sickle-shaped, broader near

basal third, with tip often slightly recurved. Clasper: Broadest at base; spiniform,

about 0.45 of clasper. Phallosome: Aedeagus large, broadly obovate, with a median

dorsal carina, tip produced forward, broad, strongly sclerotized, terminating in a

strongly recurved beak; venter broadly open except on basal third, with heavily

sclerotized ridges lateroventrally which are expanded basally and joined at base.

Proctiger: Cercal setae about 6; apical knob of paraproct with about 15 serrations.

PUPA (fig. 30). Abdomen: about 2.90 mm. Trumpet: 0.40 mm. Paddle: 0.70

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C weakly

developed, single or double. Hair 7-C moderately developed, single. Trumpet: Me-

dium brown, lighter apically; reticulate sculpturing moderate. Abdomen: Weakly

pigmented, darker mesally at base of anterior segments. Float hair (1-1) with 10-

12 primary branches and about 4-8 secondary branches. Hair 1-II-VII weakly

developed, usually single. Hair 2-VII short, within own length of base of 1-VII.

Hair 3-IL,II] usually subequal, shorter than 5-IV,V which are usually subequal.

Hair 6-VII weak, single or double, subequal to 6-III-VI. Hair 9-VII near caudo-

lateral margin of tergite, single or double; 9-VIII 4-7b. Terminal Segments: Male.

genital lobe about 1.4 of tergite VIII. Paddle: Weakly pigmented; apex broadly

rounded to slightly acuminate. Hair 1-P single.

LARVA (fig. 31). Head: 0.80 mm. Siphon: 0.88 mm. Anal Saddle: 0.35 mm.

Head: Hair 1-C conspicuously serrate laterally near apex. Hair 5-C single. Hair

6-C double. Thorax: Integument without spicules. Tubercles of hairs 5-7-P weakly

sclerotized and not connected. Abdomen: Hair 12-I present. Segment VIII: Comb

scales 8-10(5-12), with single broadly rounded to spatulate, minutely fringed spine,

in single row. Siphon: Index about 2.8(2.6-3.3). Pecten teeth about 11-14(8-17)

extending to about basal 0.55 of siphon. Hair 1-S double (double or triple), inserted

at basal 0.60 of siphon. Anal Segment: Spines on caudal margin of saddle moderately

well developed. Hair 4a-X double or triple, long, about 0.67-0.75 of hair 4b-X.

Boss weakly sclerotized or absent.

SYSTEMATICS. Haemagogus soperi can be distinguished: in the adults from all

Haemagogus except acutisentis by the combination of (1) male palpus long (0.65

of proboscis), (2) knee spots of silver scales at apex of midfemur and hindfemur,

(3) proboscis long, 1.30-1.40 of forefemur in female and 1.45-1.55 of forefemur

in male, (4) R243 short, 0.70-0.95 of R, and (5) dorsal silver scales at base of

only abdominal tergites VI and VII; in the male genitalia by the combination of

(1) tergite VIII with deep declivity mesally on distal margin, (2) apical process of

aedeagus relatively broad and truncated and (3) bare space mesally on sidepiece

distad of basal sternomesal area; and in the larva by the combination of (1) integu-

ment without spicules, (2) hair 12-I present, (3) hair 1-C serrate laterally and (4)

comb scales broadly rounded or spatulate but never pointed.

Soperi is closely related to acutisentis with which it forms the soperi complex.

The 2 species can be separated by the distribution of setae on the sidepiece of

the male genitalia and the shape of the comb scales in the larva, and with a fair

degree of certainty in the pupa by hair 3-II,III of soperi being subequal and shorter

than hair 5-IV,V which are subequal. The 2 species are sympatric over the greater

part of their ranges and often found together in the same locality.

The soperi complex is allied to equinus, the adults and immatures differing only

58 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

in relatively minor details, the male genitalia diverging more. The soperi complex

may have been isolated from equinus by the rise of the Andes, and is now only a

remnant of a once more extensive population.

BIONOMICS. Haemagogus soperi is most often found in broken or cut bamboo

internodes but has also been taken from leaf axils. It apparently attacks man readily.

I have examined several collections containing both soperi and the closely related

acutisentis, all taken from bamboo, and with water from several internodes com-

bined. It is probable that the 2 species occur in different internodes separated by

physical or temporal factors as 2 such closely related species in the same container

would undoubtedly compete with each other.

Levi-Castillo (1952; 1955) reports soperi as a vector of sylvan yellow fever on

the coast of Ecuador but this report has not been confirmed.

DISTRIBUTION (fig. 5). Haemagogus soperi is known from the Pacific coastal

lowlands of Ecuador. Material examined: 463 specimens; 69 males, 137 females,

165 pupae, 92 larvae; 79 individual rearings (17 larval, 61 pupal, 1 incomplete).

ECUADOR. Bolivar: Echeandia, R. Levi-Castillo, 1 M gen [USNM]. Canar: Cochancay (Hwy 8,

km 86), 280 m, 13 Feb 1966, J. Belkin and E. Gerberg (ECU 167), 30 F [UCLA]. Guayas:

Empalme (3 km S), 6 Feb 1966, J. Belkin et al. (ECU 120), 13 F [UCLA]. Los Rios: Juan

Montalvo, R. Levi-Castillo, 2 M gen, 2 p, 3 1 [USNM]; same data, 8 Feb 1966, J. Belkin and

E. Gerberg (ECU 121), 1 IpM (121-13), 13 IpF (121-10-12,14-19,40-43), 18 pM (121-61,64,66-

69,75 ,78,80,81 ,83,85,101,102,109,111,112,114), 18 pF (121-60,62,63,65 ,71-74,76,77,82,100,

105-108,110,113), 29 M, 29 F, 45 p, 20 P, 2 1, 42 L [UCLA]; same data (ECU 124), 3 M,1F

[UCLA]. Juan Montalvo, Hacienda Mora, 8 Feb 1966, J. Belkin and E. Gerberg (ECU 125,

125A), 1 IpM (125-17), 2 IpF (125-10,18), 4 pM (125-102,105,108,109), 5 pF (125-103,107,

113;125A-103,105), 1 IP (125-16), 4 M, 9 F, 16 P, 28 L; same data (ECU 126), 5 F [UCLA].

Pichilingue, 6 Feb 1966, J. Belkin et al. (ECU 118), 1 pM (118-100), 1 pF (118-103), 1 M,

1 F, 3 p [UCLA]. Valencia (4 km W), 6 Feb 1966, J. Belkin et al. (ECU 114), 4 pM (114-77,

78,95,101), 8 pF (114-71,75,92,94,97,109,112,114) [UCLA]. Manabi: Bolivar (?Calceta), R.

Levi-Castillo, 1 M gen [UCLA].

13. Haemagogus (H.) acutisentis Arnell, n. sp.

Figs. 5,32,33

TYPES: Holotype male with associated larval and pupal skins and genitalia slide (ECU 112-

12), 1 km E of Valencia, Los Rios, Ecuador, elev. ca 100 m, larva from fallen cacao pod,

6 Feb 1966, J.N. Belkin et al. [USNM]. Allotype female with associated larval and pupal

skins (ECU 112-13), same data as holotype [USNM]. Paratypes: 1 pF (ECU 112-103), 5 M, 4 F,

7 P (ECU 112), same data as holotype; 1 pM (ECU 114-105), 2 L(ECU 114), same data as holo-

type except 4 km W of Valencia, larvae and pupa taken from cut bamboo internode [UCLA, BM].

Haemagogus (Longipalpifer) soperi in part of Levi-Castillo (1955:480-484); Stone, Knight and

Starcke (1959:216); Forattini (1965:53).

FEMALE. Wing: 2.90 mm. Proboscis: 2.75 mm. Forefemur: 2.20 mm. Abdo-

men: 3.00 mm. Apparently indistinguishable from soperi.

MALE (fig. 32). Wing: 2.60 mm. Proboscis: 2.85 mm. Forefemur: 1.95 mm.

Abdomen: 2.80 mm. Apparently indistinguishable from soperi.

MALE GENITALIA (fig. 32). Segment VIII: Tergite about 0.70-0.75 of sternite,

distal margin declivous mesally, with enlarged setae laterally and about 20-25

lanceolate scales mesally. Segment IX: Tergite deeply emarginate and weakly scle-

rotized mesally; lobes with 1 or 2 setae. Sidepiece: Conical, length about 3 times

median width; basal tergomesal lobe slightly enlarged, with numerous setae, the

Arnell: Genus Haemagogus 59

more proximal elongate, flattened and attenuate; mesal half of sidepiece between

basal sternomesal area and distal fourth with uniformly distributed, relatively

short setae; distal fourth with well developed setae; apical sternomesal scales

lanceolate to oblanceolate with acuminate tips. Claspette: Stem bowed inward near

middle in dorsal aspect, angled dorsad near middle; filament striate, broadly sickle

shaped, with tip often recurved. Clasper: Broadest at base; spiniform about 0.45

of clasper. Phallosome: Aedeagus large, broadly obovate, with a median dorsal

carina, tip produced forward, broad, strongly sclerotized, terminating in a strongly

recurved beak; venter broadly open except on basal third, with heavily sclerotized

ridges lateroventrally which are expanded basally and joined at base. Proctiger:

Cercal setae about 5 or 6; apical knob of paraproct with about 15 serrations.

PUPA (fig. 32). Abdomen: about 3.25 mm. Trumpet: 0.45 mm. Paddle: 0.80

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C weakly

developed, single or double, weaker than 7-C which is single. Trumpet: Medium

brown, lighter distally; reticulate sculpturing moderate. Abdomen: Weakly pig-

mented, genital lobe and anterior segments darker. Float hair (1-[1) with 8-12 pri-

mary branches and usually 2-6 secondary branches. Hair 1-]]-VII moderately devel-

oped, branched or dendritic. Hair 2-VII short, usually considerably cephalad of

1-VII. Hairs 3-II,III,5-IV,V subequal or with 3-III slightly weaker. Hair 6-VII

single or double, subequal to 6-III-VI which are moderately well developed. Hair

9-VII slightly cephalad of caudolateral margin of tergite, single or double; 9-VIII

4-6b. Terminal Segments: Male genital lobe about 1.3 of tergite VIII. Paddle:

Weakly pigmented; relatively narrow, apex acuminate. Hair 1-P single.

LARVA (fig. 33). Head: 0.80 mm. Siphon: 0.85 mm. Anal Saddle: 0.35 mm.

Head: Hair 1-C conspicuously serrate laterally near apex. Hair 5-C single. Hair

6-C double. Thorax: Integument without spicules. Tubercles of hair 5-7-P weakly

to moderately sclerotized, not connected. Abdomen: Hair 12-1 present. Segment

VIII: Comb scales 9-13, with single short, sharply pointed, minutely. fringed spine,

in single row. Siphon: Index about 2.9(2.5-3.6). Pecten teeth about 12-20, extend-

ing to about basal 0.50 of siphon. Hair 1-S double (double or triple), inserted at

basal 0.55 of siphon. Anal Segment: Spines on caudal margin of saddle moder-

ately well developed. Hair 4a-X 3-4b, long, about 0.67 length of hair 4b-X. Boss

weakly sclerotized or absent.

SYSTEMATICS. Haemagogus acutisentis is apparently indistinguishable from

soperi as adults. It can be distinguished from soperi and the remainder of the

Albomaculatus Section: in the male genitalia by the combination of (1) tergite

VIII with deep declivity mesally on distal margin, (2) apical process of aedeagus

relatively broad and truncated and (3) mesal half of sidepiece distad of basal

tergomesal area with setae short and evenly distributed, never with a bare space;

and in the larva by the combination of (1) integument without spicules, (2) hair

12-I present, (3) hair 1-C serrate laterally and (4) comb scales sharply pointed.

Closely allied to soperi, acutisentis can be separated from it by the distribution

of setae on the sidepiece of the male genitalia, the sharply pointed comb scales,

and in the pupa with a fair degree of certainty by hairs 3-II,III,5-IV,V being sub-

equal or with 3-III slightly weaker. The relationships of acutisentis are discussed

above under soperi.

BIONOMICS. Larvae of acutisentis have been taken from cut bamboo, fallen

cacao pods and treeholes. Further discussion of the bionomics of this species is

presented above under soperi.

DISTRIBUTION (fig. 5). Haemagogus acutisentis is known from the Guayas

60 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

River basin and El Oro province of Ecuador. Material examined: 64 specimens;

12 males, 12 females, 22 pupae, 18 larvae; 11 individual rearings (5 larval, 6 pupal).

ECUADOR. El Oro: El Guabo, R. Levi-Castillo, 1 M gen [USNM]. Guayas: Guayaquil (Hwy 3,

km 12), 11 Feb 1966, Hjort and Schroeder (ECU 141), 1 pF (141-102), 1 F, 1 P, 1 L; same data

(ECU 147), 1 L [UCLA]. Guayaquil (Hwy 3, km 14), 11 Feb 1966, Hjort and Schroeder (ECU

135), 1 lpF (135-11) [UCLA]. Guayaquil Country Club, J. Murdock, 1 M [UCLA]. Pascuales

(Hwy 2, km 9.5), 5 Feb 1966, Hjort (ECU 105), 1 pM (105-101) [UCLA]. Los Rios: Juan

Montalvo, 8 Feb 1966, J. Belkin and E. Gerberg (ECU 122), 1 pF (122-100) [UCLA]. Juan

Montalvo (Hacienda Mora), 8 Feb 1966, J. Belkin and E. Gerberg (ECU 125), 2 IpF (125-13,15),

12 L [UCLA]. Valencia (1 km E), type series, see above. Valencia (4 km W), type series, see

above. Manabi: Guale, R. Levi-Castillo, 1 M gen [USNM].

14. Haemagogus (H.) equinus Theobald

Figs. 5,34,35

1903. Haemagogus equinus Theobald, 1903b:282-283. TYPE: Holotype female, lower end of

Old Pound Road, Kingston, St. Andrew, Jamaica, taken feeding on a horse, 24 Aug,

M. Grabham [BM].

1906. Aedes philosophicus Dyar and Knab, 1906a:195. TYPE: Lectotype larval skin (295b),

with associated male and genitalia slide (330), Tehuantepec, Oaxaca, Mexico, 1 July

1905, F. Knab [USNM; selection of Dyar, 1921:103; see Stone and Knight, 1955:

288-289] . Synonymized with equinus by Howard, Dyar and Knab (1917:874-875).

1906. Aedes affirmatus Dyar and Knab, 1906b:164. TYPE: Lectotype female, Salina Cruz,

Oaxaca, Mexico, 15 July 1905, F. Knab [USNM 10023; selection of Dyar, 1921:103;

see Stone and Knight, 1955:287]. Synonymized with equinus by Howard, Dyar and

Knab (1917:817,875).

Haemagogus (Stegoconops) equinus of Dyar (1921a:102; 1923:183, in part; 1928:134, in part);

Bonne and Bonne-Wepster (1925:430); Edwards (1932:179, in part); Martini (1935:57);

Anduze (1941b:13; 1947:353).

Haemagogus (Longipalpifer) equinus of Levi-Castillo (1951b:12,31-33); Stone, Knight and Starcke

(1959:215-216, in part); Fauran (1961:30); Forattini (1965:48-53); Cova Garcia, Sutil and

Rausseo (1966a:61-62, fig. 114; 1966b:114, fig. 202); Porter (1967:38); Belkin, Heinemann

and Page (1970:187-190); Bertram (1971:745,756); Diaz Najera (1971 :82-90).

Haemagogus (Cyanoconops) equinus of Lane (1939:121, in part; 1953:802-806); Diaz Najera

(1963:191; 1966:61).

Haemagogus equinus of Theobald and Grabham (1905:37); Theobald (1905:37); Howard, Dyar

and Knab (1917:870-871, in part); Johnson (1919:424); Dyar (1925:138-139, in part);

Gowdey (1926:74); Shannon and Del Ponte (1928:68); Kumm and Zuniga (1942:404);

Boshell-Manrique and Osorno-Mesa (1944:173); Osorno-Mesa (1944a:39); Hill and Hill (1945:

2; 1948:49); Waddell (1945:329; 1949:568); Hill and Taylor (1945:226; 1947:472); Anderson

and Osorno-Mesa (1946:613); Hovanitz (1946:35); Kumm, Osorno-Mesa and Boshell-Manrique

(1946:19-20); Thompson (1947:79); Woke (1947:365); Arnett (1949:240; 1950:114); Galindo,

Carpenter and Trapido (1949:278; 1951a:116-117; 1955:158); Galindo, Trapido and Carpenter

(1950:546); Levi-Castillo (1951a:14; 1954:83); Carpenter, Galindo and Trapido (1952:162);

Vargas and Martinez Palacios (1953:38); Komp (1954a:51-53; 1955f:163); Barreto Reyes

(1955:79); Galindo and Trapido (1955:548; 1957:147); Horsfall (1955:535); Trapido (1955:

629); Trapido and Galindo (1955:669; 1956a:304; 1956b:634; 1957:122); Trapido, Galindo

and Carpenter (1955:528); Galindo, de Rodaniche and Trapido (1956:1022); Galindo, Trapido,

Carpenter and Blanton (1956:544); Eads and Strom (1957:86); de Rodaniche and Galindo

(1957:235); de Rodaniche, Galindo and Johnson (1957:682); Boshell-Manrique and Bevier

(1958:27); Breland (1958:217); Foote and Cook (1959:141); Vargas and Diaz Najera (1959:

Arnell: Genus Haemagogus 61

362); Diaz Najera (1960:185); Kerr, Roca-Garcia and Bugher (1960:26); Groot, Morales and

Vidales (1961:399).

Stegoconops equinus of Howard, Dyar and Knab (1913:fig. 162).

Cacomyia equinus of Coquillett (1906:25). .

Cacomyia equina of Theobald (1907:554-556; 1910:494).

Haemagogus affirmatus of Busck (1908:64); Aiken (1909:3,4,24).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870).

Haemagogus regalis in part of Dyar and Knab (1906b:167); Theobald (1910:493-494).

Haemagogus cyaneus of Aiken and Rowland (1906:121); Aiken (1907:77); Aiken (1909:24).

FEMALE. Wing: 3.20 mm. Proboscis: 2.90 mm. Forefemur: 2.30 mm. Abdo-

men: 3.55 mm. Dark scales of proboscis, palpus, wing and legs dark blue to violet

with some purple reflections. Head: Eyes narrowly separated above antennae

(2 ommatidial diameters). Decumbent scales of vertex and occiput blue to light

green. Proboscis relatively tong, about 1.25-1.40 of forefemur; palpus about 0.17

of proboscis; antenna 0.61-0.72 of proboscis. Thorax: Scales of mesonotum dark

green to dark bluish green, blue over supraalar bristles, silver in antealar area; scales

on scutellum dark green to blue. Apn lobes moderately enlarged, scales dark blue,

often with silver scales laterally and anteriorly; ppn scales dark blue to bluish green

with silver scales ventrally; lower stp with single well developed bristle and 1 to 3

smaller bristles. Legs: Moderately long, forefemur length about 1.38-1.48 of dis-

tance from top of thorax to apex of midcoxa; knee spots present on midfemur

and hindfemur. Wing: R,,3 about 1.05-1.25 of R,. Abdomen: Dark scales blue

to purple; silver scales in basal bands on tergites IV-VII, usually continuous with

lateral silver patches on VI and VII.

MALE (fig. 34). Wing: 2.65 mm. Proboscis: 2.90 mm. Forefemur: 2.00 mm.

Abdomen: 3.40 mm. Head: Proboscis about 1.35-1.50 of forefemur; palpus long,

slender, straight throughout, about 0.58-0.70 of proboscis; 5-segmented, segments

2 and 3 ankylosed and long, making up about 0.58-0.65 of palpus, segment 4

making up about 0.15-0.18 of palpus, segment 5 making up about 0.12-0.14 of

palpus; segments 3-5 with 1 to several stout bristles arising near apex of each

segment; antenna about 0.55-0.63 of proboscis. Legs: Larger claw of foreleg and

smaller claw of foreleg and midleg with acute subbasal tooth, larger claw of mid-

leg simple.

MALE GENITALIA (fig. 34). Segment VIII: Tergite about 0.55 of sternite,

distal margin broadly emarginate, with enlarged setae laterally and dense cluster

of 30-40 lanceolate to obovate scales mesally. Segment IX: Tergite declivous and

moderately sclerotized mesally; lobes with 1 to 3 setae. Sidepiece: Conical, length

3.5-4.0 times median width; basal tergomesal lobe not enlarged, with numerous

setae, the more proximal ones much enlarged and flattened; mesal half distad of

basal tergomesal area with setae rather evenly distributed, short, longer on distal

fourth; apical sternomesal scales lanceolate to oblanceolate. Claspette: Stem narrow-

est at base, slightly curved dorsad on basal third, distal half with membrane pro-

jecting anteriorly and posteriorly and enclosing basal portion of filament; filament

with basal supporting ribs, a long, narrow, slightly convoluted flap dorsad of stem,

slightly expanded at apex with a recurved tip; a broad, flattened flap ventrad

of stem, the margin of which may be rounded, angled, or with variously developed

pointed process. Clasper: Broadest near middle, spiniform about 0.40-0.45 of

clasper. Phallosome: Aedeagus large, broadly obovate, tip produced distally into

a strongly sclerotized dorsal carina proximad to a small beak; venter open except

62 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

on basal fourth, with heavily sclerotized ridges lateroventrally which are expanded

basally but not meeting at midline. Proctiger: Cercal setae 2-7; apical knob of

paraproct with about 15 serrations.

PUPA (fig. 34). Abdomen: about 3.30 mm. Trumpet: 0.45 mm. Paddle: 0.70

mm. Cephalothorax: Usually rather weakly pigmented, darker dorsally. Hairs 4,5-C

moderately developed, usually subequal, single to triple, and often subequal to

7-C, which is usually single. Trumpet: Medium brown, reticulate sculpturing mod-

erate. Abdomen: Weakly to moderately pigmented, genital lobes and anterior seg-

ments darker. Float hair (1-I) with about 8-12 primary branches and 2-6 secondary

branches. 1-II-VII weakly to moderately developed, single, branched or dendritic.

Hair 2-VII variable in position, usually considerably cephalad of, but occasionally

close to 1-VII. Hairs 3-II,III,5-IV,V subequal or with 3-III weaker. Hair 6-VII

usually weak, subequal to 6-III-VI, but occasionally considerably stronger than

6-III-VI. Hair 9-VII near or somewhat cephalad of caudolateral margin of tergite,

single to triple; 6-VIII 3-6b. Terminal Segments: Male genital lobe about 1.3 of

tergite VIII. Paddle: Weakly pigmented, usually relatively narrow with acuminate

apex, occasionally rather broad with broadly rounded apex. Hair 1-P single.

LARVA (fig. 35). Head: 0.75 mm. Siphon: 0.65 mm. Anal Saddle: 0.30 mm.

Head: Hairs 5,6-C single (single or double). Thorax: Integument without spicules.

Tubercles of hairs 5-7-P connected in hairy forms, free in non-hairy forms. Abdo-

men: Hair 12-I present. Segment VIII: Comb scales 7-10(5-13), with single sharply

pointed, minutely fringed spine, in single row. Siphon: Index 2.5-3.0(2.1-3.3).

Pecten teeth 11-16(8-22), extending to about basal 0.45 of siphon. Hair 1-S double

(1-4b), inserted at basal 0.50 of siphon. Anal Segment: Spines on caudal margin

of saddle well developed. Hair 4a-X 3-5b, long. Boss weakly sclerotized or absent.

SYSTEMATICS. Haemagogus equinus can be distinguished from all remaining

members of the Albomaculatus Section: in the adults by the combination of (1)

long male palpus (about 0.65 of proboscis), (2) conspicuous knee spots on apex

of midfemur and hindfemur, (3) R243 about 1.05-1.25 of R,, (4) silver scales

dorsally on abdominal tergites IV-VII and (5) apn and posterior ppn usually with

silver scales; in the male genitalia by claspette stem with distal membrane which

projects dorsally and ventrally and encloses basal portion of filament, and filament

with conspicuous basal supporting ribs; and in the larva by the: combination of

(1) integument without spicules, (2) hair 12-I present, (3) weakly sclerotized boss,

(4) hair 1-C without serrations and (5) comb scales usually 7-10 in single, regular

row.

There is variation in equinus in the shape of the ventral portion of the claspette

filament of the male genitalia and in the number of silver scales on the apn and ppn

of the adults. Hairy forms of the larvae and pupae are common in this species

throughout its range. Correlated with the increase in length and branching of the

hairs in the hairy forms is an increase in sclerotization of the tubercles of some of

the thoracic and abdominal hairs and the boss of the ventral brush. Hairy, nonhairy

and intermediate forms are occasionally found together in the same collection.

Equinus, and the closely allied soperi and acutisentis constitute one of the domi-

nant elements of the Haemagogus fauna in terms of range, plasticity of habitat

and number of individuals. This dominance may indicate equinus to be one of the

more modern Haemagogus species, however the complex distribution pattern and

complex geological history of the area now occupied by this species may indicate

considerable age.

A relatively recent land connection between Central America and the Greater

Arnell: Genus Haemagogus 63

Antilles, at least the island of Jamaica, is suggested by the distribution pattern

of equinus. A land connection in this area during middle Tertiary, called Caribbean

Land, has been proposed by paleogeologists (Maldonado-Koerdell, 1964:15-18).

A similar distribution pattern is shown by several other culicine species (Belkin,

Heinemann and Page, 1970:9; Belkin and Hogue, 1959:430). A long-lasting dis-

junction between mainland and insular populations of equinus would likely result

in a morphological diversity between these populations which does not exist at

present. The center of dispersal of equinus may be Central America, as this.species

appears to be much more abundant in this area than elsewhere in its range. Dispersal

northward into Mexico, east to Jamaica and southeast into northern South America,

was accomplished as land connections were made to these areas. An alternate

explanation that equinus originated on the South American continent and spread

northward into Central America during middle Tertiary may be indicated by the

apparent relationship between equinus and the soperi complex of coastal Ecuador.

Such a dispersal would depend on the existence of a land connection between South

and Central America at a time prior to a land connection between Central America

and Jamaica.

BIONOMICS. Haemagogus equinus, beside being one of the most abundant and

widespread species of Haemagogus, is probably the most adaptable in terms of.

habitat utilization. In Central America it is common in the Atlantic rain forest,

is one of the most numerous species in the tropical deciduous forest of the Pacific

versant of Nicaragua and is found in considerable numbers in the deciduous forest

north along the Pacific to Sinaloa in Mexico. It is found in the thorn forest of the

Atlantic coast of northern Mexico and in coastal mangrove in many areas of Central

America although it is not common where the typical mangrove breeding species of

Haemagogus of the Splendens Section, boshelli, chalcospilans, regalis and aeritinctus

are present. It apparently reaches its maximum density in the deciduous forest of

Central America. Equinus commonly utilizes treeholes and cut and broken bamboo

as breeding sites and it is often found in peridomestic situations. It has been taken

at elevations of up to 1400 meters but is not common above 750 meters. In the

rain forest, equinus is decidedly arboreal, being taken biting above ground level

or in the forest canopy about 60 percent of the time. Most eggs hatch when first

flooded, and since the length of larval developmental time is relatively short,

populations of equinus reach a peak shortly after the beginning of the rainy season,

in contrast to other species of Haemagogus whose populations build slowly after

the onset of seasonal rains. The more important papers on the biology of equinus

are those of Galindo, Carpenter and Trapido (1951a), Trapido and Galindo (1956),

Galindo and Trapido (1957) and Hovanitz (1946).

Equinus is a proven laboratory vector of yellow fever (Waddell and Taylor,

1945, 1947; Waddell, 1949; Galindo, de Rodaniche and Trapido, 1956) and infected

females have been found in nature (de Rodaniche and Galindo, 1957; de Rodaniche,

Galindo and Johnson, 1957). Because of its arboreal habits and abundance in areas

of yellow fever epizootics in Central America, equinus is a primary suspect as a

yellow fever vector, especially where janthinomys is less abundant or absent, as in

Honduras and Guatemala (Trapido and Galindo, 1955; Boshell-Manrique and Bevier,

1958).

DISTRIBUTION (fig. 5). Haemagogus equinus extends from extreme southern

Texas and southern Sinaloa through southern Mexico, Central America, the Carib-

bean and Orinoco drainages of Colombia and Venezuela to Guyana and is also

found on the islands of Jamaica and Tobago and probably Trinidad. The records

64 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

in Stone, Knight and Starcke (1959:215) of equinus from Brazil and Bolivia are

based on misidentified janthinomys females. Material examined: 3924 specimens;

867 males, 862 females, 1079 pupae, 1116 larvae; 224 individual rearings (133

larval, 58 pupal, 33 incomplete).

BRITISH HONDURAS. Cayo: Central Farm, 70 m, 17 Aug 1967, D. Bertram (BH 479), 1 F

[UCLA]. Chiquibul Road, mile 9 from Georgeville, 300 m, 26 June 1967, D. Bertram (BH A235),

1 F; same data, 10 July 1967 (BH A270), 1 F; same data, 11 July 1967 (BH A271), 1 F; same

data. 7 Aug 1967 (BH A456), 1 F [UCLA]. Chiquibul Road, mile 8 from Georgeville, 1 July

1965, R. Disney, 1 M gen [UCLA]. Mountain Pine Ridge, nr. Augustine, 550 m, 10 Aug 1967,

P. Williams (BH 490), 3 F [UCLA].

COLOMBIA. Antioquia: Turbo, H. Kumm, 1 M, 4 M gen [USNM]. Bolivar: ?La Guinea, 1 M

gen [USNM]. Cundinamarca: Caparrapi, Volcanes Forest, 1000-1500 m, 1943, H. Kumn, 1 Ip,

1 M [USNM]. Fusagasuga, 1746 m, 1943, H. Kumm, | | [USNM]. Malta, 280 m, 25 Nov 1941,

E. Osorno-Mesa, 1 L; same data, 14 Apr 1942, H. Kumm and E. Osorno-Mesa, 1 M gen, 3 1

[USNM]. Utica, H. Kumm, 1 M [USNM]. Meta: Villavicencio, 470 m, 5 M, 1 M gen [USNM].

COSTA RICA. Alajuela: Esparta (11.8 km E), 500 m, 19 June 1963, C. Hogue (CR 97), 1 F

[UCLA]. Higuito, nr. San Mateo, 200 m, P. Schild, 3 F [USNM]. Cartago: Turrialba, 610 m,

June 1924, P. Buxton, 1 F [BM]. Guanacaste: El Coco, 19 July 1962, F. Truxal, 1 M, 7 F

[LACM]. Las Canas, H. Kumm, 1 F [UCLA]. Liberia (Finca Coyolar), 1 Aug 1964, C. Hogue,

1 F [UCLA]. (?Hacienda) Miravalles, 20 July 1922, A. Alfaro, 1 F [USNM]. Samara, 5 m,

23 Aug 1964, C. Hogue and Miranda (CR 196), 1 L [UCLA]. Limon: La Bomba, nr. Limon,

3 Oct 1971, D. Schroeder (CR 470), 1 lpM (470-10) [UCLA] . Puntarenas: Boca del Rio Barranca,

10 m, 20 June 1963, C. Hogue (CR 103), 1 F [USNM]. Esparta (3 km E), 250 m, 13 Aug 1971,

D. Schroeder (CR 357), 2 IpF (357-10,11), 2 pM (357-100,101), 5 pF (357-102-106); same data

(CR 358), 1 pM (358-101) [UCLA] . Macacona, nr. Esparta, 1 June 1943, T. Aitken, 1 F [UCLA].

San Jose: San Isidro del General, 21 Nov 1962, C. Hogue and W. Powder (CR 35), 1 P, 1 LIL

[UCLA].

EL SALVADOR. Liberdad: La Liberdad, 15 June 1953 (GML), 1 lpM (01417), 1 pM (01423),

1 pF (01429) [UCLA]. Zaragoza, H. Kumm, 3 F [UCLA]. San Miguel: San Miguel, 2M [USNM];

same data, H. Kumm, 2 M [UCLA]. San Salvador: Los Planes, H. Kumm, 2 M, 11 F [UCLA,

USNM]. Parque Balboa, 980 m, 5 Nov 1971, J. Belkin (SAL 49), 1 pF (49-101) [UCLA]. San

Salvador, H. Kumm, 3 M, 2 F [UCLA]. Sonsonate: Izalco, F. Knab, 1 F [USNM]. Izalco, Parque

Atecozol, 430 m, 6 Nov 1971, J. Belkin (SAL 53), 2 IpF (53-60,61), 1 P, 1 1, 1 L [UCLA].

Nahuilingo, H. Kumm, 1 M [UCLA]. San Antonio del Monte, 4 Aug 1964, A. Quinonez (SAL 8),

1 L; same data (SAL 10), 2 IpF (10-13,17), 2 pM (10-100,102), 2 pF (10-107,108), 1 p, 3 P,

30 L [UCLA]. Sonsonate, F. Knab, 1 M, 4 F [USNM]; same data, 2 Aug 1964, A. Quinonez

(SAL 4), 1 IpM (4-15), 2 pM (4-100,106), 1 pF (4-101), 1 p, 2 P, 22 L [UCLA]; same data,

3 Aug 1964 (SAL 6), 1 IF (6-14), 1 pM (6-101), 8 P, 61, 22 L [UCLA]; same data (SAL 7),

1 IP (7-10), 3 L [UCLA]. Locality unspecified: 1 L [USNM].

GUATEMALA. Alta Verapaz: Trece Aguas, Barber and Schwarz, 1 F [USNM]. Escuintla:

Escuintla, 326 m, Schaus and Barnes, 1 F [USNM]. El Salto, 1 Nov 1954 (GML) 1 1 [UCLA].

Izabal: Livingston, H. Barber, 1 F [USNM]. Mojaca Village (ca 8 km W Morales), 50 m, 28 June

1964, T. Zavortink, 2 F [UCLA]. Retalhuleu: Champerico, 0 m, Nov 1965, 2 M [USNM].

Suchitepequez: Patulul, 250 m, 20 July 1964, P. Cowsill (GUA 56), 2 IpF (56-10,12), 4 L

[UCLA]. Department unknown: San Jose, 12 May 1943, D. Hall, 3 M [USNM]. Locality un-

specified: R. Morales, 1 F [USNM]; 1954 (GML), 2 pM (02080, 02087) [UCLA].

GUYANA. Berbice: Courantyne, Benab Village 63, 6 Oct 1962. T. Aitken (6/X/62), 1 pM (1);

same data, 8 Oct 1962, (8/X/62), 2 IpM (2,3), 4 IpF (1,4-6), 1 IP (7) [UCLA]. Demerara:

Georgetown, 7 Nov 1967, Hansell and Rauch (GUY 34), 3 IpM (34-10-12); same data (GUY 35),

2 IpM (35-10,11), 1 pM (35-101), 1 pF (35-100), 18 L; same data, 12 Mar 1968 (GUY 73), 1 pF

(73-100), 1 L; same data, 12 Apr 1968, Ramjattan (GUY 74), 1 lpF (74-14), 1 pM (74-101), 1 pF

(74-100), 1 lp (74-13), 1 IP (74-10), 3 L [UCLA].

HONDURAS. Atlantida: Lancetilla, nr. Tela, 18 Nov 1954 (GML), 1 pF (01979), 11 [UCLA].

Lancetilla Valley, nr. Tela, 19 Aug 1964, A. Quinonez (HON 49), 3 L; same data (HON 52), 1 lpM

Arnell: Genus Haemagogus 65

(52-14), 5 IpF (52-10-13,15), 2 pM (52-100,101), 1 IP (52-16), 3 F, 3 P, 4 L; same data (HON 54),

1 pF (54-100); same data (HON 55), 1 lp (55-20) [UCLA]. Las Metalias, nr. Tela, 21 Aug 1964,

A. Quinonez (HON 57), 1 1 [UCLA]. Rio Mezapa, nr. Rio Lean, 4 Sept 1954 (GML), 21 [UCLA].

JAMAICA. St. Andrew and Kingston: Constant Spring, May 1945, R. Hill, 1 M, 1 F,3 L

[USNM]; same data, 7 Sept 1965, J. Belkin (JA 327) 2 IpF (327-14,16), 2 p, 3 L [UCLA] ; same

data (JA 328), 1 pM (328-101) [UCLA]; same data, 16 Nov 1966, O. Berlin and Watson (JA 696),

1 F [UCLA]; same data (JA 699), 1 pM (699-105) [UCLA]. Hermitage Gate, 29 Oct 1965,

W. Page (JA 391), 1 pM (391-108) [UCLA]. King’s House, Kingston, Nov 1924, G. Strathairn,

1 M [BM]. Kingston, May 1945, R. Hill, 1 M gen, 2 F [USNM]. Kingston (Pound Road), 24 Aug,

M. Grabham, 1 F (holotype) [BM]. Rockfort, 2-30 m, 21 Aug 1968, Hochman (JA 948), 9 F;

same data (JA 951), 3 IpF (951-11,13,15), 1 IP (951-12), 3 L; same data, 22 Aug 1968 (JA 952),

1 M, 6 F; same data, 24 Aug 1968 (JA 957), 4 F; same data (JA 958), 1 lpF (958-10), 5 L; same

data, 25 Aug 1968 (JA 959), 1 F [UCLA]. St. Mary: Broadgate, 28 Nov 1965, W. Page (JA 407),

1 pF (407-114) [UCLA]. St. Thomas: Grant’s Pen, 10 Dec 1965, W. Page (JA 410), 2 IP (410-

12,13), 5 L; same data (JA 412), 1 IlpM (412-10), 1 pM (412-100), 1 pF (412-101), 1 p, 11

[UCLA].

MEXICO. Campeche: Campeche (20 km SW), 2 m, 21 July 1970, D. Schroeder (MEX 586),

1 F [UCLA]. Colima: Manzanillo (ca 23 km NW), 22 July 1963, S. Telford (MT 9), 1 F [UCLA].

Manzanillo (ca 26 km SE Hwys 80,200), 200 m, 21 July 1963, 8S. Telford (MT 6-3), 1 F [UCLA].

Guerrero: Acapulco, F. Knab, 5 M, 4 F [USNM]. Acapulco (GML), 1 lpM (02524), 1 lpF (02522)

[UCLA]. Puerto Marquez (?El Marques), 30 Aug 1964, E. Fisher and D. Verity (MEX 141), 1 M;

same data (MEX 143), 1 M, 3 F [UCLA]. Jalisco: Las Penas, 18-21 July 1903, A. Duges, 5 F;

same data, June 1906, 3 F [USNM]. Nayarit: San Blas, 26 June 1956, W. McDonald (UCLA 198),

1 IpM (198-110), 2 IpF (198-106,107), 6 lp (198-103,113-117), 4 p, 4.1; same data, 27 June 1956

(UCLA 203), 4 pM (203-104,106,108,112), 1 pF (203-103), 1 lp (203-105), 4 p, 14 L; same data,

25-29 June 1956, 5 M, 13 F; same data, 25 July 1963, P. Spangler, 1 F [UCLA]. San Blas

(6 km SE), 10 July 1963, E. Fisher (MF 2), 1 lpM (2-19), 6 IpF (2-10,13,15-18), 1 pM (2-12),

1 pF (2-10), 1 L; same data (MF 3), 7 M, 6 F [UCLA]. Tepic (ca 30 km NW), 300 m, 7 June 1971,

T. Zavortink and L. Nielsen (MEX 658), 3 IpM (658-10,11,31), 12 lpF (658-12-20,27,30,32),

30 M, 17 F, 46 p, 6 1, 27 L; same data (MEX 659), 7 F, 7 P; same data (MEX 660), 4 M, 3 F, 7 p;

same data (MEX 661), 2 lpM (661-10,11), 4 lpF (661-12-14,17), 1 M, 1 p, 1 1; same data

(MEX 662), 1 IpM (662-10) [UCLA]. Oaxaca: Salina Cruz, F. Knab, 7 F [USNM]. Tehuantepec,

5M,1Mgen, 3 F, 2 p, 1 1 [USNM]. San Luis Potosi: Saketepan, nr. Tamazunchale, 200 m, 20 July

1965, D. Schroeder (MEX 220), 1 F; same data, 21 July 1965 (MEX 222), 1 lpM (222-10); same

data (MEX 228), 2 lpF (228-11,12) [UCLA]. Sinaloa: Hwy 15, 1 km NW road to La Cruz, 6 June

1971, T. Zavortink and L. Nielsen (MEX 650), 1 F, 1 p [UCLA]; same data (N-13-71), 6 IpF (6-

12), 20 M, 2 F [Utah]; same data (N-14-71), 6 M, 9 F [Utah]. Palmillas (11 km NW),40M,7 June

1971, T. Zavortink and L. Nielsen (MEX 655), 5 M, 6 F [UCLA]. Tabasco: Comalcalco (7 km N),

40 m, 12 July 1970, D. Schroeder (MEX 553), 1 IpM (553-20), 1 IpF (553-21), 1 pM (553-100)

[UCLA]. Frontera, Apr 1928, Townsend, 1 F [USNM]. Veracruz: Amatlan de los Reyes (2kmE),

800 m, 28 July 1965, D. Schroeder (MEX 241), 1 IpM (241-10) [UCLA]. Amatlan de los Reyes

(3 km S), 800 m, 28 July 1965, D. Schroeder (MEX 243), 1 lpM gen (243-10), 1 lpF (243-11), 2 L

[UCLA]. Boca del Rio, 0 m, 28 July 1964, E. Fisher (MEX 84), 3 F [UCLA]. Cerro Guzman

(?Guzmantla), 9 July 1970, D. and K. Schroeder (MEX 539), 1 IpM (539-80), 1 1 [UCLA].

Cordoba (2.5 km E), 920 m, 12 July 1964, E. Fisher and D. Verity (MEX 32), 1 F; same data,

16 July 1964 (MEX 37), 1 F; same data, 17 July 1964 (MEX 42), 2 IpM (42-10,15), 9 IpF

(42-11-14,16-19,21), 4 M, 4 F, 2 L; same data, 18 July 1964 (MEX 53), 1 IpF (53-10), 1 lp

(53-14), 2 L; same data (MEX 54), 1 IpF (54-10), 5 L; same data, 22 July 1964 (MEX 71),

1 IpF (71-16); same data (MEX 72), 2 IpM (72-11,12), 1 lpF (72-13); same data (MEX 74),

2 IpM (74-11,13), 2 IpF (74-10,14); same data (MEX 75), 3 L; same data (MEX 76), 2 IpF

(76-10,11), 1 L [UCLA]. Cuitlahauc (18 km E), 410 m, 15 July 1964, E. Fisher and D. Verity

(MEX 35), 1 F [UCLA]. Santa Lucrecia (Jesus Carranza), F. Knab, 1 F [USNM]. Veracruz,

Om, July 1928, A. Iglesias, 1 M [USNM]; same data, 12 Aug 1965, D. and K. Schroeder (MEX

295), 1 lpM (295-11), 1 IpF (295-10), 1M, 4 L [UCLA].

66 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

NICARAGUA. Chinandega: Corinto, 6 Sept 1944, H. Crowell, 10 F [UCLA]; same data,

15 Sept 1944, 1 M, 3 F [UCLA]; same data, 24 Oct 1944, 3 M, 3 F [USNM]; same data, 24 May

1945, 10 F [UCLA]; same data, 10 June 1945, 7 F [UCLA]; same data, 29 Sept 1945, 7 F

[UCLA]. Leon: Simonello (nr. Nagarote), 14 June 1964, A. Quinonez (NI 8), 1 F; same data,

16 June 1964 (NI 12), 1 pM (12-104) [UCLA]. Zelaya: Bluefields, W. Thornton, 1 F [USNM].

El Aserrio (nr. Bluefields), 14 July 1964, A. Quinonez (NI 47), 1 pM (47-100), 1 L [UCLA].

Locality unspecified: P. Woke, 1 M, 4 L [USNM].

PANAMA AND CANAL ZONE. Bocas del Toro: Almirante, 29 Apr 1963, A. Quinonez (PA

282), 1 M, 1 F [UCLA]. Almirante, mile 2, 16 Apr 1963, A. Quinonez (PA 254), 1 M; same data,

9 May 1963 (PA 342), 4 F [UCLA]. Nigua Creek, nr. Almirante, 4 May 1963, A. Quinonez (PA

312), 1 M gen [UCLA]. Canal Zone: Barro Colorado Island, 7 May 1943, W. Komp, 2 IF (43-80,

106) [UCLA, USNM]; same data, 15 May 1945, 2 IpM (5-129A,134), 2 IpF (5-132,133), 1 lp G-

138), 5 M, 2 F,11 [UCLA, USNM]; same data, 22 May 1945, 1 IF (S-206),4M,8F, 2p [UCLA,

USNM]; same data, 23 May 1945, 1 lpM (5-224), 1 lpF (5-175), 1 F, 2 1 [UCLA, USNM] ; same

data, 26 June 1945, 1 IM (5-426), 2 IF (5-418,430) [UCLA, USNM]; same data, 3 Dec 1965, A.

Quinonez (PA 856), 1 lpM (856-10), 2 p [UCLA] ; same data, 1 July 1967, W. Wirth, 1 M [USNM].

Camacho, 1 June 1922, J. B. Shropshire, 1 M, 4 F [USNM]. Chiva Chiva, 1945, 1 M [USNM];

same data, 12 Nov 1965, R. Schick and A. Quinonez (PA 777), 3 M, 3 p, 31 [UCLA]. Contractor’s

Hill (0.8 km N), 13 Dec 1965, R. Schick (PA 897), 1 pF (897-100) [UCLA]. Corozal, 29 Jan

1943, W. Komp, 1 1M (43-45), 3 M, 1 F; 4 1; same data, 6 May 1943, 1 M, 2 F; same data, July

1949, 1 M [UCLA]. Empire, 3 Apr 1922, J. Shropshire, 5 M [USNM, BM]; same data, 10 June

1922, 1 M [USNM]. Ft. Kobbe, 18 May 1950, S. Carpenter, 2 F [UCLA]. Ft. Sherman, 23 Sept

1949, 1 L [UCLA]; same data, 24 Feb 1950, 1 1 [UCLA]; same data, 6 May 1949, 5S. Carpenter,

2 M, 9 F [UCLA, Utah]. France Field, 7 June 1949, 1 L [UCLA]. Gamboa (6 km NW on Pipeline

Rd.), 30 m, 15 July 1971, H. Arnell (PA 1121), 2 M, 2 F [UCLA] . Madden Forest Preserve, Las

Cruces Trail, 100 m, 24 June 1972, H. Arnell (PA 1061), 1 L [UCLA]. Madden Forest Preserve,

0.5 km W George Green Park, 100 m, 10 July 1972, H. Arnell (PA 1096), 1 IpM (1096-10)

[UCLA]. Miraflores, 9 May 1908, A. Jennings, 2 M, 1 F; same data, 20 May 1922, J. Shropshire,

1 F [USNM]. Red Tank, 14 Nov 1949, S. Carpenter, 1 M [UCLA]. Summit, 26 May 1953, 2M,

1 F; same data, 17 July 1938, 3 M [UCLA]. Summit, W. Komp, 1 M [UCLA]; same data,

25 May 1945, 2 lpM (5-218,238), 2 Ip (5-228,235), 3 M, 1 M gen, 1 F, 11 [UCLA, USNM].

Tabernilla, A. Busck, 1 F [USNM]. Chiriqui: Potrerillos, 12 Nov 1949, P. Galindo, 1 1 [UCLA].

Cocle: El Valle, 5 June 1945, R. Arnett and W. Komp (ASM 611), 4 M [UCLA]; same data,

W. Komp, 1 IM (5-304), 1 Ip (5-350), 5 M, 3 F, 1 1 [UCLA, USNM]; same data, Nov 1946,

N. Krauss, 1 F [USNM]. Colon: Portobelo, Mar 1911, A. Busck, 3 M, 2 F [USNM]. Portobelo,

Caldera Island, A. Jennings, 1 M [USNM]. Darien: Rio Tuira, 3 Mar 1958 (GG 69), 8 M [UCLA].

Rio Tuira at mouth of Rio Paya, 2 Mar 1958 (GG 59), 6 M [UCLA] . Panama: Archipielago de las

Perlas, Isla San Jose, 26 July 1944, W. Komp, 1 IF (4-54) [UCLA, USNM]. Cerro Campana,

750 m, 6 Dec 1950, 1 L; same data, 28 Aug 1963, A. Quinonez (PA 537), 4 M; same data

(PA 538), 1 F; same data, 29 Aug 1963 (PA 539), 5 M; same data (PA 540), 1 F [UCLA].

Cerro La Victoria, 11 May 1949, P. Galindo, 4 1 [UCLA]; same data, 1949, H. Trapido, 10 F

[UCLA, USNM]; ?same data, GML Lab Colony (PA 21), 523 M, 16 M gen, 424 F, 1 F gen,

746 P, 679 L [UCLA]. El Victoria (?La Victoria), 29 June-7 July 1949, 4 M [UCLA]. La Jolla,

nr. Pacora, 6 Mar 1951, 1 F [UCLA]. La Zumbadora (nr. Cerro Azul), 600 m, 15 Feb 1963, A.

Quinonez (PA 94), 1 lpM (94-102) [UCLA]. Nuevo Emperador, 23 Nov 1965, A. Quinonez

(PA 832), 1 lpM (832-20), 1 IlpF (832-11) [UCLA]. Pacora, 29 June 1949, 1 M gen [UCLA].

Panama, Paitilla Pt., 14 July, 2 M; same data, 14 May 1949, W. Komp, 3 M [UCLA]. Tocumen,

1 June 1949, S. Carpenter, 2 F [UCLA].

TRINIDAD AND TOBAGO. TOBAGO: St. Andrew: Government Stock Farm, Scarborough,

75 m, 15 Nov 1965, T. Aitken et al. (TOB 9), 2 lpM (9-20,21), 1 pF (9-100), 3 L [UCLA].

Orange Hill, 90 m, 24 Nov 1965, R. Martinez and A. Guerra (TOB 99), 1 pM (99-103); same data,

30 Nov 1965 (TOB 138), 1 IpF (138-21), 1 L; same data (TOB 139), 1 lpF (139-30), 1 L [UCLA].

Scarborough, 85 m, 21 Nov 1965, R. Martinez and A. Guerra (TOB 73), 6 L [UCLA]. St. George:

Caledonia (5 km E Mason Hall) 300 m, 17 Nov 1965, T. Aitken et al. (TOB 49), 1 F [UCLA].

St. John: Little Tobago Island, 1 Aug 1963, T. Aitken, 2 F [UCLA]. St. Patrick: Carnbee, 75 m,

Arnell: Genus Haemagogus 67

19 Nov 1965, R. Martinez and A. Guerra (TOB 61), 1 lpM (61-10), 1 pF (61-100), 4 L; same data

(TOB 62), 1 F [UCLA]. Mt. Irvine Bay, 20 Nov 1965, R. Martinez and A. Guerra (TOB 70), 7 M,

3 F, 10 p [UCLA]. Locality unspecified: July 1905, A. Busck, 1 M [USNM]. TRINIDAD: St.

George: Monos Island, Grand Fond Bay Valley, 0-60 m, 3 Aug 1964, R. Manuel (TR 596), 6 F

[UCLA]. . ,

UNITED STATES. Texas: Brownsville, 6 Sept 1955 (GML), 2 lpF (02136,02138), 2 pF

(02134,02135), 1 M, 1 p [UCLA, USNM]; same data, 12 June 1955, K. Kajihiro, 1 P [UCLA];

same data, 13 Apr 1957, Reimann, 1 p [UCLA].

VENEZUELA. Aragua: Cata, 0 m, 21 Aug 1969, J. Valencia and Clarijo (VZ 388), 1 pF (388-

100), 1 M, 8 F, 5 p [UCLA]. Choroni (8.7 km S), 300 m, 16 July 1969, T. Zavortink et al.

(VZ 228), 1 IpF (228-10), 1 pM (228-102) [UCLA]. Maracay, 600 m, 3 Sept-26 Oct 1926,

M. Nunez-Tovar, 9 F [USNM, BM]. Ocumare de la Costa, 100 m, 5 Aug 1969, J. Valencia

(VZ 306), 2 IpM (306-10,11), 3 pM (306-100-102), 1 1; same data (VZ 308), 1 pF (308-103)

[UCLA]. Turiamo, 14 Sept 1944, 1 M [UCLA]. Carabobo: San Esteban, Aug 1940, P. Anduze,

1 M [USNM]. Sucre: Isla Patos, 5 Aug 1962, R. Manuel, 1 F [UCLA].

Additional Records From the Literature

VENEZUELA. Bolivar: Piar and Roscio districts (Anduze, 1947:353). Tachira: Jauregui district

(Anduze, 1947:353). Trujillo: Betijogue district (Anduze: 1947:353).

TROPICALIS SECTION

FEMALES. Head: Eyes narrowly separated above antennae (1 ommatidial dia-

meter). Proboscis 1.10-1.15 of forefemur. Thorax: Lower stp bristles weakly devel-

oped. Legs: Length of forefemur about 1.50 of distance from top of thorax to

apex of midcoxa. Knee spots absent. Claws of foreleg and midleg with subbasal

tooth. Wing: Ratio of R,,3; to R, varied, R, 43 about 0.40-1.10 of R,. Postnotum:

Bristles absent.

FEMALE GENITALIA. Tergite IX: Setae apparently absent.

MALES. Head: Proboscis about 1.20 of forefemur. Palpus long. Antenna 0.60

of proboscis, torus much enlarged, flagellum densely plumose, segments 1-12 with

very numerous moderate to long bristles. Legs: Claws of foreleg enlarged, unequal;

claws of midleg subequal, small.

MALE GENITALIA. Segment IX: Lobes usually with 1 or 2 setae. Sidepiece:

Apical lobe absent. Claspette: Filament modified. Clasper: Simple, cylindrical,

curved inward at apex; seta present near apex of inner surface; spiniform apical,

0.40 of clasper.

LARVAE. Head: Hair 5-C single. Hair 6-C single (single or double). Thorax:

Integument without spicules. Tubercles of hairs 5-6-P joined and hair 7-P free.

Abdomen: Hair 12-1 present. Segment VIII: Comb scales 24-28, with single broadly

rounded, minutely fringed spine, in irregular double to triple row. Anal Segment:

Spines of caudal margin of saddle reduced. Ventral brush (4-X) with 6 pair of hairs

arising from weakly sclerotized boss.

DISCUSSION. In the adults, Haemagogus tropicalis exhibits many of the dis-

tinguishing characters of the Albomaculatus Section, toothed claws in the female,

the postnotum without setae, closely approximate eyes, the male palpus long as

in 4 species of the Albomaculatus Section, while the larva is more typical of the

larvae of the Splendens Section having 6 pair of hairs in the ventral brush and 24

to 28 comb scales in a triangular patch. The female genitalia apparently lack setae

on tergite IX, a Splendens Section character, while the male genitalia exhibit char-

acters of both sections. 7ropicalis is a relict species, having been found only at a

68 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

single locality at the mouth of the Amazon. Since it is annectent between the

Albomaculatus and Splendens Sections, I have placed it in the monotypic Tropic-

alis Section.

15. Haemagogus (H.) tropicalis Cerqueira & Antunes

Figs. 5,36,37

1938. Haemagogus tropicalis Cerqueira and Antunes, 1938:1-9. TYPE: Holotype male, Curra-

linho, Para, Brazil, Jan-May 1936, H.W. Kumm and A. Rabello [LU].

Haemagogus (Longipalpifer) tropicalis of Levi-Castillo (1951b:12,35-37); Stone, Knight and

Starcke (1959:216); Forattini (1965:53).

Haemagogus (Cyanoconops) tropicalis of Lane (1939:121; 1953:807-810).

Haemagogus tropicalis of Kumm and Novis (1938:501); Kumm and Cerqueira (1951a:174).

FEMALE. Wing: 3.25 mm. Proboscis: 2.70 mm. Forefemur: 2.45 mm. Abdo-

men: 3.15 mm. Dark scales of proboscis, palpus, wing and legs purple to violet.

Head: Eyes narrowly separated above antennae (1 ommatidial diameter). Decum-

bent scales of vertex and occiput greenish blue anteriorly becoming light green

posteriorly and laterally, with some purple reflections. Proboscis about 1.10-1.15

of forefemur; palpus about 0.13 of proboscis; antenna about 0.65 of proboscis.

Thorax: Scales of mesonotum usually greenish bronze, becoming blue laterally and

posteriorly, occasionally entire mesonotum blue, scales silver in antealar area and

usually deep blue to purple over supraalar bristles and on scutellar lobes. Apn

lobes moderately enlarged, scales blue to greenish blue; ppn apparently bare; lower

stp without well developed bristle. Legs: Long, length of forefemur about 1.50

of distance from top of thorax to apex of midcoxa. Wing: Ratio of R243 to R,

variable, R343 about 0.40-1.10 of R,. Abdomen: Dark scales of tergites greenish

blue to violet, with some purple reflections, often with median basal patches of

silver scales on segments III-VII; sternites blue to purple..

MALE (fig: 36). Wing: 1.95 mm. Proboscis: 1.85 mm. Forefemur: 1.60 mm.

Abdomen: 2.30 mm. Head: Proboscis about 1.20 of forefemur; palpus long, slender,

straight throughout, about 0.55 of proboscis, 5-segmented, segments 2 and 3

ankylosed and long, making up about 0.62 of palpus length; segment 4 making up

about 0.20 of palpus length; segment 5 making up about 0.12 of palpus length;

antenna about 0.60 of proboscis. Legs: Claws of foreleg unequal, larger claw with

submedian tooth, smaller claw with acute subbasal tooth; claws of midleg sub-

equal, small, with acute subbasal tooth.

MALE GENITALIA (fig. 36). Segment VIII: Tergite about 0.80 of sternite,

distal margin straight, with enlarged setae laterally and several large spiniform

setae mesally. Segment IX: Tergite broadly emarginate and weakly sclerotized

mesally; lobes usually with 1 or 2 poorly developed setae. Sidepiece: Conical, length

about 2.5 times median width; basal tergomesal lobe poorly developed but extend-

ing distally as slightly raised area to slightly distad of middle of sidepiece, with short

setae arising from raised tubercles distally and enlarged setae proximally; apical

lobe poorly developed, represented by small cluster of moderately developed setae

dorsad of apical sternomesal scales on distal fourth; apical sternomesal scales

lanceolate to oblanceolate with acuminate tips. Claspette: Stem bowed inward.

near middle in dorsal aspect, slightly narrowed near middle, sharply angled dorsad

near distal third; filament broad, leaflike, attached to stem near middle, ventral

Arnell: Genus Haemagogus 69

half erect, rugose, divided into 2 irregular lobes, dorsal half triangular, apex form-

ing a slightly recurved beak. Clasper: Broadest near base, spiniform about 0.40 of

clasper. Phallosome: Aedeagus large, broadly obovate, tip produced dorsad into

sclerotized, beaklike process; venter broadly open except on basal half; with heav-

ily sclerotized ridges lateroventrally which are weaker basally. Proctiger: Cercal

setae 6-8; apical knob of paraproct with 15-20 serrations.

PUPA. Unknown.

LARVA (fig. 37). Head: 0.80 mm. Siphon: 0.65 mm. Anal Saddle: 0.30 mm.

Head: Hair 5-C single. Hair 6-C single (single or double). Thorax: Integument with-

out spicules. Tubercles of hairs 5,6-P connected, hair 7-P free. Abdomen: Hair 1 2-I

present. Segment VIII: Comb scales 24-28, with single broadly rounded, minutely

fringed spine, in irregular double to triple row. Siphon: Index about 2.5. Pecten

teeth 10-17, extending to about basal 0.40-0.45 of siphon. Hair 1-S triple. Anal

Segment: Spines on caudal margin of saddle much reduced, very short, each with

multiple teeth. Hair 4a-X 5b, long. Boss weakly sclerotized.

SYSTEMATICS. Haemagogus tropicalis can be distinguished from the remaining

members of the genus: in the adults by the combination of (1) subbasal tooth

on claws of female foreleg and midleg and small and subequal claws of male mid-

leg, (2) ppn apparently without scales and (3) long male palpus, 0.62 of proboscis,

and (4) lower stp without well developed bristle; in the male genitalia by claspette

filament with ventral half erect, rugose and bilobed; and in the larva by the com-

bination of (1) 24 to 28 comb scales in a triangular patch, (2) weakly sclerotized

boss and (3) hair 6-C usually single.

The affinities of tropicalis are discussed above under the Tropicalis Section.

BIONOMICS. Haemagogus tropicalis larvae have been taken from treeholes. The

adults are reported by Kumm and Cerqueira (1951:174) not to normally attack

man.

DISTRIBUTION (fig. 5). Haemagogus tropicalis is known only from one local-

ity on the Isla do Marajo at the mouth of the Amazon River. Material examined:

20 specimens; 9 males, 9 females, 2 larvae.

BRAZIL. Para: Curralinho, 2 M, 2 M gen, 3 F,2 L [USNM, BH, GML]; same data, Dec 1935,

3 F [USNM]; same data, 1936, 2M [USNM]. Curralinho, Rio Cupijo, 11 Apr 1936, A. Azeredo,

2 M, 2 F [USNM, GML]. Curralinho, Rio Pagao, 16 Apr 1936, A. Azeredo, 1 M, 1 F [USNM].

SPLENDENS SECTION

FEMALES. Head: Eyes moderately to broadly separated above antennae (2-5

ommatidial diameters). Proboscis 0.80-1.20 of forefemur. Thorax: Ppn with or

without scales; lower stp bristles weakly developed. Legs: Midcoxa with single

well developed bristle laterally; forefemur about 1.40-1.80 of distance from top

of thorax to apex of midcoxa. Knee spots absent. Claws of foreleg and midleg

simple. Wing: R,,3 about 0.32-0.50 of R,. Postnotum: Two small bristles present

posteriomesally.

FEMALE GENITALIA. Tergite IX: Setae absent.

MALES. Head: Proboscis about 0.95-1.40 of forefemur. Palpus short, 0.16-0.21

of proboscis; 4-segmented with segment 4 small to minute. Antenna 0.44-0.59 of

proboscis with torus moderately enlarged, flagellum sparsely to moderately plumose,

segments 1-12 with 6 to about 22 moderately long bristles. Legs: Claws of foreleg

and midleg enlarged, unequal; larger claw of foreleg with large acute subbasal or

70 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

blunt median tooth, smaller claw simple; larger claw of midleg simple or with

large, acute subbasal tooth, smaller claw simple.

MALE GENITALIA. Segment IX: Tergite without setae. Sidepiece: Apical lobe

usually present. Claspette: Filament highly modified. Clasper: Usually modified;

spiniform short, 0.18-0.32 of clasper.

LARVAE. Head: Hair 5-C single. Hair 6-C double. Thorax: Integument without

spicules. Tubercles of hairs 5-6-P joined, hair 7-P free. Abdomen: Hair 12-I present.

Segment VIII: Comb scales 10-50(6-75), with single broadly rounded to spatulate,

minutely fringed spine, in irregular double row to large triangular patch. Anal

Segment: Spines on caudal margin of saddle relatively weakly developed, short,

usually with multiple teeth. Ventral brush (4-X) with 6 pair of hairs, all hairs

but proximal | or 2 arising from strongly sclerotized boss. Hair 4a-X 3-8b, long.

DISCUSSION. The Splendens Section can be distinguished from the remainder

of the subgenus Haemagogus: in the adults by (1) the simple female claws, (2)

the postnotum with 2 small setae posteriorly and (3) vein R243 less than 0.50

of R,; in the male genitalia by (1) tergite IX without setae and (2) either a well

developed apical lobe on the sidepiece or the apical lobe represented by a dense

cluster of setae; and in the larvae by the combination of (1) the comb scales

numerous and in a triangular patch, (2) 6 pair of hairs in the ventral brush, (3)

a strongly sclerotized boss and (4) hair 6-C single.

The Splendens Section appears to be the most highly derived in the adults while

its larvae have retained many primitive characters. The setae on the adult post-

notum, the simple female claws, the short male palpi, the reduction of bristles

on the male antennae, and the unusual development of many structures of the

male genitalia are among the more conspicuous departures from the more primitive

aedine type and must be considered to be derived characters. In the larvae, however,

the greater development of the ventral brush and the large number of comb scales

appear to be primitive.

Five phyletic lines are recognizable within the Splendens Section. The species

of the splendens complex, splendens and celeste, differ from each other primarily

in the plumosity of the male antenna, with relatively minor differences in the

adults and larvae. This complex is probably closest to the ancestral stock which

gave rise to the Splendens Section. The 2 species have a simple clasper and relatively

simple sidepiece in the male genitalia and a large patch of scales on the adult ppn,

characters typical of the Albomaculatus and Tropicalis Sections. The splendens

complex is found on the periphery of the range of the Splendens Section, in

northern South America and the adjacent Windward Islands.

The regalis complex, the second phyletic line, contains the closely allied species

iridicolor, regalis, aeritinctus and lucifer. The adults of this complex can be sepa-

rated from each other primarily on scale color. The male genitalia differ only in

details of the clasper and claspette, and the larvae are virtually indistinguishable.

These species probably arose in Central America at a time when this area was under-

going major geological changes. This is discussed further under iridicolor. Argyro-

meris shows affinities to the regalis complex in some characters of the male genitalia

and in the larva but is not placed in the complex primarily because of differences

in the clasper and claspette. It is probably an early offshoot of the line that gave

rise to the regalis complex.

The unrelated chalcospilans and boshelli exhibit some extreme derivations, espe-

cially in the male genitalia, but show primitive characters in the stronger ventral

brush and large number of comb scales in the larvae.

Arnell: Genus Haemagogus 71

16. Haemagogus (H.) splendens Williston

Figs. 6,38,39,40

1896. Haemagogus splendens Williston, 1896:271-272. TYPE: Lectotype female, St. Vincent,

elev. 1000 ft, H.H. Smith [BM; selection of Belkin, 1968b:22].

Haemagogus (Haemagogus) splendens of Bonne and Bonne-Wepster (1925:435); Dyar (1928:

139-140); Edwards (1932:179, in part); Lane (1939:120, in part; 1953:779-781, in part);

Levi-Castillo (1951b:11,16-17, in part); Stone, Knight and Starcke (1959:218, in part);

Forattini (1965:47-48, in part).

Haemagogus splendens of Theobald (1903b:283), Dyar and Knab (1906b:166); Howard, Dyar

and Knab (1917:867, in part); MacDonald (1917:262); Dyar (1921a:114); Komp (1954:49-

51, in part).

Haemagogus cyaneus in part of Theobald (1901:239-241; 1903a:308; 1905:37; 1907:550; 1910:

493); Giles (1902:485); Blanchard (1905:412); Coquillett (1906:25).

FEMALE (fig. 38). Wing: 2.75 mm. Proboscis: 2.35 mm. Forefemur: 2.00 mm.

Abdomen: 2.80 mm. Dark scales of proboscis, palpus, wing and legs violet to

purple. Head: Eyes widely separated above antennae (4 ommatidial diameters).

Decumbent scales of vertex and occiput copper to light green. Proboscis about

1.10-1.20 of forefemur; palpus about 0.15 of proboscis; antenna about 0.55-0.60

of proboscis. Thorax: Scales of mesonotum usually copper, occasionally bronze;

bluish green over supraalar bristles, silver in antealar area; scales on scutellum

bluish green. Apn lobes with scales greenish blue to copper, often with silver scales

dorsally and mesally; ppn with large patch of silver or copper scales posteriorly.

Legs: Hindcoxa enlarged, its base slightly above upper margin of meron; length of

forefemur about 1.50-1.65 of distance from top of thorax to apex of midcoxa.

Wing: R,,3 about 0.35-0.45 of R,. Abdomen: Dark scales purple to violet; usually

with silver scales dorsally on tergites II-VII in rather broad basal band, occasionally

continuous with lateral silver patches.

MALE (fig. 38). Wing: 2.35 mm. Proboscis: 2.40 mm. Forefemur: 2.00 mm.

Abdomen: 2.55 mm. Head: Proboscis about 1.20-1.37 of forefemur; antenna

sparsely plumose, flagellar segment 4 with about 8-10 moderately developed bristles.

MALE GENITALIA (fig. 39). Segment VIII: Tergite about 0.85-0.95 of sternite,

distal margin slightly rounded, with enlarged setae and with a few large narrowly

lanceolate scales mesally. Segment IX: Tergite broadly emarginate and weakly

sclerotized mesally. Sidepiece: Conical, broad, length about 2.5 times median

width; basal tergomesal area not enlarged but extending distad as raised area with

numerous short setae to near distal third of sidepiece, basally with numerous

setae, the more proximal ones much enlarged, flattened and attenuate; apical lobe

undeveloped but represented by cluster of relatively long, curved setae; apical

sternomesal scales lanceolate to oblanceolate. Claspette: Stem bowed inward slightly

at distal third in dorsal aspect, stout, angled sharply dorsad at distal third, with

seta and spicules at angle; filament striate, broadly sickle-shaped and terminating

in slightly recurved tip dorsally, ventral portion elongate, narrow, curved away

from stem, tip bifurcate or trifurcate, pointed. Clasper: Simple, cylindrical, thickest

at base, curved sharply inward at apical third; spiniform inserted apically, about

0.33 of clasper, with bifid tip. Phallosome: Aedeagus large, more or less broadly

obovate; tip produced dorsad into rather heavily sclerotized carina terminating in

beaklike process; venter broadly open except on basal third, with sclerotized ridges

72 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

lateroventrally which are expanded basally. Proctiger: Cercal setae 3-5; apical knob

of paraproct with about 10-15 lateral serrations.

PUPA (fig. 39). Abdomen: about 3.15 mm. Trumpet: 0.40 mm. Paddle: 0.75

mm. Cephalothorax: Lightly to moderately pigmented, darker dorsally. Hairs 4,5-C

weakly developed, subequal, single to triple, weaker than 7-C which is single.

Trumpet: Light brown, slightly lighter distally. Abdomen: Lightly to moderately

pigmented, genital lobe and anterior segments somewhat darker. Float hair (1-I)

with about 8-12 primary branches and 2-6 secondary branches. Hair .1-II-VIl

weakly to rather strongly developed, single, multiple or dendritic, more or less sub-

equal on all segments. Hair 2-VII short, considerably cephalad of 1-VII. Hairs

3-ILIII,5-IV,V somewhat variable in length but more or less subequal. Hair 6-VII

usually moderately well developed, somewhat stronger than 6-III-VI which are

subequal. Hair 9-VII considerably cephalad of caudolateral margin of tergite, 1-4b;

9-VIII 4-8b. Terminal Segments: Male genital lobe about 1.0 of tergite VIII.

Paddle: Weakly pigmented, midrib slightly darker; apex broadly rounded or slightly

acuminate. Hair 1-P single.

LARVA (fig. 40). Head: 0.85 mm. Siphon: 0.77 mm. Anal Saddle: 0.34 mm.

Segment VIII: Comb scales 10-13(6-20), with single broadly rounded, minutely

fringed spine, in irregular double row. Siphon: Index about 2.5(2.2-2.7). Pecten

teeth 12-16(10-21), extending to about basal 0.40-0.45 of siphon. Hair 1-S 4b.

Anal Segment: Hair 4a-X 3-4b.

SYSTEMATICS. Haemagogus splendens can be separated from the remaining

species of the Splendens Section: in the adults by (1) large patch of silver or copper

scales on ppn, (2) broad basal bands of silver scales dorsally on abdominal tergites

II-VII, (3) scales of vertex copper to light green and (4) male antenna sparsely

plumose; in the male genitalia, from all species except celeste by (1) simple, cylin-

drical clasper with apical spiniform, (2) characteristic claspette filament with ventral

half narrow, pointed and curved away from stem, and (3) all apical sternomesal

scales lanceolate to oblanceolate; and in the larva by the combination of (1) few

comb scales, usually 10-13, (2) hair 7-III-VI subequal to hair 5 of corresponding

segment, (3) hair 4a-X with 3 or 4 branches and (4) pecten extending only to

basal 0.40-0.45 of siphon.

Closely allied to the mainland celeste, the other member of its complex, splen-

dens is indistinguishable in the male genitalia, but can be separated with reasonable

certainty in all other stages, in the female by copper to light green scales on the

vertex and usually copper scales on the ppn; in the male by the very sparsely

plumose antenna with only 8 to 10 bristles on each segment; in the pupa by sub-

equal hairs 3-II,III,5-IV,V; and in the larva by usually 10 to 13 comb scales and

hair 4a-X with 3 or 4 branches. The conspicuous difference in the structure of the

male antenna possibly indicates a difference in mating behavior.

The splendens complex is the most generalized component of the Splendens

Section sharing several characters with members of the Tropicalis and Albomacu-

latus Sections such as scales on the ppn, possibly the more densely plumose male

antenna in celeste, a simple clasper and poorly developed apical lobe on the side-

piece in the male genitalia, and fewer comb scales in the larva.

BIONOMICS. Haemagogus splendens breeds primarily in treeholes but has taken

from other plant containers such as cut bamboo internodes and bromeliads. Females

bite man readily. It has not been incriminated in disease transmission.

DISTRIBUTION (fig. 6). Haemagogus splendens is known from the Windward

Islands: Martinique, St. Vincent, the Grenadines and Grenada. Material examined:

Arnell: Genus Haemagogus 73

210 specimens; 50 males, 70 females, 48 pupae, 42 larvae; 48 individual rearings

(26 larval, 18 pupal, 4 incomplete).

GRENADA. St. Andrew: Belair School, 8 July 1929 (GRR 21), 1 F [UCLA]. Lower Mt. St.

Catherine, 600 m, 25 Oct 1963, R. Martinez (GR 89), 1 F [UCLA]. St. George: Anandale Water-

fall, 210 m, 28 Oct 1963, R. Martinez (GR 95), 2 IpF (95-101,110), 1 IP (95-104), 1 F [UCLA].

Richmond Hill, 25 July 1942, E. Cochrane, 4 L [USNM]. Tempe, 180 m, 9 Oct 1963, R. Martinez

(GR 14), 3 L [UCLA]. Woborn, 15 m, 10 Oct 1963, R. Martinez (GR 19), 1 IpF (19-111), 2 L

[UCLA]. St. Patrick: River Antoine Estate, 60 m, 29 Oct 1963, R. Martinez (GR 99), 2 IpF (99-

109,113), 2 L [UCLA]. Locality unspecified: 10 July 1929 (GRR 31), 10 F; same data (GRR 36),

7 M; same data, 17 July 1929 (GRR 58), 1 M; same data (GRR 58A), 1 F; same data, 22 July

1929 (GRR 66), 1 M; same data, 23 July 1929 (GRR 73,73A), 3 M, 1 F [UCLA]; same data,

10 July 1929, F. Root, 2M, 2 F [BM]. ia th ies |

THE GRENADINES. Bequia Island, 16 Aug 1929, W. Hoffman (LAR 33B), 4 F [UCLA].

Union Island, 20-29 Aug 1929, W. Hoffman (LAR 33F), 6 F; same data, 20 Aug 1929, W.

Hoffman (LAR 49), 2 M [UCLA]. Mayreau Island, 20 Aug 1929, W. Hoffman (LAR 38), 2 M

[UCLA]. Carriacou Island, 21 Aug 1929, W. Hoffman (LAR 36A), 1 F; 21 Aug 1929, W. Hoffman

(LAR 67), 3 M, 1 F; 1 Aug 1929 (GRR 102), 3 M, 3 F [UCLA].

MARTINIQUE. Fort de France (18-22 km N), 8-9 Aug 1929, W. Hoffman (LAR 284A), 1 F

[UCLA].

ST. VINCENT. Charlotte: Byera, 120 m, 25 Nov 1963, A. Guerra (VT 24), 1 IpF (24-102), 1 pF

(24-103), 1 lp (24-101) [UCLA]. Chapmans, 120 m, 5 Dec 1963, A. Guerra (VT 53), 1 IpF (53-

105), 1 pF (53-103) [UCLA]. Diamond, 60 m, 7 Dec 1963, A. Guerra (VT 58), 1 IpM (58-104), 1

IpF (58-105), 2 pM (58-101,102), 1 pF (58-103) [UCLA]. Mt. William, 120 m, 14 Dec 1963, A.

Guerra (VT 88) 1 pM (88-103) [UCLA]. Three Rivers, 300 m, 20 Nov 1963, A. Guerra (VT-8), 1

IpM (8-101) [UCLA]. St. Andrew: Lowmans, 180 m, 5 Jan 1944, W. Leslie Webb, 1 M [USNM];

same data, 11 Dec 1963, A. Guerra (VT 77), 2 lpM (77-104,113), 4 lpF (77-105,106,110,112),

1 pM (77-103), 4 pF (77-101,102,108,109), 1 L [UCLA]; same data (VT 78), 1 lpF (78-105),

1 pM (78-103), 3 pF (78-101,102,104) [UCLA]. St. David: Chateaubelair (1.5 km NE), 30 m,

19 Nov 1963, A. Guerra (VT 1), 1 IpM (1-101), 1 IP (1-102); same data (VT 5), 1 IP (5-101)

[UCLA]. Crater Lake, 850 m, 9 Dec 1963, A. Guerra (VT 63), 2 IpF (63-101,105), 1 pF (63-103)

[UCLA]. St. George: Kings Hill, 120 m, 12 Dec 1963, A. Guerra (VT 80), 1 IpM (80-104),

1 pM (80-102), 2 pF (80-101,103) 3 F; same data (VT 81), 4 IpF (81-104-106,108), 2 pM

(101,102), 8 M [UCLA]. St. Patrick: Layou, 120 m, 4 Dec 1963, A. Guerra (VT 50), 1 F

(50-101). Wallilabou, 120 m, 27 Nov 1963, A. Guerra (VT 30), 1 IpM (30-101) [UCLA]. Locality

unspecified: 6 Oct 1899, H. Powell, 2 F [BM]; 300 m, H. Smith, 3 F (holotype) [BM]; Low, 1 F

[BM].

17. Haemagogus (H.) celeste Dyar & Nunez Tovar

Figs. 6,41,42

1927. Haemagogus celeste Dyar and Nunez Tovar, 1927:152. TYPE: Lectotype male (2270,

“No. 3”), with pupal skin (2270) [uncertain association] and genitalia slide (2270),

Maracay, Aragua, Venezuela, 11 Nov 1926, M. Nunez Tovar [USNM; selection of

Stone and Knight, 1955:287] . Synonymized with splendens by Edwards (1932:179).

Haemagogus (Haemagogus) celeste of Dyar (1928:141-142); Anduze (1941a:829).

Haemagogus (Haemagogus) splendens in part of Lane (1939:120; 1953:779-781); Anduze (1941b:

13; 1947:354); Levi-Castillo (1951b:11,16-17); Stone, Knight and Starcke (1959:218); Fora-

ttini (1965:47-48); Cova Garcia, Sutil and Rausseo (1966a:60-61, fig. 120; 1966b:46,112-114,

fig. 199); Aitken, Worth and Tikasingh (1968:257).

Haemagogus splendens in part of Anderson and Osorno-Mesa (1946:613); Hovanitz (1946:35);

Kumm, Osorno-Mesa and Boshell-Manrique (1946:20); Osorno-Mesa (1947:453); Waddell

74 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

(1949:568,572); Levi-Castillo (1951:14); Komp (1954a:50; 1955e:277); Barreto Reyes (1955:

78).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870).

Haemagogus regalis in part of Dyar and Knab (1906b:167).

FEMALE. Wing: 3.00 mm. Proboscis: 2.45 mm. Forefemur: 2.20 mm. Abdomen:

about 3.05 mm. Dark scales of proboscis, palpus, wing and legs violet to purple.

Head: Eyes widely separated above antennae (5 ommatidial diameters). Decumbent

scales of vertex blue to violet, becoming green on occiput. Proboscis about 1.15-

1.20 of forefemur; palpus about 0.14 of proboscis; antenna about 0.52-0.58 of

proboscis. Thorax: Scales of mesonotum copper to bronze, bluish green to greenish

blue over supraalar bristles, silver in antealar area; scales on scutellum usually greenish

blue, rarely violet. Apn lobes with scales greenish blue to bluish green, often with

silver scales dorsally and mesally; ppn with large patch of scales posteriorly, usually

silver, rarely purple. Legs: Hindcoxa enlarged, its base slightly above upper margin

of meron; length of forefemur about 1.50-1.60 of distance from top of thorax to

apex of midcoxa. Wing: R,,3 about 0.35-0.45 of R,. Abdomen: Dark scales purple

to violet; usually with silver scales dorsally on tergites II-VII in rather broad basal

band, occasionally continuous with lateral silver patches.

MALE (fig. 41). Wing: 2.40 mm. Proboscis: 2.50 mm. Forefemur: 1.95 mm.

Abdomen: 2.55 mm. Head: Proboscis about 1.25-1.35 of forefemur; antenna mod-

erately plumose, flagellar segment 4 with about 18-22 moderately developed bris-

tles.

MALE GENITALIA (fig. 41). Apparently indistinguishable from splendens.

PUPA (fig. 41). Abdomen: about 3.75 mm. Trumpet: 0.50 mm. Paddle: 0.75

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

weakly to moderately developed, subequal, single to triple, weaker than 7-C which

is single. Trumpet: Medium brown, reticulate sculpturing moderate. Abdomen:

Weakly to moderately pigmented, genital lobe and anterior segments usually darker.

Float hair (1-1) with about 8-14 primary branches and 2-6 secondary branches. Hair

1-II-VII weakly to moderately developed, single to triple, subequal on all segments.

Hair 2-VII usually considerably basad of 1-VII. Hairs 3-II,III,5-IV,V stronger on II,

becoming slightly shorter on each succeeding segment. Hair 6-VII subequal to or

stronger than 6-III-VI which are subequal. Hair 9-VII slightly to considerably

cephalad of caudolateral margin of tergite, 1-4b; 9-VIII usually 5-8b. Terminal

Segments: Male genital lobe about 1.0 of tergite VIII. Paddle: Weakly pigmented,

midrib slightly darker; apex broadly rounded or very slightly acuminate. Hair 1-P

single.

LARVA (fig. 42). Head: 0.90 mm. Siphon: 0.80 mm. Anal Saddle: 0.37 mm.

Segment VIII: Comb scales 17-22(12-27), with single broadly rounded minutely

fringed spine, in irregular double to triple row. Siphon: Index about 2.4(2.2-2.6).

Pecten teeth 13-16(10-22), extending to about basal 0.50 of siphon. Hair 1-S triple

(2-4). Anal Segment: Hair 4a-X 4-5b.

SYSTEMATICS. Haemagogus celeste can be separated from the remaining species

of the Splendens Section: in the adults by (1) large patch of silver or occasionally

purple scales on ppn, (2) broad basal bands of silver scales on abdominal tergites

II-VII, (3) blue to violet scales on vertex and (4) moderately plumose male antenna;

in the male genitalia as in splendens; and in the larva by the combination of (1)

comb scales usually 17-22 and in double or triple row, (2) hair 7-III-VI subequal

to hair 5 of corresponding segment, (3) usually 4 or 5 branched hair 4a-X and

(4) pecten extending to about basal 0.50 of siphon.

Arnell: Genus Haemagogus 75

Though indistinguishable from the closely related splendens in the male genitalia,

celeste can be separated from splendens in the female by the blue to violet scales

on the vertex and usually silver ppn scales; in the male by the moderately plumose

antenna, each segment with about 18 to 22 bristles; in the pupa by hairs 3-II,III,

5-IV,V becoming weaker on each succeeding segment; and in the larva by usually

17 to 22 comb scales and the 4-or 5-branched hair 4a-X. The affinities of celeste

to other species of Haemagogus are discussed above under splendens.

BIONOMICS. Haemagogus celeste usually breeds in treeholes and cut or broken

bamboo internodes and in Venezuela it is often found in bromeliads. T.H.G. Aitken

reports (personal communication) celeste to be found almost exclusively in man-

grove treeholes on Trinidad, a somewhat different ecological situation than this spe-

cies inhabits on the mainland, and unusual in that the other mangrove inhabit-

ing species of the Splendens Section are seldom found outside that habitat. Females

apparently attack humans readily. This species (as splendens) has been shown to

be capable of transmitting yellow fever in the laboratory (Anderson and Osorno-

Mesa, 1946) although it is a relatively inefficient vector (Waddell, 1949). It is not

known to be a vector under natural conditions.

DISTRIBUTION (fig. 6). Haemagogus celeste is known from the lower Magdalena

and Zulia Rivers of Colombia, the Orinoco basin, the Caribbean drainage of

Venezuela and the islands of Trinidad and Tobago. Material examined: 951 speci-

mens; 163 males, 222 females, 276 pupae, 290 larvae; 257 individual rearings

(156 larval, 48 pupal, 53 incomplete).

COLOMBIA. Boyaca: Remolinos, 120 m, 2 July 1965, E. Osorno-Mesa et al. (COB 54), 3 lpF

(54-10-12), 1 P, 1 1 [UCLA]; same data (COB 55), 5 lpM (55-12-15,17), 3 IpF (55-10,11,16), 2 P,

7 L [UCLA]; same data (COB 56), 1 lpM (56-20), 2 IpF (56-21,22), 1 L [UCLA]. Magdalena:

El Retiro, 2 M gen [USNM]. Santander del Norte: Zulia, Feb 1944, H. Kumm, 3 M, 2 M gen, 2 F,

8 1 [UCLA, USNM].

TRINIDAD AND TOBAGO. TOBAGO: St. Patrick: Bon Accord Estate, 0 m, 15-16 Nov 1965,

T. Aitken et al. (TOB 19), 1 F [UCLA]. Pigeon Point, 2-3 Aug 1957, T. Aitken; 5 F [UCLA].

TRINIDAD: Caroni: Caroni Swamp, 18 Oct 1956, H. Trapido et al., 2 F [UCLA]. Mayaro:

Guayaguayare, 29 Oct 1964, A. Guerra (TR 790), 2 IpF (790-101,102) [UCLA]; same data

(TR 802), 3 F [UCLA]. Nariva: Bush Bush Forest, Nariva Swamp, 11 Mar 1964, TRVL (TR 171),

2 IpM (171-113,192), 1 lpF (171-114), 1 lp (171-121) [UCLA] ; same data, 27 May 1964 (TR 427),

2 IpM (427-138,149), 3 lpF (427-122,128,134), 1 F [UCLA]; same data (TR 432), 1 IpM (432-

146), 2 L [UCLA]; same data, 21 June 1965 (TR 1224), 1 IpM (1224-11), 1 lpF (1224-10)

[UCLA]; same data, 7 Sept 1961, T. Aitken (2-7/IX/61), 7 lpF (1,3-8), 1 lp (2) [UCLA]; same

data, 26 Sept 1961 (9-26/IX/61), 2 IpM (1,2), 3 IpF (3-5) [UCLA]; same data, 30 Jan 1963

(1-30/1/63), 1 IpF (1) [UCLA]; same data, 14 Feb 1963 (12-14/II/63), 2 lpM (1,2), 1 lpF (4),

1 IP (3), 1 M [UCLA]; same data, 5 Mar 1963 (14-5/III/63), 3 lpM (1-3) [UCLA]; same data,

1 Dec 1960, 2 P [UCLA]; same data, 1959-1963, 3 M, 2 F, 36 lp, 8 p, 6 L [UCLA]. Cocal,

beginning of boat line, 1 Sept 1960, T. Aitken (1/IX/60), 1 lpM (2) [UCLA]; same data, 1 Nov

1960 (1/XI/60), 1 lpM (2), 1 L [UCLA]. St. Andrew: North Manzanilla, 13 Sept 1957, T. Aitken,

!'7 M, 28 F [UCLA]. St. George: Chaguaramas, U.S. Naval Station, 23 Jan 1964, TRVL

(TR 48), 1 IP (48-126), 1 L [UCLA]; same data, 6 Dec 1960, T. Aitken, 1 lp [UCLA]; same

data, 2 Jan 1957, 1 M [UCLA]. Huevos Island,Tortue Bay, 75 m, 5 June 1965, R. Manuel

(TR 1202), 1 IP (1202-10) [UCLA]; same data, 6 June 1965 (TR 1203), 2 lpM (1203-15,16),

6 IpF (1203-11-14,17,18), 1 pM (1203-100), 4 L [UCLA]; same data (TR 1204), 8 lpM (1204-

11,13-15,17,19,20,22), 4 IpF (1204-10,16,18,21), 1 pM (1204-100), 1 pF (1204-101), 4 L

[UCLA]; same data (TR 1205), 7 lpF (1205-11-17), 5 pM (1205-100-104), 3 L [UCLA]; same

data (TR 1208), 6 lpM (1208-11-14,17,20), 9 IpF (1208-15,16,18,19,21-25), 1 pM (1208-100),

5 L [UCLA]; same data (TR 1209), 5 lpM (1209-12-16), 2 IpF (1209-17,18), 7 L [UCLA];

same data, 7 June 1965 (TR 1210), 2 IpM (1210-14,21), 7 lpF (1210-10-13,15-20), 4 pM (1210-

100-102,104), 1 pF (1210-103), 7 L [UCLA] ; same data, 5-7 June 1965 (TR 1211), 2 F [UCLA].

76 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Monos Island, 260 m, 17 May 1964, R. Manuel (TR 409), 1 F [UCLA]. Monos Island, Grand

Fond Bay Valley, 0 m, 2 Aug 1964, R. Manuel (TR 585), 1 IpF (585-122) [UCLA]; same

data (TR 586), 2 lpM (586-124,128), 8 IpF (586-118,119,121,125,126,136,139), 1 pF (586-

102), 2 IP (586-106,109) [UCLA] ; same data (TR 590), 2 IpF (590-128,131) [UCLA]; same data

(TR 591), 3 IpM (591-118,136,137), 5 IpF (591-124,127,131,132,134) [UCLA]; same data

(TR 596), 1 F [UCLA]. Locality unspecified: F. Urich, 1 M, 1 M gen, 1 F [USNM]; June 1905,

A. Busk, 1 F [USNM]; 28 June 1914, J. Dickson, 1 F [BM].

VENEZUELA. Anzoategui: Puerto La Cruz, 27 Sept 1944, 1 F [UCLA]. Aragua: Cagua,

400 m, 12 Aug 1969, J. Pulido and J. Valencia (VZ 339), 2 lpM (339-101,106), 2 lpF (339-100,

107), 1 IP (339-21), 1 1 [UCLA]; same data (VZ 340), 1 lpM (340-40), 2 lpF (340-10,11), 1 M

[UCLA]; same data (VZ 341), 1 lpM (341-40), 7 IpF (841-10-15,41), 2 pM (841-101,113), 6 pF

(341-90,100-102,105,112,114), 1 IP (341-62), 13 M, 6 F, 3 1 [UCLA]; same data (VZ 343),

1 IpF (343-10) [UCLA] . Guayabita, 14 July 1927, M. Nunez-Tovar, 3 M, 11 F [USNM]. Maracay,

600 m, 15 July 1969, J. Pulido and J. Valencia (VZ 204), 3 IpF (204-11,13,14), 2 pM (204-100,

101), 1 IP (204-16), 4 M, 3 F, 1 1 [UCLA]; same data (VZ 205), 1 IpF (205-11) [UCLA]; same

data (VZ 207), 1 lpF (207-20) [UCLA]; same data, 15 July 1969 (VZ 241), 1 IpF (241-20) 2 1

[UCLA]; same data, 20 Aug 1969 (VZ 381), 3 lpM (381-11,12,70), 1 IpF (381-13), 2 pM (381-

102,104), 1 pF (381-101), 3 lp (881-20,21) [UCLA]; same data, July 1929, F. Root, 5 M, 1 F

[BM]; same data, 24 Aug-15 Nov 1926, M. Nunez-Tovar, 12 F, 1 M gen [USNM]. Ocumare de la

Costa, 21 July 1927, M. Nunez-Tovar, 4 M, 5 F [USNM, BM]. Turmero, 560 m, 30 Aug 1966,

Vasquez (VZ 28), 1 pM (28-102), 2M, 2 P, 1 L [UCLA]; same data (VZ 29), 2 pM (29-108,109),

2 pF (28-106,107), 1 M, 1 F, 3 L [UCLA]; same data (VZ 30), 1 lpF (30-23), 5 pF (30-105,107-

109,111), 1 p, 1 L [UCLA]; same data, 9 Sept 1966 (VZ 37), 1 pM (37-27), 2 IP (37-25,29), 8 L

[UCLA]; same data (VZ 38), 1 pM (38-105), 2 pF (38-102,103), 1 P [UCLA]; same data, 14 Sept

1966 (VZ 41), 2 L [UCLA]; same data, 27 Oct 1966 (VZ 44), 1 lpF (44-13), 1 pM (44-12), 4 pF

(44-15,17-19) [UCLA] ; same data, 1 Nov 1966 (VZ 50), 1 p, 1 L [UCLA] ; same data(VZ 51),1P

[UCLA]; same data, 2 Nov 1966 (VZ 52), 1 pF (52-101), 1 IP (52-59) [UCLA]. Carabobo:

Carabobo, Oct 1941, P. Anduze, 1 lp [USNM]. Mariara, 400 m, 19 July 1969, J. Pulido and J.

Valencia (VZ 245), 1 lpM (245-10) [UCLA]. Guarico: Santa Maria de Ipire, Sept 1936, P. Anduze,

1 L [USNM]. Locality unspecified, July 1944, P. Anduze, 1 M gen [USNM]. Monagas: Maturin

(42 km SE), 28 June 1958, A. Menke, 2 F [LACM]. Quiriquire, 12 June 1935, W. Komp, 1 F

[UCLA]. Rancho Cachipo, Quiriquire, Sept 1936, W. Komp, 2 M gen [USNM]. Yaracuy: Boca de

Yaracuy, Jan 1938, P. Anduze, 1 M, 1 F [USNM]. San Felipe, May 1937, P. Anduze, 1 M

[USNM]. Zulia: Machiques, 1 June 1938, P. Anduze, 1 M gen [USNM]. State unknown: La

Trinidad, 14 July 1927, M. Nunez-Tovar, 5 M, 4 F [USNM]. Locality unspecified: July 1948,

P. Anduze, 1 M [USNM];P. Anduze, 2 M gen [USNM].

Additional Records From the Literature

COLOMBIA. Atlantico: Barranquilla (Kumm, Osorno-Mesa and Boshell-Manrique, 1946:20).

Arauca: Arauca (Kumm, Osorno-Mesa and Boshell-Manrique, 1946:20).

VENEZUELA. Bolivar: Heres, Roscio and Piar districts (Anduze, 1947:354).

18. Haemagogus (H.) iridicolor Dyar

Figs. 7,43,44

1921. Haemagogus (Haemagogus) iridicolor Dyar, 1921a:106-107. TYPE: Lectotype male

(1468) with genitalia slide, Higuito, nr. San Mateo, Alajuela, Costa Rica, P. Schild

[USNM; selection of Komp, 1955:29].

Haemagogus (Haemagogus) iridicolor of Bonne and Bonne-Wepster (1925:433); Dyar (1928:

136); Edwards (1932:179); Lane (1939:119); Stone, Knight and Starcke (1959:217); Forattini

(1965:39-41); Galindo and Trapido (1967:107).

Arnell: Genus Haemagogus VW

Haemagogus (Stegoconops) iridicolor of Lane (1953:789-790).

Haemagogus iridicolor of Kumm, Komp and Ruiz (1940:398); Arnett (1949:240); Galindo,

Carpenter and Trapido (1949:278); Horsfall (1955:535-536); Komp (1955a:30); Trapido,

Galindo and Carpenter (1955:529); Trapido and Galindo (1956a:304; 1957:126); Galindo

and Trapido (1955:548; 1957:147).

Haemagogus splendens in part of Howard, Dyar and Knab (1917:867).

FEMALE. Wing: 2.90 mm. Proboscis: 2.30 mm. Forefemur: 2.20 mm. Abdo-

men: 2.90 mm. Dark scales of proboscis, palpus, wing and legs purple and violet.

Head: Eyes widely separated above antennae (4 ommatidial diameters). Decumbent

scales of vertex and occiput blue to violet with some purple reflections. Proboscis

about 1.00-1.10 of forefemur; palpus about 0.16 of proboscis; antenna about 0.60-

0.66 of proboscis. Thorax: Scales of mesonotum copper to dark bronze, usually

dark green in fossa, greenish blue above supraalar bristles, silver in antealar area;

scales on scutellum greenish blue. Apn lobes with violet scales; ppn bare or with

small patch of silver scales posteriorly. Legs: Length of forefemur about 1.50-1.70

of distance from top of thorax to apex of midcoxa. Wing: R,,3; about 0.40-0.50

of R,. Abdomen: Dark scales violet or purple with some violet reflections; silver

scales present on tergites VI and VII in small median basal patch or indistinct basal

band.

MALE (fig. 43). Wing: 2.70 mm. Proboscis: 2.45 mm. Forefemur: 2.30 mm.

Abdomen: 3.65 mm. Head: Proboscis about 1.05-1.20 of forefemur; antenna short,

about 0.50 of proboscis, sparsely plumose, flagellar segment 4 with 10-14 moder-

ately developed bristles.

MALE GENITALIA (fig. 43). Segment VIII: Tergite about 0.85 of sternite,

distal margin broadly emarginate, with numerous well developed setae, setae longer

laterally. Segment IX: Tergite straight and weakly sclerotized mesally. Sidepiece:

Conical, truncate, length about 2.5 times median width; basal tergomesal lobe

enlarged, extending distally as raised area to near middle of sidepiece, with numer-

ous setae, the distal ones short, becoming longer proximally, the more proximal

ones enlarged, flattened and attenuate; a multiple row of short setae extending

obliquely from distal margin of basal lobe to near distal third of sidepiece; setae

absent between this row and apical lobe; apical lobe prominent, bearing about

20 moderately developed setae; apical sternomesal scales lanceolate to oblanceolate

with acuminate tips. Claspette: Stem bowed inward near middle in dorsal aspect,

thick basally, sharply angled dorsad near middle and narrowed on distal half;

filament a broad, cuneiform leaf attached at apex of stem, somewhat rounded at

dorsal angle. Clasper: Flattened, curved, expanded considerably on apical fourth

and tapering to rather sharp point; spiniform short, spatulate, inserted subapically

on inner surface. Phallosome: Aedeagus large, obovate, tip produced dorsally into

small serrated carina, venter broadly open except on basal third, with sclerotized

ridges lateroventrally. Proctiger: Cercal setae 3-5; apical knob of paraproct with

about 20 serrations.

PUPA (fig. 43). Abdomen: about 3.25 mm. Trumpet: 0.40 mm. Paddle: 0.70

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

rather weakly developed, subequal, single to triple, weaker than 7-C which is usu-

ally single. Trumpet: Medium to dark brown, lighter distally. Abdomen: Weakly

to moderately pigmented, genital lobe and anterior segments slightly darker. Float

hair (1-I) with about 8-14 primary branches and 2-6 secondary branches. Hair

1-II-VII rather weakly developed, usually single (1-4b) and subequal on all seg-

ments. Hair 2-VII short, considerably cephalad of 1-VIII. Hair 3-III slightly weaker

78 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

than 3-II,5-IV,V which are more or less subequal. Hair 6-VII weakly to moder-

ately developed, subequal to or considerably stronger than 6-HI-VI which are sub-

equal. Hair 9-VII usually considerably cephalad of caudolateral margin of tergite,

2-5b; 9-VIII 5-10b. Terminal Segments: Male genital lobe about 1.4 of tergite VIII.

Paddle: Weakly pigmented, midrib weakly pigmented; relatively broad; apex broadly

rounded. Hair 1-P single.

LARVA (fig. 44). Head: 0.75 mm. Siphon: 0.65 mm. Anal Saddle: 0.30 mm.

Segment VIII: Comb scales 25-28, with single broadly rounded, minutely fringed

spine, in irregular triple row. Siphon: Index about 2.5(2.3-2.7). Pecten teeth about

12-15(8-16), extending to about basal 0.50-0.60 of siphon. Hair 1-S 4-5b. Anal

Segment: Hair 4a-X 4-5b(4-7b).

SYSTEMATICS. Haemagogus iridicolor can be distinguished from most of the

remaining species in the Splendens Section by the following characters: in the

adults, from all species except lucifer by (1) dark coxal integument, (2) proboscis

slightly longer (1.00-1.10) than forefemur, (3) scales of mesonotum usually copper

to dark green but rarely blue and (4) dark abdominal scales purple or violet; in

the male genitalia by the claspette filament which is a broad, cuneiform leaf inserted

distally on claspette stem; and in the larva from all species except regalis, lucifer

and argyromeris by the combination of (1) 25 to 28 comb scales in irregular triple

row, (2) usually 4-or 5-branched hair 4a-X, (3) hair 7-III-VI subequal to hair 5 of

corresponding segment, (4) relatively long spines on caudal margin of saddle and

(5) pecten extending to about basal 0.50-0.60 of siphon.

This species can be determined with certainty only in the male genitalia as the

adults resemble lucifer closely, except in specimens with silver ppn scales. The

larva is indistinguishable from lucifer, regalis and argyromeris, and closely resembles

aeritinctus except in the relative development of the spines of the caudal margin

of the saddle, a character difficult to use and possibly not constant. The pupa

apparently has no demarcating characters. 3

Haemagogus iridicolor, aeritinctus, regalis and lucifer form a closely related spe-

cies complex, the regalis complex, marked by similarities in structure in all stages,

similar or related ecological requirements and, for the most part, allopatric distri-

bution. The 4 species probably arose from a parental stock in lower Central America,

possibly as a result of geographical isolation. Geohistorical evidence as interpreted

by Maldonado-Koerdell (1964:15-18) indicates the subsidence during middle Terti-

ary of parts of Caribbean Land, a Middle American early and middle Tertiary land

mass which included much of present Central America and the West Indies, form-

ing a series of small land masses in what is now southeastern Central America.

The 4 species of this complex may have evolved at this time, each on a separate

land mass. Iridicolor is probably autochthonous to the Atlantic rain forests of

Nicaragua and Costa Rica as it reaches its greatest density in this area.

BIONOMICS. Haemagogus iridicolor breeds primarily in treeholes and cut or

broken bamboo internodes, and has been found occasionally in terrestrial and

epiphytic bromeliads and fallen fruits. In the rain forests of the Atlantic slope

it is found primarily in treeholes but is more often taken in bamboo in the drier

Pacific side of Costa Rica. It is found commonly from sea level to elevations of

over 1000 meters. One collection was taken at over 2400 meters in a cloud forest

in Costa Rica. Both iridicolor and lucifer occur together on the Atlantic side of

western Panama. Although iridicolor reaches its maximum density in the rain forest

of Costa Rica and Nicaragua, it is more abundant than lucifer in the mangrove

swamps of Bocas del Toro province while lucifer is more common than iridicolor

Arnell: Genus Haemagogus 79

in the forest. Iridicolor has never been incriminated in the transmission of yellow

fever and nothing is known of its capability as a vector.

DISTRIBUTION (fig. 7). Haemagogus iridicolor extends from the Atlantic ver-

sant of Nicaragua and possibly Honduras (see aeritinctus) through both the Atlan-

tic and Pacific versants of Costa Rica to extreme western Panama. Material exam-

ined: 1549 specimens; 242 males, 250 females, 404 pupae, 653 larvae; 378 indi-

vidual rearings (239 larval, 104 pupal, 35 incomplete).

COSTA RICA. Alajuela: Alajuela, 925 m, 3 Nov 1971, D. and K. Schroeder (CR 508), 5 lpM

(508-13,15-17,107), 5 IpF (508-10-12,14,18), 2 pM (508-102,106), 1 pF (508-105), 5 L [UCLA];

same data (CR 509), 2 lpM (509-13,14), 3 lpF (509-10-12), 2 pM (509-102,103), 2 pF (509-101,

104), 2 P, 7 L [UCLA]; same data, (CR 514), 1 lpM (514-12), 2 IpF (514-10,11), 1 L [UCLA];

same data, 14 Nov 1971 (CR 528), 2 lpM (528-22,23), 7 lpF (528-11,20,21,24,25,27,28), 9 pF

(528-61,82,99,102,103,107,108,111,113), 7 pF (528-83,91,92,95 96,109,112), 1 F,6 L [UCLA];

same data (CR 531), 2 IpM (531-10,23), 1 IpF (531-11), 3 pM (531-50,106,112), 8 pF (531-51,100,

103-105,109-111), 1 p, 6 P, 51,17 L [UCLA]; same data, 26 May 1921-Apr 1922, A. Alfaro, 28

M, 16 F [USNM]. Alajuela (2 km S), 860 m, 27 July 1971, S. Heinemann (CR 295), 1 lpM (295-

21), 1 IpF (295-20), 1 pF (295-100), 1 L [UCLA]. Alajuela (3 km S), 880 m, 13 Nov 1962,

C. Hogue and W. Powder (CR 18), 1 IpM (18-109), 1 lpF (18-102), 1 pF (18-108), 1 IP (18-104)

[UCLA]. Alajuela (3 km SE), 880 m, 18 July 1971, D. Heinemann (CR 268), 2 L [UCLA] ; same

data (CR 271), 2 lpM (271-32,34,35), 5 IpF (271-30,31,33,36,37), 1 pM (271-101), 1 IP (271-60),

2 L [UCLA]; same data, 13 Nov 1971, D. and K. Schroeder (CR 527), 1 IpM (527-10), 2 pM (527-

101,102), 1 pF (527-100) [UCLA]. Atenas (6 km W), 950 m, 2 Aug 1971, S. Heinemann (CR 320),

4 IpM (320-10,14,15,18) 5 IlpF (320-11,13,16,17,19) 1 pM (320-100,101), 1 IP (320-12) [UCLA];

same data (CR 321), 15 M, 1 F [UCLA]; same data (CR 322), 3 lpM (322-10,11,14), 6 IpF (322-

12,13,15,17-19), 1 pF (322-100), 1 Ip (322-16), 1 L [UCLA]; same data (CR 323), 4 L [UCLA] ;

same data (CR 324), 1 lpM (324-11), 1 IP (324-10), 2 L [UCLA]. Higuito, P. Schild, 9 M, 1M

gen, 7 F [USNM]. La Fortuna, 25 Oct 1951, W. Komp, 3 M, 3 F [UCLA]. San Mateo (3 km E),

300 m, 1 Nov 1971, D. and K. Schroeder (CR 498), 1 lpF (498-11), 1 IP (498-10) [UCLA]. San

Ramon (3 km N), 1000 m, 12 Aug 1971, S. Heinemann (CR 343), 1 lpM (343-11), 3 lpF (343-12-

14), 2 pM (343-100,101), 1 IP (343-10), 5 L [UCLA]; same data (CR 344), 1 IpM (344-10,11),

1 pM (344-104), 4 pF (344-100-103), 2 IP (344-33,36), 4 P, 3 L [UCLA]; same data (CR 345),

2 IpM (345-18,20), 9 IlpF (345-10-17,19), 1 pM (345-105), 5 pF (345-100-104) [UCLA]. Volcan

Poas Summit (3 km S), 2400 m, 14 Nov 1962, C. Hogue and W. Powder (CR 20), 1 IpF (20-101)

[UCLA]. Guanacaste: Cerro Maravilla, 20 May 1921, A. Alfaro, 1 M [USNM]. Heredia: Puerto

Viejo, 100 m, 15 Aug 1964,C. Hogue (CR 191), 5 L [UCLA]; same data, 8 Jan 1972, D. Schroeder

(CR 585), 1 lpM (585-10) [UCLA]. Limon: Barra de Parismina, H. Kumm, 1 F [USNM]. La

Bomba, 3 Oct 1971, D. Schroeder (CR 470), 2 lpM (470-11,12), 1 L [UCLA]. Limon, F. Knab,

2 F [USNM]. Limon (6 km SW), 14 Dec 1971, D. Schroeder (CR 550), 1 IpF (550-40) [UCLA].

Portete (S km NW Limon), 2 m, 5 Oct 1971, D. Schroeder (CR 476), 5 IpM (476-12,14,18,19,60),

9 IpF (476-10,11,13,17,61-65), 5 pM (476-91,102,106,110,112), 5 pF (476-100,103-105,107),

2 IP (476-15,16), 6 P, 65 L [UCLA]; same data (CR 478), 7 lpM (478-11-13,15,17,19,32), 5 lpF

(478-10,14,16,30,31), 1 pM (478-101), 1 pF (478-102), 1 IP (478-18), 1 P, 4 L [UCLA].

Puntarenas: Dominical, 30 May 1943, T. Aitken, 1 M, 1 F [UCLA]. Esparta (5 km E), 250 m,

13 Aug 1971, S. Heinemann and D. Schroeder (CR 356), 5 IpF (356-10,12-15), 2 pM (356-100,

101), 1 IP (356-11), 1 P, 1 1, 1 L [UCLA]; same data (CR 358), 5 IpF (358-14,15,20,21,23),

2 pM (358-102,105), 4 pF (358-103,106,109,111), 16 L [UCLA]. Golfito (GML), 1 lpM (03189)

[UCLA]. Villa Niely, 8 Aug 1963, C. Hogue (CR 167), 7 L [UCLA]. San Jose: San Isidro del

General, 21 Nov 1962, C. Hogue and W. Powder (CR 35), 2 IP (35-201,203), 1 F, 2 L [UCLA];

same data, 14 July 1963, C. Hogue (CR 149), 1 IpF (149-103), 3 IP (149-102,105,108), 3 M, 3 F,

4 p, 24 L [UCLA]; same data (CR 150), 1 M [UCLA] ; same data, 21 June 1964 (CR 177), 2 pM

(177-203,205), 1 pF (177-201), 1 lp (177-101) [UCLA]; same data (CR 178), 1 M gen, 1 L

[UCLA]; same data, 24 June 1964 (CR 183), 14 L [UCLA]. Santa Ana (1 km N), 850 m,

25 July 1971, S. Heinemann (CR 293), 7 IpM (293-10-16), 4 IpF (293-17-19,21), 2 pM (293-

103,104), 1 pF (293-101), 1 IP (293-20), 3 L [UCLA]. Province unknown: Suerre (Atlantic side),

80 Contrib. Amer. Ent. Inst., vol. 10, no. 2; 1973

22 July 1923, A. Alfaro, 2 F [USNM].

NICARAGUA. Chontales: Villa Somoza, June, July 1953 (GML), 2 lpM (01471,01518), 4 IpF

(01453,01468,01522,01523), 1 pF (01521), 3 1 [UCLA]. Zelaya: Bluefields, W. Thornton, 6 F

[USNM] ; same data, 11 July 1964, A. Quinonez (NI 36), 2 F [UCLA] ; same data (NI 38), 1 IpM

(38-12), 2 pF (38-100,105), 1 pF (38-104), 1 P, 55 L [UCLA]; same data, 13 July 1964 (NI 42),

1 1F (42-11), 1 pF (42-12), 12 L [UCLA] ; same data (NI 43), 4 IpF (43-10-13), 3 pM (43-100-102),

11, 11 L [UCLA]; same data (NI 47), 7 L [UCLA] ; same data, 20 July 1964 (NI 57), 1 IpM (57-

30), 1 L [UCLA] ; same data, 23 Nov 1971, D. and K. Schroeder (NIC 82), 2 F [UCLA] ; same data,

26 Nov 1971 (NIC 108), 1 lpF (108-20) [UCLA]; same data, 27 Nov 1971 (NIC 112), 2 F

[UCLA]; same data (NIC 115), 6 IpM (115-10-12,17,18,20), 6 IpF (115-13-16,19,21), 7 L

[UCLA] ; same data (NIC 117), 12 IpM (117-10,12-15,30,31,33,34,37,38,42), 7 IpF (117-11,16-

19,36,39), 2 L [UCLA]; same data (NIC 118), 3 IpM (118-14,15,17), 6 IpF (118-10-12,16,18,19),

1 pM (118-100), 1 lp (118-13), 9 L [UCLA]; same data (NIC 126), 3 IpM (126-11,12,15), 3 IpF

(126-13,14,16), 1 IP (126-10) [UCLA]; same data, 28 Nov 1971 (NIC 128), 6 IpM (128-10-12,

15-17), 3 IpF (128-13,14,18), 11 L [UCLA]; same data (NIC 129), 8 IpM (129-13-19,22), 5 IpF

(129-10-12,20,21), 2 pM (129-100,101), 13 L [UCLA]; same data (NIC 131), 5 IpM (131-10-12,

14,15), 3 lpF (131-13,16,17), 2 L [UCLA] ; same data (NIC 134), 2 IpM (134-11,12), 1 IpF (134-

13), 1 pM (134-100), 1 P, 1 1 [UCLA]; same data (NIC 135), 2 IpF (135-10,11), 1 L [UCLA].

El Puente, nr. Bluefields, 12 Sept 1967, A. Adames and Herrara (NIC 75), 1 IpM (75-13), 2 IpF

(75-10,16), 3 Ip (11,14,15) [UCLA]. Rio Curinhuas (100 km from mouth), 29 Nov 1971, I.

Sanderson, 2 F [UCLA].

PANAMA AND CANAL ZONE. Bocas del Toro: Almirante, 29 Dec 1928, 1 M gen [USNM];

same data, Feb 1931, W. Komp, 1 M gen [USNM]; same data, 27 Apr 1963, A. Quinonez

(PA 264), 1 IpM (264-111), 1 lpF (264-104), 3 pM (264-101,107,108), 1 IP (264-110) [UCLA];

same data, 29 Apr 1963 (PA 282), 1 L [UCLA]; same data, 4 May 1963 (PA 317), 1 pM (317-

102), 1 1 [UCLA]. Almirante (crematory), 30 m, 30 Apr 1963, A. Quinonez (PA 290), 1 IpF

(290-106), 11, 8 L [UCLA]. Almirante, Bocas-Chiriqui Rd, 6 May 1963, A. Quinonez (PA 324),

1 lpF (324-101), 1 IP (324-102), 2 L [UCLA]; same data (PA 326), 1 pF (326-101) [UCLA].

Almirante, Nigua Creek, 4 May 1963, A. Quinonez (PA 311), 1 IpM (311-105), 1 IpF (311-103),

4 pM (311-101,102,104,105), 6 L [UCLA]; same data (PA 312), 1 IpF (312-102), 3 pM (312-

110,112), 1 pF (312-101), 2 L [UCLA]. Bocas del Toro, 1947, P. Galindo, 6 lp [USNM] ; same

data, 4 Sept 1957 (GML), 1 IpM (03239), 1 1 [UCLA]. Bocas del Toro, Big Creek, 9 Apr 1964,

A. Quinonez (PA 653), 1 lpM (653-104), 2 IpF (653-103,105), 2 pM (653-106,108), 7 L [UCLA].

Secretarro, 7 June 1949, 12 M, 9 F [UCLA]. Chiriqui: Concepcion, 10 Nov 1949 (GML), 21

[UCLA]. Locality unspecified: S. Carpenter, 1 M [UCLA].

‘19. Haemagogus (H.) aeritinctus Galindo & Trapido

Figs. 7,45,46

1967. Haemagogus (Haemagogus) aeritinctus Galindo and Trapido, 1967:103-111. TYPE: Holo-

type male with associated larval and pupal skins and genitalia slide, vicinity of Stann

Creek, British Honduras, larva collected in Rhizophora mangle treehole, 5 Oct 1955,

P. Galindo and H. Trapido [USNM].

Haemagogus (Haemagogus) aeritinctus of Stone (1970:157).

Haemagogus (Haemagogus) regalis in part of Dyar (1921a:105); Martini (1935:56); Stone, Knight

and Starcke (1959:217).

Haemagogus regalis in part of Dyar and Knab (1906a:167).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870).

FEMALE. Wing: 2.15 mm. Proboscis: 1.70 mm. Forefemur: 1.45 mm. Abdo-

men: 2.20 mm. Dark scales of proboscis, palpus, wing and legs primarily purple,

with some violet reflections. Head: Eyes widely separated above antennae (4 omma-

Arnell: Genus Haemagogus 81

tidial diameters). Decumbent scales of vertex and occiput purple and violet ante-

riorly and laterally, light yellowish brown posteriorly. Proboscis about 1.10-1.20

of forefemur; palpus about 0.15 of proboscis; antenna about 0.55-0.60 of proboscis.

Thorax: Scales of mesonotum copper with some purple reflections, bluish green

over supraalar bristles, silver in antealar area; scales on scutellum bluish green.

Apn lobes with scales purple and violet, a few silver scales mesally; ppn bare.

Legs: Integument of apex of coxae, trochanters and base of femora yellow; length

of forefemur about 1.50 of distance from top of thorax to apex of midcoxa.

Wing: R43 about 0.50 of R,. Abdomen: Dark scales predominantly purple, with

some violet and copper reflections; silver scales dorsally in indistinct basal band on

tergites VII and VIII. ;

MALE (fig. 45). Wing: 2.30 mm. Proboscis: 2.05 mm. Forefemur: 1.80 mm.

Abdomen: 2.85 mm. Head: Proboscis about 1.15 of forefemur; antenna sparsely

plumose, flagellar segment 4 with about 6 moderately long bristles.

MALE GENITALIA (fig. 45). Segment VIII: Tergite about 0.85 of sternite,

distal margin straight or slightly rounded, with numerous strong setae, those mesad

stout and short. Segment IX: Tergite straight and unsclerotized mesally. Sidepiece:

Conical, truncate, length about 2.5 times median width; basal tergomesal lobe

enlarged, extending distad as raised area past middle of sidepiece, with numerous

setae, short distally, becoming longer proximally, the more proximal ones much

enlarged, flattened and attenuate; distal third of sidepiece without setae on mesal

half; apical lobe prominent, bearing 10-15 moderately developed setae; apical

sternomesal scales lanceolate to oblanceolate with acuminate tips. Claspette: Stem

narrow, bent sharply dorsad at apical third and slightly expanded beyond; filament

a flat, expanded leaf inserted distally on stem and terminating in a slightly recurved

point. Clasper: Flattened, curved, expanded at apical fourth and tapering to a rather

sharp point; spiniform short, spatulate, inserted subapically on inner surface. Phallo-

some: Aedeagus moderately large, obovate, tip expanded dorsally into serrated

carina terminating in small beak; venter broadly open except on basal third, with

sclerotized ridges lateroventrally which are expanded basally. Proctiger: Cercal

setae 5; apical knob of paraproct with about 15 serrations.

PUPA (fig. 45). Abdomen: about 2.80 mm. Trumpet: 0.45 mm. Paddle: 0.55

mm. Cephalothorax: Moderately pigmented, darker dorsally. Hair 4-C weak, usu-

ally double, subequal to 5-C which is usually single and weaker than 7-C which is

single. Trumpet: Light brown, reticulate sculpturing moderate. Abdomen: Rather

weakly pigmented, genital lobe and anterior segments darker. Float hair (1-I) with

about 8-10 primary branches and 2-6 secondary branches. Hair 1-II-VII weak,

single to triple, subequal on all segments. Hair 2-VII considerably cephalad of

I-VI. Hair 3-If stronger than 3-III but weaker than 5-IV,V which are subequal.

Hair 6-VII weaker than 6-III-VI which are subequal. Hair 9-VII near caudolateral

margin of tergite, usually double; 9-VIII 4-7b. Terminal Segments: Male genital

lobe about 1.4 of tergite VIII. Paddle: Weakly pigmented; relatively broad; apex

broadly rounded. Hair 1-P single.

LARVA (fig. 46). Head: 0.70 mm. Siphon: 0.53 mm. Anal Saddle: 0.26 mm.

Segment VIII: Comb scales 20-28, with single broadly rounded, minutely fringed

spine, in irregular double row. Siphon: Index about 2.5. Pecten teeth 10-13, ex-

tending to about basal 0.45 of siphon. Hair 1-S 4b. Anal Segment: Spines on

caudal margin of saddle reduced, very short, each with multiple teeth. Hair 4a-X 5b.

SYSTEMATICS. Haemagogus aeritinctus can be separated from the remaining

species of the Splendens Section: in the adults by the combination of (1) bare ppn,

82 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

(2) scales of mesonotum copper, (3) proboscis about 1.10-1.20 of forefemur and

(4) yellow integument of apex of coxae and trochanters; in the male genitalia by

(1) claspette filament shaped like a flat, expanded leaf and inserted distally on stem

and (2) flattened and sharply pointed clasper; and in the larva by the combination

of (1) comb scales 20-28 and in irregular double row, (2) pecten extending to about

basal 0.45 of siphon, (3) spines on caudal margin of saddle much reduced, and (4)

hair 7-III-VI subequal to hair 5 of corresponding segment. The development of the

spines on the caudal margin of the saddle is a difficult character to use and may be

variable. If this character state breaks down, the larva of aeritinctus would be

impossible to distinguish from iridicolor, regalis, lucifer and argyromerts.

This species is closely related to iridicolor, regalis and lucifer, with which it

forms the regalis complex. The evolution of this group is discussed under iridicolor.

BIONOMICS. Haemagogus aeritinctus breeds in treeholes and is restricted to

mangrove associations. It has never been taken inland. Females attack man readily

and bite around the head, in contrast to most other species of Haemagogus which

tend to bite low on the body. This species is not known to be involved in the

transmission of yellow fever.

DISTRIBUTION (fig. 7). Haemagogus aeritinctus is known only from the Atlan-

tic coast of British Honduras, Guatemala and northern Honduras. The two records

from Honduras are based on immature stages and one or both of these may

actually be iridicolor. Material examined: 46 specimens; 6 males, 6 females, 14

pupae, 20 larvae; 14 individual rearings (9 larval, 3 pupal, 2 incomplete).

BRITISH HONDURAS. Belize: Belize, 5 Oct 1955 (GML), 1 lpF (02157), 1 pF (02155)

[UCLA]. Stann Creek: Stann Creek, 5 Oct 1955 (GML), 4 IpM (02180,02186,02189,02192),

4 IpF (02169,02173-02175), 2 pM (02187,02193), 1 lp (02166), 61 [UCLA, USNM].

HONDURAS. Colon: Puerto Castilla, 9 Aug 1964, A. Quinonez (HON 14), 1 L; same data

(HON 15), 2 L [UCLA]. Cortes: Puerto Cortes, Rio Mar, 10 Aug 1968, A. Adames (HON 64),

1 IP (64-10) [UCLA].

Additional Record From the Literature

GUATEMALA. Izabal: Puerto Barrios (Galindo and Trapido, 1967:111).

20. Haemagogus (H.) regalis Dyar & Knab

Figs. 7,47,48

1906. Haemagogus regalis Dyar and Knab, 1906a:167. TYPE: Holotype male (330 v) with

associated larval (fragment) and pupal skins and genitalia slide (36.1.8b), Sonsonate,

Sonsonate, El Salvador, 30 Aug 1905, F. Knab [USNM, 10024].

Haemagogus (Haemagogus) regalis of Dyar (1921a:105, in part; 1928:136); Edwards (1932:179);

Martini (1935:56, in part); Lane (1939:120, in part; 1953:787-789); Stone, Knight and Starcke

(1959:217, in part); Diaz Najera (1966:61); Galindo and Trapido (1967:109-110).

Haemagogus (Stegoconops) regalis of Bonne and Bonne-Wepster (1925:430).

Haemagogus regalis of Theobald (1910:493-494, in part); Kumm and Zuniga (1942:404); Trapido

and Galindo (1956a:305); Vargas and Diaz Najera (1959:361); Diaz Najera (1960:187).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870). |

Stegoconops albomaculatus of Howard, Dyar and Knab (1913:fig. 163).

Aedes cyaneus in part of Dyar and Knab (1906a:191,202).

FEMALE. Wing: 3.25 mm. Proboscis: 2.50 mm. Forefemur: 2.45 mm. Abdo-

men: 3.50 mm. Dark scales of proboscis, palpus, wing and legs purple and violet.

Arnell: Genus Haemagogus 83

Head: Eyes widely separated above antennae (4 ommatidial diameters). Decumbent

scales of vertex and occiput blue with silver reflections. Proboscis about 1.00-1.10

of forefemur; palpus about 0.17 of proboscis; antenna about 0.70 of proboscis.

Thorax: Scales of mesonotum copper to greenish blue or blue, bluish green over

supraalar bristles, silver in antealar area; scales on scutellum greenish blue. Apn

lobes with scales blue; ppn bare. Legs: Length of forefemur 1.50-1.60 of distance

from top of thorax to apex of midcoxa. Wing: R,43; about 0.40-0.45 of Ro».

Abdomen: Dark scales usually blue with some purple reflections; usually silver

scales present dorsally in basal band or median basal patch on tergites III or IV-VII.

MALE (fig. 47). Wing: 2.20 mm. Proboscis: 2.15 mm. Forefemur: 1.85 mm.

Abdomen: 2.90 mm. Head: Proboscis about 1.12-1.20 of forefemur; antenna

sparsely plumose, flagellar segment 4 usually with 12 moderately developed bristles.

MALE GENITALIA (fig. 47). Segment VIII: Tergite about 0.90 of sternite,

distal margin broadly slightly emarginate, with long, stiff setae, more numerous

and stronger mesally. Segment IX: Tergite slightly acclivous and weakly sclerotized

mesally. Sidepiece: Conical, truncate, length about 2.5 times median width, basal

tergomesal area enlarged, extending distad as raised area past middle of sidepiece,

with numerous setae, short distally, becoming longer proximally, the more prox-

imal ones considerably enlarged; distal third of sidepiece without setae on mesal

half; apical lobe prominent, bearing 10-15 moderately developed setae; apical sterno-

mesal scales lanceolate to oblanceolate, the more dorsal projecting more or less

dorsad, those ventral projecting mesad. Claspette: Stem bowed inward near apex

in dorsal aspect, narrow, bent sharply dorsad at apical third and tapering to rather

acute apex; filament a flat, expanded leaf terminating in an acute point, inserted

distally on stem and extending ventrally over apical third of stem as an attached,

folded membrane. Clasper: Flattened, curved, more or less arcuate and terminating

in a rounded point; spiniform short, spatulate, inserted subapically on inner surface.

Phallosome: Aedeagus moderately large, obovate, tip expanded dorsally into a small

serrated carina terminating in a beaklike process; venter broadly open on distal half,

closed basally, with sclerotized ridges lateroventrally which are expanded basally.

Proctiger: Cercal setae 4-6; apical knob of paraproct with about 15 serrations.

PUPA (fig. 47). Abdomen: about 2.95 mm. Trumpet: 0.35 mm. Paddle: 0.60

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hairs 4,5-C

more or less subequal, single to triple, weaker than 7-C which is single. Trumpet:

Medium brown, lighter distally. Abdomen: Weakly to moderately pigmented, genital

lobe darker, anterior segments slightly darker. Float hair (1-1) with about 10-14 pri-

mary branches and 2-6 secondary branches. Hair 1-II-VII weakly to strongly devel-

oped single, multiple or dendritic, more or less subequal on all segments. Hair 2-VII

short, considerably cephalad of 1-VII. Hair 3-III weaker than 3-II which is subequal

to or slightly weaker than 5-IV,V which are subequal. Hair 6-VII often moderately

developed, stronger than or occasionally subequal to 6-III-VI. Hair 9-VII slightly to

considerably cephalad of caudolateral margin of tergite, 3-4b; 9-VIII about 5-10b.

Terminal Segments: Male genital lobe about 1.3 of tergite VIII. Paddle: Usually

weakly pigmented, midrib weakly pigmented; relatively broad, apex usually broadly

rounded, occasionally somewhat acuminate. Hair 1-P single.

LARVA (fig. 48). Head: 0.85 mm. Siphon: 0.70 mm. Anal Saddle: 0.32 mm.

Segment VIII: Comb scales 22-26(18-29), with single broadly rounded minutely

fringed spine, in irregular double to triple row. Siphon: Index about 2.5(2.1-2.7).

Pecten teeth about 13-18, extending to about basal 0.45-0.50 of siphon. Hair 1-S

4-5b(3-6b). Anal Segment: Hair 4a-X 5b(4-6b).

84 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

SYSTEMATICS. Haemagogus regalis can be distinguished from most other species

in the Splendens Section: in the adults by the combination of (1) bare ppn, (2)

proboscis slightly longer (1.00-1.10) than forefemur, (3) abdomen with dark scales

usually blue with silver scales dorsally at base of tergites III-VII and (4) coxae with

dark integument; in the male genitalia by (1) arcuate clasper, (2) claspette stem

uniformly narrow on basal half and (3) claspette filament extending posteriorly

over distal third of stem as an attached membrane; and in the larva from all species

except iridicolor, lucifer and argyromeris, by the combination of (1) usually 22 to

26 comb scales, (2) usually 5-branched hair 4a-X, (3) pecten extending to about

basal 0.50 of siphon, (4) hair 7-III-VI subequal to hair 5 of corresponding segment,

and (5) relatively long spines on caudal margin of saddle.

Closely allied to iridicolor, aeritinctus and lucifer, regalis forms the regalis com-

plex with these 3 species. The possible evolution of this complex is discussed under

iridicolor. The habitat of regalis is restricted to coastal areas, primarily mangrove

swamps. It reaches its greatest density on the Pacific coast from El Salvador to

Mexico but is also found on the Atlantic side of the Isthmus of Tehuantepec.

This distribution pattern would tend to support the proposed Atlantic-Pacific con-

nection at the Isthmus of Tehuantepec during middle Tertiary (Maldonaldo-Koerdell,

(1964:15). A similar distribution pattern is found in Deinocerites pseudes Dyar

and Knab, 1909, also a littoral species (Adames, 1971:76).

BIONOMICS. Little is known of the bionomics of regalis. It breeds mainly in

treeholes but occasionally in other plant containers. It seems to be mostly restricted

to mangrove swamps and adjacent coastal areas, although it is found inland at low

elevations in swampy areas on the Caribbean coast of Mexico. It apparently is not

involved in disease transmission.

DISTRIBUTION (fig. 7). Haemagogus regalis is known from the Atlantic coastal

states of Tabasco and southern Veracruz, Mexico and the Pacific coast from south-

ern Chiapas, Mexico to Honduras. The records in Stone, Knight and Starcke (1959:

217) of regalis from British Honduras and Panama and Colombia undoubtedly refer

to aeritinctus and lucifer, respectively. Material examined: 153 specimens; 47 males,

33 females, 23 pupae, 50 larvae; 23 individual rearings (10 larval, 9 pupal, 4 incom-

plete).

EL SALVADOR. La Liberdad: Estero Ticuisiapa, H. Kumm, 5 M, 7 F, 71 [UCLA, USNM].

Sonsonate: San Antonio, 4 Aug 1964 (SAL 10), 1 IP (10-105), 5 L [UCLA]. Sonsonate, 1 Aug

1964 (SAL 2), 2 L [UCLA]; same data (SAL 4), 1 pM (4-103), 1 IP (4-10), 17 L [UCLA]; same

data, F. Knab, 3 M, 1 M gen, 4 F, 1 lp [USNM]. Usultan: Isla Espiritu Santo, H. Kumm, 3 M, 1 M

gen, 2 1 [USNM]. Peninsula de San Juan del Gozo, 30 July 1941, H. Kumm, 2 M gen, 2 F, 1 1

[USNM]. Locality unspecified: 2 L [USNM].

GUATEMALA. Escuintla: Puerto de Iztapa, 17,18 July 1953, P. Galindo and H. Trapido

(GML), 4 lpM (01114,01123,01125,01126), 3 IpF (01113,01116,01121), 3 pM (01130,01133,

01621), 3 pF (01129,01131,01132), 1 lp (01127), 10 M [UCLA, GML].

MEXICO. Chiapas: Puerto Madero, 3 Aug 1953 (GML), 1 M, 8 F [GML] . Tabasco: Tenosique

de Pino Suarez, 2 M gen [USNM]. Veracruz: Coatzacoalcos, 2 Aug 1953, P. Galindo and H.

Trapido (GML), 2 IpM (0998,03218), 1 IpF (01141), 2 pM (01145,01149), 8 M, 4 F [UCLA,

GML]. Santa Lucrecia (Jesus Carranza), F. Knab, 1 F [USNM].

21, Haemagogus (H.) lucifer (Howard, Dyar & Knab)

Figs. 7,49,50

1913. Stegoconops lucifer Howard, Dyar and Knab, 1913:fig. 164. TYPE: Lectotype male

Arnell: Genus Haemagogus 85

(299) on slide 309, Tabernilla, Canal Zone, Panama, 14 Apr 1909, A.H. Jennings

[USNM; selection of Dyar, 1921a:107; see Stone and Knight, 1955:288] . Synonymized

with regalis by Komp (1954c:193-195); resurrected by Trapido and Galindo (1956a:

305).

Haemagogus (Haemagogus) lucifer of Dyar (1921a:107; 1923:183; 1928:138-139); Bonne and

Bonne-Wepster (1925:434); Edwards (1932:179); Lane (1939:119; 1953:781-782); Levi-

Castillo (1951b:11,22-23); Stone, Knight and Starcke (1959:217); Forattini (1965:35-39);

Cova Garcia (1966a:61-62, fig. 115; 1966b:112, fig. 200); Galindo and Trapido (1967:

109-110).

Haemagogus lucifer of Dyar (1925b:139); Kumm, Komp and Ruiz (1940:398); Boshell-Manrique

and Osorno-Mesa (1944:173); Osorno-Mesa (1944a:45); Hovanitz (1946:35); Kumm, Osorno-

Mesa and Boshell-Manrique (1946:20); Arnett (1949:240; 1950:107); Galindo, Carpenter and

Trapido (1949:278; 1951a:118-119; 1955:158); Galindo, Trapido and Carpenter (1950:546);

Carpenter, Galindo and Trapido (1952:162); Barreto Reyes (1955:78); Trapido, Galindo and

Carpenter (1955:528); Galindo, de Rodaniche and Trapido (1956:1024); Galindo, Trapido,

Carpenter and Blanton (1956:544); Trapido and Galindo (1956a:304; 1957:122); de Rodaniche

and Galindo (1957:236); de Rodaniche, Galindo and Johnson (1957:682); Galindo, de

Rodaniche and Johnson (1959:557); Kerr, Roca-Garcia and Bugher (1960:26); Groot, Morales

and Vidales (1961:399); Galindo and de Rodaniche (1964:846).

Haemagogus regalis in part of Komp (1954c:193-195; 1956:37).

Haemagogus splendens in part of Busck (1908:64); Howard, Dyar and Knab (1917:867).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870).

FEMALE. Wing: 2.80 mm. Proboscis: 2.35 mm. Forefemur: 2.15 mm. Abdo-

men: 2.80 mm. Dark scales of proboscis, palpus, wing and legs purple and violet.

Head: Eyes widely separated above antennae (4 ommatidial diameters). Decumbent

scales of vertex and occiput violet with some purple reflections. Proboscis about

1.05-1.15 of forefemur; palpus about 0.15 of proboscis; antenna about 0.60-0.65

of proboscis. Thorax: Scales of mesonotum dark green to copper, bluish green over

supraalar bristles, silver in antealar area; scales on scutellum bluish green but occa-

sionally purple on scutellar lobes. Apn lobes with scales violet to purple; ppn bare.

Legs: Length of forefemur about 1.60-1.75 of distance from top of thorax to apex

of midcoxa. Wing: R,43 about 0.30-0.40 of R,. Abdomen: Dark scales usually

purple, occasionally blue to violet; usually with silver scales dorsally in small basal

patch on tergites VI and VII.

MALE (fig. 49). Wing: 2.45 mm. Proboscis: 2.45 mm. Forefemur: 2.05 mm.

Abdomen: 2.90 mm. Head: Proboscis about 1.15-1.20 of forefemur; antenna

about 0.50 of proboscis, sparsely plumose, flagellar segment 4 with 12 moderately

developed bristles.

MALE GENITALIA (fig. 49). Segment VIII: Tergite about 0.80 of sternite,

distal margin broadly emarginate, slightly acclivous at midline, with numerous

long stiff setae, generally longer and with slightly hooked tips laterally; a few

large, cuneate scales laterally. Segment IX: Tergite acclivous and weakly sclerotized

on midline. Sidepiece: Conical, truncate, length about 2.5 times median width;

basal tergomesal area enlarged, extending distad as raised area to near distal third

of sidepiece, with numerous setae, short distally, becoming longer proximally, the

more proximal ones much enlarged. flattened and attenuate, distal third of side-

piece without setae on mesal half; apical lobe prominent, beating 10-15 moder-

ately developed setae; apical sternomesal scales lanceolate to oblanceolate with

acuminate tips, the more dorsal projecting more or less dorsad, those ventral pro-

jecting mesad. Claspette: Stem bowed inward near apex in dorsal aspect, stout,

86 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

expanded slightly at basal third, then constricted slightly beyond middle, angled

sharply dorsad at apical third and tapering to acute apex; filament a flat expanded

leaf terminating in an acute point, inserted distally on stem and extending poste-

riorly over apical third of stem as attached folded membrane. Clasper: Flattened,

curved, expanded on distal fourth and tapering to rather sharp point; apical spini-

form short, spatulate, inserted subapically on inner surface. Phallosome: Aedeagus

moderately large, obovate, tip expanded dorsally into a small serrated carina

terminating in a beaklike process; venter broadly open on distal half, closed basally,

with sclerotized ridges lateroventrally. Proctiger: Cercal setae 4 or 5; apical knob

of paraproct with about 20 serrations.

PUPA (fig. 49). Abdomen: about 3.20 mm. Trumpet: 0.40 mm. Paddle: 0.65

mm. Cephalothorax: Usually weakly pigmented, occasionally moderately pigment-

ed, somewhat darker dorsally. Hairs 4,5-C rather weakly developed, subequal, 1-4b,

weaker than 7-C which is single. Trumpet: Medium brown, reticulate sculpturing

moderate. Abdomen: Weakly pigmented, genital lobe and anterior segments slight-

ly darker. Float hair (1-I) with about 10-15 primary branches and 2-6 secondary

branches. Hair 1-II-VII weakly to strongly developed, single, multiple or dendritic,

subequal on all segments. Hair 2-VII short, considerably cephalad of 1-VII. Hair

3-III usually weaker than 3-II,5-IV,V which are usually subequal. Hair 6-VII sub-

equal to or stronger than 6-IIJ-VI which are subequal. Hair 9-VII slightly to

considerably cephalad of caudolateral margin of tergite, 2-5b; 9-VIII 4-8b. Terminal

Segments: Male genital lobe about 1.3 of tergite VII. Paddle: Weakly pigmented,

midrib weakly pigmented; relatively broad, apex usually broadly rounded. Hair

1-P single.

LARVA (fig. 50). Head: 0.80 mm. Siphon: 0.75 mm. Anal Saddle: 0.31 mm.

Segment VIII: Comb scales 25-28(21-33), with single broadly rounded, minutely

fringed spine, in irregular double to triple row. Siphon: Index about 2.4(2.0-2.6).

Pecten teeth about 13-15(9-17), extending to about basal 0.50 of siphon. Hair 1-S

4b(3-5b). Anal Segment: Hair 4a-X 4-6b.

SYSTEMATICS. Haemagogus lucifer can be distinguished from most of the spe-

cies of the Splendens Section by the following characters: in the adults, from all

except iridicolor, by (1) dark coxal integument, (2) proboscis about 1.05 to 1.15

of forefemur, (3) dark green to copper scales of mesonotum, and (4) dark scales

of abdomen usually purple, occasionally blue to violet; in the male genitalia by

(1) clasper expanded on distal fourth and tapered to sharp point beyond, (2) stout

claspette stem which is expanded at basal third and (3) claspette filament extending

posteriorly over distal third of stem as an attached membrane; and in the larva, from

all except iridicolor, regalis and argyromeris, by the combination of (1) usually 25 to

28 comb scales, (2) hair 4a-X usually 4 to 6 branched, (3) pecten extending to about

basal 0.50 of siphon, (4) hair 7-III-VI subequal to hair 5 of corresponding segment

and (5) relatively long spines on caudal margin of saddle.

Haemagogus lucifer, iridicolor, aeritinctus and regalis form the regalis complex,

the evolution of which is discussed under iridicolor. The present distribution of

lucifer indicates that it probably arose on a separate land mass near the present

Panamanian Isthmus area and spread into northwestern South America when that

land mass became connected to South America. Lucifer seems to be widespread,

though relatively uncommon, in the drainage of the Magdalena River with pene-

trations into the upper Orinoco and Amazon systems.

BIONOMICS. Haemagogus lucifer reaches its maximum density in primary tropi-

cal rain forest in Panama but is found in large numbers in second growth and

Arnell: Genus Haemagogus 87

occasionally in peridomestic situations in areas of high rainfall on the Atlantic

side of Panama. It breeds almost exclusively in treeholes. As most eggs hatch when

first flooded, lucifer is abundant soon after the onset of seasonal rains and main-

tains relatively high populations throughout the rainy season. It is arboreal in

habit, though not as much as janthinomys. Females bite man readily, usually around

the lower part of the body. Males have been observed near the host where they

await females. Copulation takes place in flight. Where lucifer and iridicolor occur

together on the Atlantic coast of western Panama, lucifer is more common in the

forest and iridicolor predominates in coastal mangrove.

Lucifer has been found to be capable of harboring yellow fever virus in the

laboratory (Galindo, de Rodaniche and Trapido, 1956) and has been reported to

be naturally infected with this virus (de Rodaniche, Galindo and Johnson, 1957).

A thorough discussion of the bionomics and disease potential of lucifer may be

found in Galindo, Trapido and Carpenter (1950) and Galindo, Carpenter and

Trapido (1951; 1955).

DISTRIBUTION (fig. 7). Haemagogus lucifer is known from the Atlantic versant

of Costa Rica and western Panama, the Atlantic and Pacific versants of central and

eastern Panama and the Caribbean drainage of Colombia, with one record from

the upper Orinoco basin of Colombia and one record from the upper Amazon basin

of Ecuador. Material examined: 1246 specimens; 273 males, 348 females, 247

pupae, 378 larvae; 247 individuals (107 larval, 68 pupal, 72 incomplete).

COLOMBIA. Antioquia: Casabe, 3 IM gen [USNM]. Turbo, 1 M gen [USNM]. Cundinamarca:

Caparrapi, Volcanes Forest, 1000-1500 m, 25 May 1943, H. Kumm, 1 Ip, 2 M, 1 M gen, 1 F [UCLA,

USNM]. Malta, 280 m, 18 Feb 1943, E. Osorno-Mesa, 1 M, 1 M gen, 1 F [UCLA, USNM]. Meta:

Villavicencio, 1944, M. Bates, 1 M, 1 M gen, 1 1 [USNM]. Santander: Barranca (nr. Velez), H.

Kumm, 1 M [USNM]. Pescadera, Rio Horta, H. Kumm, 1 M [USNM].

COSTA RICA. Limon: Guapiles, H. Kumm, 2 M, 1 F, 11 [USNM].

ECUADOR. Napo-Pastaza: Ila, R. Levi-Castillo, 4 M gen [USNM].

PANAMA AND CANAL ZONE. Bocas del Toro: Almirante, 9 May 1963, A. Quinonez (PA 342),

2 F [UCLA]; same data, Feb 1931, W. Komp, 1 F [UCLA]; same data, 20 Aug 1944, 11,1 L

[UCLA]. Almirante, mile 2, 26 Apr 1963, A. Quinonez (PA 254), 1 IpF (254-104), 2 p [UCLA] ;

same data, 27 Apr 1963 (PA 259), 1 IpM (259-113), 3 IpF (259-102,115,117), 5 pM (259-101,

106,107,109,110), 2 pF (259-103,108), 2 IP (259-112,114), 1 P, 10 L [UCLA] .Chiriquisito, 19 Apr

1963, A. Quinonez (PA 234), 1 IpM (234-108), 1 pM (234-105), 6 L [UCLA] ; same data (PA 235),

1 Ip (235-103) [UCLA]; same data, 20 Apr 1963 (PA 245), 4 lpM (245-101,102,104,106), 5 IpF

(245-108-111,113), 1 lp (245-103), 1 IP (245-112), 1 P, 2 L [UCLA]. El Guabo, nr. Chriquisito,

15 Apr 1963, A. Quinonez (PA 205), 1 IpM (205-108), 1 lpF (205-107), 1 IM (205-103) 1 pM (205-

101), 5 L [UCLA]; same data (PA 206), 1 L [UCLA]. Nigua Creek, nr. Almirante, 27 Apr 1963, A.

Quinonez (PA 257), 1 lpM (257-113) [UCLA]; same data, 28 Apr 1963 (PA 277), 3 lpF (277-

105-107), 7 pM (277-102-104,108,111-113), 2 pF (277-109,110) [UCLA]; same data, 2 May

1963 (PA 300), 5 L [UCLA] . Canal Zone: Barbacoas Island, 14 Dec 1965, A. Quinonez (PA 904),

1 pM (904-104), 2 pF (904-100,103) [UCLA]. Barro Colorado Island, 7 May 1943, W. Komp,

3 IM (43-68,83,101), 3 IF (43-73,77,99), 2 Ip (43-103,108), 8 M, 10 F, 221 [UCLA, USNM];

same data, 21 May 1943, G. Fairchild, 7 IM (43-141,141,147,147,147,147,191), 3 lpF (43-201),

2 IF (43-157,159), 12 M, 1 M gen, 3 F, 6 1 [UCLA, USNM]; same data, 23 May 1943, W. Komp,

2 M, 2 F [UCLA]; same data, 31 May 1943, 2 IF (43-225,233), 1 M, 1 F, 51 [UCLA, USNM];

same data, 15 May 1945, 1 IpM (5-143), 2 IpF (5-142,143), 1 M [UCLA, USNM]; same data,

22 May, 2 IF (5-220,244) [UCLA, USNM]; same data, 23 May 1945, 2 IF (5-226,237) [UCLA] ;

same data, 26 May 1945, 1 IM (5-427), 1 F [UCLA, USNM]; same data, 3 Dec 1945, A. Quinonez

(PA 856), 1 lp (856-11), 1 F [UCLA] ; same data (PA 857), 2 IpM (857-10,15), 4 IpF (857-11,14),

3 pM (857-102,103,106), 8 pF (857,100,100a,101,104,105,108-110), 3 M, 2 F,6 p, 6 P [UCLA];

88 Contrib. Amer. Ent. Inst., vol. 10, no. 2; 1973

same data (PA 860), 1 pM (860-100), 1 1 [UCLA]; same data, 4 Dec 1965 (PA 861), 2 IpM (861-

17,20), 3 lpF (861-21,23,23a), 1 IP (861-22), 2 M, 2 F,3 p, 1 L [UCLA]. Camacho, 22 Apr 1922,

J. Shropshire, 1 F [USNM]; same data, 1 June 1922, 1 F [USNM]. Chagres Camp, 25 July 1921,

thru C. Ludlow, 1 M [USNM]. Chagres River, 2 June 1952, S. Carpenter, 2 F [UCLA] . Chagres

River at Pina, 4 Aug 1943, W. Komp, 1 pM (43-275) [UCLA]; same data, 5 Aug 1943, 1 Ip (43-

273), 1 1 [USNM]. Chiva Chiva, nr. Miraflores, May 1945, W. Komp, 1 M [UCLA]; same data,

11 Nov 1965, A. Quinonez (PA 768), 1 IpM (768-10), 2 IpF (768-21,22), 1 pM (768-100), 2 M,

2 P, 31 [UCLA]; same data (PA 771), 1 IpM (771-24), 1 IpF (771-25), 2 pM (771-21,23), 3 pF

(771-22,26,100), 3 M, 2 F, 3 p [UCLA]; same data (777), 1 F [UCLA]. Contractor’s Hill (0.8

km N), 13 Dec 1965, A. Quinonez (PA 898), 1 IpM (898-11), 1 pF (898-10), 2M, 2p,11,1L

[UCLA]. Corozal, 20 June 1920, J. Zetek, 1 F [USNM]; same data, 15 Jan 1943, W. Komp, 1 F

[UCLA]; same data, 29 Jan 1943, 1 M, 1 F [UCLA]; same data, 30 Jan 1943, 21 [UCLA] ; same

data, 2 Feb 1943, 1 1 [UCLA]; same data, 18 Feb 1943,2M [UCLA] ; same data, 31 Aug-7 Sept

1943, 41 [USNM] ; same data, 26 Mar 1945, 1 pM (5-63), 18 Ip (5-47,48,50,53,55,57,58,58,59,61,

62,65-67,82,82,84,84), 6 1 [UCLA, USNM] ; same data, 3-4 May 1945, 5 IpF (5-109,117,119,123,

125), 1 IF (5-121), 1 pM (5-118), 1 Ip (5-110), 7 F, 8 M, 1 p, 61 [UCLA, USNM] ; same data,

13 June 1945, 41 [UCLA, USNM]. Corozal Damsite, 2 Mar 1943, W. Komp, 2M, 1 F [UCLA];

same data, 6 May 1943, 1 IF (43-83), 3 M, 3 F [UCLA, USNM]; same data, 28 July 1943, 11M

(43-255), 1 F [UCLA]; same data, 18 Sept 1943, 4 IM (43-317A) [UCLA] ; same data, 12 Jan

1944, 1 M [UCLA]. Empire, 8 July 1922, J. Shropshire, 1 M gen [USNM]; same data, 7 Aug

1944 (ASM 91-1), 1 F [UCLA]. Farfan, 24 June 1949, 1 F [UCLA]. Ft. Clayton, 11 Nov 1944,

Adams and Van Doren (ASM 165-1), 1 L [UCLA]; same data, 28 Nov 1944, K. Frick (ASM 300-

3), 1 L [UCLA]; same data, 19 Dec 1951-12 June 1952, S. Carpenter, 8 F [UCLA]. Ft. Davis,

27 Dec 1951-10 Nov 1953, S. Carpenter, 5 F [UCLA]; same data, 22 Nov 1965, R. Schick and

A. Quinonez (PA 814), 1 IpF (814-10), 1 M, 1 p, 1 1 [UCLA]. Ft. Gulick, 17 Dec 1951, S.

Carpenter, 3 F [UCLA]. Ft. Kobbe, 4 June 1952, S. Carpenter, 2 F [UCLA]. Ft. Randolph,

9-19 July 1938, Richardson, 3 M, 5 F [UCLA]; same data, Aug 1938, W. Komp, 2 M gen

[USNM]; same data, 8 Aug 1939, 1 M, 6 F [UCLA]. Ft. Sherman, 3 M, 1 L [UCLA]; same

data, 5 May-4 Nov 1949, 11 M, 1 M gen, 10 F [UCLA, Utah]; same data, 2 May 1951 (GML),

1 IM (02121) [UCLA]; same data, 11 July 1956 (GML), 1 lpM (02975), 1 IpF (02963) [UCLA];

same data, 1916, L. Dunn, 1 F [USNM]; same data, 16 July 1920, J. Zetek, 2 F [USNM]. France

Field, 2 Aug 1972, H. Arnell and M. Boreham (PA 1157), 4 IpM (1157-10-12,18), 7 IpF (1157-13-

17,19,20) [UCLA]. Gatun, 15 Sept 1926, D. Curry, 1 lp [USNM]. Gatun Farm, 17,18 Aug 1953,

S. Carpenter, 10 F [UCLA]. Juan Mina (6 km NE Gamboa), 25 m, 18 Jan 1963, A. Quinonez

(PA 5), 1 lpM (5-105), 1 lpF (5-103), 2 IP (5-107,108), 1 P, 1 L [UCLA] ; same data, 20 July 1972,

H. Arnell and R. Hinds (PA 1138), 1 pF (1138-100) [UCLA]. Juan Mina (2 km W), 19 July 1972,

H. Arnell and R. Hinds (PA 1130), 1 pM (1130-100), 1 1 [UCLA]. Largo Remo Island, 27 July

1926, D. Curry, 2 M, 2 F [UCLA, USNM]. Lion Hill, A. Busck, 1 M, 2 F [USNM]. Madden Forest

Preserve, George Green Park (0.5 km W), 100 m, 10 July 1972, H. Arnell (PA 1096), 5 IP (1096-11-

15), 1 L [UCLA]. Madden Forest Preserve, Las Cruces Trail, 100 m, 16 Nov 1965, A. Quinonez

and Moody (PA 792), 2 IpF (792-10,12) [UCLA]; same data, 10 July 1971, H. Arnell (PA 1098),

1 IpM (1098-14), 2 IpF (1098-11,13), 1 pM (1098-100), 2 IP (1098-10,12), 2 L [UCLA] . Margarita,

14 June 1922, J. Shropshire, 1 M [USNM]; same data, 8 Oct 1964, A. Quinonez (PA 714), 1 IpF

(714-12), 1 IP (714-16), 4 L [UCLA]; same data (PA 716), 1 lpM (716-10) [UCLA]. Miraflores

(Hwys C-2, C-21), 10 Nov 1965, R. Schick and A. Quinonez (PA 767), 1 IpF (767-10), 2 pF (767-

100,101), 1 F, 2 L [UCLA] . Mojinga Swamp, 10 m, 8 July 1949, 1 lp [UCLA] ; same data, 21 June

1934, W. Komp, 1 1 [USNM]; same data, 8 July 1949, L. Rozeboom (PAR 9.1-9.3), 2 M, 1 F

[UCLA]; same data, 13 Oct 1964, A. Quinonez (PA 721), 4 pM (721-11,14,15,100), 2 pF (721-

10,101), 2 L [UCLA]; same data (PA 724), 1 lpM (724-10), 1 pM (721-102), 1 pF (721-11); same

data (PA 727), 2 pM (727-101,103) [UCLA] ; same data, 29 July 1972, H. Arnell (PA 1153), 1 IpF

(1153-20), 2 pM (1153-100,101), 2 L [UCLA]. Mt. Hope, 12 July 1922, J. Shropshire, 1 M, 1M

gen, 1 F [USNM]. Summit, 11 Oct 1939, W. Komp, 1 M [UCLA]; same data, 1-3 Nov 1939, 1 M,

2 F [UCLA]; same data, 12 Nov 1943, 21 [UCLA]; same data, 30 Aug 1944, 1 lp [UCLA] ; same

Arnell: Genus Haemagogus 89

data, 25 May 1945, 21F, 1 M [UCLA]; same data, Nov 1946, N. Krauss, 1 F [UCLA]. Tabernilla,

A. Busck, 3 M, 2 F [UCLA, USNM]; same data, 24 July 1908, A. Jennings, 7 M, 1 M gen, 20 F, 2 p

[USNM]; same data, 30 July 1908, 1 M, 1 M gen, 1 F [USNM]; same data, 22 Dec 1908, 1 M, 1 M

gen [USNM]; same data, 14 Apr 1909, 1 M, 1 M gen, 1 F [USNM]. Toro Point (Ft. Sherman),

7 Jan 1922, J. Shropshire, 1 M [USNM]. Atlantic side (locality unspecified), June 1938, W. Komp,

5 M, 4 F [UCLA]. Canal Zone (locality unspecified), A. Jennings, 5 F [USNM]. Cocle: El Valle,

600 m, 4 Sept 1938, W. Komp, 5 F [UCLA]; same data, 5 June 1945, 1 IM (5-322), 1 IF (5-330),

1 pM (5-371A), 1 lp (5-387), 2 M, 1 1 [UCLA, USNM]; same data, 5 June 1945, R. Arnett and

W. Komp (ASM 613), 3 M, 10 F [UCLA]; same data (ASM 615), 2 M, 4 F [UCLA]; same data,

10 July 1949, L. Rozeboom (PAR 13.2, 13.4), 2 M, 1 F [UCLA]; same data, 13 Aug 1963, A.

Quinonez (PA 502), 1 IpF (502-101), 1 IF (502-102), 1 IP (502-103), 6 L [UCLA]; same data

(PA 507), 4 L [UCLA] ; same data, 19 Aug 1963 (PA 522), 2 IpM (522-101,102) [UCLA] . Colon:

Buena Vista, 28 Sept 1949, 1 M, 1 F [UCLA]; same data, 24 Sept 1964, A. Quinonez (PA 710),

1 IpF (710-12), 1 pM (710-101), 1 L [UCLA]. Cativa, 18 Nov 1949, 1 M, 6 F [UCLA] ; same data,

22 Nov 1965, R. Schick and A. Quinonez (PA 807), 1 lpF (807-10), 1 pM (807-100), 2 F,1 L

[UCLA]. Pina, 11 July 1956 (GML), 3 IpF (02961,02970,02972), 1 pM (02962); same data,

30 Nov 1963, A. Quinonez (PA 578), 2 lpM (578-101,102), 2 IpF (578-103,104), 1 IP (578-105),

17 L [UCLA]. Portobelo (Caldera Island), 12-16 Aug 1908, A. Jennings, 1 M, 1 M gen [USNM].

Portobelo, 23 May 1908, A. Jennings, 1 F [UCLA]; same data, 13 Mar 1911, A. Busck, 1 F

[UCLA] ; same data, 4 Dec 1963, A. Quinonez (PA 581), 1 p, 5 L [UCLA] ; same data, 9 Dec 1963

(PA 600), 1 pM (600-101), 3 pF (600-102,104,107), 1 IP (600-106), 33 L [UCLA]; same data

(PA 601), 1 IpM (601-103), 2 L [UCLA]. Salud, 11 July 1956 (GML), 1 lpM gen (02967), 2 pM

(0295 1,02952) [UCLA] . Darien: El Real, H.Kumm, 1 F [UCLA]. La Palma, 120 m, 26 Nov 1966,

O. Berlin (PA 951), 2 IpM (951-10,11), 1 pM (951-114) [UCLA]; same data, 60 m, 7 Dec 1966,

O. Berlin and R. Hinds (PA 988), 1 IpF (988-10) [UCLA]; same data (PA 991), 1 IpM (991-10)

[UCLA]. Paya, 25 Feb 1958 (GG 1), 4 lpM (1-103,105,106,109), 6 IpF (1-101,107,108,110,112,

116), 1 pM (1-121) 4 pF (1-104,111,113,123), 1 F [UCLA]; same data, 21 Feb 1958 (GG 1),

1 IpF (1-131) [UCLA]. Pucro, 8 July 1958 (GG 109), 2 P [UCLA]. (?Rio) Tuira, 26 Feb 1958

(GG 31), 3 M [UCLA]. Rio Tuira at mouth of Rio Paya, 1 Mar 1958 (GG 49), 4 M [UCLA] ; same

data, 2 Mar 1958 (GG 59), 2 M [UCLA]; same data, 3 Mar 1958 (GG 69), 2 M [UCLA];

Panama: Cerro Azul, May-Dec 1954 (GML), 8 1 [UCLA]. Cerro Campana, 760 m, 28 Aug

1963, A. Quinonez (PA 537), 1 M [UCLA]; same data, 29 Aug 1963 (PA 539), 1 F [UCLA];

same data (PA 540), 5 F [UCLA]. Cerro La Victoria, 1949, H. Trapido, 1 M, 22 F [UCLA].

El Capesito (mr. San Carlos), 150 m, 13 Aug 1963, A. Quinonez (PA 503), 1 pF (503-126)

[UCLA]. El Llano, Rio Bayano, July 1942, H. Kumm, 1 M gen [UCLA]; same data, 1943,

6 M [UCLA]. La Chorrera, 4 July 1944 (ASM 18), 1 L [UCLA]. Lagarto (nr. Bejuco), 8 June

1945, R. Arnett (ASM 619), 3 M, 3 F [UCLA]; same data, 9 June 1945 (ASM 624), 2 M,

6 F [UCLA]; same data, 12 Aug 1963, A. Quinonez (PA 500), 1 lpF (500-103), 3 IP (500-101,

102,104) [UCLA]. La Joya, (?La Jolla, nr. Pacora) 21 June 1944, P. Adams (ASM 9-1), 2 F

[UCLA]. La Zumbadora (nr. Cerro Azul), 600 m, 15 Feb 1963, A. Quinonez (PA 94), 1 P,

1 1, 1 L [UCLA]; same data, 18 Feb 1963 (PA 114), 1 L [UCLA]. Nuevo Emperador, 23 Nov

1964, A. Quinonez (PA 824), 1 M, 1 P [UCLA]; same data (PA 825), 1 lpF (825-10), 2 M,

2 F, 2 p, 2 1 [UCLA]. Pacora, 9 June-13 Sept 1949, 2 F [UCLA]; same data, 10 Nov 1953,

S. Carpenter, 5 F [UCLA]. Paja (Nuevo Emperador), 15 Apr 1941, G. Fairchild, 1 F [USNM].

Panama, 1 F [USNM]; same data (GML), 2 lpM (02124,02125) [UCLA] ; same data, A. Jennings,

1 Ip [USNM]; same data, 10 May 1941, Middlecauff, 2 L [UCLA]. Rio Pequini, patio del

canpamento, 10 Dec 1965, J. Mena (PA 924), 1 IpF (924-10), 1 L [UCLA]. Taboga Island,

A. Busck, 1 F [USNM]. Tocumen, 1 June 1949, S. Carpenter, 1 M [UCLA] . Province unknown:

Cabima, 24,26 May 1911, A. Busck, 2 F [USNM]. Locality unknown: W. Komp, 1 M, 1 F, 41

[UCLA, USNM].

Additional Record From the Literature

COLOMBIA. Cordoba: Monteria (Kumm, Osorno-Mesa and Boshell-Manrique, 1946:20).

90 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

22. Haemagogus (H.) argyromeris Dyar & Ludlow

Piss 6.51.52

1921. Haemagogus argyromeris Dyar and Ludlow, 1921:679-680. TYPE: Lectotype male with

genitalia slide (1456), Corozal, Canal Zone, Panama, larva taken in a “container”,

27 Oct 1920 [USNM; selection of Stone and Knight, 1955:287].

1921. Haemagogus (Haemagogus) gladiator Dyar, 1921a:108-109. TYPE: Holotype male (39)

with genitalia slide (1488), Corozal, Canal Zone, Panama, larva from “‘tree hole near

Kraft’s house”, 30 Nov 1909, A.H. Jennings [USNM, 24340]. Synonymized with

argyromeris by Dyar (1925:139-140).

Haemagogus (Haemagogus) argyromeris of Dyar (1921a:109, in part; 1923:183, in part; 1928:

137-138); Bonne and Bonne-Wepster (1925:434, in part); Edwards (1932:179); Lane (1953:

786-787); Stone, Knight and Starcke (1959:217); Forattini (1965 :43-45).

Haemagogus argyromeris of Dyar (1925b:139-140); Arnett (1949:239; 1950:107); Komp (1949:

72; 1955:179-180); Galindo, Trapido and Carpenter (1950:546); Galindo, Carpenter and

Trapido (1951a:118-119; 1955:158); Carpenter, Galindo and Trapido (1952:160); Trapido,

Galindo and Carpenter (1955:306); Trapido and Galindo (1956a:306).

Haemagogus (Haemagogus) gladiator of Dyar (1923:183).

Haemagogus regalis in part of Busck (1908:64).

Haemagogus albomaculatus in part of Howard, Dyar and Knab (1917:870).

Stegoconops albomaculatus in part of Howard, Dyar and Knab (1913:fig. 439).

FEMALE. Wing: 3.05 mm. Proboscis: 2.05 mm. Forefemur: 2.55 mm. Abdomen:

3.30 mm. Dark scales of proboscis, palpus, wing and legs dark blue to violet with

purple reflections. Head: Eyes widely separated above antennae (4 ommatidial di-

ameters). Decumbent scales of vertex and occiput greenish blue to bluish green,

more bluish green laterally. Proboscis short, 0.90-0.95 of forefemur; palpus about

0.20 of proboscis; antenna about 0.71 of proboscis. Thorax: Scales of mesonotum

bronze to greenish blue, blue to violet in fossa, bluish green over supraalar bristles,

silver in antealar area; scales on scutellum bluish green. Apn lobes with scales

greenish blue; ppn bare. Legs: Length of forefemur about 1.53-1.59 of distance

from top of thorax to apex of midcoxa. Wing: R43 about 0.35-0.55 of R,.

Abdomen: Dark scales blue to violet; silver scales in basal bands on tergites V

or VI-VIII, occasionally IV-VIII, often continuous with basolateral silver patches

on VI and VII.

MALE (fig. 51). Wing: 2.30 mm. Proboscis: 1.85 mm. Forefemur: 1.80 mm.

Abdomen: 2.90 mm. Head: Proboscis about 0.95 of forefemur; antenna mod-

erately plumose, flagellar segment 4 with 16-20 bristles.

MALE GENITALIA (fig. 51). Segment VIII: Tergite about 0.75 of sternite;

distal margin acclivous toward midline, with well developed setae, more numerous

mesally, and scales of two types, a few elongate, lanceolate scales near midline and

enlarged cuneate scales along entire margin. Segment IX: Tergite unsclerotized on

midline. Sidepiece: Conical, truncate, length about 3.0 times median width; basal

tergomesal area slightly enlarged, extending distad as slightly raised area to distal

third of sidepiece, with numerous setae which are short distally, becoming longer

proximally, the more proximal ones much enlarged, flattened, attenuate; apical

lobe well developed, bearing numerous moderately large setae; apical sternomesal

scales lanceolate to oblanceolate with acuminate tips. Claspette: Stem bowed in-

ward apically in dorsal aspect, narrow, curved dorsad on basal two-thirds, apical

third expanded posteriorly into spatulate knob which is spiculose posteriorly and

with a seta on posterior margin; filament a striate, narrow, convoluted leaf forming

Arnell: Genus Haemagogus 91

a beaklike anterior process, inserted apically on stem. Clasper: Curved, flattened,

gradually expanded to apex which is more or less bulbous; apical spiniform about

0.25 of clasper, flattened, peglike, inserted subapically on inner surface of clasper.

Phallosome: Aedeagus moderately large, broadly obovate; apex produced dorsad

into serrated carina terminating in beaklike process; venter broadly open except

on basal third, with sclerotized ridges lateroventrally which are expanded basally.

Proctiger: Cercal setae 4; apical knob of paraproct with about 15-20 serrations.

PUPA (fig. 51). Abdomen: about 3.20 mm. Trumpet: 0.35 mm. Paddle: 0.65

mm. Cephalothorax: Weakly pigmented, slightly darker dorsally. Hairs 4,5-C sub-

equal, single or double, weaker than 7-C which is single. Trumpet: Light brown,

reticulate sculpturing rather weak. Abdomen: Weakly to moderately pigmented,

genital lobe and anterior segments often darker. Float hair (1-I) with about 8-10

primary branches and 2-6 secondary branches. Hair 1-II-VII weakly developed,

single or double, more or less subequal on all segments. Hair 2-VII short, within

own length of 1-VII. Hair 3-II,III usually subequal or with 3-III slightly weaker,

and slightly weaker than 5-IV,V which are usually subequal. Hair 6-VII weak,

subequal to 6-III-VI. Hair 9-VII often somewhat cephalad of caudolateral margin

of tergite, single or double; 9-VIII 5-7b. Terminal Segments: Male genital lobe about

1.2 of tergite VIII. Paddle: Weakly pigmented; relatively broad, with broadly round-

ed apex. Hair 1-P double (single or double).

LARVA (fig. 52). Head: 0.75 mm. Siphon: 0.70 mm. Anal Saddle: 0.33 mm.

Segment VIII: Comb scales 23-27(16-23), with single broadly rounded, minutely

fringed spine, in irregular triple row. Siphon: Index about 2.6-2.8(1.5-3.0). Pecten

teeth 10-12(8-13), extending to about basal 0.50-0.55 of siphon. Hair 1-S 3-4b.

Anal Segment: Hair 4a-X 3-4b.

SYSTEMATICS. Haemagogus argyromeris can be distinguished from the other

species of the Splendens Section: in the adults by (1) bare ppn, (2) short proboscis,

about 0.90 of forefemur, (3) dark coxal integument, (4) dark scales of abdomen

blue or violet with dorsal silver scales at bases of tergites V to VII; in the male

genitalia by (1) clasper flattened, curved and expanded in width to apex and (2)

claspette stem expanded into spatulate knob on distal third; in the pupa by

usually double hair 1-P; and in the larva, from all species except iridicolor, regalis

and lucifer, by the combination of (1) usually 23 to 27 comb scales, (2) 3-or

4-branched hair 4a-X, (3) pecten extending to about basal 0.50 or 0.55 of siphon,

(4) hair 7-III-VI subequal to hair 5 of corresponding segment and (5) relatively long

spines on caudal margin of saddle.

The affinities of argyromeris are not clear, although close resemblance of the

larva and similarity in the male genitalia to the regalis complex probably indicates

argyromeris to be an early offshoot of this species complex. Its extremely limited

distribution in the Panamanian Isthmus and abundance in marginal habitats suggest

that argyromeris is a relict species, able to survive competition with other species

of Haemagogus because of its ability to utilize habitats not available to the others.

BIONOMICS. Haemagogus argyromeris is a mosquito of second growth and open

tropical deciduous forest, rarely found in primary forest. It is more closely associ-

ated with man than any other Haemagogus species and it utilizes the greatest

variety of habitats: treeholes, bamboo, fallen fruits, terrestrial bromeliads, artifi-

cial containers and occasionally rockholes and ground pools. Many eggs do not

hatch when first flooded, consequently populations are slow in building up after

rains. Females apparently attack man readily only in the absence of other preferred

hosts and usually bite low on the body. This species is probably not involved in

92 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

the transmission of sylvan yellow fever because of its absence in forest situations.

DISTRIBUTION (fig. 6). Haemagogus argyromeris extends along the Pacific

coast from western to central Panama, crossing to the Atlantic at the Panama

Isthmus. Material examined: 1552 specimens; 234 males, 839 females, 158 pupae,

321 larvae; 153 individual rearings (65 larval, 10 pupal, 78 incomplete).

PANAMA AND CANAL ZONE. Canal Zone: Ancon, 23 Nov 1907-22 July 1908, A. Jennings,

5 M, 4 F [USNM]. Barro Colorado Island, 31 May 1943, W. Komp, 2 M, 2 F [UCLA]. Bella Vista,

7 July 1926, D. Curry, 1 F [USNM]; same data, 16 Aug 1943, H. Crowell (Komp) 2 IF (43-277,

278), 2 lp (43-276), 1 p [UCLA, USNM]. Camacho, 15-22 June 1922, J. Shropshire, 7 M, 1 F

[UCLA]. Chiva Chiva, 10 Nov 1965, A. Quinonez (PA 764), 2 IpM (764-20,21), 1 IpF (764-22);

same data, 11 Nov 1965 (PA 768) 1 lpM (768-20); same data (PA 773), 2 F, 2 p; same data (PA

770), 1 P [UCLA]. Cocoli Naval Hospital, 25 June 1945, W. Komp, 1 1M (5-417), 4 IF (5-417,

417,424,425), 1 F [UCLA, USNM] . Corozal, 30 Nov 1907, A. Jennings, 1 IpF (39.3), 1 IM (38.2),

2 IF (38.4,39.8), 1 M [USNM]; same data, 27 Oct 1920, C. Ludlow, 1 F [USNM]; same data,

July 1941, W. Komp, 41 [UCLA]; same data, 14 Aug 1941, 13 M [UCLA]; same data, 22 Sept

1941, 1M, 1 F[UCLA]; same data, 8 Jan 1943, 1 M, 21 [UCLA, USNM]; same data, 29 Jan 1943,

9 M, 7 F [UCLA]; same data, 18 Feb 1943, 4 M, 6 F [UCLA]; same data, 6 June 1943, 1 M gen

[UCLA] ; same data, 30 Aug 1943, 1 lpM gen (43-286), 2 IM (43-284,288), 4 IF (43-290,290,290,

296), 5 lp (43-279,282,282,282,283), 2 F, 71 [UCLA, USNM]; same data, 6, 7 Sept 1943, 3 Ip

(43-309), 14 1 [UCLA, USNM]; same data, 1943, 3 M [UCLA] ; same data, 14 July 1944, 6p, 51

[UCLA, USNM]; same data, 21 Feb 1945, 13 lp (5-8-12,14-19, [20,23], [22,27]), 8 p, 191, 14L

[UCLA, USNM]; same data, 13,14 Mar 1945, 18 L [UCLA]; same data, 26 Mar 1945, 2 lp (5-68,

69), 2 p, 41, 4 L[UCLA, USNM] ; same data, 3, 4 May 1945, 3 pM (5-103), 1 IF (5-103), 1 F[UCLA,

USNM]; same data, 13 June 1945, 2 IpM (5-368,372), 2 IpF (5-283,283) 2 IM (5-335,376), 1 M,

1 F, 1 p [UCLA, USNM]; same data, 23 June 1945, 1 IF (5-405), 21 [UCLA, USNM] ; same data,

21 Dec 1943, H. Crowell (Komp), 2 M, 1 F [UCLA]. Corozal Damsite, 28 July 1943, W. Komp,

1 M [UCLA]. Empire, 2 July 1921, C. Ludlow, 1 M [USNM]; same data, 8 Aug-6 Oct 1921,4M

[USNM]; same data, 6 July 1922, J. Shropshire, 4 M, 1 F [USNM]. Ft. Amador, 14 Sept 1949,

1 IF [UCLA]; same data, 4 Dec 1949, 2 M, 4 F [UCLA]; same data, 5 Jan 1950, 53 F [UCLA];

same data, 9 Jan 1950, 1 M [UCLA]; same data, 27 Sept 1949 (GML), 2 Ip (03394,03411)

[UCLA]; same data, 27 Nov 1949 (GML), 2 IpF (00444,00445), 1 lp (00446) [UCLA]; same

data, 2 Dec 1949-12 Jan 1950, S. Carpenter, 1 M, 551 F [UCLA]; same data, 18-29 Dec 1949,

J. Duncan, 16 F [Utah]. Ft. Clayton, 13 Jan, 1 F [USNM]; same data, 24 Apr 1944, P. Adams,

1 L [USNM]; same data, 7 Oct 1944, K. Frick (ASM 200), 1 M, 1 L [UCLA]; same data, 5 Nov

1944 (ASM 256), 1 F,3 L [UCLA] ; same data, 28 Nov 1944 (ASM 301), 5 M, 3 F [UCLA] ; same

data, 17 Dec 1944, (ASM 346), 1 L [UCLA]; same data (ASM 347), 10 M, 5 F [UCLA]; same

data, 7 Jan 1945 (ASM 367), 9 M, 3 F [UCLA]; same data, 29 Jan 1945 (ASM 386), 2 M, 1 F

[UCLA] . Frijoles, 1 Dec 1965, R. Schick and A. Quinonez (PA 844), 1 p [UCLA]. Gatun, 25 Aug

1926, 51; same data, 20 Oct 1926, 5 F; same data, 22 July 1908, A. Jennings, 1 M; same data, Dec

1913, H. Trask, 1 M [USNM]. La Pita, 20 June 1921, C. Ludlow, 1 M [USNM]. Las Cascadas,

A. Jennings, 2 M, 2 F [USNM]. Matachin, 5 May 1908, A. Jennings, 1 F [USNM]. Miraflores,

9 May 1908, A. Jennings, 1 M, 1 M gen, 3 F [USNM]; same data, 7 Jan 1922, J. Shropshire, 2 M,

1 F [UCLA]; same data, 10 Nov 1965, R. Schick and A. Quinonez (PA 767), 1 lpF (767-12),

1 pM (767-102) [UCLA]. Miraflores Lake, 8 Nov 1965, R. Schick and A. Quinonez (PA 762),

2 IpF (762-14,16), 1 pM (762-100), 2 M, 3 p, 1 P, 21 [UCLA]. Mojinga Swamp, 13 Oct 1964,

A. Quinonez (PA 723), 1 P [UCLA]. Monte Lirio, 14 Jan 1922, J. Shropshire, 1 F [UCLA]. Mt.

Hope, 5 June 1943, W. Komp, 1 M [UCLA]. San Pablo, 14 May 1908, A. Jennings, 1 F [USNM].

Paraiso, 1 M gen, 3 P, 2 L [UCLA, USNM]; same data, J. Zetek, 1 M, [USNM]. Rio Cardenas,

3 Mar 1922, J. Shropshire, 1 F; same data, 12 Nov 1965, A. Quinonez (PA 783), 1 F, 1 P; same

data (PA 785), 1 lpM (785-10), 1 M, 1 p, 11 [UCLA]. Rio Chagres, A. Busck, 1 M [USNM]. Rio

Chagres, 30 Dec 1931, 1 F [UCLA]. Summit, 25 May 1935, W. Komp, 1 M gen [USNM]; same

data, 26 May 1935, 2 M, 2 F [UCLA]; same data, May 1935, 21 [USNM]; same data, 3 Nov 1937,

11 [USNM] ; same data, 12 Sept 1939, 1 M, 2 F [UCLA]; same data, 27 Sept 1939, 1 M, [UCLA];

same data, 1 Oct 1939, 2 F,41 [UCLA, USNM] ; same data, 11 Oct 1939, 11 [USNM] ; same data,

Arnell: Genus Haemagogus 93

1 Nov 1939, 1 M, 1 F [UCLA]; same data, 3 Nov 1939, 1 M [UCLA]; same data, 17 Aug 1941, 1

IM, 1 IF, 1 F, 1 1 [UCLA, USNM]; same data, 12 Jan 1943, 1 F [UCLA]; same data, July 1946,

N. Krauss, 1 M, 1 F [USNM]. Tabernilla, A. Busck, 1 M, 2 F [USNM]. Atlantic side (locality un-

specified), June 1938, W. Komp [UCLA]; same data, 20 June 1945, 1 lpM (5-404), 2 IF (5-403),

1 M, 1 F [UCLA, USNM]; same data, 1945, Henderson, 1 1 [UCLA]. Pacific side (locality unspeci-

fied), July 1944 (ASM 24-6), 2 L [UCLA]. Locality unspecified, A. Jennings, 2 F [USNM].

Locality unspecified, Feb 1944, 1 F [USNM]; same data, 1916, Dunne, 1 M gen [USNM]; same

data, Jan 1945, S. Carpenter, 3 M, 7 F [UCLA]. Chiriqui: David, F. Snyder, 1 F [UCLA]. Cocle:

El Valle, 600 m, 31 Aug 1938, W. Komp, 1 M, 1 F [UCLA]; same data, 8 Oct 1939, 2M [UCLA];

same data, 5 June 1945, 4 IpM (5-274,340,349,377), 11 IpF (5-298,312,320,321,341-343,346,

348,353,354), 5S IM (5-361,373,373,373,400), 13 IF (5-291,313,324,333,344,388,394,395,397,

407,407,407,407), 1 Ip (5-379), 1M, 3 F, 61 [UCLA, USNM] ; same data, 5 June 1945, R. Arnett

and W. Komp (ASM 613), 1 F [UCLA]; same data (ASM 615), 1 M [UCLA]. Colon: Buena Vista,

24 Sept 1964, A. Quinonez (PA 710), 4 IpM (710-13,30,31,38), 2 lpF (710-17,39), 1 pF (710-

102), 18 L [UCLA]. Pina, 2 Sept 1930, D. Curry, 2 M, 2 F [UCLA]; same data, 5 Aug 1943,

W. Komp, 1 p [USNM]. Los Santos: Tonosi, 5 May 1949 (GML), 1 M [UCLA]. Panama: (?Cerro)

Campanita, 21 Sept 1950, 1 lp [UCLA]. Chame, 8-11 Oct, M. Matthes, 1 lpM, 1 F, 51 [UCLA,

USNM]. Chepo, 15 Oct 1939, W. Komp, 3 M, 2 F [UCLA]. El Libano, 22 m, 9 Aug 1963, A.

Quinonez (PA 487), 2 IpM (487-102,103), 1 pM (487-104), 1 lp (487-101), 6 L [UCLA]. El

Victoria (?7La Victoria), 29 June 1949, S. Carpenter, 1 F [UCLA]. Juan Diaz, 4 Nov 1965, R.

Schick and A. Quinonez (PA 747), 1 pF (747-100) [UCLA]. La Chorrera, 21 Oct 1944,Van Doren

(ASM 277), 1 M [UCLA]. Nueva Gorgona, 20 m, 13 Dec 1966, O. Berlin (PA 1003), 2 M, 1 F,3 p,

6. P [UCLA]. Nuevo Emperador, 23 Nov 1965, A. Quinonez (PA 825), 2 F, 4 p [UCLA]. Pacora,

19 Nov 1939, M. Matthes, 1 M, 1 F [UCLA]; same data, 24 Oct 1949 (GML), 21 [UCLA]. Panama,

A. Jennings, 2 M gen, 1 F [USNM]; same data, 1 July 1926, D. Curry, 7 L [USNM]; same data, 5

May 1944 (ASM 5-1), 5 M, 1 L [UCLA]. Panama, Paitilla Pt., 5 Aug 1928, H. Dyar and R.

Shannon, 1 M [USNM] ; same data, 14 May 1939, W. Komp, 1 M, 1 F [UCLA]; same data, 11 Sept

1939, 5 M, 1 F [UCLA]; same data, 1939, 3 F [UCLA]; same data, 24 Aug 1941, 1 M [UCLA];

same data, 28 May 1958 (GML), 1 IpM (03389), 1 IpM gen (03384), 2 IpF (03213,03382), 1 pM

(03386), 1 L [UCLA]. Panama Viejo, 23 Oct 1934, L. Rozeboom (PAR 67), 1 M, 2 F; same data

(PAR 68), 1 M, 1 F; same data, 27 July 1935 (PAR 82), 1 M, 2 F; same data, 16 Oct 1944, Adams

and Van Doren (ASM 205-1), 1 L [UCLA]. Rio Teta, San Carlos, 22 Aug 1963, A. Quinonez

(PA 531), 5 IP (531-118-122); same data, 31 Aug 1963 (PA 546), 6 IpM (546-101,102,104-106,

109), 6 IpF (546-107,110,111-114), 1 pM (546-108), 1 IP (546-103), 7 L [UCLA]. Taboga Island,

A. Busck, 1 M gen, 3 F [USNM]; same data, A. Jennings, 1 M [USNM]. Tocumen, 6 Sept 1963,

A. Quinonez (PA 549), 4 IpM (549-101 ,103-105), 4 IpF (549-106-109) [UCLA] . Locality unspeci-

fied: 1 M, 1 F [USNM]; 1 M [UCLA]; J. B. Shropshire, 2 F [USNM]; L. H. Dunn, 1 lp [USNM];

(ASM 93), 1 F [UCLA]; (ASM 99-1), 1 M [UCLA]; (ASM 182), 1 F [UCLA]; L. Rozeboom

(PAR 136), 3 M, 2 F [UCLA]; same data, 23 Nov 1934 (PAR 72), 8 M, 9 F [UCLA].

23. Haemagogus (H.) chalcospilans Dyar

Figs. 6,53,54

1921. Haemagogus (Haemagogus) chalcospilans Dyar, 1921a:110-112. TYPE: Holotype male

(247) and genitalia slide (1481), Caldera Island, Portobelo Bay, Colon, Panama, larva

from salt pools in rocks near seacoast, 22 Mar 1908, A.H. Jennings [USNM].

Haemagogus (Haemagogus) chalcospilans of Dyar (1923:183; 1928:139-140); Bonne and Bonne-

Wepster (1925:434); Edwards (1932:179); Lane (1939:119; 1953:782-784); Levi-Castillo

(1951b:11,25-26), Stone, Knight and Starcke (1959:217); Forattini (1965:45-47); Galindo

and Trapido (1967:107).

Haemagogus chalcospilans of Dyar (1925b:140); Kumm, Komp and Ruiz (1940:398); Kumm,

94 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Osorno-Mesa and Boshell-Manrique (1946:19); Woke (1947:365); Arnett (1949:239; 1950:

107), Galindo, Trapido and Carpenter (1950:546); Galindo, Carpenter and Trapido (195la:

119; 1955:163); Carpenter, Galindo and Trapido (1945:160); Barreto Reyes (1955:78); Komp

(1955:181); Trapido and Galindo (1956a:305).

Haemagogus splendens in part of Howard, Dyar and Knab (1917:867).

FEMALE. Wing: 2.70 mm. Proboscis: 1.70 mm. Forefemur: 2.05 mm. Abdo-

men: 2.80 mm. Dark scales of proboscis, palpus, wing and legs purple with some

violet reflections. Head: Eyes broadly separated above antennae (4 ommatidial

diameters). Decumbent scales of vertex and occiput purple. Proboscis short, about

0.80-0.90 of forefemur; palpus about 0.18 of proboscis; antenna about 0.74-0.77

of proboscis. Thorax: Scales of mesonotum copper to bronze, conspicuously purple

or occasionally dark blue in fossa and blue to greenish blue near scutal angle,

bluish green over supraalar bristles, silver in antealar area; scales on scutellum

bluish green. Apn lobes with purple scales and a few silver scales mesally; ppn

bare. Legs: Integument of apex of coxae and trochanters yellow; length of fore-

femur about 1.65-1.80 of distance from top of thorax to apex of midcoxa. Wing:

R243 about 0.33-0.40 of R,. Abdomen: Dark scales purple to violet with some

copper reflections; silver scales dorsally on tergites V or VI and VII in rather

broad basal band, often continuous with lateral silver patches.

MALE (fig. 53). Wing: 2.30 mm. Proboscis: 1.80 mm. Forefemur: 1.95 mm.

Abdomen: 2.90 mm. Head: Proboscis short, about 0.90-0.95 of forefemur; antenna

sparsely plumose, flagellar segment 4 with 8 moderately long bristles. Legs: Larger

claw of foreleg with short, blunt median tooth; smaller claw of foreleg and claws

of midleg simple.

MALE GENITALIA (fig. 53). Segment VIII: Tergite short, about 0.60 of

sternite, distal margin broadly, shallowly emarginate, straight or acclivous mesally,

with dense setae, those laterad much elongate and bent mesad at apex, those mesad

shorter, 0.25-0.67 of lateral setae. Segment IX: Tergite broadly emarginate and

weakly sclerotized mesally. Sidepiece: Conical, truncate, length about 2.5 times

median width; basal tergomesal lobe slightly enlarged, extending distad as raised

area to near distal third of sidepiece, with numerous setae, the distal ones short,

becoming longer proximally, the more proximal ones elongate and slightly flattened

basally; apical lobe prominent, bearing about 20 moderately developed setae; apical

sternomesal scales of several types, a more dorsal, dense cluster of nearly circular and

lanceolate scales oriented more or less dorsad, and ventrally narrow sinuous scales

proximally and large obovate to lanceolate scales distally oriented mesad; a cluster

of moderately developed setae just distad of sternomesal scales. Claspette: Stem

bowed inward near middle in dorsal aspect, curved dorsad, narrowed slightly on

_ basal third, expanded ventrally on apical 0.20 into large rounded or sharply pointed

knob, often with serrations on point, usually with a large seta ventrally at base of

knob; filament a convoluted membranous flap, beak shaped anteriorly, attached

at apex of stem. Clasper: Rounded, broadest at base and somewhat sinuous, a

small seta inserted on inner surface proximad of spiniform; spiniform short, cylin-

drical, inserted apically. Phallosome: Aedeagus moderately large, obovate, tip pro-

duced dorsad into serrated carina terminating in beaklike process; venter broadly

open except on basal half, with sclerotized ridges lateroventrally. Proctiger: Cercal

setae 4-6; apical knob of paraproct with 10-15 serrations laterally and conspicuous

knob mesally.

PUPA (fig. 53). Abdomen: about 2.95 mm. Trumpet: 0.30 mm. Paddle: 0.60

mm. Cephalothorax: Weakly to moderately pigmented, darker dorsally. Hair 5-C

Arnell: Genus Haemagogus 95

moderately developed, 2-4b, stronger than 4-C and subequal to 7-C, which is

single or double. Trumpet: Light to medium brown, reticulate sculpturing moder-

ate. Abdomen: Weakly to moderately pigmented, genital lobes and anterior seg-

ments slightly darker. Float hair (1-I) with 12-20 primary branches and 2-6 second-

ary branches. Hair 1-II-VII weakly to very strongly developed, single, multiple or

dendritic, usually becoming weaker on posterior segments. Hair 2-VII short, con-

siderably cephalad of 1-VII. Hairs 3-II,II,5-IV,V, subequal, with 3-III somewhat

weaker, or 3-II,III subequal and weaker than 5-IV,V, which are subequal. Hair 6-

VII usually weakly developed, subequal to 6-III-VI, single or double. Hair 9-VII

moderately to considerably cephalad of caudolateral margin of tergite, 2-6b; 9-VIII

8-12b. Terminal Segments: Male genital lobe about 1.5 of tergite VIII. Paddle:

Weakly pigmented; relatively broad, apex broadly rounded. Hair 1-P 3-4b.

LARVA (fig. 54). Head: 0.50 mm. Siphon: 0.65 mm. Anal Saddle: 0.33 mm.

Abdomen: Hair 7-III-VI weaker than hair 5 of corresponding segment. Segment

VIII: Comb scales 45-55(40-75), with single spatulate, minutely fringed spine;

in large triangular patch, distal scales considerably larger than proximal scales.

Siphon: Index about 2.4(1.9-2.6). Pecten teeth 12-17(10-21), extending to about

basal 0.45-0.50 of siphon. Hair 1-S 4-5b. Anal Segment: Spines on caudal margin

of saddle reduced, very short, each with multiple teeth. Hair 4a-X 6-9b.

SYSTEMATICS. Haemagogus chalcospilans can be readily distinguished from the

other species of the Splendens Section: in the adults by (1) bare ppn, (2) short

proboscis, 0.80 of forefemur in female and 0.90-0.95 of forefemur in male,

(3) yellow integument of apex of coxae and trochanters, (4) scales of fossa purple

or occasionally dark blue, contrasting with predominantly bronze or copper scales

on remainder of mesonotum and (5) larger claw of male foreleg with blunt submedian

tooth and smaller claw of foreleg and claws of midleg of male simple; in the male

genitalia by (1) lateral setae on distal margin of tergite VIII much elongate and bent

mesad at apex, (2) clasper rounded and somewhat sinuous, with apical spiniform

and (3) circular, lanceolate, and sinuous apical sternomesal scales on the sidepiece;

in the pupa by 3-or 4-branched hair 1-P; and in the larva by (1) large number of

comb scales, usually 45-55 but occasionally as many as 75, in large triangular patch,

(2) 6-to 9-branched hair 4a-X and (3) single hair 5-M.

_ Haemagogus chalcolpilans exhibits what appears to be clinal variation in char-

acters of the male genitalia and adult. The apical ventral portion of the claspette

filament differs in shape, the northern populations tending to be more broadly

rounded than the typical form, and in the presence or absence of a ventral seta

(see fig. 53). The mesal setae of the distal margin of tergite VIII are considerably

longer in northern Costa Rican populations than in the typical form from Panama,

although populations from intermediate localities in southern Costa Rica have much

shorter setae than does the typical form. Scales on the fossa tend to be more blue

in the northern populations than the distinct purple scales of the southern popula-

tions.

This seems to be one of the more highly derived species of the subgenus in the

adult, several characters being unique to chalcospilans, among them several male

genitalic characters and the male claws. However the larva exhibits the primitive

characters of a very well developed ventral brush and a large patch of up to 75 comb

scales.

BIONOMICS. Haemagogus chalcospilans is a littoral species, found primarily in

mangrove. It is found in tremendous numbers in mangrove treeholes and occasion-

ally inhabits artificial containers and rarely ground pools. Females are fierce biters,

96 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

attacking readily in the shade and usually about the head and upper body. This

species apparently is not involved in the transmission of yellow fever.

DISTRIBUTION (fig. 6). Haemagogus chalcospilans is known from the Pacific

coast of Central America from near the Gulf of Fonseca in Honduras where it

was collected with regalis by Pedro Galindo (personal communication) to eastern

Panama and the Atlantic coast from central Panama to northwestern Colombia.

Material examined: 3297 specimens; 478 males, 565 females, 680 pupae, 1574

larvae; 384 individual rearings (201 larval, 122 pupal, 61 incomplete).

COLOMBIA. Antioquia: Turbo, 1 M, 1 M gen [USNM].

COSTA RICA. Puntarenas: Dominical, 28 May 1943, T. Aitken, 16 M, 15 F; same data, 14 June

1943, 1 M, 2 F; same data, 18 June 1943, 8 M, 12 F [UCLA]. Golfito, H. Kumm, 9 M, 20 F

[UCLA]. Pitahaya (1.5 km S), 29 Dec 1971, D. Schroeder (CR 579), 2 IpM (579-11,18), 8 IpF

(579-12-17,19,20), 1 pM (579-100), 1 pF (579-101), 1 IP (579-10), 1 P, 2 1, 9 L; same data

(CR 580), 1 lpM (580-20), 2 pM (580-83,84), 2 pF (580-101,102), 5 P, 2 1, 5 L; same data

(CR 581), 1 lpF (581-10), 2 1; same data (CR 582), 7 IpM (582-20-26), 3 IpF (582-27-29), 3 pF

(582-100,102), 7 p, 8 1, 4 L; same data (CR 583), 17 L [UCLA]. Puerto Jimenez, H. Kumm,

1 M [UCLA]. Puntarenas, H. Kumm, 16 M, 28 F; same data, 1938, 3 M gen, 1 F [UCLA].

Puntarenas, 8 June 1954 (GML), 1 IpM (01544), 1 lpF (01554) [UCLA] . Puntarenas (12 km E),

3 m, 14 Aug 1971, D. Schroeder (CR 360), 1 IpF (360-10), 1 pF (360-100); same data (CR 362),

1 lpF (362-10); same data (CR 363), 3 lpM (363-11,12,16), 4 lpF (363-10,13-15), 8 pM (363-101,

103-106,108,110,111), 4 pF (363-100,107,109,112), 1 p, 3 P, 2 L; same data (CR 364), 1 IpM

(364-10); same data (CR 366), 1 pM (366-100); same data (0.5 km N) (CR 370), 1 IpM (370-10),

1 IpF (370-11), 1 L [UCLA]. Rincon, 26 June 1963, C. Hogue (CR 127), 2 IpF (127-101,102),

1 pF (127-201); same data, 29 June 1963 (CR 130), 3 IpM (130-101,102,105), 2 IpF (130-104,

107), 1 IP (130-103), 2 M, 2 F, 10 p, 1 P, 56 L; same data (CR 131), 4 IpF (131-102-105), 30 M,

42 F, 59 p, 28 P, 2 1, 160 L; same data (CR 132), 2 p; same data (CR 134), 2 IpM (134-103,105),

3 IpF (134-101,102,104), 3 M, 4 F, 7 P, 21, 41 L; same data (CR 137), 1 IP (137-105), 2 P, 47 L;

same data (CR 138),-1 F; same data (CR 138), 1 F; same data, 29 July 1963, 2M [UCLA]. San

Jose: San Isidro del General, 28 May 1943, T. Aitken, 3 M, 9 F [UCLA].

NICARAGUA. Rivas: San Juan del Sur, 20 Sept 1943, P. Woke, 4 F [UCLA].

PANAMA AND CANAL ZONE. Canal Zone: Albrook AFB, 2 Nov 1965, R. Schick and A.

Quinonez (PA 738), 4 lpM (738-12,13,15,24), 7 lpF (738-11,14,16,17,19,21,23), 7 pM (738-100,

102-104,107-109), 4 pF (738-22,101,105,106), 32 M, 22 F, 46 p, 4 P, 5 1, 150 L; same data

(PA 739), 2 IpM (739-11,13), 2 lpF (739-10,14), 1 pM (739-100), 3 pF (739-101-103), 1 IP

(739-12), 2 P, 40 L; same data (PA 740), 9 IpM (740-10-17,91), 1 pM (740-100), 1 pF (740-90),

3 M, 6F,9 p, 41, 8 L; same data (PA 741), 1 lpM (741-11), 3 IlpF (741-10,12,13), 2 pM (741-102,

103), 2 pF (741-100,101), 2M, 5 F, 8 p, 3 P, 51, 28 L; same data (PA 742), 4 IpM (742-10,12,13,

15), 5 IpF (742-11,14,16-18), 1 pM (742-101), 2 pF (742-100,102), 3 F, 7 p, 30 L; same data

(PA 743), 3 F [UCLA]. Ancon, 15 Aug 1918, thru C. Ludlow, 4 M, 3 F; same data, 10 Feb 1908,

A. Jennings, 2 M [USNM]. Bella Vista, 7 July 1926, D. Curry, 3 M, 1 F [USNM]. Diablo, 20 Oct

1943, H. Crowell, 3 M, 4 F [UCLA]. Ft. Kobbe, Kobbe Beach, 12 Aug 1972, H. Arnell (PA 1170),

2 IpM (1170-18,38), 16 IpF (1170-11,13-17,23-25,33-35,37,39-41), 10 pM (1170-100-105,107,

108,110,111), 6 pF (1170-90,106,109,112-114), 1 lp (1170-26), 13 IP (10,12,19-22,27-32,36),

17 M, 3 F, 19 p, 4 P, 301, 20 L; same data (PA 1171), 4 IpM (1171-12,14,16,21), 6 IpF (1171-11,

13,15,17-19), 6 pM (1171-101-106), 1 pF (1171-100), 2 1P(1171-10,20), 8 L; same data (PA 1173),

2 IpM (1173-10,11), 16 IpF (1173-13-16,18-29), 10 pM (1173-100-109), 2 IP (1173-12,17), 2 P,

91,9 L; same data (PA 1174), 2 IpM (1174-10,16), 12 IpF (1174-11-15,17-23), 11 pM (1174-30,

32,102-106,108,111-113), 7 pF (1174-31,100,101,107,109,110,114), 28 P, 25 L [UCLA]. Ft.

Kobbe, Venado Beach, W. Komp, 1 M, 2 F; same data, 21 Feb 1939, 5 M; same data, 20 Oct 1939,

3 M, 3 F; same data, 8 M, 4 F; same data, 23 Oct 1939, 8 M, 3 F [UCLA]. Ft. Sherman, 26 June

1958, 1 L [UCLA]; same data, 16 July 1920, J. Zetek, 1 F [USNM] ; same data, 12-19 Nov 1920,

1 F [USNM]; same data, Aug 1938, W. Komp, 1 M, 1 F [UCLA]; same data, 30 Nov 1939, 1 M,

2F [UCLA]. Gatun, 15 Sept 1926, D. Curry, 8 Ip, 1 1 [USNM]. Margarita, 14 June 1922, J.

Shropshire, 1 M gen [USNM]; same data, 8 Oct 1964, A. Quinonez (PA 716), 1 pM (716-101)

Arnell: Genus Haemagogus 97

[UCLA]. Miraflores Lake, 9 Nov 1965, A. Quinonez (PA 762), 4 IpM (762-12,15,18,19), 3 lpF

(762-10,11,13), 1 pM (762-101), 1 lp (762-17), 4M, 6 F, 10 p, 1 P, 21, 10 L; same data (PA 763),

1 lpM (763-15), 4 IpF (763-10,11,17,18), 3 pM (763-100,101,104), 5 pF (763-103,105-108), 2 IP

(763-14,16), 3 F, 30 L [UCLA]. Pina Guapa Island, W. Komp, 1 M, 5 F [UCLA]. Sweetwater

Reservoir (nr. Ft. Sherman), 10 Nov 1964, A. Quinonez (PA 731), 1 IpF (731-10), 1 lp (731-14),

2 IP (731-11,17), 19 L; same data (PA 732), 1 pM (732-11), 1 pF (732-13), 1 P, 5 L; same data,

14 Nov 1964 (PA 734), 2 IpF (734-11,15), 1 lp (734-12), 1 L [UCLA]. Atlantic side (locality un-

specified), Brown, 4 M, 4 F; same data, June 1937, W. Komp, 3 M, 5 F; same data, June 1938,

2 M, 3 F [UCLA]. Canal Zone (locality unspecified), A. Jennings, 3 F [USNM]. Colon: Colon,

A. Jennings, 1 F [USNM]. Portobelo, 4 Dec 1963, A. Quinonez (PA 580), 1 IpM (580-106), 3 lpF

(580-102,109,110), 2 pM (580-104,107), 4 pF (580-101,103,108,111), 1 1, 240 L; same data,

9 Dec 1963 (PA 600), 1 pF (600-103); same data, 10 Dec 1963 (PA 603), 1 F [UCLA]. Portobelo

(Caldera Island), 21 Jan 1908, A. Jennings, 2 M; same data, 22 Jan 1908, 2 p, 1 1; same data,

22 Mar 1908, 2 F; same data, 8 Apr 1908, 1 F; same data, 14 Feb 1909, 6 M, 10 F [USNM].

Portobelo (Rio Caldera), 6 Dec 1963, A. Quinonez (PA 592), 1 p, 5 L [UCLA]. Puerto Pilon,

27 July 1956 (GML), 1 IpM (02988) [UCLA]. Darien: Piriaque, nr. El Real, 13 Jan 1964

(PA 622), 29 L [UCLA]. Panama: Archipielago de las Perlas, Isla del Rey, H. Kumm, 8 M, 4 F

[UCLA, USNM]; Isla San Jose, 26 July 1944, W. Komp, 8 IM (4-44,46,46,47,47,49,52,55), 6 IF

(4-41 ,42,45 46.49.53), 1 pM (4-41A), 1 lp (4-40), 28 M, 26 F, 91 [UCLA, USNM]; Isla San Jose,

Playa Grande (GML), 5 Aug 1956, 4 IpF (03018,03024,03038,03041) [UCLA]; Isleta Trapiche,

8 July 1936, L. Rozeboom (PAR 102), 32 M, 10 F [UCLA]. Chiman, Oct 1942, W. Komp, 5 M,

2 F [UCLA, USNM]. El Corozal, nr. El Libano, 9 Aug 1963, A. Quinonez (PA 486), 1 lpM (486-

104), 3 IpF (486-103,107,108), 1 pM (486-111), 1 pF (486-113), 4 IP (486-102,105,109,110),

1p, 1 P, 11, 11 L [UCLA]. El Libano, 27 Aug 1963, A. Quinonez (PA 535), 1 F; same data,

26 Nov 1963 (PA 565), 1 L; same data (PA 567), 5 IpM (567-104-106,111,122), 5 IpF (567-108,

109,113,114,118), 4 IP (567-102,103,107,123), 66 L [UCLA]. Juan Diaz, 29 Nov 1963, A.

Quinonez (PA 569), 1 pF (569-101) [UCLA]. La Chorrera, 5 Sept 1944, Adams (ASM 156), 1M

[UCLA]. Pacora, 8 Nov 1965, R. Schick (PA 761), 1 IpM (761-13) [UCLA]. Panama, 2 M, 3 F

[USNM] ;same data, A. Jennings, 1 M, 3 F [UCLA]. Panama, Paitilla Pt., 1930, 2 M,3 F [USNM] ;

same data, June 1938, 2 F [UCLA]; same data, 14 July 1938, W. Komp, 3 F [UCLA]; same data,

1939, 10 M, 5 F [UCLA]; same data, 11 Apr 1939, 1 M, 3 F [UCLA]; same data, 14 May 1939,

2M, 2 F [UCLA]; same data, 7 Oct 1939, 2 F, 1 1 [UCLA, USNM]; same data, 24 Aug 1941,

1 M, 3 F [UCLA]. Panama, Rio Abajo, 6 July 1944, K. Frick (ASM 55), 1M [UCLA]. Panama,

San Francisco de la Caleta, May 1938, W. Komp, 1 M gen [USNM]. Panama Viejo, G. Wade,

1 M [UCLA]. Panama Viejo, 28 May 1936, 1 M gen [USNM]. Panama Viejo, 1 July 1949, 15 F,

1 L; same data, 21 Oct 1949, 3 F; same data, 27 July 1935, L. Rozeboom (PAR 82), 10M, 6 F;

same data, 16 Nov 1935 (PAR 32), 1 M, 2 F; same data, 6 July 1944, R. Arnett et al. (ASM 58),

1 F; same data, 29 Nov 1963, A. Quinonez (PA 570), 2 IpM (570-102,110), 7 IpF (570-101,103-

108), 2 IP (S70-109,111), 213 L [UCLA]. San Blas: Mandinga, 15 Oct 1956 (GML), 3 IpM

(03063,03068,03069), 1 IpF (03071) [UCLA]. Locality unspecified: 20 Aug 1935, L. Rozeboom

(PAR 84), 9M, 5 F [UCLA].

24. Haemagogus (H.) boshelli Osorno-Mesa

Figs. 6,55,56

1944. Haemagogus boshelli Osorno-Mesa, 1944b:165-170. TYPE: Holotype male with genitalia

slide, Bahia de Solano, Choco, Colombia, elev. 2-20 m, larva from tree hole or coconut

shell on ground, May 1942, E. Osorno-Mesa [USNM].

1955. Haemagogus (Osornomyia) garciai Levi-Castillo, 1955a:359-364. TYPE: Holotype male

genitalia slide (male lost), Isla de Changuaral, Bahia de Ancon de Sardinas, Esmeraldas,

Ecuador [USNM]. NEW SYNONYMY.

Haemagogus (Haemagogus) boshelli of Levi-Castillo (1951b:11,30-31); Lane (1953:784-786);

98 Contrib. Amer. Ent. Inst., vol. 10, no. 2,197 3

Stone, Knight and Starcke (1959:217); Galindo and Trapido (1967:107).

Haemagogus boshelli of Hovanitz (1946:35); Kumm, Osorno-Mesa and Boshell-Manrique (1946:

17); Barreto Reyes (1955:78); Komp (1955:181); Trapido and Galindo (1956a:305).

Haemagogus (Osornomyia) boshelli in part of Levi-Castillo (1955a:361).

Haemagogus (Haemagogus) garciai in part of Stone, Knight and Starcke (1959:217).

FEMALE. Wing: 2.70 mm. Proboscis: 1.90 mm. Forefemur: 1.95 mm. Abdo-

men: 2.80 mm. Dark scales of proboscis, palpus, wing and legs purple with some

violet and bronze reflections. Head: Eyes rather narrowly separated above antennae

(1-2 ommatidial diameters). Decumbent scales of vertex and occiput purple to

violet anteriorly, light yellowish brown posteriorly. Proboscis short, about 0.95-

1.05 of forefemur; palpus about 0.17 of proboscis; antenna about 0.70 of proboscis.

Thorax: Scales of mesonotum copper to bronze, bluish green to greenish blue above

supraalar bristles, silver in antealar area; scales on scutellum bluish green to greenish

blue. Apn lobes with scales violet to purple, silver scales mesally; ppn bare. Legs:

Length of forefemur about 1.40-1.50 of distance from top of thorax to apex

of midcoxa. Wing: R,,3; about 0.40-0.50 of R,. Abdomen: Dark scales purple

with copper reflections; usually with silver scales dorsally on tergites VI,VII and

occasionally VIII in basal band or median basal patch.

MALE (fig. 55). Wing: 2.20 mm. Forefemur: 1.75 mm. Proboscis: 1.70 mm. Abdo-

men: 2.80 mm. Head: Proboscis about 0.95-1.05 of forefemur; antenna sparsely

plumose, flagellar segment 4 with 6-8 moderately long bristles.

MALE GENITALIA (fig. 55). Segment VIII: Tergite about 0.90 of sternite,

distal margin rounded, densely covered with enlarged setae. Segment IX: Tergite

straight, moderately sclerotized except on midline. Sidepiece: Conical, truncate,

length about 2.5 times median width; basal tergomesal lobe considerably enlarged,

densely covered with setae, short and curved distally, becoming stronger and hooked

proximally; a large oval area near middle of sidepiece giving rise to numerous thick,

short, blunt, peglike setae; an irregular row of fine, curved setae extending from

near mesal edge of this oval area obliquely to near distal margin of basal tergo-

mesal lobe; apical lobe prominent, thumb-shaped, with numerous fine setae; apical

sternomesal scales lanceolate, ovate or obovate. Claspette: Setae ventrally near base;

filament inserted apically on stem, large, striate, leaflike, bent at right angle near

middle with upright half narrow and pointed and dorsal half broad and rounded.

Clasper: Flattened, angled sharply inward near middle, an elbowlike flap on outer

surface distad of angle; tip expanded slightly on outer edge and angled inward;

spiniform inserted proximad of distal expansion, relatively long, flattened and

expanded distally. Phallosome: Aedeagus moderately large, more or less pear-shaped,

the apex produced dorsad forming small serrated carina terminating in a sharp point;

venter open except at basal third, a small lobe projecting dorsad from point of

closure, sclerotized ridges laterally, expanded basally and meeting at midline. Proc-

tiger: Cercal setae 3 or 4; paraproct narrowed apically; apical knob with about

10 serrations.

PUPA (fig. 55). Abdomen: about 2.95 mm. Trumpet: 0.40 mm. Paddle: 0.55

mm. Cephalothorax: Usually very weakly pigmented, occasionally moderately pig-

mented, darker dorsally. Hairs 4,5-C moderately developed, 2-5b, usually subequal

and slightly weaker than 7-C which is usually double. Hair 8-C usually double

(single or double). Trumpet: Medium brown, lighter distally; reticulate sculpturing

moderate. Abdomen: Very weakly to moderately pigmented, anterior segments

slightly darker. Float hair (1-I) with about 10-14 primary branches and 2-6 second-

ary branches. Hair 1-I]-VII weakly developed, 1-6b, more or less subequal on all

Arnell: Genus Haemagogus 99

segments. Hair 2-VII considerably cephalad of 1-VII. Hairs 3-II,5-IV,V subequal

and slightly stronger than 3-IIJ. Hair 6-VII weak, single or double, subequal to

6-III-VI. Hair 9 considerably cephalad of caudolateral margin of tergite, 1-4b;

9-VIII about 8-12b. Terminal Segments: Male genital lobe about 1.8 of tergite

VIII. Paddle: Weakly pigmented, midrib lightly pigmented; broad, inner part con-

siderably larger than outer part; apex broadly emarginate. Hair 1-P usually double.

LARVA (fig. 56). Head: 0.70 mm. Siphon: 0.65 mm. Anal Saddle: 0.30 mm.

Thorax: Hair 5-M double. Abdomen: Hair 7-III-VI weakly developed, smaller than

hair 5 of corresponding segment. Segment VIII: Comb scales 35-42(29-49), with

single broadly rounded to spatulate, minutely fringed spine, in irregular triple row

to large triangular patch. Siphon: Index about 2.5(2.2-2.6). Pecten teeth 12-15

(9-17), extending to about basal 0.50 of siphon. Hair 1-S 4b(4-7b). Anal Segment:

Spines on caudal margin of saddle reduced, most very short with multiple teeth.

Hair 4a-X 6b(4-7b).

SYSTEMATICS. Haemagogus boshelli can be distinguished from the other species

of the Splendens Section by the following characters: in the adults by (1) bare ppn,

(2) eyes narrowly separated above antennae (1 or 2 ommatidial diameters), (3) pro-

boscis about equal (0.95-1.05) in length to forefemur and (4) dark scales of abdo-

men purple; in the male genitalia by (1) unique clasper development, flattened,

bent at right angle near middle with elbowlike flap distad of angle and with bulbous

apex and (2) large oval area of peglike setae near middle of sidepiece; and in the

larva by (1) double hair 5-M, (2) numerous, usually 35-42 comb scales, (3) usually

6 branched hair 4a-X and (4) hair 7-IIJ-VI weaker than hair 5 of corresponding

segment.

This species is among the most highly derived in Haemagogus as indicated by

the most unusual male genitalia as well as several other characters of the adult.

The male genitalia are considerably enlarged, as indicated by the genital lobe of

the male pupa, with an extreme development of the clasper and sidepiece. As

these structures have no counterpart among the remaining Haemagogus, it is diffi-

cult to determine its affinities or derivation. The larva has a well developed ventral

brush and a large number of comb scales.

BIONOMICS. Haemagogus boshelli is found primarily in littoral situations from

San Miguel Bay in Panama to Ecuador. Like several other species in the Splendens

Section, boshelli inhabits mostly mangrove treeholes. It penetrates inland into rain

forest situations in areas of heavy rainfall in northern Colombia where there is no

competition from other species of Haemagogus.

DISTRIBUTION (fig. 6). Haemagogus boshelli extends along the Pacific coast

of South America from eastern Panama to Ecuador, with one record from the

Atlantic drainage of northwestern Colombia. Material examined: 836 specimens;

100 males, 93 females, 108 pupae, 535 larvae; 80 individual earings (49 larval,

22 pupal, 9 incomplete).

PANAMA. Darien: Bahia Pina, Ensenada Molino, 2 Aug 1956 (GML), 3 lpM (03096,03105,

03117), 2 lpF (03106,03118), 2 lp (03092,03101) [UCLA]. Jaque, 17 Dec 1963 (PA 608), 4 pF

(608-102,103,105,106), 5 IP (608-101,107-111),1 p, 85 L [UCLA]; same data, 18 Dec 1963

(PA 610), 4 lpF (610-101,103,104,106), 5 P, 32 L [UCLA]; same data, 6 May 1950, P. Galindo

(GML), 1 IpF (00424) [UCLA]; same data, 3-4 July 1945, W. Komp, 38 M, 1 M gen, 36 F, 2 p,

19 1, 4 L [UCLA, USNM]. Rio Jaque, 19 Dec 1963 (PA 612), 3 IpM (612-104,115,116), 3 lpF

(612-108,110,111), 1 pM (612-109), 3 pF (612-101,102,106), 245 L [UCLA]. Rio Pina, 2 Aug

1956 (GML), 1 IpM (03088), 1 lpF (03113) [UCLA].

COLOMBIA. Choco: Bahia Solano, 8 M, 3 M gen, 7 F, 21, 2 L [USNM]; same data, 1943, 3 M

[UCLA]; same data, 18 July 1942, E. Osorno-Mesa, 1 M [UCLA]. Narino: Isla Gorgona, July

100 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

1924, L. Cheesman, 2 M, 2 F [BM]. Valle del Cauca: Buenaventura, 1944, E. Osorno-Mesa, 2 M,

2 F, 41 [UCLA, USNM]. Rip Raposo Virus Field Station, nr. Buenaventura, 30 m, 13 Jan 1965,

V. Lee (COL 41), 2 lpM (41-10,11), 3 IpF (41-12-14), 1 pM (41-100), 4 L; same data (COL 42),

1 pM (42-12), 5 L; same data, 23 Feb 1965 (COL 56), 3 lpM (56-10,15,17), 4 lpF (56-10,16,18,

19), 5 M,3 F,6P,41, 19 L; same data, 17 Mar 1965 (COL 87), 1 pM (87-100); same data, 31 Mar

1965 (COL 89), 1 IpM (89-20); same data, 5 May 1965, P. Barreto and V. Rivas (COL 110), 1 IpM

(110-13), 1 IpF (110-14), 1 pM (110-106), 4 pF (110-101,102,104,109), 1 IP (110-15), 2 p (110-

105,107), 8 L [UCLA]. Rio Raposo, brackish water zone, 0 m, 14 Oct 1965, V. Lee (COL 94),

1 IpM (94-14), 6 IpF (94-10,11,13,16-18), 5 pM (94-100-104), 5 P, 21 L; same data (COL 95),

1 IpM (95-10), 1 pM (95-101), 2 1, 2 L; same data, 17 Dec 1965 (COL 144), 1 IpF (144-14), 6 M,

3 F, 7 p, 8 L; same data (COL 143), 2 IpM (144-11,17), 1 lp (144-19), 1 P, 23 L [UCLA]. Rio

Raposo, mangrove zone, 0 m, 4 Apr 1966, P. Orjuela (COL 221), 3 IpM (221-10,17,20), 1 pM

(221-102), 7 L [UCLA]. :

ECUADOR. Esmeraldas: Changuaral Island, Bahia de Ancon de Sardinas, R. Levi-Castillo, 1 M

gen (garciai holotype) [USNM]; same data, 14-18 Aug 1972, M. Arzube R. (ECU 179), 2 F

[UCLA].

Additional Record From the Literature

COLOMBIA. Choco: Napipi (Osorno-Mesa, 1944:169).

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ama in the fall of 1956. Am. J. Trop. Med. Hyg. 6:681-685.

110 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

Rosen, Leon and L.E. Rozeboom

1954. Morphologic variations of larvae of the scutellaris group of Aedes (Dip-

tera, Culicidae) in Polynesia. Am. J. Trop. Med. Hyg. 3:529-538.

Shannon, Raymond C.

1931. The environment and behavior of some Brazilian mosquitoes. Entomol.

Soc. Wash., Proc. 33:1-27.

1939. Methods for collecting and feeding mosquitoes in jungle yellow fever

studies. Am. J. Trop. Med. 19:131-140.

Shannon, Raymond C. and E. Del Ponte

1928. Los culicidos en la Argentina. Inst. Bacteriol. Malbran, Rev. 5:29-140.

Shannon, Raymond C., L. Whitman and M. Franca

1938. Yellow fever virus in jungle mosquitoes. Science 88:110-111.

Sheath, PAT,

1939. Yellow fever in British Guiana. R. Soc. Trop. Med. Hyg., Trans. 33:241-

252:

1940. Yellow fever in British Guiana. Further Observations. R. Soc. Trop. Med.

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Soper, Fred L.

1935. Rural and jungle yellow fever: A new public health problem in Colombia.

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1957. Corrections in the taxonomy and nomenclature of mosquitoes (Diptera,

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Stone, Alan and K.L. Knight

1955. Type specimens of mosquitoes in the United States National Museum: I,

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1901. A monograph of the Culicidae or mosquitoes. Vol. 2. Lond., Br. Mus.

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1972. Mosquito Studies (Diptera, Culicidae). XXVIII. The New World species

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FIGURES

1. Distribution of Haemagogus (Haemagogus), Albomaculatus Section, Tropicalis

Section and Splendens Section

2. Distribution of anastasionis, chrysochlorus, spegazzinii and baresi

3. Distribution of janthinomys, capricornii and mesodentatus

4. Distribution of andinus, nebulosus, albomaculatus and panarchys

5. Distribution of soperi, acutisentis, equinus and tropicalis

6. Distribution of splendens, celeste, argyromeris, chalcospilans and boshelli

7. Distribution of iridicolor, aeritinctus, regalis and lucifer

8. Haemagogus (H.) anastasionis; male genitalia, male claws, pupa

9. Haemagogus (H.) anastasionis; larva

10. Haemagogus (H.) spegazzinii; male genitalia, male claws, pupa

11. Haemagogus (H.) spegazzinii; larva

12. Haemagogus (H.) baresi; male genitalia, male claws, pupa

13. Haemagogus (H.) baresi; larva

14. Haemagogus (H.) andinus; male genitalia, male claws, pupa

15. Haemagogus (H.) andinus; larva

16. Haemagogus (H.) nebulosus; pupa

17. Haemagogus (H.) nebulosus; larva

18. Haemagogus (H.) janthinomys; male and female heads and claws; male fore-

tarsus 5; female thorax, wing, legs and genitalia

19. Haemagogus (H.) janthinomys; male genitalia, pupa

20. Haemagogus (H.) janthinomys; larva

21. Haemagogus (H.) capricornii; male genitalia, male claws, pupa

22. Haemagogus (H.) capricornii; larva

23. Haemagogus (H.) mesodentatus; male genitalia, male claws, pupa

24. Haemagogus (H.) mesodentatus; larva .

25. Haemagogus (H.) albomaculatus; male genitalia, male claws, pupa

26. Haemagogus (H.) albomaculatus; larva

27. Haemagogus (H.) panarchys; male and female heads and claws; male foretarsus

5; female thorax, wing, legs and genitalia

28. Haemagogus (H.) panarchys; male genitalia, pupa

29. Haemagogus (H.) panarchys; larva

30. Haemagogus (H.) soperi; male genitalia, male claws, pupa

31. Haemagogus (H.) soperi; larva

32. Haemagogus (H.) acutisentis; male genitalia, male claws, pupa

33. Haemagogus (H.) acutisentis; larva

34. Haemagogus (H.) equinus; male genitalia, male claws, pupa

35. Haemagogus (H.) equinus; larva

36. Haemagogus (H.) chrysochlorus; male genitalia, male claws

Haemagogus (H.) tropicalis; male genitalia, male claws

37. Haemagogus (H.) tropicalis; larva

38. Haemagogus (H.) splendens; male and female heads and claws; male foretarsus

5; female thorax, wing, legs and genitalia

Arnell: Genus Haemagogus 113

. Haemagogus (H.) splendens; male genitalia, pupa

. Haemagogus (H.) splendens; larva

. Haemagogus (H.) celeste; male genitalia, male claws, pupa

. Haemagogus (H.) celeste; larva

. Haemagogus (H.) iridicolor; male genitalia, male claws, pupa

. Haemagogus (H.) iridicolor; \arva

. Haemagogus (H.) aeritinctus; male genitalia, male claws, pupa

. Haemagogus (H.) aeritinctus; larva

. Haemagogus (H.) regalis; male genitalia, male claws, pupa

. Haemagogus (H.) regalis; larva

. Haemagogus (H.) lucifer; male genitalia, male claws, pupa

Haemagogus (H.) lucifer; larva

. Haemagogus (H.) argyromeris; male genitalia, male claws, pupa

. Haemagogus (H.) argyromeris; larva

. Haemagogus (H.) chalcospilans; male genitalia, male claws, pupa

. Haemagogus (H.) chalcospilans; larva

. Haemagogus (H.) boshelli; male genitalia, male claws, pupa

. Haemagogus (H.) boshelli; larva

‘la & .

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NORTH & SOUTH AMERICA

KILOMETERS

STATUTE MILES

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MOSQUITOES OF MIDDLE AMERICA

NORTH & SOUTH AMERICA

KILOMETERS

1500 2000

STATUTE MILES

UCLA - ZOOLOGY - ENT ~ 1971

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00s Oot os 0

0001 10S oor os 0 A

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mac,

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SN}IUI}IIOv=

eS:

F '

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Guanacaste

HAEMAGOGUS

eee a ee

13 1

6 3

10 | ar =

S rE NIN

Vill

=> 3 =

Me uN ta

es DS

anastasionis

NI 12

Leon

Nicaragua

Fig.10

HAEMAGOGUS

Ceara

Brazil

LA |

ANITA

ne OE LU IGTE ox

Dy eee”

‘ v v ‘ i i as

‘ iG ‘ v [a ]

! dj v ue op v4

< v boy ™ Y

v ‘G aes v i

sf v

i. .

a eR

Santa Cruz

Bolivia

Ze sw SC

SR NS

RIS NS = SN

SS =

HAEMAGOGUS

ao.

bao

ms ©

spegazzin

Ceara

Brazil

iS}

= i)

Wr 4,

Vill

‘Hh?

Aa:

yy, \\\ Gat

VII

Ai\

pm 1

son

—-o

Fig.12

HAEMAGOGUS

Amazonas

Brazil

Eee ar re Me

CG \tal

Loreto

“

Vali

0.3 « \ ’ 2s :

baresi

Fig. 14

HAEMAGOGUS

—— —>

—————

———

—————

ESS

———

————

—_ ia —— :

=~" —SS—

Wy WAT

andinus

Colombia

Fig. 16

HAEMAGOGUS

nebulosus

VZ 151

Venezuela

—SS

——

pc Ee

—S—

= =

——-

——

Ss;

Zvi

GGR23

Darien

Reine

SN S

WE,

Uh rf

WARE ~— °

ARN "

janthinomys

VZ 312

Aragua

Venezuela

\ NW — .

Colombia

Fig. 19

HAEMAGOGUS

TR 300

Trinidad

Trinidad and Tobago

je $

Wr >

<= ae

< SS

TR 299

Trinidad COB 40

Trinidad and Tobago Meta

Or ae Colombia

\; i/

\ AN (War)

Minas Gerais

Brazil

; janthinomys

f co

f “N

| Y

Li) Aa

!), Z a

eae

3 Eon f

un oe

fon)

Yf»

14 ¢ |

Vif 4

$ y :

| : Gaal

’ uy

8 2 V ee q Y

ss ee $4 2 9 ZT 4e Aig) \

. ;

i, 11 Wd Ve, Wy 1

0, 3 2 | ‘eh

sp |

8/ 9

8 2 VI , ,

6 AS " 11 1° ;

in | | Ms y"s

; 4 ‘i - i | |

/ janthinomys | i

7 TR 300 :

" Trinidad

Trinidad and Tobago

Fig. 21

HAEMAGOGUS

Rio de Janeiro

Brazil

4 7 LZ \\

| a \

W422 \ 18

TO |

ANG

. tS LICUISY BSE i

| \\ eo)

i sou y > t

\\ i;

7 a 1

\ y

\ sp

= Ss \ M

pee

8 Cee erie

\ WY WW T

Se 1

capricornii

Sao Paulo

Brazil

HAEMAGOGUS my

GUA 88

Igabal

Guatemala

| \

fl iI i

Way Uf

OY) Vol |

ae ! nates

S .

BE Laorey © i

CAC)

, ra

W\ | da VP

"ae oP :

‘

v vf

Ip mesodentatus

aH les

mesodentatus

CR 88

San Jose

Costa Rica

Fig. 25

HAEMAGOGUS

dul

SSS

alt

m Ll

‘|

FGC 3947

Guyane

French Guiana

a ae

masnacocus 4

i \ 7

FGC 3947

Guyane

‘\a French Guiana

Fig. 27

HAEMAGOGUS

AUT iL Se ii

if ri AN AD

i Vivaah f

AN iH AN MONI | IN

HN RNA aN

1 a ie

HAAN ai AN da WN

ae ae

i il\\ Fan Pi Nw if iti Ny

10 ¥ iN CONN, CL ae

iN PPR Wh

Hy pil Whi \ f Y

Ain in

waa

\\ wi ECU 136

iS, A ngs 78: h

im Zt ZH

ih Uy i Jab Guayas

ae Ont yi!

KN On as" \\

ys Wh Ae

| a: a

yi At N Wy \ HN

YS ik \ ] I) \

a a

a Ge ioe

A \\\ ah My, \

PIN ZN LINN

BAN Th yf \

AY 2

f\) { U\\\ Af \

Wy \\ yj) \ | (t

/\ a {inn ‘

Aw ie Hi

Mir AN TAN : Hh

FAN BAAN rh

ihe Gi. eh

ji a ah

aN Ge UR

HNN i:

1 JAN pithy ® Hi

WN Y cy i ; \

i 7M Ws

l di \ i F

A HN

SS.

SSS

SNSSSSS

Se SS

ha Ze

=: =

SSS

ECU 161 Q

Fig. 28

HAEMAGOGUS

ECU 135

LEFTY)

YS —

\ ©

“AN

aes

(

\ hb. (,

Cie: A

\ 7

1.0 \. i SSS

N oral Se

x a f 17: ~ ¥

Pad

Ven eal

Ps PE

| rat

a at

~\ At panarchys

LX 7 :

HAEMAGOGUS

Fig. 30

HAEMAGOGUS

\ |

\ MUNI LY.

op" aa wil Ss ie

4 | es

| ih Poh We 7 v

y v = Wi) ys Cie)

Niu & A v S VOW us,’ Wi viy “y te

2) é vv@v

v vu (rege . ’

vy v U v 0 v Vi eae

Canty vv v v Cah vy

\ j , ¥ eintey, v ARNT ghar sies Fale

‘

v ¥ cf td

v v Ut va Cowal P v red

v - 4 v “oA

v v voy v v Ln aa f}

i 3 Werner v Aes id Piegety.

0.3

QL “C4 &

CZ==a <Z ° ©

PS pA 2 ic = ie

Q ——__ <—— ; x Xe

e AWN WS

Q s \ CRRRSSSSS

3 Q 1 K\R SSS

. AS KINA R

é e a g ( A) RIN SS

Oy = XQ SS

a GN Z NA \\\

a AN RAK

soperi

ECU 121

Los Rios

Ecuador

zou 328 Fy g. 32

Ecuador

HAEMAGOGUS

ECU 112

Los Rios

Ecuador

FORE

ine

A | W | pia 4 AN

, ee _ te y r fi y

i »)

H i] S

S ' §

’

AMIE

ey Vi

—_—a

LZ gE.

CLE,

ra = >

OzE__=

Vill

BE €

17 2)

—]

pe ©

7 2)

acutisentis

ECU 125

Los Rios

Ecuador

- Fig. 34

Jamaica

HAEMAGOGUS

JA 391

Jamaica

“iy

AW :

Jamaica

JA 328

equinus

chrysochlorus

Curacao

FORE.

\ a t MT] i i ] cS) fe

\V ae | : i Ry /

——

= —=— -

R~ 2@ « wy =

a Ds Lv Ss

PTS, hE SSSS&

el SS SSS

yy S .

E———

— 7 Zz

SSSSS—

é gi

fi Zz <

LE AAA —>

EE ce

gE FP

——S = = = 40s

w= =—=<—<

ORE

VInisT

tropicalis 4

MID

FORE aed

nt i Ys ij}.

me HNN Uff))ij\\\)' eee

a A iii — Me

‘ik mS iii. ~ em a ESSN

1) a SS i HAUTE eet Ne ARAN

it i HAEMAGOGUS Mi ATE TENANT

Hh LSS

in ik

Aw sis

ie | a

Bias ALVIN ANANTH SR

GS ENN

i NN Nn

Ae AR

f IN. \ i ve \

“iy Ay iv NY

Ais i ih iN

a ih

| ALS

\ A Mints

i \

Hi uN \ \

BAN

Pen : VT 81

Wi A \ St. Vincent

i Q

y) 7

a | \ yh iy

hen Ai ; }—_—_____ 1.0 —_—_——__1

A AVR f Y Zp QGEGLE

a LIE

| a

AN AHS ANN \ A

ql \ v\ I N VN \ S S hi

mi) \ am RSSERSAKS tug

in HN RSS Ja

1] |) I S y

ni \

ATS

SASS.

A OSs:

HIND i

Q / VT 78 Gh

St. Vincent jy

CCK

VT 81

‘ot St, Vincent

—SS=

SS SE

= =

——_

= S—

———

i BY

| r SSS

===> 2 EZ

SS -=— nnn -

SSS > aS Lo

SP Ree Go

i ge ee

SSS

——————

Zoe SZ

S50. SSeS =

SSSs SS

Lz,

—

——

VIG?

_ St. Vincent

0.3

Fig. 39

HAEMAGOGUS

= - oe

MY A

il ¥ me 4

D4 N yaa

Pe - i

Op) i} |

a ae Ly

% } >| SE Dol

NNR ie A Vf od

\ N J "A: 1h 5 LY

\ 6 bp 5 up F ij

Ai P fk Ud

I7\ ~~ splendens | |

\ ‘ Vis VT 81

St. Vincent

Lao ieee

=> ee

HAEMAGOGUS

eee? 8

— J

5 &

4 SS

meen

splendens

Vine

St. Vincent

= Fig. 41

_- HAEMAGOGUS NN

VZ 341

Aragua 4 Sani

Venezuela SS |

6 —— =) I

—< <A :

nee 4 F \

7 s | ie

3y \

ORE J

Ysp

Il 8

1.0 6

4 2 7

3 11

a ne |

Cae

1} |

1 5

At)

mt } + ball [ UF

og ti Ka i fj )

. ie Y, & a — i Oo" y.

S__

—

AN MM teat |

1 are alge es A VR EATS

HAEMAGOGUS

: ‘iy Iw?

(| / ~N : | \

ff i \N S

Fig. 43

HAEMAGOGUS aw \

Costa Rica

CR 320

Alajuela

Costa Rica

j\\\

FORE

IN AT,

age y i | ‘I y i j y

PT Ie

! 9

“3 i

Dal oF v//]y

“$e Cvs wey)

a-———-4

iridicolor

vill

d a A

iridicolor

CR 322

Alajuela

Costa Rica

Fig. 45

HAEMAGOGUS

aeritinctus ates

aeritinctus

British Honduras

Fig. 47

HAEMAGOGUS

El Salvador

HAEMAGOGUS

12°

13 |

; MP

™ an

ie—— ay & al

(| p\ = x CA ge NS

Pa NG

" | | v ;

onsonate

‘

| HAEMAGOGUS x :

Fig. 49

PA 861

Canal Zone iN

lucifer

PA 861

Canal Zone

= Fig. 1

Panama

HAEMAGOGUS

PA 767

Canal Zone

Panama

Canal Zone

Panama

ASM 386

Canal Zone

Panama

RIN

SST

1.0

VII

~sp

. 2

12 5 eee

“5 My

I NH i

SS ql if \ \

0-7) fe" )

argyromeris a

PA 764

Canal Zone

Panama

PA 716 ]

Canal Zone 8

Panama

HAEMAGOGUS

PA 738

Canal Zone

Panama

Canal Zone

Panama

Puntarenas

Costa Rica

CR 363

Puntarenas

Costa Rica

oN Fig.55

Valle del Cauca

Colombia

HAEMAGOGUS

COL 221

Valle del Cauca

_ Colombia

Valle del Cauca

Colombia

\\ \ {

x \ \ AB 4!

(

boshelli vi .

boshelli

COL 56

il Valle del Cauca

Colombia

170 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

TABLE OF DISTRIBUTIONS

2 - Bs

ee |e Se a Shae

— Chal ‘ OS

= Ss a 2 S a c 3

fo “a 0) n S = i — Oo o)

n >> od o chen ~ = 72) Ne

Se} = oO ™ Xe) = o. a> 7

cS a oy - = © oS o) = oo

3 oO n = < : en oO ~

att N a) rt ora) on pert

USA Soh OR Rae HS na ok A

JAMAICA ae ae

ores linia

Atlantic

SinTewna wae ie:

: Atlantic

BRIT. HONDURAS _ —E oO

ELSALVADOR _ a wae

woos eT TT TT Tet TST

Atlantic

NICARAGUA /2%H¢ oe

Bete]

COSTA RICA 4 Boe eae

ier aban

West

PANAMA Central

soo eet | tel | tel

a Y

Pacific

COLOMBIA Carib.

Orino co-Amazon

erm S| | 1 | [*isl | le

Orinoco

NETH. ANTILLES ey se

LESS. ANTILLES oon Pe te

TRINIDAD-TOBAGO TS ee eee

GUIANAS oa a ae

ame ee a: a Pa

Southeast

PARAGUAY

ARGENTINA

BOLIVIA

PERU eee Oe ee ee

Arnell: Genus Haemagogus 171

TABLE OF DISTRIBUTIONS

18. iridicolor

19. aeritinctus

23. chalcospilans

24. boshelli

20. regalis

21. lucifer

9°)

se)

—

13. acutisentis

ies:

90)

—<

oe)

Sey

bie y

WwW

14. equinus

O88 @@

172

Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

CONSPECTUS OF TAXONOMIC CHANGES

NEW TAXA

acutisentis

chrysochlorus .

nebulosus

CHANGES IN TAXONOMIC STATUS

alticola, synonymy .

celeste, to specific rank

dominguezi, synonymy

falco, synonymy .

garcial, synonymy

gorgasi, synonymy .....

janthinomys, to specific rank .

Longipalpifer, synonymy

obscurescens, change in synonymy .

petrocchiae, synonymy

Stegoconops, synonymy .

Arnell: Genus Haemagogus 173

INDEX TO SCIENTIFIC NAMES

acutisentis, 13k, 16k, 19k, 23, 57, 58-60, 62;

figs. 5, 32-33

Aedes, 1,3, 6, 19, 21

Aedini, 3, 6, 7

aeritinctus, 7, 14k, 17k, 19k, 63, 70, 78, 79,

80-82, 84, 86; figs. 7, 45, 46

affirmatus, 60, 61 |

albomaculatus, 2, 3, 7, 13k, 16k, 18k, 20, 24,

50-53; figs. 4, 25, 26

albomaculatus of authors, 38, 61‘, 74, 80, 82,

85, 90

Albomaculatus Section, 1,4, 6, 12k, 15k, 18k,

22, 22-24, 25, 30, 32; 34, 35, 36,39, 46,

52, 59, 62.67, 68; 70, f2ifae t

albopictus (Aedes), 5

alticola, 47

anastasionis, 4, 7, 14k, 15k, 18k, 23, 24-27,

28, 30,31, 33,37, fs 2, 0, 9

anastasionis of authors, 27, 38

anastasionis complex, 23, 25, 28, 30, 33, 35

andinus, 13k, 16k, 18k, 23, 33, 33-35, 36;

figs. 4, 14, 15

andinus complex, 23, 33, 35

argyromeris, 7, 14k, 17k, 19k, 70, 78, 82,

84, 86, 90-93; figs. 6, 51, 52

baresi, 2, 13k, 16k, 18k, 23, 31-33, 35; figs.

Pee Oe

boshelli, 7, 14k, 17k, 19k, 63, 70, 97-100;

figs. 6, 55, 56

Cacomyia, 2, 20, 21

capricornii, 3, 7, 13k, 16k, 18k, 20, 23, 28,

30, 36, 37, 40, 44-46; figs. 3, 21, 22

capricornii of authors, 37, 38

capricornii complex, 23

celeste, 7, 14k, 17k, 19k, 70, 72, 73-76; figs.

6, 41, 42

chalcospilans, 7, 14k, 17k, 19k, 63, 70, 93-

97; figs. 6, 53, 54

chrysochlorus, 4, 14k, 15k, 17k, 23, 25, 26,

27-28, 30, 33; figs. 2, 36

Conopostegus, 1, 3, 4, 5, 6, 8, 9, 12k, 15k,

18k, 19-20, 22, 24, 55

cyaneus (Culex), 2

cyaneus of authors, 38, 51, 61, 82

Cyanoconops, 20, 21

dominguezi, 24, 26

equinus, 2, 3, 7, 13k, 16k, 19k, 23, 28, 40,

44, 46, 57, 58, 60-67; figs. 5, 34, 35

equinus of authors, 38, 45

falco, 37, 40

falco of authors, 56

Finlaya, 1,3, 6

garciai, 97,98, 100

gladiator, 90

gorgasi, 46, 47

Haemagogus, J, 2, 3, 4, 5, 6, 7, 8-12, 12k,

15k, 18k, 19, 20520-22,223% 24:°53,-55,

5762, 63,'70.:75, 82, 91, 99: fie J

Heizmannia, 6, 7

Inga, 33

iridicolor, 15k, 17k, 19k, 70, 76-80, 82, 84,

86, 87, 91; figs. 7, 43, 44

janthinomys, 7, 13k, 16k, 18k, 23, 26, 28,

30, 31, 36-44, 45, 46, 49, 52, 63, 64, 87;

Figs. 3, 18-20

Kochi Group (Aedes), 6

leucocelaenus (Haemagogus), 3, 7, 8, 19

leucomelas (Haemagogus), 3

lindneri, 2, 28, 29

lindneri of authors, 29

Longipalpifer, 3, 5,6, 20, 22, 23

lucifer, 3, 7, 15k, 17k, 19k, 70, 78, 82, 84,

84-89, 91; figs. 7, 49, 50

mangle (Rhizophora), 80

mesodentatus, 7, 12k, 16k, 18k, 23, 46-50;

figs. 3, 23, 24

nebulosus, 13k, 15k, 18k, 23, 35, 35-36; figs.

4,16, 17

obscurescens, 2, 26, 37, 40, 43

Osornomyia, 20

174 Contrib. Amer. Ent. Inst., vol. 10, no. 2, 1973

panarchys, 3, 12k, 15k, 19k, 20, 23, 53-56;

figs. 4, 27-29

petrocchiae, 37, 40

philosophicus, 60

pseudes (Deinocerites), 84

regalis, 3, 7, 15k, 17k, 19k, 63, 70, 78, 82,

82-84, 85, 86, 91, 96; figs. 7, 47, 48

regalis of authors, 38, 61, 74, 80, 85, 90

regalis complex, 70, 78, 82, 84, 86

Sabethini, 7

scintillans (Heizmannia), 7

scutellaris (Aedes), 5

spegazzinil, 14k, 15k, 18k, 23, 25, 28, 28-31,

33, 36, 40, 46; figs. 2, 10, 11

spegazzinii of authors, 38

splendens, 2, 3, 8, 14k, 17k, 19k, 20, 70, 71-

73, 74, 75; figs. 6, 38-40

splendens of authors, 38, 51, 73, 77, 85, 94

splendens complex, 70, 72

Splendens Section, 1,4,6, 12k, 14k, 15k, 17k,

18k, 19k, 22, 63, 67, 68, 69-70, 72, 74,

75, 78, 81, 84, 86, 91, 95, 99; fig. 1

soperi, 7, 13k, 16k, 19k, 23, 56-58, 59, 62;

figs. 5, 30, 31

soperi of authors, 58

soperi complex, 23, 57, 58, 63

Stegoconops, 3,5, 6, 20, 21, 22, 23

Toxorhynchitini, 7

tropicalis, 6, 12k, 15k, 18k, 22, 23, 67, 68-69;

figs. 5, 36, 37

Tropicalis Section, 1,6, 12k, 15k, 18k, 22, 67-

68, 69, 70, 72; fig. 1

uriartei, 28, 30,31, 40

uriartei of authors, 24, 28, 29

Contributions

of the

American Entomological Institute

Volume 10, Number 3, 1973

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. -XIX.

Bothaella, A New Subgenus of Aedes Meigen

John F. Reinert

CONTENTS

Rear es eo a

INTRODUCTION «© oo RAPReD Sweet TANS Gwe See res

GENUS AEDES MEIGEN, BOTHAELLA, NEW SUBGENUS ........

KEYS TO THE SPECIES OF AEDES (BOTHAELLA)............

Pe eee ee a ee

a ae Ta ES MN A Sa Fe a ew Cae

POUR Ho PAGE EARVAE. &.o . Sees tek be 6 ee secle do

DESCRIPTION OF THE SPECIES OCCURRING IN SOUTHEAST ASIA...

WelCHGe NEW EDECICS 6. ae a Te eerie aie ee ea 8 es

PIGHIGGE7 ROW SOCCIOS 0k hee we acle chine ie SE ee Bele

Piet NCW SDECIOS ocr ol kee 8 ao eyes Wa eye ne TR ee we

ACK MOW SGT MEN TS ok ois ce Be a ee eo wien eee 6 ere ee ee

Te Pie a ae a ee ee eae pe ee we ee ee ee be |

APPENDICES

TABLE 1. RECORD OF THE BRANCHING OF THE SETAE ON

THE PUPAE OF AEDES (BOTHAELLA) HELENAE . .

TABLE 2. RECORD OF THE BRANCHING OF THE SETAE ON

THE PUPAE OF AEDES (BOTHAELLA) ELDRIDGE! .

TABLE 3. RECORD OF THE BRANCHING OF THE SETAE ON

THE LARVAE OF AEDES (BOTHAELLA) HELENAE .

TABLE 4. RECORD OF THE BRANCHING OF THE SETAE ON

THE LARVAE OF AEDES (BOTHAELLA)

Ps ae, © rs eS I 8

ODOMDADADOAONKHrF

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA, -XIX.

BOTHAELLA, A NEW SUBGENUS OF AEDES MEIGEN!

John F, Reinert”

ABSTRACT

A new subgenus, Bothaella, of Aedes Meigen is defined, discussed and

compared to related taxa. Descriptions, illustrations and keys are presented

. for the known stages of the 3 included species, helenae, eldridgei and kleini,

all of which are new.

INTRODUCTION

During the course of revisionary work on the subgenera of Aedes

Meigen, material of 3 new species was discovered which differed significantly

from the other known subgenera. It therefore became necessary to define the

present new subgenus, Bothaella, and describe the 3 new species, helenae,

eldridgei and kleini. Keys are given for the known stages of these species

and their geographical ranges are plotted on the map in figure 1.

Abbreviations used in literature cited conform to the World List of

Scientific Periodicals, 4th edition, Butterworth, Washington, 1963. The

abbreviations used in the distribution sections are as follows: o= male;

¢ = female; L = larva; p = pupal skin; and 1 = larval skin. Pupal descriptions,

tables and key utilize the following abbreviations: C = cephalothorax; I-VIII =

abdominal segments 1 through 8; and P = paddle. In larval descriptions, tables

and key the following abbreviations signify: A = antenna; C = head; I-VIII, X =

abdominal segments 1 through 8 and 10; M = mesothorax; P = prothorax; S =

siphon; and T = metathorax. When possible, 10 specimens were used in deter-

mining the range, mode and mean seta branching in pupal and larval descrip-

tions and tables. In the pupal descriptions and tables, the number of branches

on abdominal hair 1-I was measured on the basal 0.33 of the seta. Measurement

scales on the illustrations are in millimeters. Distribution records are indi-

cated as follows: countries are in capital letters; provinces are in italics; and

place names have the first letter capitalized. The number of specimens ex-

amined from each province follows the last place name of the province in the

distribution sections. The spelling of provinces and locality names in Thailand

[This work was supported in part by Research Contract No. DA-49-193-MD-

2672 from the U. S. Army Medical Research and Development Command,

Office of the Surgeon General and carried out at the Southeast Asia Mosquito

Project, Smithsonian Institution, Washington, D. C. 20560.

2Major, Medical Service Corps, U. S. Army, Department of Entomology,

Walter Reed Army Institute of Research, Washington, D. C. 20012.

> Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

is taken from the Official Standard Names Gazetteer No. 97, prepared by the

Office of Geography, Department of the Interior, Washington, D. C., April

1966; and those from Cambodia from the Official Standard Names Gazetteer,

2nd Edition, prepared by the Geographic Names Division, U. S. Army Topo-

eraphic Command, Washington, D. C., November 1971. Locality names which

do not appear in the gazetteers are spelled according to the data sheets and

labels on the specimens.

Nomenclature and chaetotaxy used for males, females and male genitalia

follow Knight (1970) and Knight and Laffoon (1970a, 1970b, 1971) and those for

the pupae and larvae follow Belkin (1962). The terminology of the female

genitalia is taken from Laffoon and Knight (1971) except for additional ter-

minology as given by Reinert (1973c) and Jones and Wheeler (1965).

GENUS AEDES MEIGEN

SUBGENUS BOTHAELLA, NEW SUBGENUS

Type species: Aedes helenae Reinert

The 3 species assigned to the subgenus share the following combination

of characters.

MALE. Head. Antenna brown, plumose, pedicel with a patch of over-

lapping, broad, silvery scales on mesal and ventral surfaces, flagellomere 1

with a few brown scales, total length 0. 73-0. 85 length of proboscis; clypeus

blackish-brown, bare; maxillary palpus short, dark brown scaled, 4 segmented,

total length 0. 18-0.19 length of proboscis; proboscis dark brown scaled (kleini

with a few pale scales ventrally), total length 1.08-1.13 length of femur I;

eyes contiguous; vertex and lateral areas covered with overlapping, broad,

decumbent scales, a dorsomedian anterior triangular patch of silvery scales;

occiput with numerous erect forked scales; several dark, long ocular setae;

2 dark, long interocular setae. Thorax. Scutal integument dark; scutum

covered with narrow, curved, dark scales with prescutellar space bare; scu-

tellum with broad dark scales on lateral lobe, median lobe with broad silvery

scales basally and broad dark ones apically; following areas with setae which

are dark and well developed; 2 median anterior promontory, numerous dorso-

central (anterior and posterior), scutal fossal (2-4 anterior, 2-3 lateral and 1

posterior), numerous supra-alar, 5-8 posterior medial scutal (pale), 1 postalar

callar and scutellar (3 long and 3-4 short lateral; 4-5 long and 2-4 short median);

pleural integument dark; antepronotum with a large patch of overlapping broad,

silvery scales, 7-9 dark setae; postpronotum without scales, 2-4 dark posteri-

or setae; propleuron with a large patch of overlapping, broad silvery scales,

2-3 dark setae; prosternum, subspiracular area, paratergite, mesomeron and

metepisternum bare; postspiracular area with a large patch of overlapping,

broad, silvery scales, 1-2 dark setae; mesepisternum with an upper and a

posterior patch of overlapping broad silvery scales, 1-2 upper and 1-2 lower

setae; prealar knob with 3-8 dark setae; mesepimeron with a patch of overlap-

ping, broad, silvery scales on upper area, 3-5 setae on upper area posterior

to scale patch and 1 long seta on lower area. Legs. Coxae I-III each with

several setae and a patch of broad scales which are silvery on II and III; fe-

mora I-II. dark brown scaled, with white markings on posterior surfaces of

I-III and anterior surfaces of II and III, I with or without anterior white scales,

I length greater (1.09-1.11) than length of II; tibiae I-III dark brown scaled

(a few apical white scales in helenae); tarsi dark brown scaled with or without

Reinert: Bothaella, new subgenus 3

white bands; posttarsi I-III each with 2 ungues, I with ungues unequal, both

with a tooth, II with ungues unequal, larger one simple and smaller one with a

tooth, II with ungues equal and simple. Wing. Veins covered with dark

scales dorsally and ventrally; alula with several narrow, dark scales along

margin; upper calypter with numerous dark setae along margin; 1 remigial

seta. Halter. Pedicel pale; capitellum silvery scaled with dark scales at

base. Abdomen. TergaI-VI each blackish-brown scaled with a large latero-

basal silvery scaled spot; sterna II-VII each blackish-brown scaled with a

basal silvery scaled band; terga and sterna with numerous setae, mostly

along posterior margins. Genitalia. Tergum IX band-like, moderately pig-

mented with 3-9 stout setae on each side of midline; gonocoxite short, broad,

heavily pigmented, mesal surface membranous, dorsal surface with numerous

moderately long or long thin setae, ventral surface with numerous short

setae and a large circular apical mesal area which extends tergally and is

covered with either short setae or moderately pigmented, short, broadly

flattened, scale-like setae; gonostylus long, apical 0.42-0.46 expanded with

several small fine setae and a heavily pigmented gonostylar claw which is

short, bluntly rounded apically and attached apically or subapically to gono-

stylus; basal mesal lobe with apical 0.50 large, expanded and with 28-244

short setae, basal portion covered with small spicules and connected mesally

with its mate; proctiger long, paraproct moderately pigmented with apex blunt

and heavily pigmented, cercus membranous, cercal setae absent; tergum X

narrow, heavily pigmented; phallosome with aedeagus divided into 2 lateral,

moderately pigmented plates which are connected at base and apex, each

lateral plate with outer lateral area curved tergally and with a mesal, longi-

tudinal, heavily pigmented area terminating in 3-8 short teeth, paramere

moderately long, 0. 72-0. 82 length of aedeagus, heavily pigmented, parameral

apodeme long, 1.25-1.42 length of paramere, heavily pigmented; sternum IX

large with basal, lateral and median areas moderately pigmented, 4-11 short

to moderately long setae near center.

FEMALE. Similar to male in general habitus but with the following

differences. Head. Antenna with basal 0.75-0. 85 of flagellomere 1 pale,

total length 0. 77-0.95 length of proboscis; maxillary palpus total length 0. 17-

0.19 length of proboscis; proboscis total length 0.95-1.05 length of femur I.

Thorax. 2-4 median anterior promontory setae; scutellum with 3-4 long and

2-6 short setae on lateral lobe and 4-6 long and 2-5 short setae on median

lobe; antepronotum with 6-11 setae; postpronotum with 2-5 posterior setae;

propleuron with 2-4 setae; prealar knob with 4-9 setae; mesepimeron with 3-8

setae on upper area and 1 lower one. Legs. Femur I length greater (1. 07-1. 15)

than length of II; posttarsi I-III each with 2 ungues, I with ungues equal, both

with a tooth, II with ungues equal, both with a tooth, IIT with ungues equal and

simple. Genitalia. Tergum VIII with basal 0.60-0.90 retracted into segment

VII, wide, base and apex slightly concave, several long stout setae along apical

margin, moderately long thin setae and numerous broad dark scales on apical

0.35-0.55, index 0.47-0.60; sternum VIII large, wide, base very slightly con-

cave, apex with a small median indentation and a small lobe on each side of mid-

line, numerous setae and broad dark scales on apical 0. 70-0. 80, apical inter-

segmental membrane very lightly pigmented mesally, index 0. 63-0. 73; tergum

IX wide, moderately pigmented with a shallow median apical indentation, 1-6

setae apically on each side of midline, index 0. 42-0. 64; insula tongue-like, long,

moderately pigmented, covered with minute spicules and with 2-5 tiny tuberculi

on apical 0. 20; lower vaginal lip, narrow, moderately pigmented, covered with

small hair-like spicules; upper vaginal lip narrow, posterior margin somewhat

. Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

flattened, moderately to heavily pigmented, covered with small hair-like

spicules, upper vaginal sclerite large, moderately to heavily pigmented; post-

genital lobe moderately long, apex with a small to moderately deep median

indentation, 7-14 setae on each side of midline, completely covered with minute

hair-like spicules, dorsal PGL index 0. 79-1.10, ventral PGL index 1. 47-2. 03;

cercus short, apex bluntly rounded, mesal margin slightly concave, completely

covered with minute spicules, scales absent, dorsal surface with 4-5 long stout

setae apically and several moderately long and short setae on remainder of

apical 0.40-0.50, ventral surface with 7-14 short setae on apical margin,

index 2.05-2.69, cercus/ dorsal PGL index 2. 58-3. 64; 3 heavily pigmented,

spherical seminal capsules, 1 large and 2 slightly smaller ones; base of

accessory gland duct heavily pigmented.

PUPA. Cephailothorax. Setae 1, 7, 8-C single to triple; 2, 9-C single

or double; 3, 6-C single; 4-C single to 6 branched; 5-C double to 6 branched; 1,

2-C short; 3, 6-C very long and stout; 4, 5, 7-9-C moderately long. Melano-

tum. Seta 10-C single to 5 branched; 11-C single or double; 12-C single to

4 branched. Respiratory trumpet. Moderately pigmented; a number of scat-

tered, short setae on inner surface of meatus; pinna 0. 29-0. 38 length of meatus;

index 4.46-5.77. Abdomen. Setae 0-II-VII single; 0-VIII single or double; 1-I

with 11-34 branches on basal 0. 33; 1-II single to 6 branched; 1-III, IV single to

triple; I-V, VII single; 1-VI single or double; 2-I, II single to triple; 2-II,

IV-VII single or double; 3-I, II single; 3-II, VI single or double; 3-IV double

to 6 branched; 3-V single to triple; 3-VII single to 4 branched; 4-I triple to 8

branched; 4-II, V double to 7 branched; 4-III, IV single to 6 branched; 4-VI

double to 5 branched; 4-VII single to triple; 4-VIII single or double; 5-I single to

5 branched; 5-II single or double; 5-III single to triple; 5-IV-VII single; 6-I,

II, V single; 6-III, IV, VI single or double; 6-VII double to 5 branched; 7-I, VII

single to triple; 7-II single to 4 branched; 7-III single to 7 branched; 7-IV, VI

single to 5 branched; 7-V double to 8 branched; 8-III single to 4 branched; 8-IV

single to 5 branched; 8-V, VII double to 5 branched; 8-VI double to 6 branched;

9-I single or double; 9-II-VI single; 9-VII double to 10 branched; 9-VIII with 5-12

branched; 10-I, II single to triple; 10-IIl, V-VII single or double; 10-IV single to

4 branched; 11-II[-VI single; 11-VII single or double; 14-III-VII single; 14-VIII

single or double; 3-I-III and 5-III-V very long and stout; 6-VII short, slightly

cephalad and mesad to 9-VII; 9-VII, VIII long, stout and barbed. Paddle. Ovoid,

with minute spicules on apical 0. 80-0. 89 of outer and apical 0. 15-0. 29 of inner

margins; midrib moderately pigmented and reaches apex; 1-P moderately long,

barbed, double to 14 branched; index 1.30-1. 76.

LARVA. Head. Moderately pigmented; setae 0, 1, 3, 5, 6, 13-C

single; 4-C with 4-14 branches; 7-C with 6-10 branches; 8, 10, 12-C single or

double; 9-C single to triple; 11-C with 4-8 branches; 14-C double to 8 branched;

15-C double to 4 branched; basal maxillary hair double to 15 branched; 4-C

moderately long, mesad and slightly cephalad to 6-C; 5-C long, well caudad of

6-C; 6-C long, well mesad to 7-C; 7-C moderately long, cephalad and well

laterad to 5-C; mental plate heavily pigmented with 18-22 teeth; mouth brush with

mesal hairs pectinate apically. Antenna. Short, moderately pigmented, a very

few minute spicules on shaft; 1-A short, double or triple, attached 0. 51-0. 62

from base; 2-A long; 3-A short, 0.31-0.38 length of 2-A; 4-A moderately long,

0. 60-0. 83 length of 2-A; 5-A short, flattened; 6-A short, peg-like, 0. 29-0. 33

length of 2-A; 2, 3-A attached subapically. Thorax. Seta 0-P with 6-14

branches; 1-P triple to 5 branched; 2, 4-6, 10, 12-P single; 3-P with 4-15

branches; 7, 11-P double or triple; 8-P with 6-18 branches; 9-P double to 7

branched; 13-P with 4-21 branches; 14-P triple to 6 branched; 1-M with 5-9

Reinert: Bothaella, new subgenus D

branches; 2-M double to 4 branched; 3-5, 7, 10-12-M single; 6-M triple to 6

branched; 8-M with 4-6 branches; 9-M double to 6 branched; 13-M with 8-19

branches; 14-M with 6-16 branches; 1-T with 4-12 branches; 2, 6, 10-12-T

single; 3-T double to 5 branched; 4-T single to triple; 5-T double to 9 branched;

7-T with 5-8 branches; 8-T with 5-14 branches; 9-T double to 4 branched; 13 _T

with 5-16 branches. Abdomen. 0-II-VIII single: 1-I with 5-10 branches; 1-1

with 4-9 branches; 1-III with 4-10 branches; 1-IV triple to 9 branched; 1-V

triple to 7 branched; 1-VI triple to 6 branched; 1-VII triple to 5 branched; 1-

VII double to 11 branched: 2-I triple to 10 branched; 2-II triple to 14 branched:

2-I0 triple to 11 branched: 2-IV, V single to 11 branched; 2-VI single to 9

branched; 2-VII with 5-10 branches; 2-VII single; 3-I-VI single; 3-VII single to

5 branched; 3-VIII double to 8 branched; 4-I with 4-9 branches; 4-II double to

6 branched; 4-III double to 8 branched; 4-IV double to 9 branched; 4-V triple to

7 branched; 4-VI, VIII single; 4-VI single or double; 5-I triple to 8 branched;

5-II triple to 6 branched; 5-II[ triple to 7 branched; 5-IV, VI double to 5

branched; 5-V, VI triple to 5 branched; 5-VIII double to 7 branched; 6-I triple

to 6 branched; 6-II triple or 4 branched; 6-III double or triple; 6-IV-VI double;

6-VII with 4-7 branches; 7-I single to triple; 7-II double; 7-III triple to 9

branched; 7-IV with 4-10 branches; 7-V triple to 8 branched; 7-VI double to 4

branched; 7-VII single or double; 8-II single to 4 branched; 8-III, V single or

double; 8-IV single; 8-VI triple to 8 branched; 8-VI with Bes 14 branches; 9-I

single to triple; 9-II double to 7 branched; 9 -TIl, VI single to 6 branched: 9-IV,

V single to 5 branched; 9-VII double to 4 branched: 10-I, IV-VI single; 10-11,

III single or double; 10-VII single to 4 branched; 11-1 with 5-16 branches; ve

II-(V, VI double to 5 branched; 11-V double to 4 branched; 11-VII single to

triple; 12-II single or double; 12-III single to triple; 12-IV double or triple; 12-

V, Vi single; 12-VII single to 4 branched; 13-I with 4-15 branches; 13-II triple

to 16 branched; 13-III triple to 11 branched; 13-IV triple to 9 branched; 13-V

double to 8 branched; 13-VI with 4-13 branches; 13-VIL with 4-9 branches; 14-

II, V, VID single; 14-III, IV, VI, VII single or double; VIII with comb com-

posed of 15-26 scales arranged in 2 irregular rows; 1-X long, stout, barbed,

double to 4 branched, attached near posterior margin of saddle; 2-X long, stout,

with 4-6 branches; 3-X very long, stout, single; ventral brush composed of 8-9

setae on grid, each single to triple, no precratal setae, basal 1-2 setae moder-

ately long, remainder long; saddle moderately pigmented, with large spicules

on posterior margin, acus absent; 4 anal papillae, long and slender. Siphon.

Heavily pigmented, acus well developed, index 2. 90-4. 89; pecten on basal 0.36

of siphon, composed of 12-17 evenly spaced teeth, each tooth with a slender

attenuated filament with small denticles on ventral margin; 1-S long, barbed,

double to 5 branched, base attached on basal 0. 43 of siphon and distad to last

pecten tooth; 2-S short, single and on apical margin of siphon; 6, 8, 9, 13-S

short and single.

EGG. Not known.

DISTRIBUTION. This subgenus is currently known from Thailand

from the following 4 provinces: Chiang Mai, Kanchanaburi, Lampang and Nan

and from Cambodia from 2 provinces: Kompong Som and Kompong Speu.

The known distributions of the 3 included species of Bothaella are

plotted on the map in Figure 1.

TAXONOMIC DISCUSSION. The subgenus Bothaella is characterized

and separated from the other subgenera of Aedes by the following: in the adults

by the combination of (1) head with only decumbent broad scales on vertex and

erect forked scales restricted to occiput, (2) antennal pedicel with a large

patch of overlapping, broad, silvery scales on mesal and ventral areas, short

6 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

fine hairs absent, (3) dorsocentral (anterior and posterior) setae well developed,

(4) acrostichal setae absent, (5) prosternum bare, (6) propleuron with 2-4

setae, (7) mesepisternum with 1-2 upper and 1-2 posterior setae, (8) mese-

pimeron with 3-8 upper and 1 lower setae, (9) male maxillary palpus short

(0.18-0.19 length of proboscis) and (10) male posttarsi I-III each with 2 ungues,

I with ungues unequal, each with a tooth, II with ungues unequal, smaller one

with a tooth and III with ungues equal and simple; in the male genitalia by the

combination of (1) development of the aedeagus which is composed of 2 lateral

plates each with lateral surfaces curved tergally and presence of poorly to

well developed short distal teeth, (2) proctiger without cercal setae, (3) de-

velopment of basal mesal lobe into a large expanded distal lobe bearing 28-244

short setae, (4) development of gonocoxite and (5) development of gonostylus;

in the female genitalia by the combination of (1) tergum VIII shape and number of

scales, (2) sternum VII shape and number of scales, (3) tergum IX shape and

number of setae, (4) insula tongue-like with distal tuberculi, (5) development of

postgenital lobe, (5) cercus short, mesal margin slightly concave and absence

of scales and (6) 1 large and 2 slightly smaller spherical, seminal capsules; in

the pupae by the combination of (1) setae 1, 2-C short and usually single, 3-C

very long, stout and single, (2) setae 4, 5-C approximately equally developed

in length and branching, (3) seta 6-C very long, stout and single and 7-C short

to moderately long and single to triple, (4) setae 3-II-III and 5-IV-V very long,

stout and single, (5) 6-VII short, double to 5 branched and slightly cephalad and

mesad to 9-VII, (6) 9-VII-VII long, stout, barbed, double to 10 branched on VII

and 5-12 branched on VIII and (7) paddle with small spicules on outer and inner

margins but without hair-like fringe and seta 1-P multiple branched and barbed;

and in the fourth stage larvae by the combination of (1) setae 5, 6-C stout, long

and single, 4-C moderately long, multiple branched (4-14 branched), mesad

and slightly cephalad to 6-C, 5-C well caudad to 6-C, (2) antenna short with

seta 1-A short and double or triple, (3) seta 13-P well developed with 4-21

branches, (4) setae 6-I-VI long, barbed and double on IV-VI, (5) seta 1-VII long,

stout and triple to 5 branched and 3-VII short, (6) seta 13-VII cephalad and well

separated from 10-VI, (7) seta 1-X long, stout, barbed, double to 4 branched

and attached near caudal margin of saddle, (8) saddle incompletely rings segment

X and ventral brush composed of 8-9 setae each of which is single to triple and

(9) siphon with pecten of 12-17 evenly spaced teeth and no detached ones, 1-S

long, barbed, double to 5 branched and attached on basal 0.45 of siphon distad

to last pecten tooth.

The adult habitus and chaetotaxy of the subgenus Bothaella is most

similar to those of the subgenera Stegomyia Theobald and Christophersiomytia

Barraud of Aedes and the genus Heizmannia Ludlow. This subgenus can be

separated from Stegomyia which is outlined by Reinert (1973b) by the following

characters: prosternum bare; propleuron with 2-4 setae; mesepimeron with 1

lower seta; remigial setae present and maxillary palpus of male short, total

length 0. 18-0.19 length of proboscis while Stegomyia possesses: prosternum

with broad scales, at least on dorsolateral areas; propleuron with 5-7 setae;

mesepimeron without lower setae; remigial setae absent and maxillary palpus

of male usually approximately equal in length to proboscis (some members of the

albolineatus group have palpi 0.48 length of proboscis). The subgenus Christo-

phersiomyia possesses the following which differ from Bothaella: prosternum

with white scales; propleuron with 6-14 (usually 3-10) setae; paratergite and

postpronotum with numerous scales; scutum with a large patch of narrow,

curved, white scales on anterior 0.50 and proboscis with a broad band of white

scales near middle. Bothaella resembles the genus Heizmannia but lacks the

Reinert: Bothaella, new subgenus 7

following features of this genus (see Reinert 1973a): prosternum covered with

broad silvery scales; mesepisternum without setae on upper area; anterior

dorsocentral setae absent (setae anterior to prescutellar space); prescutellar

space completely covered with scales; mesopostnotum with setae (subgenus

Heizmannia only) and antepronota large, nearly contiguous or only moderately

separated.

The short maxillary palpus of the male is also similar to those of the

following subgenera of Aedes: Aedes Meigen, Cancraedes Edwards, Christo-

phersiomyia, Geoskusea Edwards, Huaedes Huang, Leptosomatomyia Theobald,

Neomacleaya Theobald, Nothoskusea Dumbleton, Pavaedes Edwards, Rhinosku-

sea Edwards and Verrallina Theobald. Members of the subgenus Bothaella do

not resemble any of these subgenera in the adult habitus and are easily separated

from them by one or more of the following characters: absence of acrostichal

setae; number of setae on propleuron; number of setae on mesepisternum; ab-

sence of short fine hairs on antennal pedicel and development of the female and

male ungues of the posttarsi.

Male genitalia of Bothaella have a resemblance to those of the subgenus

Leptosomatomyia and to a lesser extent to the subgenus Huaedes in the develop-

ment of tergum IX, gonostylus and basal mesal lobe but differ in the development

of the gonocoxite, proctiger and aedeagus. The female genitalia also show

similarities to Leptosomatomyia and Huaedes but differ in the development of

tergum IX, absence of scales on the cercus and in the latter subgenus by the

number of tuberculi on the insula. Female genitalia of Leptosomatomyia also

differ from both Bothaella and Huaedes by having the apical intersegmental

membrane of sternum VIII moderately to heavily pigmented.

Pupae of Bothaella are most similar to members of the subgenus Stegomyia

but are easily differentiated by a combination of characters 1, 3, 6 and 7 as listed

above.

The fourth stage larvae of Bothaella resemble species of Stegomyia,

Finlaya, Howardina Theobald and Heizmannia. From the Stegomyia they are

easily separated by: the number of setae in the ventral brush; presence of seta

13-P, however, this seta is present and well developed in hoguei Belkin, upolensis

Marks, vobinsoni Belkin and tulagiensis Edwards; and setae 10-VII and 13-VII

are well separated (these 2 setae are also well separated in robinsoni and tulagi-

ensis). The larvae of these 4 species of Stegomyia are similar in many respects

to the subgenus Bothaelia but are unlike it in the development of the ventral brush

and seta 3-X which is not single in these species. From the Finlaya they are

separated by the following combination of characters: seta 6-C single; seta 10-P

single; seta 13-P present; and setae 6, 8-S short. Larvae of the sexlineatus

section and 2 species of the walkeri section (i.e., cozumelensis Diaz Najera and

toliota Dyar and Knab) of Howardina (subgenus as defined by Berlin, 1969) have

a number of similarities to those of Bothaella especially in the presence of seta

13-P; however, Howardina differ by possessing the following features: setae 5,

6-C not both single (except 2 species of walkeri section); 3-VII very long and

single; 13-VII close to and not well cephalad of 10-VII; and shape of the comb

scales. Bothaella larvae are very Similar in many respects to those of the

genus Hezzmannia but are differentiated by the following combination of features

of the latter genus: seta 5-C barbed and multiple branched (3-16 branched, usually

with 5 or more branches except in complex Theobald in which 5-C is single to

triple, of 14 larvae and larval skins in the SEAMP collection 5-C was single in

_ 2, double in 11 and triple in 1); absence of seta 13-P; seta 13-VII attached close

to seta 10-VII; and seta 2-X usually double or triple (occasionally 4 branched in

achaetae (Leicester), catesi (Lien), communis (Leicester), demeilloni Mattingly,

8 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

macdonaldi Mattingly and triple to 7 branched in reidi Mattingly).

This subgenus is dedicated to Dr. Botha de Meillon, Principal Investi-

gator, Southeast Asia Mosquito Project, Smithsonian Institution, in acknow-

ledgement of his many contributions to the study of the world Culicidae and his

suidance and inspiration.

MEDICAL IMPORTANCE. Nothing is known about the medical signifi-

cance of the species included in this subgenus.

BIOLOGY. The most common habitat of fitiniatar’ stages is water in

small rock pools, however, occasionally they have been found in bamboos.

Larvae and pupae have been collected from clear or colored, fresh, temporary

water in small rock pools (a number of times from volcanic rock) and several

times from split bamboo, bamboo stumps and cups, located in partial or heavy

shade, situated in primary and secondary rainforests, secondary deciduous

forests and bamboo groves, and always in mountainous terrain between altitudes

of 75 and 1520 meters.

Adults of kleini were collected in a small dry forest and 1 female was

taken biting man between 1800 and 1900 hours in a littoral rainforest.

KEYS TO SPECIES OF AEDES (BOTHAELLA)

ADULTS

2. Mesepisternum with 2 setae; ee with 2-3 posterior setae.

eldridgei

Mesepisternum with 3 setae; Ee with 3-5 posterior setae.

helenae

MALE GENITALIA

1. Basal mesal lobe with 28-37 short setae; gonostylus with 5-8 tiny setae on

SSNANGCE DOVGON se ' goping ost Seb gs vations a He SEH. EAE E ne awe helenae

Basal mesal lobe with 70-244 short setae; gonostylus with 10-14 tiny setae

On ORDaKdeU POFtlONjes “essere: Vocal she a eke ee SO a ea wud See 2

2. Basal mesal lobe with 70-78 setae and with a ventromesal finger-like pro-

jection terminating with a broad apically flattened seta. . . . eldridgei

Basal mesal lobe with 201-244 setae and without a finger-like projection.

kleini

PUPAE*

Paddle hair 1 double to 5 branched; abdominal hair 9-VII double to 4 branched.

eldridgei

Paddle hair 1 with 6-14 branches; abdominal hair 9-VII with 5-10 branches.

helenae

*Pupae of kleini are not known.

Reinert: Bothaella, new subgenus 9

FOURTH STAGE LARVAE*

Head hair 4-C with 4-8 branches; abdominal hair 2-II with 7-14 branches.

helenae

Head hair 4-C with 9-14 branches; abdominal hair 2-II with 3-5 branches.

eldridgei

DESCRIPTION OF THE SPECIES OCCURRING IN SOUTHEAST ASIA

AEDES (BOTHAELLA) HELENAE, NEW SPECIES

(Pigas (2) 45.5) 8) 44.713)

MALE (Fig. 2). Head. Antenna brown, plumose, pedicel with a patch

of overlapping broad, silvery scales on mesal and ventral surfaces, flagellomere

1 with a few brown scales, total length approximately 0.81 length of proboscis;

clypeus blackish-brown, bare; maxillary palpus short, dark brown scaled, 4

segmented, segments 1 and 2 short, segment 3 approximately 0.48 length of

segment 4 which is long and slightly longer than combined length of basal 3

segments, total length approximately 0.19 length of proboscis; proboscis dark

brown scaled, total length approximately 1.08 length of femur I; eyes contiguous;

vertex and lateral areas covered with overlapping, broad, decumbent scales, a

dorsomedian triangular patch of silvery scales anteriorly with apex extending

onto dorsal portion of interocular space, a small patch of silvery scales on

postgena next to eye, remainder of scales blackish-brown; occiput with numerous

blackish-brown, erect, forked scales; several dark, long ocular setae; 2 dark,

long interocular setae. Thorax. Scutal integument blackish-brown; scutum

covered with narrow, curved, blackish-brown scales with prescutellar space bare;

scutellum with a patch of broad blackish-brown scales on lateral lobe, median

lobe with a basal patch of broad silvery scales and broad blackish-brown ones

apically; following areas with setae which are dark and well developed: 2 median

anterior promontory, numerous dorsocentral (anterior and posterior), scutal

fossal (2-3 anterior, 2-3 lateral and 1 posterior), numerous supra-alar, 6-8

posterior medial scutal (pale), 1 postalar callar and scutellar (3 long and 2-4

short lateral; 4-5 long and 2-4 short median); pleural integument blackish-

brown; antepronotum with a large patch of overlapping, broad, silvery scales,

7-8 long dark setae; postpronotum with 3-4 long, dark, posterior setae; pro-

pleuron with a large patch of overlapping, broad, silvery scales, 2-3 long,

dark setae; prosternum, subspiracular area, paratergite, mesomeron and

metepisternum bare; postspiracular area with a large patch of overlapping,

broad, silvery scales, 1-2 long, dark setae; mesepisternum with an upper and

a posterior patch of overlapping, broad, silvery scales, 2 long, dark, upper

setae and 1 long, dark lower one; prealar knob with 5-8 moderately long, dark

setae; mesepimeron with a patch of overlapping, broad, silvery scales on upper

area, a patch of 3-5 moderately long, dark setae posterior to scale patch on

upper area and 1 long, dark seta on lower area. Legs (Fig. 4). Coxae I-III

each with several moderately long and long setae, I with anterior and lateral

areas covered with broad blackish-brown scales and a dorsal and an antero-

ventral patch of broad silvery ones, II and III each with an anterior patch of

broad silvery scales; trochanters I-III each with broad silvery scales; femora

*Larvae of kleini are not known.

10 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

I and II each covered with broad blackish-brown scales, I with an anteroapical

silvery scaled spot, afew similar scales at base and forming a narrow postero-

dorsal stripe on basal 0.40-0.50, II with a dorsoapical silvery scaled spot, a few

similar scales at base and forming an indistinct patch on basal 0. 60-0. 70 of

posterior surface, III with anterior and posterior surfaces with basal 0. 75-0. 80

and apex silvery scaled, remainder blackish-brown scaled, a few brown scales

on dorsal margin of basal silvery area, total length of I approximately 1.11

length of II; tibiae I-III each blackish-brown scaled, II with a few dorsoapical

silvery scales, III with a small ventroapical patch of silvery scales; tarsus I-III

each blackish-brown scaled, III with tarsomeres 1 and 2 each with a narrow

basal and apical white scaled band, tarsomeres 3 and 4 each with a small

dorsobasal and dorsoapical white scaled spot, tarsomere 5 pale scaled; post-

tarsi I-III each with 2 ungues, I with ungues unequal, both with a tooth, II with

ungues unequal, larger one simple and smaller one with a tooth, III with ungues

equal and simple. Wing. Veins covered with blackish-brown scales dorsally and

ventrally; alula with several narrow blackish-brown scales along margin; upper

calypter with numerous dark setae; 1 remigial seta. Halter. Pedicel pale;

capitellum silvery scaled with dark brown scales at base. Abdomen. TergalI-

VII each blackish-brown scaled with a large laterobasal silvery scaled spot;

sterna II-VII each blackish-brown scaled with a basal silvery scaled band;

terga and sterna with numerous golden setae, mostly along posterior margins.

Genitalia (Fig. 8). Tergum IX band-like with lateral caudal areas broad and

eradually tapering mesally, moderately pigmented, 3-8 stout setae more or less

in a longitudinal line on each side of midline, a number of small spicules scat-

tered mostly on basal area; gonocoxite short, broad, heavily pigmented, mesal

surface membranous, entire surface covered with minute spicules, dorsal sur-

face with numerous moderately long thin setae, lateral margin with a few long

stout setae, ventral surface with numerous short thin setae over entire area

except for a large circular apical mesal area which is covered with a tightly

packed patch of moderately pigmented, short, broadly flattened, erect, scale-

like setae, lateral and ventral surfaces with scattered broad dark scales;

gonostylus long, total length, minus claw, approximately 0.80 length of gono-

coxite, apical 0.45 expanded but sloping to a blunt apex and with 4-7 short fine

setae along outer margin and 1-2 similar ones on inner margin, gonostylar claw

short, apex broad and rounded, heavily pigmented and attached apically; basal

mesal lobe with apical portion large, expanded with 28-37 short, stout, evenly

spaced setae, those along apical margin stouter, basal portion covered with

small spicules and connected mesally with its mate; proctiger long, paraproct

moderately pigmented with apex blunt and heavily pigmented, cercus membranous,

cercal setae absent; tergum X narrow, heavily pigmented; phallosome with

aedeagus divided into 2 lateral, moderately pigmented plates which are connected

basally and narrowly at apex, each lateral plate with lateral area curved ter-

gally and with a mesal, longitudinal, heavily pigmented scleritized area termina-

ting in 3-4 poorly developed, short teeth (teeth visible in lateral view), paramere

moderately long, approximately 0. 82 length of aedeagus, heavily pigmented,

parameral apodeme long, approximately 1.25 length of paramere, heavily pig-

mented; sternum IX large, basal, lateral and median areas moderately pigmented,

entire surface covered with minute spicules, 4-7 short to moderately long setae

near center.

FEMALE (Fig. 2). Similar to male in general habitus but with the

following differences. Head. Antenna with basal 0. 85 of flagellomere 1 pale,

total length approximately 0.83 length of proboscis; maxillary palpus approxi-

mately 0.19 length of proboscis; proboscis approximately 1.05 length of femur I.

Reinert: Bothaella, new subgenus 11

Thorax. 2-4 median anterior promontory setae; 3-4 anterior scutal fossal

setae; scutellar setae (3 long and 3-6 short lateral; 4-5 long and 3-5 short

median); antepronotum with 6-9 setae; postpronotum with 3-5 setae; prealar

knob with 5-9 setae; propleuron with 2-4 setae; mesepimeron with 4-8 upper

setae and 1 lower one (1 specimen with 1 long and 1 short lower setae).

Legs (Fig. 4). Femur I total length approximately 1.07 length of II; tarsus I

with tarsomere 1 with a narrow, basal, white scaled band, tarsomeres 2 and

3 each with a small dorsobasal white scaled spot, II with tarsomeres 1-4 each

with a narrow, basal white scaled band and a small, dorsoapical, white scaled

spot, III with tarsomeres 1-4 each with a moderately broad basal and apical

silvery scaled band, tarsomere 5 completely silvery scaled; tarsilI and II of some

specimens with pale scaled areas reduced; posttarsi I-III each with 2 ungues,

I with ungues equal, both with a tooth, II with ungues equal, both with a tooth,

III with ungues equal and simple. Genitalia (Fig. 5). Tergum VIII with basal

0.60-0. 80 retracted into segment VII, wide, base and apex slightly concave,

several long stout setae along apical margin, moderately long thin setae and

numerous broad dark scales on apical 0.35, scales extending 0.50 basally along

lateral margins, index 0. 51-0. 60; sternum VIII large, wide, base slightly

concave, apex with a small lobe on each side of midline, numerous setae and

broad dark scales on apical 0. 80, index 0. 63-0. 73; tergum IX wide, moderately

pigmented with a median apical indentation, 3-6 setae apically on each side of

midline, index 0.42-0. 59; insula tongue-like, long, moderately pigmented,

covered with minute spicules and with 3-5 tiny tuberculi on apical 0.20; lower

vaginal lip moderately pigmented, covered with hair-like spicules; narrow and

with base broader; upper vaginal lip narrow, moderately to heavily pigmented,

covered with hair-like spicules; upper vaginal sclerite large, moderately to

heavily pigmented; postgenital lobe moderately long, apex with a moderately

deep median indentation, 7-10 setae on each side of midline, completely covered

with minute hair-like spicules, dorsal PGL index 0.92-1.10, ventral PGL index

1, 83-2.03; cercus short, apex bluntly rounded, mesal margin slightly concave,

completely covered with minute spicules, scales absent, dorsal surface with 4-5

long stout setae apically and several moderately long and short setae on remainder

of apical 0.45, ventral surface with 8-12 short setae on apical margin, index

2.34-2.58, cercus/dorsal PGL index 2.66-3.24; 3 heavily pigmented, spherical

seminal capsules, 1 large and 2 slightly smaller ones; base of accessory gland

duct heavily pigmented.

PUPA (Fig. 11). Chaetotaxy as figured and recorded in table 1.

Respiratory trumpet. Moderately pigmented; a number of scattered, short

setae on inner surface of meatus; index 4.46-5.24, average 4.95. Abdomen.

Seta 4-III triple to 6 branched; 4-V triple to 7 branched; 4-VI double to 5

branched; 7-VII single to triple; 9-VII with 5-10 branches; 9-VIII with 6-12

branches. Paddle. Ovoid, with minute spicules on apical 0. 85-0. 89 of outer

and apical 0. 15-0. 24 of inner margins; midrib moderately pigmented and

reaches apex; 1-P moderately long, barbed, 6-14 branched; index 1. 46-1. 76,

average 1.62.

LARVA (Fig. 13). Chaetotaxy as figured and recorded in table 3.

Head, Seta 4-C moderately long, 4-8 branched; 5, 6-C long, single; 7-C

moderately long, 7-10 branched; 14-C moderately long to long, 5-8 branched;

basal maxillary hair 9-15 branched; mental plate with 18-20 (usually 19) teeth;

mouth brush with mesal hairs pectinate apically. Antenna. Short, moderately

pigmented; seta 1-A short, double, attached 0.51- 0.62 from base. Thorax.

Seta 3-P with 11-15 branches; 8-P with 12-18 branches; 9-P with 4-7 branches;

13-P with 13-21 branches; 14-P with 4-6 branches; 13-M with 13-19 branches;

12 | Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

14-M with 10-16 branches; 5-T with 6-9 branches; 8-T with 10-14 branches;

13-T with 10-16 branches. Abdomen. Seta 1, 2-I with 7-10 branches; 4-I with

5-7 branches; 11-I with 10-16 branches; 13-I with 8-15 branches; 2-II with 7-14

branches; 4-II with 4-6 branches; 13-II with 8-16 branches; 2-IIT with 5-10

branches; 4-III with 5-8 branches: 9-III triple to 6 branched: 13-III with 7-11

branches; 2-IV with 8-11 branches; 4-IV with 4-9 branches; 9-IV triple to 5

branched; 2-V with 8-11 branches; 5-V with 4-5 branches; 9-V triple to 5

branched; 13-V with 4-8 branches; 2-VI with 6-9 branches; 5-VI with 4-5

branches; 8-VI with 4-8 branches; 9-VI triple to 6 branched; 2-VII with 7-10

branches; 1-VIII with 8-11 branches; 3-VII with 5-8 branches; 1-X long, barbed,

double to 4 branched; 2-X long, 4-6 branched; 3-X long, single: comb with 18- 25

(usually 22) scales arranged in 2 irregular rows; saddle incompletely rings seg-

ment X, acus absent, moderately to heavily pigmented with long spine-like

spicules on posterior margin; ventral brush of 8 setae on grid, each single to

triple; 4 anal papillae, long and slender. Siphon. Heavily pigmented, acus well

developed, index 4. 08-4. 89; pecten on basal 0.31 of siphon, composed of 13-16

(usually 15) evenly spaced teeth, each with a few denticles on ventral margin;

seta 1-S long, barbed, double or triple, base attached on basal 0.40 of siphon

and distal to last pecten tooth.

EGG. Not known.

TYPE-DATA. Type series includes holotype male, allotype and 33 para-

types (15 males, 18 females). Holotype male with associated pupal and larval

skins, THAILAND, Chiang Mai, Amphoe Chiang Dao, Ban Tham Kraeb, 7

September 1970, Kol Mongkolpanya collector, collection number 04770-9, SEAMP

accession number 271, genitalia preparation number T73.138, collected as a

larva from fresh colored water in a partially shaded, small, rock pool located

in a deciduous forest in mountainous terrain at an altitude of 1000 meters; allo-

type with associated pupal and larval skins, collection number 04770-14, other

data as in holotype; paratypes 12 males with associated pupal and larval skins

(collection numbers: 04770-1, 04770-2, 04770-4, 04770-5, 04770-7, 04770-11,

04770-19, 04765-3, 04765-4, 04766- t: 04766-3, 04766-9), 2 males with

associated pupal skins (collection numbers: 04770- 100, 04765-100), 15 females

with associated pupal and larval skins (collection numbers: 04770-8, 04770-12,

04770-13, 04770-15, 04770-16, 04770-20, 04770-21, 04765-1, 04765- 2, 04765-6,

04765-7, 04765-8, 04765- 9; 04.766- 2, 04766- rL),o2 females with associated pupal

skins (collection numbers: 04770- 101, 04766- 100), 1 male (collection number

04765-10) and 1 female (collection number 04770- 6), all with same data as holo-

type. All type material is deposited in the U. S. National Museum (Natural

History), Washington, D. C., except 1 male and 1 female paratypes (both with

associated skins) which will be deposited in the British Museum (Natural History),

London, England, and 2 male and 2 female paratypes (all with associated skins)

which will be sent to the Department of Entomology, SEATO Medical Research

Laboratory, Bangkok, Thailand.

DISTRIBUTION. Material examined: 700, 1119, 85 L, 177 with associ-

ated skins (46 p, 1311).

THAILAND. Chiang Mai, Ban Huai Kaeo, Ban Huai Tat, Ban Tham

Kraeb, Chiang Dao, Doi Pha Daeng; 510, 779, 127p, 46 L, 113 1. Lampang,

San Chao Pho Phatu Pha; 1% 79, 8p, 17 L. Nan, Doi Chik Chong; 18¢, 279,

42 p, 22 L, 181.

TAXONOMIC DISCUSSION. The adult habitus of helenae, eldridgei

and kleini are very similar but can be separated in both sexes by the following

list of features, helenae possesses: mesepisternum with 2 upper and 1 posteri-

or setae, tarsus II with basal white scaled bands on tarsomeres 1-4 and post-

Reinert: Bothaella, new subgenus 13

pronotum with 3-4 (male) or 3-5 (female) posterior setae, eldridgei possesses:

mesepisternum with 1 upper and 1 posterior setae, tarsus III with basal white

scaled bands on tarsomeres 1-4 and postpronotum with 2 (male) or 2-3 (female)

posterior setae, and kleini possesses: mesepisternum with 1-2 upper and 1-2

posterior setae, tarsus III completely brown scaled and postpronotum with 2

(male and female) posterior setae. Adults of helenae and eldridgei each have

an upper and a lower patch of scales on the mesepisternum while those of kleini

_ possess a single large patch of scales extending over upper, posterior and lower

portions of this structure.

Male genitalia of helenae show close similarities in morphological

development to both eldridgei and kleini but each species has diagnostic differ-

ences as follows--helenae possesses: basal mesal lobe with 28-37 setae,

sternum IX with 4-6 setae, tergum IX with 3-8 setae on each side of midline,

gonostylus with 4-7 short setae on outer and 1-2 similar ones on inner surfaces,

and gonocoxite with a closely packed patch of short, broad, scale-like setae distally

on sternomesal area; eldridgei possesses: basal mesal lobe with 70-78 setae

and a ventromesal finger-like projection at apex terminating in a flattened seta,

sternum IX with 8-9 setae, tergum IX with 3-4 setae on each side of midline,

gonostylus with 7-8 short setae on outer and 3-6 similar ones on inner surfaces,

and gonocoxite with a group of short setae distally on sternomesal area; and

kleini possesses: basal mesal lobe with 201-244 setae, sternum IX with 9-11 setae,

tergum IX with 7-9 setae on each side of midline, gonostylus with 9-11 short

setae on outer and 1 similar one on inner surfaces, and gonocoxite with a closely

packed patch of short, broad, scale-like setae distally on sternomesal area.

The female genitalia of helenae are very similar to both eldridgei and

kleini but are separated by the following, helenae with: tergum IX with 3-6

setae on each side of midline, postgenital lobe with 7-10 setae on each side of

midline, and cercus index 2,34-2.58; eldridgei with: tergum IX with 3-5 setae

on each side of midline, postgenital lobe with 11-14 setae on each side of mid-

line, and cercus index 2.05-2.20; and kleini with: tergum IX with 1-2 setae on

each side of midline, postgenital lobe with 7-8 setae on each side of midline,

and cercus index 2. 46-2. 69.

Pupae of helenae and eldridgei are very similar but can be distinguished

from each other by seta 9-VII which is 5-10 branched in helenae and double to 4

branched in eldridgei and seta 1-P is 6-14 branched in the former species and

double to 5 branched in the latter species.

The larvae of helenae and eldridgei are similar in many features but

are easy to separate by using the setae mentioned in the key and species descrip-

tions. Some of the most distinctive setae are as follows, in helenae: 4-C with

4-8 branches; 3-P with 11-15 branches; 5-T with 6-9 branches; 13-T with 10-16

branches; 11-I with 10-16 branches; 2-II with 7-14 branches; 9-III triple to 6

branched; 2-IV, V, 1-VIII with 8-11 branches; 9-IV triple to 5 branched and 2-VI

with 6-9 branches, while eldridgei has: 4-C with 9-14 branches; 3-P with 4-8

branches; 5-T double to 4 branched; 13-T, 11-I with 5-7 branches; 2-II triple to

9 branched; 9-III-V single; 2-IV single to 4 branched; 2-V, VI single to triple

and 1-VII double to 5 branched. The siphonal index of helenae is 4. 08-4. 89

as compared to 2.90-3. 63 for eldridgei.

The larval skin of the allotype female (04770-14) has 2 setae 4-C on the

left side, their bases are side by side and they are both 5 branched, seta 4-C on

the right side is 4 branched.

This species is dedicated to my wife, Mary Helen (Mollie), for her con-

tinued encouragement and assistance in typing my manuscripts.

BIOLOGY. The immature habitat is water in small rock pools. Im-

14 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

matures were collected from clear or colored, fresh, temporary water (usually

containing dead leaves) in small rock pools, small rock pools in volcanic rock,

twice each from small split bamboo and small bamboo stumps, and once from a

bamboo cup, all located in mountainous terrain. Collection sites of the im-

matures were in partial or heavy shade and most often in secondary deciduous

forests (15 collections) but also in primary and secondary rainforests (10 col-

lections) and at altitudes between 380 and 1520 meters (most often at or near

1000 meters). Larvae were collected once from water in a small rock pool in

association with Aedes eldridgei. :

One larva (04766-D) was heavily infected with Coelomomyces stegomyiae

Keilin. Identifitication of this parasite was made by Dr. J. N. Couch.

AEDES (BOTHAELLA) ELDRIDGEI, NEW SPECIES

(Figs. 3, 4, 6, 9, 12, 14)

MALE (Fig. 3). Head. Antenna brown, plumose, pedicel with a patch of

overlapping, broad, silvery scales on mesal and ventral surfaces, flagellomere

1 with a few brown scales, total length approximately 0.73 length of proboscis;

clypeus blackish-brown, bare; maxillary palpus short, dark brown scaled, 4

segmented, segments 1 and 2 short, segment 3 approximately 0.50 length of

segment 4 which is long and slightly longer than basal 3 segments, afew short

dark brown setae at apex of segments 2, 3 and 4, total length approximately

0.18 length of proboscis; proboscis dark brown scaled, total length approxi-

mately 1.13 length of femur I; eyes contiguous; vertex and lateral areas covered

with overlapping, broad, decumbent scales, a dorsomedian triangular patch of

silvery scales anteriorly with apex extending onto dorsal portion of interocular

space, afew pale scales on postgena next to eye, remainder of scales blackish-

brown; occiput with numerous black, erect, forked scales; several dark, long

ocular setae; 2 long, dark interocular setae. Thorax. Scutal integument

blackish-brown; scutum covered with narrow, curved reddish-brown scales

with prescutellar space bare; scutellum with a patch of broad blackish-brown

scales on lateral lobe, median lobe with a patch of broad silvery scales and

broad blackish-brown ones apically; following areas with setae which are dark

and well developed: 2 median anterior promontory, numerous dorsocentral

(anterior and posterior), scutal fossal (2-3 anterior, 2-3 lateral and 1 posterior),

numerous supra-alar, 6-7 posterior medial scutal (pale), 1 postalar callar

and scutellar (3 long and 2-3 short lateral; 4 long and 2-3 short median); pleural

integument blackish-brown; antepronotum with a large patch of overlapping,

broad, silvery scales, 7 long dark setae; postpronotum with 2 dark posterior

setae, upper one short and lower one long; propleuron with a large patch of

overlapping, broad, silvery scales, 3 long dark setae; prosternum, subspiracu-

lar area, paratergite, mesomeron and metepisternum bare; postspiracular area

with a large patch of overlapping, broad, silvery scales, 1-2 dark setae; mese-

pisternum with an upper and a posterior patch of overlapping, broad, silvery

scales, 1 long, dark upper seta and 1 long, dark lower one; prealar knob with

5-6 moderately long, dark setae; mesepimeron with a patch of overlapping,

broad, silvery scales on upper area, a patch of 3-4 moderately long, dark

setae posterior to scale patch on upper area and 1 long dark seta on lower

area. Legs (Fig. 4). Coxae I-III each with several moderately long and long

setae, I with anterior and lateral areas covered with broad, blackish-brown

scales and a dorsal and an anteroventral patch of broad silvery scales, I and

III each with an anterior patch of broad silvery scales; trochanters I-III each

Reinert: Bothaella, new subgenus 15

covered with broad blackish-brown scales and a few white ones dorsally at

base, I with white scales on posterior surface at base and forming a narrow

posterodorsal stripe on basal 0.40, one specimen with a small dorsoapical

silvery scaled spot, II with white scales on posterior surface at base and form-

ing a narrow posteroventral stripe on basal 0.55, II also with an anteroapical

small patch of silvery scales, III covered with broad silvery scales on basal

0.62 and with a narrow dorsal stripe of broad blackish-brown scales on basal

0.47, stripe overlaps slightly onto anterior and posterior surfaces, remainder

covered with broad blackish-brown scales except apex of anterior and dorsal sur-

faces which are covered with broad silvery scales, total length of I approxi-

mately 1.10 length of II; tibiae I-III each blackish-brown scaled; tarsi I-III each

blackish-brown scaled, III with tarsomeres 1 and 2 each with a narrow basal

and apical white band, tarsomeres 3 and 4 each with a small dorsobasal and

dorsoapical white scaled spot, tarsomere 5 pale scaled with a few pale brown

scales at middle; posttarsi I-III each with 2 ungues, I with ungues unequal,

both with a tooth, II with ungues unequal, larger one simple and smaller one

with a tooth, III with ungues equal and simple. Wing. Veins covered with dark

brown scales dorsally and ventrally; alula with several narrow, dark brown

scales along margin; upper calypter with numerous dark setae; 1 remigial seta.

Halter. Pedicel pale; capitellum silvery scaled with dark brown scales at base.

Abdomen. TergaI-VII each covered with blackish-brown scales with a latero-

basal silvery scaled spot; sterna II-VII each blackish-brown scaled with a basal

silvery scaled band; terga and sterna with numerous dark setae, mostly along

posterior margins. Genitalia (Fig. 9). Tergum IX band-like with lateral caudal

areas broad and gradually tapering mesally, moderately pigmented, 3-4 stout

setae in a small patch on each side of midline, a number of minute spicules

scattered mostly on basal area; gonocoxite short, broad, heavily pigmented,

mesal surface membranous, entire surface covered with minute spicules, dor-

sal surface with numerous moderately long and long thin setae forming 4 rows

on mesal area from base to apex, lateral margin with several moderately long

to long stout setae, ventral surface with numerous short thin setae over entire

area including a large apical mesal area, lateral and ventral surfaces with

scattered broad dark scales; gonostylus long, total length, minus claw, approxi-

mately 0.70 length of gonocoxite, apical 0.46 expanded but sloping to a blunt apex

and with 7-8 short fine setae along outer margin and 3-6 similar ones on inner

margin, a patch of spicules on tergal surface near middle of outer margin of

expanded portion, gonostylar claw short, apex broad and rounded, heavily

pigmented and attached subapically; basal mesal lobe with apical portion large,

expanded, with 70-78 short, thin, evenly spaced setae and a ventromesal finger-

like projection at apex which terminates in a heavily pigmented, long, apically

expanded claspette filament, basal portion of lobe connected mesally with its

mate, small spicules covering basal portion and finger-like projection; proctiger

long, paraproct moderately pigmented with apex blunt and heavily pigmented,

cercus membranous, cercal setae absent; tergum X narrow, heavily pigmented;

phallosome with aedeagus divided into 2 lateral, moderately pigmented plates

which are connected basally and apically, each plate with lateral area curved

tergally and with a mesal, longitudinal, heavily pigmented area terminating

in 6-8 well developed, heavily pigmented, short, tergally curved teeth, paramere

moderately long, approximately 0.75 length of aedeagus, heavily pigmented,

parameral apodeme long, approximately 1.35 length of paramere, heavily pig-

mented; sternum IX large, basal, lateral and median areas moderately pigmented,

entire surface covered with minute spicules, 8-9 short to moderately long setae

near center.

16 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

FEMALE (Fig. 3). Similar to male in general habitus but with the

following differences. Head. Antenna with basal 0.85 of flagellomere 1 pale,

total length approximately 0.95 length of proboscis; maxillary palpus approxi-

mately 0.19 length of proboscis; proboscis approximately 0.95 length of femur I;

lateral surface of head with a small patch of silvery scales located next to eye

at level of the ventral margin of the antepronotum. Thorax. 3 median anterior

promontory setae; 2-4 anterior scutal fossal setae; scutellum (3-4 long and 3 -4

short setae on lateral lobe; 4-6 long and 3-4 short setae on median lobe); ante-

pronotum with 9-11 setae; postpronotum with 2-3 setae; propleuron with 2-3

setae; postspiracular area with 1 seta; prealar knob with 6-9 setae. Legs (Fig.

4), Femora I and II without basal white scales on anterior surfaces but with

white scales at base on posterior surfaces, I-III each with an apical silvery

scaled spot extending over anterior, dorsal and posterior surfaces, total length

of I approximately 1.08 length of II; tibia III with a few dorsoapical pale scales;

tarsus I with tarsomere 1 with a narrow, basal, white scaled band, tarsomeres

2 and 3 each with a small dorsobasal white scaled spot, II with tarsomeres 1-4

each with a narrow, basal white scaled band and a small, dorsoapical, white

scaled spot; III with tarsomeres 1-4 each with a moderately broad basal and

apical silvery scaled band, tarsomere 5 completely silvery scaled; tarsi I and

II of some specimens with pale scaled areas reduced; posttarsi I-III each with

2 ungues, I with ungues equal, both with a tooth, II with ungues equal, both with

a tooth, III with ungues equal and simple. Genitalia (Fig. 6). Tergum VIII with

basal 0. 80-0. 90 retracted into segment VII, wide, base and apex slightly con-

cave, several long stout setae along apical margin, moderately long thin setae and

numerous broad dark scales on apical 0.50, index 0.47-0.49; sternum VIII large,

wide, base very slightly concave, apex with a small lobe on each side of midline,

numerous setae and broad dark scales on apical 0. 80, usually a few broad pale

scales medially, index 0.65-0. 70; tergum IX wide, moderately pigmented with

a median apical indentation, 3-5 setae apically on each side of midline, index

0. 47-0. 64; insula tongue-like, long, moderately pigmented, covered with minute

spicules and with 2-4 tiny tuberculi on apical 0.20; lower vaginal lip moderately

pigmented, covered with hair-like spicules, narrow and with base broader; upper

vaginal lip narrow, moderately to heavily pigmented, covered with hair-like

spicules; upper vaginal sclerite large, moderately to heavily pigmented; post-

genital lobe moderately long, apex with a moderately deep median indentation,

11-14 setae on each side of midline, completely covered with minute hair-like

spicules, dorsal PGL index 0. 79-0. 87, ventral PGL index 1.47-1.51; cercus

short, apex bluntly rounded, mesal margin slightly concave, completely covered

with minute spicules, scales absent, dorsal surface with 4-5 long stout setae

apically and several moderately long and short setae on remainder of apical 0. 50,

ventral surface with 8-14 short setae on apical margin, index 2. 05-2. 20, cercus/

dorsal PGL index 2.58-2. 81; 3 heavily pigmented, spherical seminal capsules, 1

large and 2 slightly smaller ones; base of accessory gland duct heavily pigmented.

PUPA (Fig. 12). Chaetotaxy as figured and recorded in table 2. Res-

piratory trumpet. Moderately pigmented; a number of scattered, short setae on

inner surface of meatus; index 4.63-5.77, average 5.22. Abdomen. Seta 4-III

single to triple; 4-V double to 5 branched; 4-VI double or triple; 7-VII single;

9-VII double to 4 branched; 9-VIII with 5-9 branches. Paddle. Ovoid, with

minute spicules on apical 0. 80-0. 85 of outer and apical 0. 25-0. 29 of inner mar-

gins; midrib moderately pigmented and reaches apex; 1-P moderately long,

barbed, double to 5 branched; index 1.30-1.66, average 1.45.

LARVA (Fig. 14). Chaetotaxy as figured and recorded in table 4. Head.

Seta 4-C moderately long, 9-14 branched; 5, 6-C long, single; 7-C moderately

Reinert: Bothaella, new subgenus 17

long, 6-10 branched; 14-C moderately long, double to 4 branched; basal maxil-

lary hair double to 8 branched; mental plate with 19-22 (usually 20) teeth; mouth

brush with mesal hairs pectinate apically. Antenna. Short, moderately pig-

mented; seta 1-A short, double or triple, attached 0. 55-0. 58 from base.

Thorax. Seta 3-P with 4-8 branches; 8-P with 6-12 branches; 9-P double or

triple; 13-P with 4-12 branches; 14-P triple to 5 branched; 13-M with

8-14 branches; 14-M with 6-11 branches; 5-T double to 4 branched; 8-T with

9-8 branches; 13-T with 5-7 branches. Abdomen. Seta 1-I with 5-7 branches;

2-I triple to 5 branched; 4-I with 4-5 branches; 11-I with 5-7 branches; 13-I

with 4-9 branches; 2-II triple to 5 branched; 4-II double to 4 branched; 13-II

triple to 7 branched; 2-III triple or 4 branched; 4-III double or triple; 9-II

single; 13-III triple to 8 branched; 2-IV single to 4 branched; 4-IV double or

triple; 9-IV single; 2-V single to triple; 5-V triple; 9-V single; 13-V double to

0 branched; 2-VI single to triple; 5-VI triple; 8-VI triple or 4 branched; 9-VI

single or double; 2-VII with 5-6 branches; 1-VIII double to 5 branched; 3-VIII

double to 6 branched; 1-X long, barbed, double; 2-X long, 4-6 branched; 3-X

long, single; comb with 15-26 (usually 24) scales arranged in 2 irregular rows;

saddle incompletely rings segment X, acus absent, moderately to heavily pig-

mented with long spine-like spicules on posterior margin; ventral brush of 8-9

setae on grid, each single to triple; 4 anal papillae, long and slender. Siphon.

Heavily pigmented, acus well developed, index 2.90-3.63; pecten on basal 0.36

of siphon, composed of 12-17 (usually 14) evenly spaced teeth, each with a few

ventral and dorsal denticles; seta 1-S long, barbed, triple to 5 branched, base

attached on basal 0.43 of siphon and distal to last pecten tooth.

EGG. Not known.

TYPE-DATA. Type series includes holotype male, allotype and 7

paratypes (2 males, 5 females). Holotype male with associated pupal and

larval skins, THAILAND, Chiang Mai, Amphoe Chiang Dao, Ban Tham Kraeb,

7 September 1970, Kol Mongkolpanya collector, collection number 04761-5,

SEAMP accession number 259, genitalia preparation number T72.564, collected

as a larva from fresh colored water in a partially shaded, small, rock pool

located in a secondary deciduous forest in mountainous terrain at an altitude of

1000 meters; allotype with associated pupal and larval skins, with same data as

holotype except for the following: 26 September 1970, collection number 04845-4

SEAMP accession number 271, small rock pool on side of a cliff at an altitude of

1520 meters; paratype 1 male, THAILAND Kanchanaburi, Huai Bong Ti, 2 June

1965, Kol Mongkolpanya collector, collection number 00189-101, collected as a

pupa from fresh colored water in a heavily shaded, small rock pool on the side

of a mountain in a primary bamboo grove in mountainous terrain at an altitude of

200 meters; paratypes 3 females with associated pupal skins (collection numbers:

00225-100, "00225- 104, 00225-105), THAILAND, Kanchanaburi, Khao Na Chang,

6 June 1965, ae Peyton collector, collected as pupae from fresh clear water

ina partially shaded, small rock pool on the side of a mountain located ina

secondary rain forest in a valley at an altitude of 75 meters; paratypes 2 females,

collection number 00225, with same data as specimen 00225- 100; paratype 1

male with associated pupal and larval skins, same data as allotype except for the

following: collection number 04832-5, SEAMP accession number 241, genitalia

preparation number 71-397, collected from a small rock pool located inside

mouth of a cave. All type material is deposted in the U. S. National Museum

(Natural History), Washington, D. C., except 1 paratype female with associated

pupal skin which will be deposited in the British Museum (Natural History),

London, England, and 1 paratype female with associated pupal skin which will

be sent to the Department of Entomology, SEATO Medical Research Laboratory,

18 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

Bangkok, Thailand.

DISTRIBUTION. Material examined: 3c, 59, 20 L, 8 with associated

skins (5p, 3 1).

THAILAND. Chiang Mai, Ban Tham Kraeb, Chiang Dao; 20, 29, 4p,.

31. Kanchanaburi, Ban Sai Yok, Huai Bong Ti, Huai Mae Nam Noi, Khao Na

Chang; 1% 39, 4p, 15 L. Nan, Doi Chik Chong; 5 L.

TAXONOMIC DISCUSSION. Adults of eldridgei are very similar to those

of helenae and kleini and are discussed under the taxonomic discussion section

of helenae. | 8

Male genitalia of eldridgei are similar to the other 2 species of the sub-

genus and are discussed in the taxonomic discussion section of helenae. The

most diagnostic features of the genitalia of eldridgei are the development of the

basal mesal lobe and the aedeagal teeth.

The female genitalia are compared to the other 2 species of the subgenus

under the taxonomic discussion section of helenae.

Pupae of eldridgei and helenae can be separated by the number of branches

on setae 9-VII and 1-P as mentioned in the key. :

Larvae of eldridgei are similar to those of helenae and are discussed

under the taxonomic discussion section of that species. Setae of the latter spe-

cies tend to have more branches than those of eldridgez.

This species is dedicated to Lieutenant Colonel Bruce F. Eldridge,

Chief of the Department of Entomology, Walter Reed Army Institute of Research,

in appreciation of the counsel and support he has always given me.

BIOLOGY. The immature habitat is water in small rock pools. Im-

matures were collected from clear or occasionally colored, fresh, temporary

water (usually containing dead leaves and other dead vegetation) in small rock

pools situated on mountain sides, located in partial or heavy shade, in primary

bamboo groves, primary rainforests, and secondary deciduous and bamboo

forests. One larval collection was made from a fresh clear water rock pool

inside the mouth of a cave. All immature collections were in mountainous ter-

rain at altitudes between 75 and 1520 meters (usually near 200 meters). Larvae

were collected once in association with Aedes helenae. :

AEDES (BOTHAELLA) KLEINI, NEW SPECIES

(Figs. 4. . 7... 10)

MALE. Head. Antenna brown, plumose, pedicel with a patch of over-

lapping, broad, silvery scales on mesal and ventral surfaces, flagellomere 1

with several small dark brown scales, total length approximately 0. 85 length of

proboscis; clypeus dark brown, bare; maxillary palpus short and dark brown

scaled, 4 segmented, segments 1 and 2 short, segment 3 approximately 0,44

length of segment 4 which is long and slightly longer than basal 3 segments, a

few short dark brown setae at apex of segments 2, 3 and 4, total length approxi-

mately 0.18 length of proboscis; proboscis dark brown scaled with a few pale

scales forming an indistinct stripe on basal 0.50 of ventral surface (paratype

without pale scales), total length approximately 1.09 length of femur I; eyes

contiguous; vertex and lateral areas covered with overlapping, broad, decumbent

scales, a dorsomedian triangular patch of silvery scales anteriorly with apex

extending onto dorsal portion of interocular space and a small patch of similar

scales on postgena, remainder of scales dark brown; occiput with numerous pale

to dark, long, erect, forked scales; several dark, long ocular setae; 2 long,

dark interocular setae. Thorax. Scutal integument dark brown; scutum covered

with narrow, curved reddish-black scales with prescutellar space bare; scutellum

Reinert: Bothaella, new subgenus 19

with a patch of broad dark brown scales on lateral lobe, median lobe with a

patch of broad silvery scales and broad dark brown ones apically; following

areas with setae which are dark and well developed: 2 median anterior

promontory, numerous dorsocentral (anterior and posterior), scutal fossal

(3-4 anterior, 2-3 lateral and 1 posterior), numerous supra-alar, 5-7 posteri-

or medial scutal (pale), 1 postalar callar and scutellar (3 long and 2-3 short

lateral; 4 long and 2-3 short median); pleural integument dark brown; ante-

pronotum with a large patch of overlapping, broad, silvery scales, 7-9 long

and short dark setae; postpronotum with 1 long and 1 short to moderately long,

dark, posterior setae; propleuron with a large patch of overlapping, broad,

silvery scales, 1-2 long and 1 moderately long dark setae; prosternum, sub-

spiracular area, paratergite, mesomeron and metepisternum bare; post-

spiracular area with a large patch of overlapping, broad, silvery scales

covering most of area, 1 long dark seta; mesepisternum with a large patch of

overlapping, broad, silvery scales on upper and posterior areas, 1-2 long,

golden upper setae and 1-2 long, golden lower ones; prealar knob with 3-5

moderately long, dark setae, mesepimeron with a patch of overlapping, broad,

silvery scales on upper area, a patch of 3-5 moderately long, golden setae

posterior to scale patch on upper area and 1 long golden seta on lower median

area. Legs (Fig. 4). Coxae I-III each with several moderately long and long

brown or golden setae, I with anterior and lateral areas covered with broad

silvery scales, II and III each with an anterior patch of broad silvery scales;

trochanters I-III each with broad silvery scales; femora I and II covered with

broad dark brown scales, I and II each with a posteromedian patch of pale scales

which is on basal 0.30 of I and basal 0. 50 of II, III with a broad, longitudinal,

silvery scaled stripe on basal 0.75 of anterior surface and basal 0.65 of pos-

terior surface, stripe broad basally and tapered to a point apically, alsoa

lateroapical silvery scaled spot on anterior surface and a small spot on posterior

surface, total length of I approximately 1.09 length of II; tibiae I-III and tarsi I-

Ill each dark brown scaled; posttarsi I-III each with 2 ungues, I with ungues un-

equal, both with a tooth, II with ungues unequal, larger one simple and smaller

one with a tooth, II with ungues equal and simple. Wing. Veins covered with

dark brown scales dorsally and ventrally; alula with a row of narrow, dark

brown scales along margin; upper calypter with numerous setae; 1 remigial

seta. Halter. Pedicel pale; capitellum silvery scaled with dark brown scales

at base. Abdomen. TergalI-VII each covered with dark brown scales with a

large laterobasal silvery scaled spot; sterna II-VII each dark brown scaled

with a basal silvery scaled band; terga and sterna with numerous golden setae,

mostly along posterior margins. Genitalia (Fig. 10). Tergum IX band-like

with lateral caudal areas broad and gradually tapering mesally, moderately

pigmented, 7-9 stout setae in a more or less double longitudinal line on each

side of midline, a number of minute spicules scattered mostly on basal area;

gonocoxite short, broad, heavily pigmented, mesal surface membranous,

entire surface covered with minute spicules, dorsal surface with numerous

moderately long and long thin setae forming 3 rows on basal 0.60 of mesal

area, a few short setae at apex, lateral surface with several long stout setae

from base to apex, ventral surface with numerous short thin setae over entire

area except for a large circular apical mesal area which is covered with a

tightly packed patch of moderately pigmented, short, broadly flattened, erect,

scale-like setae, lateral and ventral surfaces with scattered broad dark scales;

gonocoxite long, total length, minus claw, approximately 0.73 length of gono-

coxite, apical 0.42 expanded with 9-11 short fine setae along outer margin and

1 similar seta on inner margin, gonostylar claw short, apex broad and rounded,

20 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

heavily pigmented and attached subapically on inner margin; basal mesal lobe

with apical portion large, expanded, with 201-244 short, thin, evenly spaced

setae, basal portion covered with small spicules and connected mesally with

its mate; proctiger long, paraproct moderately pigmented with apex blunt and

heavily pigmented, cercus membranous, cercal setae absent; tergum X narrow,

heavily pigmented; phallosome with aedeagus divided into 2 lateral, moderately

pigmented plates which are connected basally and narrowly at apex, each plate

with lateral area curved tergally and with a mesal, longitudinal, heavily pig-

mented area terminating in 4-5 short teeth (teeth visible in lateral view), para-

mere moderately long, approximately 0.72 length of aedeagus, heavily pig-

mented, parameral apodeme long, approximately 1.42 length of paramere,

heavily pigmented; sternum IX large, basal, lateral and median areas moder-

ately pigmented, entire surface covered with minute spicules, 5-6 short to

moderately long setae near center.

FEMALE (Fig. 4). Similar to male in general habitus but with the

following differences. Head. Antenna with basal 0.75 of flagellomere 1 pale,

0.77 length of proboscis; maxillary palpus 0.17 length of proboscis; proboscis

0.98 length of femurlI. Thorax. Postspiracular area with 1-2 setae; prealar

knob with 4-5 setae. Legs (Fig. 4). Femora II and III each with a dorsoapical

silvery scaled spot, large on III (spot on II represented by 1-3 white scales in

paratype), total length of I approximately 1.15 length of II; posttarsi I-III each

with 2 ungues, I with ungues equal, both with a tooth, II with ungues equal, both

with a tooth, III with ungues equal and simple. Genitalia (Fig. 7). Tergum

VIII with basal 0. 70-0. 80 retracted into segment VII, wide, base and apex

slightly concave, several long stout setae along apical margin, moderately long

thin setae and numerous broad dark scales on apical 0.55, index 0. 51-0. 56;

sternum VIII large, wide, base slightly concave, apex with a small lobe on

each side of midline, numerous setae and broad dark scales on apical 0. 70,

index 0. 66-0.67; tergum IX wide, moderately pigmented with a median apical

indentation, 1-2 setae apically on each side of midline, index 0. 51-0. 52; insula

tongue-like, long, moderately pigmented, covered with minute spicules and

with 3 tiny tuberculi on apical 0.20; lower vaginal lip moderately pigmented,

covered with hair-like spicules, narrow and with base broader; upper vaginal

lip narrow, moderately to heavily pigmented, covered with hair-like spicules;

upper vaginal sclerite large, moderately to heavily pigmented; postgenital lobe

moderately long, apex with a moderately deep median indentation, 7-8 setae

on each side of midline, completely covered with minute spicules, dorsal PGL

index 0.90-0.96, ventral PGL index 1.'74-2.04; cercus short, apex bluntly

rounded, mesal margin slightly concave, completely covered with minute

spicules, scales absent, dorsal surface with 4-5 long stout setae apically and

several moderately long and short setae on remainder of apical 0.40, ventral

surface with 7-9 short setae on apical margin, index 2.46-2.69, cercus/dorsal

PGL index 3. 56-3.64; 3 heavily pigmented, spherical seminal capsules, 1 large

and 2 slightly smaller ones; base of accessory gland duct heavily pigmented.

PUPA, LARVA and EGG. Not known.

TYPE-DATA. Type series includes holotype male, allotype, 1 paratype

male and 1 paratype female. Holotype male, CAMBODIA, Kompong Speu,

Oudong, 25 September 1970, J-M. Klein collector, collection number 808,

SEAMP accession number 336, genitalia preparation number T73.109, adult

collected at rest in a small dry forest around the well known temples containing

the king tombs of Oudong; allotype with same data as holotype except the follow-

ing: date 18 September 1970 and genitalia preparation number T73.69; para-

type male with same data as allotype except the following: collection number 809,

Reinert: Bothaella, new subgenus 21

SEAMP accession number 301, genitalia preparation number T72. 642; and

paratype female, CAMBODIA, Kompong Som, Sihanoukville, 31 March 1970,

J-M. Klein collector, SEAMP accession number 270, genitalia preparation

number T72.635, taken biting man between 1800-1900 hours in a littoral

rainforest. The holotype and allotype are deposted in the U. S. National

Museum (Natural History), Washington, D. C., and the 2 paratypes are de-

posited in the J-M. Klein Collection.

DISTRIBUTION. Material examined: holotype o% allotype 2 and 1c‘and

12 paratypes.

CAMBODIA. Kompong Som, Sihanoukville; 12. Kompong Speu, Oudong;

20;) 19.

TAXONOMIC DISCUSSION. Adults of kleini are very similar to both

helenae and eldridgei but are easily distinguished from the latter 2 species by

the dark scaled tarsi, coxa I white scaled and mesepisternum with a single

large white scaled patch.

The male genitalia of Rlezni are similar to both eldridgei and helenae

but with the closest morphological resemblance to the latter species. Aedes

kleini genitalia are easily separated from the other 2 species of the subgenus

by the number of setae on the basal mesal lobe and by the other features men-

tioned in the taxonomic discussion section of helenae.

Female genitalia are similar to helenae and eldridgei and are compared

to these 2 species in the taxonomic discussion section of helenae.

This species is dedicated to Dr. J-M. Klein, Office de la Recherche

Scientifique et Technique Outre-Mer (ORSTOM), Bondy, France, in recogni-

tion of his contributions to the study of Cambodian mosquitoes.

BIOLOGY. Adults were taken at rest in a small dry forest. One female

was collected biting man in a littoral rainforest between 1800-1900 hours.

Nothing is known about the immature habitats.

ACKNOWLEDGEMENTS

Grateful appreciation is expressed to: Dr. Botha de Meillon, Principal

Investigator, Southeast Asia Mosquito Project (SEAMP), and Lieutenant Colonel

Bruce F. Eldridge, Chief of the Department of Entomology, Walter Reed Army

Institute of Research, for critically reading the manuscript; to Dr. Douglas

J. Gould, Chief of the Department of Medical Entomology, Southeast Asia

Treaty Organization Medical Research Laboratory, Bangkok, Thailand, and

his personnel and Dr. J-M. Klein, Office de la Recherche Scientifique et

Technique Outre-Mer, Bondy, France, for the collection of the specimens; to

Mr. Young T. Sohn and Mrs. Shuling Tung, SEAMP, and K. Miyasaka, U. S.

Army 406th Medical Laboratory, Tokyo, Japan, for preparing the illustrations;

to Mr. Thomas V. Gaffigan, SEAMP, for technical assistance; to Dr. J. N.

Couch, Department of Biology, University of North Carolina, Chapel Hill,

North Carolina, for identifying the Coelomomyces specimens; to my wife,

Mollie, for typing the drafts; and to Mrs. Janet Rupp for typing the manuscript

for offset reproduction.

22 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

LITERATURE CITED

BELKIN, J. N.

1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Univ.

Calif. Press, Berkeley, 2 vols. 608 and 412 p.

BERLIN, O. G. W.

1969. Mosquito studies (Diptera, Culicidae) XII. A revision of the

neotropical subgenus Howardina of Aedes. Contr. Am. ent.

Inst. 4(2): 1-190.

JONES, J. C. and R: E. WHEELER.

1965. An analytical study of coitus in Aedes aegypti (Linnaeus).

J. Morph. 117(3): 401-423.

KNIGHT, K. L.

1970. A mosquito taxonomic glossary I. Adult head (external). Mosq.

Syst. Newsletter 2(1): 23-33.

KNIGHT, K. L. and J. L. LAFFOON.

1970a. A mosquito taxonomic glossary III. Adult thorax. Mosq. Syst.

Newsletter 2(3): 132-146.

KNIGHT, K. L. and J. L. LAFFOON.

1970b. A mosquito taxonomic glossary IV. Adult thoracic appendages.

Mosq. Syst. Newsletter 2(4): 165-177.

KNIGHT, K. L. and J. L. LAFFOON.

1971. A mosquito taxonomic glossary V. Abdomen (except female

genitalia). Mosq. Syst. Newsletter 3(1): 8-24.

LAFFOON, J. L. andK. L. KNIGHT.

1971. A mosquito taxonomic glossary VI. Female genitalia. Mosq.

Syst. Newsletter 3(2): 32-41.

REINERT, J. F.

1973a. Notes on the genus Heizmannia and the inclusion in subgenus

Mattinglyia of two species previously placed in Aedes subgenus

uncertain (Diptera: Culicidae). Mosq. Syst. 5(1): 14-23.

REINERT, J. F.

1973b. Aedes wainwrighti Baisas, a synonym of Aedes Stegomyia)

meronephada (Dyar and Shannon), with notes on the subgenus

Stegomyia Theobald (Diptera: Culicidae). Mosq. Syst. 5(1): 27-30.

REINERT, J. F.

1973c. Contributions to the mosquito fauna of Southeast Asia. -XVI.

Genus Aedes Meigen, subgenus Aedimorphus Theobald in South-

east Asia. Contr. Am. ent. Inst. 9(5): 1-218.

Reinert: Bothaella, new subgenus 23

APPENDIX: TABLE 1. Record of the branching of the

setae on the pupae of Aedes (Bothaella) helenae

‘Hair Range Mode Mean = ~—~—~—~—'iHair'~=Range Mode Mean

Cephalothorax(C) Abdomen II

1 1-2 1 14 0 1 1 1

2 1 1 1 if 1-3 2 2.1

3 1 1 1 2 1-2 1 1,3

4 2-6 a 4.2 3 1 1 1

5 2-4 3 3.6 a 2-7 4 4.8

6 1 1 1 5 1-2 1 1.2

f 1-3 2 1.9 6 1 1 1

8 1-2 1 | ae | 7 2-4 2 2.6

9 1-2 1 ee 9 if 1 1

Metanotum(C) 10 1-3 2 22

10 1-5 2 2.5 Abdomen III

11 1-2 1 1.4 0 i 1 1

12 1-4 3 2.4 1 1-2 i} 1.4

Abdomen I 2 1-3 1 i ey

1 11-34 21 17.3 3 i! 1 1

Z 1-3 1 1.6 4 3-6 3 3.8

3 1 1 1 5 1-2 1 1

4 3-8 6 5.4 6 1-2 1 2

5 2-5 3 3.2 7 2-6 5 4.5

6 1 1 1 8 2-4 3 2.9

7 1-3 2 2 9 1 1 1

9 1 1 1 10 1-2 2 1.6

10 1-2 2 1.8 11 1 1 1

14 1 1 1

24 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

TABLE 1. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen IV Abdomen V (Cont. )

0 1 1 1 10 1-2 1 a

1 1-3 1 1,4 11 1 1 1

2 1-2 1 1.2 14 1 1 1

3 3-5 3 3.9 Abdomen VI

4 2-6 3 3.4 0 1 1 1

. 1 1 1 1 1-2 1 1.1

6 1 1 1 2 1-2 1 1,2

7 2-5 4 3.5 3 1-2 1 1.1

8 2-4 3 2.6 4 2-5 2 3

9 1 1 1 5 f 1 1

10 1-4 2 2.1 6 1-2 1 Lee

11 1 1 1 7 2-5 3 2.9

14 1 1 1 8 2-5 2 3.2

Abdomen V 2, 1 1 1

0 1 1 1 10 1 1 1

1 1 1 1 11 1 1 1

2 1-2 1 a 14 1 1 1

3 1-3 1 1,4 Abdomen VIL

4 3-7 4 4.8 0 1 1 1

5 1 1 1 1 1 1 1

6 1 1 1 2 1-2 : |

7 4-8 6 6.1 3 2-4 2 2.3

8 2-5 2 2.6 4, 1-3 1 1.4

9 1 1 1 5 1 1 1

Reinert: Bothaella, new subgenus 25

TABLE 1. (Continued)

‘Hair Range Mode Mean ~—‘*iHair Range Mode Mean

6 2-5 4 3.6 Abdomen VIII

y 1-3 1 1.2 0 1-2 1 ts

8 2-5 he ee 4 1 1 1

9 5-10 a nee 9 6-12 9 9.1

10 142 1 1.4 14 {<2 1 1.4

11 1-2 1 1.3 Paddle(P)

14 1 1 1 1 6a14°. > 13 ita

APPENDIX: TABLE 2. Record of the Branching of the

setae on the pupae of Aedes (Bothaella) eldridgei

Hair Range Mode Mean Hair Range Mode Mean

Cephalothorax(C) Abdomen I (Cont. )

1 1-3 1 1. 2 1 1 1

2 1-2 1 Oe 3 1 1 1

3 1 1 1 4 3-6 4 4

4 1-5 3 2.9 5) 1-4 2 1.9

5 2-3 2 “ae 6 1 1 1

6 1 1 1 " 1-2 1 1.3

7 1-3 1 1.4 9 1-2 1 1.1

8 1-3 1 1.3 10 1-3 3 2.1

9 1-2 1 i Abdomen II

Metanotum(C) 0 1 1 1

10 1-3 2 1.9 1 1-6 2 2.7

11 1 1 1 2 1 1 4

12 1-3 2 1.8 3 1 1 1

Abdomen I 4 3-6 4 4.1

1 14-27 15 18.4 ) 1-2 1 ace

TABLE 2. (Continued)

Hair

Range

Abdomen II (Cont. )

1 1

1-3 2

1 1

1-2 1

Abdomen III

1 1

1-3 1

1 1

1-2 1

1-3 2

1-2 1

1-7 2

1-3 2

1 1

1-2 1

1 1

Abdomen IV

1 1

1-2 1

2-6 3

Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

Mode Mean

1.8

1.3

Hair

Range

Abdomen IV (Cont. )

Mode

1-4 2

ms 1

1-2 1

1-5 2

1 1

1-2 1

1 1

r 1

Abdomen V

1 1

1-2 1

2-5 4

1 1

2-7 &

2-5 &

1 1

1-2 1

1 1

Mean

2.2

3.6

oat

Reinert: Bothaella, new subgenus at

TABLE 2. (Continued)

‘Hair Range Mode Mean ~—~—~—*CHair+=Range Mode Mean

Abdomen VI Abdomen VII (Cont. )

0 1 1 1 3 1-3 1 Leo

1 1 1 1 4 1 1 1

2 1 1 1 5 1 1 1

3 1 1 1 6 2-5 2 2.9

4 2-3 2 2.2 f 1 1 1

5 1 1 1 8 2-5 3 wad

6 1 1 1 9 2-4 3 a.9

7 1-3 2 2.2 10 1 1 1

8 3-6 4 3.9 11 1-2 1 r3

9 1 1 1 14 1 1 1

10 1-2 1 : Abdomen VIII

11 1 1 1 0 1-2 1 ee

14 1 1 1 4 1-2 1 Lee

Abdomen VII 2) 9-9 7 Get

0 1 1 1 14 1 1 1

1 gee 1 1 | Paddle(P)

2 1 1 1 1 2-5 3 3.8

28 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

APPENDIX: TABLE 3. Record of the branching of the

setae on the larvae of Aedes (Bothaella) helenae

Hair Range Mode Mean Hair Range Mode Mean

| Antenna(A) . Prothorax(P)(Cont. )

1 2 2 2 4 1 1 1

Head(C) 5) 1 1 1

0 1 1 1 6 1 1 1

1 1 1 1 7 2-3 2 2.2

3 1 1 1 8 12-18 14 14,3

4 4-8 4 4.9 2 4-7 6 5. 6

3) 1 | 1 10 1 1 1

6 1 1 i 11 2-3 2 |

7 7-10 9 8.6 12 1 1 1

8 1 1 1 ious geet a GLB

9 2-3 2 2.3 14 4-6 5) 4.8

10 1 1 1 Mesothorax(M)

11 5-8 7 6.8 1 5-7 6 5.8

12 1-2 2 1,8 2 2-4 3 2.8

13 1 1 1 3 1 1 1

14 5-8 6 6.3 4 1 1 i

15 2-4 3 3 9) 1 1 1

bmh 9-15 12 12.3 6 3-5 3 3.9

Prothorax(P) 7 1 1 1

0 8-14 9 1 ae 8 4-6 4 4.9

1 4-5 5 4.7 9 4-6 5) 4.8

2 1 1 1 10 1 1 1

Reinert: Bothaella, new subgenus 29

TABLE 3. (Continued)

‘Hair Range Mode Mean = Hair Range Mode Mean

Mesothorax(M) (Cont. ) Abdomen I (Cont. )

12 1 1 1 ‘} 2 2 2

13 13-19 14 15 9 2-3 3 ya

14 10-16 10 12.3 10 1 1 1

Metathorax(T) 11 10-16 11 12.4

1 5-12 6 V2 13 8-15 13 11.6

2 1 1 1 Abdomen II

3 4-5 4 4.1 0 1 1 1

“ 2-3 2 2.1 1 6-9 9 7.6

5 6-9 7 Ye 2 7-14 11 10.6

6 1 1 1 3 1 1 1

7 5-6 5) Fe A 4 4-6 5) 5. 2

8 10-14 11 11,5 D 3-6 6 o.3

9 3-4 3 3.4 6 3 3 3

10 1 1 1 7 2 2 2

11 1 1 1 8 1-4 2 1.8

12 1 1 1 9 3-7 4 4.6

13 10-16 12 13.1 10 1-2 1 PI

Abdomen I 11 3-5 4 3.8

1 7-10 8 8.2 12 1-2 1 1,2

2 7-10 9 O48 13 8-16 9 Lt..7

3 1 1 1 14 1 1 1

‘ 5-9 5 6 Abdomen III

5 4-8 6 5.8 0 1 1 1

Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

TABLE 3. (Continued)

Hair

Range Mode Mean

Abdomen III (Cont. )

9-11 10

1 1

5-8 5

4-7 5

2-3 2

4-8 5

1-2 2

3-6 4

1 1

2-5 3

1-3 2

7-11 10

1-2 1

Abdomen IV

1 1

5-9 5

8-11 9

1 1

4-9 6

4-5 4

2 2

5-10 5

1 1

3-5 3

10.1

1.2

6.4

3.0

Hair Range Mode Mean

Abdomen IV (Cont. )

10 1 1 1

11 3-5 4. 2.9

12 2 2 2

13 5-9 7 6.9

14 1-2 cf 2

Abdomen V

0 1 1 1

1 4-6 A 4.7

2 8-11 9 9.3

3 A i if

4 3-7 5 4.9

5) 4-5 4 4.2

6 2 2 2

7 4-8 if 6

8 1-2 1 1.2

a 3-5 3 3.9

10 1 1 1

11 3-4 4, 320

12 1 1 1

13 4-8 4 3D. 9

14 1 1 1

Abdomen VI

0 1 1 1

1 3-6 4 4.3

Reinert: Bothaella, new subgenus 31

TABLE 3. (Continued)

‘Hair Range Mode Mean = ~—~—~—~—*Hair ~=Range Mode Mean

Abdomen VI (Cont. ) Abdomen VII (Cont. )

2 6-9 8 ted 8 5-14 9 8.6

3 1 1 1 9 3-4 4 3.8

4 1 1 1 10 1-4 1 a

5 4-5 4 4.3 11 1-3 2 2

6 2 2 2 12 1 1 1

7 2-4 2 2.7 13 4-8 6 6.4

8 4-8 D 5. 4 14 1-2 1 2

9 3-6 3 3.8 Abdomen VIII

10 1 1 1 0 1 1 1

Lt 2-5 4 rE 1 8-11 8 9

12 1 1 1 2 1 1 1

13 4-13 11 OA. 3 0-8 6 6.2

14 1-2 | 1,2 4 1 1 1

Abdomen VII o 3-7 D D

0 1 1 A 14 1 1 :

a 3-5 3 3.8 Abdomen X

2 7-10 8 8.3 1 2-4 2 2.4

3 4-5 4 4.3 2 4-6 5 5.3

& 1 1 1 3 1 1 1

5 4.5 4 4.6 Siphon(S)

6 4-7 3) a | 1 2-3 3 2.6

7 1 1 1 2 1 1 :

32 Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

TABLE 4. Record of the branching of the setae on the

larvae of Aedes (Bothaella) eldridgei

‘Hair Range Mode Mean ~—‘*Hair'+=Range Mode Mean

Antenna(A) 3 Prothorax(P)(Cont. )

1 2-3 2 ot 3 4-8 8 6.8

Head(C) 4 1 1 1

0 1 1 1 5 1 1 1

1 1 1 1 6 1 1 1

3 1 1 1 7 2-3 2 2.3

4 9-14 13 11.3 8 6-12 8 8.5

5 1 1 1 9 2-3 2 ya

6 1 1 1 10 1 1 1

7 6-10 7 8 ji 2-3 2 pa |

8 1-2 1 1.3 12 1 1 1

9 1-3 2 2.1 13 4-12 7 Tk

10 1-2 2 1.6 i 3-5 5 4.4

11 4-8 5 5.6 Mesothorax(M)

12 2 2 2 1 6-9 9 7.1

13 1 1 1 2 2-3 2 2.1

14 2-4 3 2.6 3 1 1 1

15 2-3 3 2.6 4 + 1 1

bmh 2-8 4 4.5 5 1 1 1

Prothorax(P) 6 3-6 4 3.9

0 6-12 9 8.8 7 1 1 1

1 3-9 5 ae 8 4-6 5 5

2 1 1 1 9 2-5 3 3.2

10 1 1 1

Reinert: Bothaella, new subgenus 33

TABLE 4. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Mesothorax(M)(Cont. ) Abdomen I (Cont. )

te. 1 1 1 6 3-6 5 4,8

12 1 1 1 7 1-3 2 £3

13 8-14 9 10.3 9 1-3 2 1.9

14 6-11 6 8.5 10 1 1 1

Metathorax(T) 11 5-7 5 5.1

1 4-8 6 6.6 13 4-9 4 5.3

2 1 1 1 Abdomen II

3 2-4 4 3.6 0 1 1 1

4 1-2 1 1.2 1 4-8 5 5. 4

5 2-4 3 i 2 3-5 4 3:9

6 1 1 1 3 1 1 1

7 6-8 7 tA 4 2-4 “ 3.4

8 5-8 5) 6.1 5) 3-4 3 3.3

9 2-4 3 3 6 3-4 3 a5

10 1 1 1 7 2 2 2

11 Uk 1 1 8 1-2 2 1.6

12 1 1 1 9 2-5 2 2.9

13 5-7 6 0.8 10 1 1 1

Abdomen I 11 2-3 2 2.4

1 5-7 7 5. 8 12 1 1 1

2 3-5 4 3.7 13 3-7 6 D. 6

3 1 1 1 14 1 1 1

‘ 4-5 4 4.3 Abdomen III

4) 3-7 4 4.5 0 1 1 1

Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

TABLE 4. (Continued)

Hair Range Mode Mean

Abdomen III (Cont. )

1 4-7 4 D.2

2 3-4 é 3.6

3 1 1 1

4 2-3 3 2.6

5 3-7 3 4,

6 2 2 2

ql 3-9 7 6

8 1-2 1 1.3

9 1 rE i:

10 1-2 1 1.2

11 2-3 3 2.6

12 1 1 1

13 3-8 5 4.5

14 1 1 1

Abdomen IV

0 1 1 1

1 3-6 5 4.7

2 1-4 2 PACS |

3 1 1 1

4 2-3 2 2.4

5 2-5 3 3.6

6 2 2 2

7 4-8 5 ot

8 1 1 1

Hair Range Mode Mean

Abdomen IV (Cont. )

2 1 1 1

10 1 1 1

11 2-3 2 221

12 2-3 2 2.3

13 3-7 5 4.4

14 1 1 1

Abdomen V

0 1 1 1

1 3-7 5 4.3

2 1-3 2 2.2

3 1 1 1

4, 3-4 3 3.4

5 3 3 3

6 2 2 2

7 3-7 5 5.4

8 1-2 1 el!

g 1 1 1

10 1 1 1

11 2-3 2 2.4

ie 2d 1 1

13 2-5 4, 3.8

14 1 1 1

Abdomen VI

0 1 1 1

Reinert: Bothaella, new subgenus 35

TABLE 4. (Continued)

‘Hair Range Mode Mean ~—~—*Hair'~=Range Mode Mean

Abdomen VI (Cont. ) Abdomen VII (Cont. )

1 3-5 3 8 7 1-2 1 1,3

2 1-3 2 1.8 8 0-6 3) D.3

3 1 1 1 9.28 O58 2 2.2

4 1 1 1 10 1 1 is

3) 3 3 3 Le 1-3 2 2.2

6 2 2 2 12 1-4 4 2.8

7 2-3 3 Ze eS: 4-9 5) 5.3

8 3-4 4 3.6 14 1 1 1

2 1-2 1 4 Abdomen VIII

10 L 1 i 0 1 1 1

11 3-4 3 3.2 1 2-5 3 3.2

12 1 1 1 2 1 1 1

13 4.9 4 6 3 2-6 4 3.9

14 1 1 1 4 1 1 1

Abdomen VII » 2-5 2 3.1

0 1 1 1 14 1 1 1

ie 3-5 3 3.6 Abdomen X |

2 5-6 6 D. 6 1 2 2 2

3 1-2 1: 1,2 2 4-6 4 4.6

4 1-2 1 Pe: 3 1 1 1

5 2-4 4 | Siphon(S)

6 4-6 6 9) 1 3-5 3 3.3

2 1 1 13

Contrib. Amer. Ent. Inst., vol. 10, no. 3, 1973

LIST OF FIGURES

Distribution of the species of Bothaella

. Aedes (Bothaella) helenae - adult morphology

Aedes (Bothaella) eldridgei - adult morphology

Aedes (Bothaella) kleini - adult morphology; helenae,

eldridgei and kleini - male and female tarsomeres 5

and posttarsi

Aedes (Bothaella) helenae - female genitalia

Aedes (Bothaella) eldridgei - female genitalia

Aedes (Bothaella) kleini - female genitalia

Aedes (Bothaella) helenae - male genitalia

Aedes (Bothaella) eldridgei - male genitalia

Aedes (Bothaella) kleini - male genitalia

Aedes (Bothaella) helenae - pupa

Aedes (Bothaella) eldridgei - pupa

Aedes (Bothaella) helenae - larva

Aedes (Bothaella) eldridgei - larva

BURMA

100

THAILAND

103

106

oocne”

CAMBODIA

*% = kleini

@ = helenae

B= e/dridge/

106

109

ig.

—_——

lpm traieeeereee te

Titers

e/ldridgei

kleini

helenae o&

kleini of

SLs ORR A helenae 2

kleini @

NS, mre

(

Mi,

, 4 /

GAN INTIS iM |

ARYL Mi, oe dae

MA ¥ WE ae |

{

bla:

See

helenae

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Brush Hairs

helenae

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Reinert: Bothaella, new subgenus a1

INDEX

Italicized pages are those which begin the primary treatment of the

taxon. Numbers in parenthesis refer to the figures illustrating the species in

question.

achaetae

Aedes

Aedes, Aedes

albolineatus group

Bothaella

Cancraedes

catesi

Christophersiomyia

Coelomomyces

communis

complex

cozumelensis

demeilloni

eldridgei

Finlaya

Geoskusea

Heizmannia

helenae

hoguei

Howardina

Huaedes

kleini

Leptosomatomyia

macdonaldi

Neomacleaya

Nothoskusea

Paraedes

reidi

Rhinoskusea

robinsoni

sexlineatus section

Stegomyia

stegomyiae

tulagiensis

upolensis

Verrallina

walkeri section

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’

Contributions

of the

American Entomological Institute

Volume 10, Number 4, 1974

eye

A Taxonomic Revision of the Termitophilous

Tribe Termitohospitini. Part III. The Subtribe Termitusina

With Notes on their Behavior and the Ultrastructure of Certain Setae

(Coleoptera: Staphylinidae)

David H. Kistner

ii Contr. Amer. Ent. Inst., vol.10, no.4, 1974

TABLE OF CONTENTS

Page

PS Sg o 0 Sie ee ee are Annes Oo ree ee ee 1

Pie EE 62.0) Gi 0S i Meee ares ont ure ne sin ere Ceara oe rer ees re 2

KEY TO THE SUBTRIBES OF TERMITOHOSPITINI ...........cceceees 2,

CORN CES 15 SURRY Sepa aE ee ae ore” Fr vere ear re Ge eer ee 2

ACKNOWLEDGMENTS 2 os oe R St a es RPS eR 3

KEY TO GENERA OF THE SUBTRIBE TERMITUSINA,...........0000- 3

eR a oh he ee erg ne er ee re ere eee ee 4

KEY PTOSBPECIES OF TERIMITUSA cee sb ees Sak ta beads es cee eae es 7

VermHUse SUGIICUS 2 oa ak ce ke wa OR eee Roo oe rk VO es ee ee ee wes 9

i a sc es eh i RS SRE COW ae ee eee ees 10

Ty BOMIEORY 8 ons ee bes oe ke AREAS Se ne ei ce ee eas 11

Os oe ve eee eee eee VAY ewok es Wa eee ek 2a oe 13

A ee a ae oe Oak OT Se ee ee ee WS wa Vis eee Se aS 13

NCS) Be er ee or 16

OT WEEMS a ok boss aoe be sls eae ade oe eee bee eae ee eee ew Oe © 16

T. GYDASL 22 eee ccc er ere c rcs c es cer creer enesesesesenecersseress 17

Ri A ee ee ee 18

Fo HORA SOC0Se yg a he ee cee betes hea eee eee neon sen nbc seas 18

PERN US Re OE ee ee ee er er re 21

pe er ee ee ae ee 22

oS AG ee EA er rn re ere 2.

CO NS ee Oe ee Cee 23

SC ee Oe ree heer ear 23

T. MalawienSisS ..... ccc ee cece ere ce rere ce reser eerceseseeeees 25

Dy TRTIOE ee a sod a ws Be eed be eee hee Ooo ce eee mene ee sees 29

T. Mixaseta ..... cee ce cece ecco c ces err crererercrcecscesenscees 26

ee er Me rr ee ee ee cee 26

Po MUNCTIS 6 ce cc ec cc eee ecm weer eenn eed eeserescsreeresee 27

fist eee ee ee ee 29

Oe NS ee ee kee cc ee ne cee eke Pears hans ee ase ee Re en 29

Te CA op ee coe be ce 4 ek be eager te et ese en 30

T. Puelle) . 2... ecw ce cee rece weet erence ences cern ereerenerones 30

Tie BUCRS OVE ce cei bce be thE ey 6 eee Oe a Oke a ee ee ee ea eee wees 34

Di SUOOSIOORD oC aAle bone oo oe PARES ae eb Deka we NS hs Ree ee eee ee eo 34

To Re PS hs he we cee eens ee he scares nals swe © 35

T. UgANdae ... cc crec creer sre cererecer eres rcrsccscercececeses 36

T. VICQASI 0... cee ees c scene ener er erence reccssecersesecceees 36

T. WASMANNE . 2... cc cece sence ewes ecw en sone nsesesesecnerences 37

De WHATS) 3.6 sc Sa bee see Nema ee ee eae RS Tee <P errr , 39

Kistner: A revision of the Subtribe Termitusina iii

Gens TROLS COIIER ek i es oe eB ee ee le eee ewes 39

KEY TO SPECIES DF THORAC TUGA ck ie ce bees we awed eee we 41

ThOracorusa AIPA CAe sek oes ak we we Oe ee oe ee a Ee ee een 41

ds; CAYPISGOCNSIS onus ec cans Ws ten eae oe RRR GA Re He ae Rene 9 43

Te KyMrae .... cee eee e cece cree cece ect ce seer ec cesses ceecessees 44

Genus Termiusodes J ov vceeals wd oh ee ee ee ee ne ee ee 45

KEY TO SPECIES OF TERME Counts 2.0 ice css hae Vad beeen een ws 48

Termitusodes COongolrewels «sie. o'e oo wo cae ea ae eee ee eee as 48

e¢ TAUPO a ca ba ee oa ee eR a ed eee ee ee ee 51

T. Leleupi 2... ccc cece cece eee e cece e cc reer e nee rcreceeseserens ae

ETA Se BI I. a eV a eK ee aw bn ww ee ee eee 54

ULTRASTRUCTURE OF SETAE. 262553 deta & iia ws ce oe See eee eae 58

RI Ee er et Le i ghee ew a ee eS AOE ak ee i a 59

Hats

Wh ae.

A Taxonomic Revision of the Termitophilous

Tribe Termitohospitini. Part II. The Subtribe Termitusina

With Notes on their Behavior and

the Ultrastructure of Certain Setae

(Coleoptera: Staphylinidae) 1

David H. Kistner

ABSTRACT

The entire subtribe Termitusina has been revised. One new genus,

Thoracotusa (type species, T. alzadae) is described while the other 2 genera,

Termitusa and Termitosodes are redescribed. Keys are provided to distin-

quish all the subtribes of the Termitohospitini. A key to the genera of Ter-

mitusina is provided. The following new species are described: Termitusa

atlanticus (Rhodesia), T. bandai (Malawi), T. boultoni (Rhodesia), T. cloveri

(Zambia), T. coatoni (Zambia), T. delignei (Gabon), T. dybasi (Cameroun),

T. hexasetosa (Zaire), T. katangensis (Zaire), T. leleupi (Zaire), T. light-

ningae (Guinea), T. malawiensis (Malawi), T. minor (Zaire), T. mixaseta

(Zaire), T. muneris (Zambia), T. pasteelsi (Gabon), T. regina (Zaire), T.

ruellei (South Africa), T. sheasbyi (South Africa), T. superhirsutus (Zambia),

T. ugandae (Uganda), T. viegasi (Angola), T. wasmanni (Rhodesia), T. wil-

liamsi (Zambia), Thoracotusa alzadae (Zambia), T. carrissoensis (Angola),

T. kymrae (Zaire), Termitusodes congolensis (Zaire), T. leleupi (Zaire).

Most of the old species were redescribed and illustrated. Keys to all species

of all genera are provided.

The behavior of the species and the relationship of the glands to their

behavior are reviewed. The interpretation is offered that the Termitusina

are adapted to the colony and to the lives of the termites but they are not in-

tegrated into the social life of the termites. The ultrastructure of certain

setae and microtrichia are presented.

I this study was financed in part by the National Science Foundation

(Grant GB-42727X). It was completed during a sabbatical leave awarded by

California State University, Chico.

2Address: Shinner Institute for the Study of Interrelated Insects, Dept.

of Biology, California State University, Chico, CA 95926. :

2 Contr. Amer, Ent. Inst., vol.10, no.4, 1974

INTRODUCTION

The subtribe Termitusina was first defined by Seevers (1957: 192) as

a subtribe of the tribe Termitohospitini, It shares with the rest of the Termi-

tohospitini the 4-4-5 tarsal formula; the labial palps with the first 2 segments

long and slender and the third segment reduced in length; the distinct men-

tum; the slender ligula, and the large and margined mesocoxal acetabulae.

The subtribes are most easily distinquished by the following key.

KEY TO THE SUBTRIBES OF TERMITOHOSPITINI

1, Body form fimuloid or Bearly milo. . 6 6 6 ee ee ee ee

Body form not limuloid, somewhat physogastric; prognathous; pro-

notum collar-like in shape. .-. « © « « © © © © © © © © oe Lermitospectrina

(including Termitoecia Bernhauer, revised by Kistner (1970b)).

2. Body form not strongly limuloid; head easily visible from the dorsal

surface, not covered by the pronotum; meso- and metasternum with

a very large intercoxal ridge separating the mesocoxae. .........

. Hetairotermitina (containing the genus Hetairotermes Cameron,

revised by Kistner (1970a)).

Body form strongly limuloid, meso- and metasternum without inter-

i a a ie gos Sede a ee ee Ue tS

3. Pronotum covers entire head; mesosternum length equal to that of

the metasternum; antennae with sides of segments expanded to cover the

Oeticles. oo eB ee Termitohospitina (only known from the New World)

Pronotum not covering entire head; metasternum much longer than

the mesosternum; antennae with distinct petioles not covered by exten-

sions of the sides of the segments. .Termitusina (only known from Africa)

METHODS

It is nearly useless to look at dried specimens of this subtribe. We

keep all specimens in 80% alcohol after they have been fixed in FAA. These

can be sorted to species in fluid. Then to determine the species, 2 males

and 2 females from each series of each species are placed intact into cold

KOH. There are no obvious secondary sex characters. Males tend to have

a slightly thicker abdomen than females. Sometimes in the callow species,

they can be sexed by shining strong light through the abdomen. These are

made into slides separating out meso- and metasternum, pronotum, elytra,

male genitalia or spermathecae, and abdominal segments II-IX. The species

can be determined from these. All other methods have been given by

Kistner (1968). All measurements are in mm unless otherwise stated.

For simplicity, certain conventions were adopted. When counting the

setae on the pronotum, I have started from the midline and designated the

number of setae in each row out to the outer edge. On the elytra the num-

bers are given in the rows from the suture to the outside edge. For conven-

ience, the rows have been marked on the photos of the pronota and elytra.

When slides are made, broken setae can be identified by the empty sockets.

Kistner: A revision of the Subtribe Termitusina 3

For the readers' convenience, these setae have been redrawn into their loca-

tions on half of the pronotum and on the elytra.

Most of this paper was written prior to a field trip to Africa in 1972-73

and all of the figures had been constructed into plates. However, the paper

could not be completed. During the field trip, 5 new species were collected

as well as numerous additional specimens of species described herein. It

then became necessary to rewrite about 50% of this paper. However, the

plates could not be conveniently changed. Thus the figures of new species

and new sexes, etc. are all collected together on separate plates in what ap-

pears to be an incomprehensible sequence without this explanation.

ACKNOWLEDGMENTS

For help in the field, Iam extremely grateful to Mr. Rudyerd Boulton,

Director of the Atlantica Ecological Research Station, Salisbury, Rhodesia;

Dr. W. G. H. Coaton, Plant Protection Research Institute, Pretoria, South

Africa; and Mr. Karl Kiihne, Director, International Red Locust Control

service, Mbala, Zambia. Without their enthusiastic help, my recent field

efforts could not have been so productive.

For help during the 1972-73 field trip, Iam extremely grateful to Dr.

A. de Barros Machado, Laboratorio de Biologia, Companhia de Diamantes de

Angola, Dundo (DIAMANG) for numerous helpful acts while in the field, Iam

also extremely grateful to Mr. Jorge Viegas, Director of DIAMANG and the

whole DIAMANG organization for making our stay in Angola so pleasant

through their logistic aid and unbounded hospitality.

Iam grateful to M. P. Basilewsky, Musée Royal de l'Afrique Centrale,

Tervuren, (M. R. A. C.) for the loan of specimens collected by Dr. N.

Leleup in the Congo. I am also grateful to be able to study specimens pro-

vided by Dr. J. M. Pasteels, Université Libre de Bruxelles, Belgium, (J.

M. P.) and by Mr. Henry Dybas, Field Museum of Natural History, Chicago,

(F. M. N. H.), and Coaton (N. C. I.). The initials given after the above

institutions are used in the text to designate where specimens are deposited.

Specimens in the collection of the author are designated (D. K.). Iam

further grateful to Dr. W. G. H. Coaton for permission to publish the excel-

lent drawing by Mr. Virgil Branco, now at the South African Museum, Cape-

town.

For host identifications, Iam deeply grateful to Mr. R. M. C. Williams

and Mr. W. A. Sands of the Termite Research Unit, British Museum

(Natural History), London and to Dr. Kumar Krishna, City University of New

York. Their identifications are individually credited to them in the text.

This paper could not have been completed so quickly were it not for the

efforts of the following Shinner Assistants of California State University,

Chico: Daniela Davison, Jan Fischer, Carolyn Gooch, Herbert Jacobson,

Nancy Piastuch, John Shower, and Riley Swift.

KEY TO GENERA OF THE SUBTRIBE TERMITUSINA

1. Mesocoxal acetabulae proceed only about half way to the posterior

4 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

edge of the metasternum, (Fig. 136)........ . -Thoracotusa (p. 39 )

Mesocoxal acetabulae proceed all the way to the posterior edge

Of the metas eri (Fie. BS we eee ke eg cg ee a re

2. Maxillary palp with the apical segment Legnaanal in position (Fig. 7);

Porm encore, (aie Oe Se ee we Termitusa (p. 4 )

Maxillary palp with the apical seoment attached to the side of seg-

ment 3, (Fig. 146); labrum long, (Fig. 153)..... Termitusodes (p. 45)

Genus Termitusa Wasmann

Termitusa Wasmann 1905: 199; 1911: 162; Fenyes 1918: 106.

Glaphyrinus Bernhauer 1941: 375; Seevers 1957: 209 (synonymized

the genus).

Termitusa, Seevers 1957: 209; Pasteels 1967: 62.

Distinguished from all other genera including Termitusodes to which it

is most closely related by the relatively short labrum and the more rapidly

tapering abdomen. Distinguished from Thoracotusa by the position of the

mesocoxal acetabulae margins which go to a point just anterior to the pos-

terior border of the metanotum.

Overall shape ovoid, as in Fig. 1. Head broadly oval in shape. Dorsal

surface of the head without further sculpture. Head without a neck and with-

out a nuchal ridge. Eyes present, reniform, with many facets directed both

anteriorly and laterally. Gula relatively short with sides straight. Submen-

tum fused to the gula; mentum distinct from the submentum. Antennae

inserted between the eyes and the insertion of the anterior arms of the ten-

torium; 11-segmented, shaped as in Fig. 2. Labrum relatively short, shaped

as in Fig. 6. Mandibles slightly asymmetrical, shaped as in Fig. 3 and 4.

Maxillae shaped as in Fig. 7, palpi 4-segmented; apical segment attached to

most distal part or parts on preceeding segment. Labium shaped as in Fig.

5, palpi 2-segmented with the lst segment about twice as long as the 2nd.

There is a very small setae-like structure at the tip of the terminal segment

of the labial palp, which may represent a very regressed third segment.

Ligula distinct. Maxillary acetabulae not distinctly margined.

Pronotum wider than long, shaped as in Fig. 1, although the shape is

somewhat variable by species. Dorsal surface of the pronotum with various

setae counts which are variable by species. Dorsal surface of the pronotum

reflexed ventrally to a variable extent depending upon the species. Proster-

num (Fig. 9) evenly rounded between the legs and with relatively short anter-

olateral articulation processes . Prothoracic coxal cavities closed behind by

membrane in which are located the entirely membranous mesothoracic per-

itremes. Elytra shaped as in Fig. 47-71, with the ventral reflection of ely-

tra at approximately a 90°angle to the dorsal surface but does not reflect

underneath the body as in Thoracotusa. Meso- and metanotum shaped as in

Fig. 10. Wings present, of normal size, with the usual staphylinid venation.

Meso- and metasternum shaped as in Fig. 8. Mesocoxal acetabulae dis-

tinctly margined, note how the posterior margin of these acetabulae extends

to very near the posterior border of the metasternum. Pro-, meso-, and

metalegs shaped as in Fig. 15, 16, and 17, respectively; tarsal formula,

4-4-5,

Kistner: A revision of the Subtribe Termitusina

Fig. 1, -- Termitusa coatoni: A, dorsal view; B, lateral view.

Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 2-14, -- Termitusa generic characters: T. atlanticus: 2, antennae; 5, labium; 9,

prosternum; 11, wing; 12, abdominal segments II-VIN; 14, abdominal segment IX. T.

muneris; 3, left manidble; 4, right mandible; 6, labrum; 7, maxillae; 8, meso- and meta-

sternum; 10, meso- and metanotum; 13, abdominal segments VII and VIII.

Kistner: A revision of the Subtribe Termitusina as

Fig. 15-17, -- Termitusa generic characters: T. muneris: 15, proleg. T. atlanticus:

16, mesoleg; 17, metaleg. —— ee ee Ue

Abdomen subconical in shape as in Fig. 12. Abdominal segment I

represented by tergite which is fused to the metanotum (Fig. 10). Segment

Il represented by tergite alone. Segments III-VI with 1 tergite, 1 sternite,

and 2 pairs of paratergites each. Inner and outer paratergites of approx-

imately equal length. The dorsal rim of the abdomen occurs between the

outer paratergites and the sternites. Segment VII with 1 tergite, 1 sternite,

and 1 pair of full length paratergites, the outers, while the inner parater-

gites are reduced to subtriangular sclerites at the anterior end of the outer

paratergites. Segment VIII represented by the tergite and sternite alone.

segment IX highly modified into a 4-valved structure, shaped as in Fig. 14.

The apodemes of the male segment IX are greatly elongated and in the male

there is an additional small sclerite ventrally which probably represents the

sternite of the segment. These are absent in the female. Male genitalia

sclerotized, variable in shape by species. Spermatheca sclerotized, shape

variable by species.

Type species. - of Termitusa: T. sjoestedti Wasmann. Blackwelder

1952: 380.

- of Glaphyrinus: G. mundus Bernhauer. Blackwelder

1952: 170 (misspelled, Glaphyrius).

KEY TO SPECIES OF TERMITUSA

F Tergite Tbwith € BACK MACrOBOtae, .. oss 60.0 -0% 064 Nike pennesceu OD

Tergite TLwitt GO INACE MACTOSCIRGs.a5 cwkeea neon cr:deenes ananon. ©

Tergite III with 2 black macrosetae........ T. lightningae (p. 23 )

2. ( 1) Elytra with 4 rows of macrosetae; with 2 or 3 macrosetae on

later ak CG Oi 0 is o.ca Hiss ase ee olin eh ann ekk a Kaine eens © od

Elytra with 5 rows of macrosetae; with 3 macrosetae on lateral ql

CORO ie oss nhs £1 SAAR RRA OAs kk ahem ede +o oe ne

ie Ae 2 Elytra with 1 seta along inner margin......... T. bandai (p.10 )

Elytra with 3 setae along inner margin..........T. lujae (p.23 )

8 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

4, (2) Elytra with 2 setae on lateral edge........ T. malawiensis (p. 25 )

Elytra with 3 setae on lateral edge......eeoees 7. boultoni (p. 11.)

S. 41) Perede ly with 4 black macrosetae. 2. Se ee ee. 10

Persie iy wii6 black macrosetac. . 6.) 6 Sk i 6

6. (5) Pronotum with 4 rows of macrosetae, middle row with

HSCINe ee ee a See a ee ees he 8

Pronotanl Wit 3 TOWws O@SElIAG | a. Se es eke)

7. (6) Pronotum with 5 rows of macrosetae, middle row with

Se SetHe 4 oh os Se i. hexasetosa (p. 18)

Pronotum with 5 rows of macrosetae, middle row with

D Selo a as T. katangensis (p. 22 )

8. (4) Elytra with 4 rows of macrosetae..... TT. Wasmapni (p. 37 :

Blytra wil O-rows of macroselae.<.o 6 ee as a ee eS

9. (8) Pronotum with many short yellow setae between the

Brjor seine. fo ee ee ~ sack. @Oatoni (p. 16. )

Pronotum with no short yellow setae between the

fe fOr Seles eG ea ee se aes T. quadricollis (p. 29 J

10. (5) Tergite V with 4 blnek Wacredeiées. a

Tergite V with 6 black macrosetae...... T. mixaseta (p. 26 7

11. (10) Microsetae so long and so numerous as to be indistin-

guishable from the macrosetae. ..... T. superhirsutus (p. 35 )

Microsetae and macrosetae easily distinguishable. ....... 12

12. (11) Pronotum with 5 rows of macrosetae from the midline

SOC CAPO OOP Ok a he as: ak Bae was eos 13

Pronotum with 4 rows of macrosetae from the midline

tote eral ede ee a ds a a es ea 20

15.472) Plvtra with 4 rows of macroseiae.... 6s a 14

Pilvirg with.o rows cl macrdsetici c,h bee ee ee ee 17

14, (13) Elytra with 2 macrosetae on the lateral edge. ......... 15

Elytra with 3 macrosetae on the lateral edge. .T. williamsi (p. 39 )

15. (14) Tergite VIII with 2 macrosetae...... Sabine ok fo Sees on «218

Tergite VIII with 4 macrosetae; median lobe of male genitalia

shaped as in Fig. 173; spermatheca unknown...I. cameroni (p.13_ )

16. (15) Second row of elytra with 2 macrosetae; spermatheca shaped

as in Fig. I H, medium lobe of male genitalia shaped as in

ie. 8G, ce ee ees oes ee Vaeeasl nn. sp... (p. 36.)

Second row of elytra with 3 macrosetae; spermatheca as

in Fig. 94, medium lobe of male genitalia as in Fig. 72...........

[Se DCS WEC wi ne poe su cee ce 2 ablanticus (9)

17.. (4) Blytra with 2 macrosetae on lateral edge. . 2... 6s we 18

Elytra with 3 macrosetae on lateral edge. ........2e-s 19

18. (17) Abdominal sternite III with 8 macrosetae; median lobe

of male genitalia shaped as in Fig. 78; spermatheca shaped

Ae i Pe GO er ee T. cloveri (p. 13 )

Abdominal sternite III with 12 macrosetae; median lobe

of male genitalia unknown; spermatheca shaped as

in Pe 1G. ee ek a ee ee ee ae ee T. regina (p. 30 )

19. (17) Abdominal sternite VI with 8 macrosetae; median lobe

of male genitalia shaped as in Fig. 89, spermatheca

shaped as in ‘Pie: 106.05 ve > eo eee % . . 1. muneris (p. 27 )

Abdominal sternite VI with 6 macrosetae; median lobe

of male genitalia shaped as in Fig. 88; spermatheca

Kistner: A revision of the Subtribe Termitusina 9

shaned.as' ia: FiG. 1 100.0. 6 4 oes Gs cep atie eee @ T;.:leleupi LAD. ie )

20,..(18).Ebytea. with Swows ol mactochactads « ...4= «66s 4s

Elytra with 4vows of Macrochactae so 6 4c awe ae weg ke -

21. (20) Pronotum with middle rows of 5 setae. ..... TY, minor. (0.29 °)

Pronotum with middle rows of 6 setae. ..... T. dybasi (p.17 )

22. (20) Elytra with 2 setae on each lateral edge. .......2...2.026-. 25

Elytra with more than 2 setae on each lateraledge....... 23

23. (22), Elytra with 4.setae.on each lateral cdee. o a....0 #2 <i wisi oie 24

Elytra with 3 setae on each lateral edge. .... T. ugandae (p. 36)

24. (23) Abdominal sternite VII with 10 macrosetae; median

lobe of male genitalia shaped as in Fig. 85; spermatheca

Shaped ae Hy rie. 110, he ot ee T. ruellei (p. 30 )

Abdominal sternite VII with 8 macrosetae, median lobe of

male genitalia shaped as in Fig. 91; spermatheca shaped

Ao a Fe ee ee eae T. sheasbyi (p. 34 )

25. (22) Abdominal sternite III with 12 macrosetae. . . T. escalarae (p.18 )

Abdominal sternite III with 8 macrosetae. ........e.46-. 26

26. (25) Pronotum with no secondary setae between the macrosetae.. 27

Pronotum with at least a few secondary setae between the

WC OOG aes whl ak 6 a ea ee we ee ee ee 29

27. (26) Abdominal sternite VI with 8 macrosetae; median lobe of

male genitalia shaped as in Fig. 86; spermatheca shaped

ASU Pie lee ee ce ag ee ee ee T. sjoestedti (p. 34 )

| Abdominal sternive VI with G6 macroseine. .°. (2). 2°. a4 eee 28

28, (27) Tergites II and III with 4 macrosetae. ..... .T. hystrix (p. 21 )

Tergite Il and III with 2 macrosetae....... I. mundus (p.26 )

29. (26) Abdominal sternite III with 8 macrosetae; median lobe of

male genitalia shaped as in Fig. 84; spermatheca shaped

Ge My ee oI ay a ae pk ee eee T. pasteelsi (p.29 )

Abdominal sternite III with 10 setae; median lobe of male

genitalia shaped as in Fig. 179; spermatheca shaped as in

Oe One aba ee Sarasa ee ee ees dk T. delignei (p.16 )

Termitusa atlanticus n. sp.

Figures '2,. 5, 9, 11,:12,:14,.16,.17, 18, 47, 72, 94

Most closely related to T. cameroni from which it is distinguished by

the chaetotaxy of tergite VIII, the shape of the male genitalia, and the shape

of the spermatheca.

Color light reddish brown throughout with the elytra and head some-

what darker than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with numerous fine yellow setae interspersed

among the macrosetae. Macrochaetotaxy of pronotum as in Fig. 18; setae

counts are 5,3,1,3,3. Macrochaetotaxy of elytra as in Fig. 47; setae counts

are 2,3,3,2. Macrochaetotaxy of abdominal tergites II-VIII as follows: 2,4,

4,4,4,4,2. Macrosetae on tergites VII and VIII are shorter and thinner than

the rest of the tergites. There are numerous light yellow setae on the apical

third of abdominal segment VIII which is only slightly indented on its poster-

ior border. Paratergites without macrosetae. Macrochaetotaxy of abdom-

10 Contr. Amer. Ent. Inst., vol.10, no.4, 4974

inal sternites III-VIII as follows: 2-8,8,8,8,6,6; the 2 on sternite VII are

lateral. Median lobe of male genitalia shaped as in Fig. 72. Spermatheca

shaped as in Fig. 94.

Measurements. - Pronotum length, 0. 43-0.51; elytra length, 0. 35-

0.40. Number measured, 10.

Holotype. - Male, No. 14617, Rhodesia, Atlantica Ecological Research

Station, nr. Salisbury, 27 Mar. 1970, ex nest T-265, Coll. J. R. Clover,

D. H., and A. C. Kistner, No. 1907. In the collection of D. H. Kistner.

Paratypes. - Rhodesia: 84, same data as holotype, (D. K.); 11, Umtali

Distr. , 62 km ex Umtali - Burma Valley, 16 Oct. 1973, ex nest TM 29021.

Coll. J. L. Sheasby, No. T-879, (N.C.I., D.K.); 4, Melsetter Distr. , 10

km ex Melsetter - Cashel via Scenic Drive, 10 Oct. 1973, ex nest TM 28731,

Coll. J. L. Sheasby, No. T-878, (N.C.I., D.K.); 1, Chipinga Distr. , 8 km

ex Mt. Selinca - Chipinga, 29 Oct. 1973, ex grey mound of nest TM 29672,

Coll. W. G. H. Coaton, No. T-881, (N.C.I.). Malawi: 99, Ntchisi Forest

Reserve, 11 Sept. 1973, ex nest T-563, Coll. D. H., A. C., and K. M. C.

Kistner, (D. K.).

Notes. - The host termites of nest T-265 were determined to be Cubi-

termes pallidiceps (Sjoestedt) by R. M. C. Williams while the host termites

of all the other colonies were determined to be the same species by compari-

son with specimens from colony T-2¢5 by Kistner. Specimens of colony T-

265 and T-583 are in the collection of the British Museum (Natural History),

London ane the Kistner collection, while specimens from the rest of the co-

lonies collected by Sheasby anc Coaton are in the Natural Isoptera Collection

Pretoria and the Kistner collection.

Termitusa bandai n. sp.

Figures I] A- E

Most closely related to T. lujae from which it is distinguished by the

chaetotaxy of the pronotum, elytra, and the shape of the male genitalia.

When females are known of T. lujae there will undoubtedly be differences

there also.

Color dark reddish brown throughout with the abdomen somewhat

lighter than the rest of the body, legs somewhat lighter than this. Dorsal

surface of the head, pronotum, and elytra smooth and shiny beneath the very

dense vestiture of lighter setae. Macrochaetotaxy of pronotum as in Fig. II

B; counts from the midline are: 4, 3, 1, 3, 3, Macrochaetotaxy of elytra as

follows: 1, 3, 3, 4, 3; as in Fig. I1 A. Macrochaetotaxy of abdominal ter-

gites II-VIII as follows: 2, 6, 6, 6, 6, 6, many. All of the macrosetae are

interspersed with numerous pale yellow setae. Setae on tergite VII are

shorter ana thinner than those on the rest of the tergites. Setae on tergite

VIII are so dense as to obscure differences between macro- and microsetae.

Paratergites without macrosetae. Macrochaetotaxy of abdominal sternites

Ili-VIli as follows: 10, 19, 19, 10, 2-10, 6-10. Median lobe of the male ge-

nitalia shaped as in Fig, II E. Lateral lobe of male genitalia area ey as in

Fig. Ii D. Spermatheca shaped as in Fig. ITC.

Measurements. - Pronotum length, 9. 48-9.50; elytra length, 0. 36-

Kistner: A revision of the Subtribe Termitusina 11

0,38, Number measured, 10.

Holotype. - Male, number 15401, Malawi, Ntchisi Forest Reserve,

12 April 1973, ex nest T-586, Coll. D. H., A. C., and K. M. C. Kistner,

No. 3039. In the collection of D. H. Kistner.

Paratypes. - 29, same data as holotype, (D.K.).

Notes. - The host termites were determined to be Cubitermes sp. by

Kistner, specimens of which are in the collection of the British Museum

(Natural History), London and the Kistner Collection. This species has a

slightly larger head than C. pallidiceps but is larger than C. sankurensis

(Wasmann). The species is named after the Life President of Malawi, Dr.

N. Hastings Banda.

Termitusa boultoni n. sp.

Figures 19, 48, 77, 95

Distinguished from T. wasmanni and T, malawiensis to which it is

most closely related, by the chaetotaxy of the pronotum, elytra, and the

shape of the male genitalia and spermatheca.

Color dark reddish brown throughout with the legs and antenna some-

what lighter than the rest of the body. Dorsal surface of the head, prono-

tum, and elytra smooth and shiny with numerous fine yellow setae inter-

Spersed among the macrochaetae. Macrochaetotaxy of the pronotum as in

Fig. 19; setae counts are 4,3,3,3. Macrochaetotaxy of elytra as in Fig. 48;

setae counts are 2,3,3,3. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 4,6,6,6,6,6,2. There are numerous light yellow setae scattered

in among the macrochaetae on the abdomen. Macrochaetae on tergites VII

and VIII are thinner and lighter in color than those on the other tergites.

Tergite VIII with many yellow setae on the apical half. Dorsal margin of

tergite VIII indented. No macrochaetae on paratergites. Macrochaetotaxy

of abdominal sternites III-VIII as follows: 12,12, 12,12,10,6-8. Median

lobe of male genitalia shaped as in Fig. 77. Spermatheca shaped as in Fig.

95.

Measurements, - Pronotum length, 0. 41-0. 45; elytra length, 0. 33-

0.35. Number measured, 3.

Holotype. - Male, No. 14551, Rhodesia, 1 mi ex Atlantica Ecological

Research Station-Saffron Walden, 29 Mar. 1970, ex nest T-267, Coll J. R.

Clover, D. H. & A. C. Kistner, No. 1921. In the collection of D. H.

Kistner.

Paratypes. - 1 male, 1 female, same data as holotype except from

nest T-268 and with collection number 1923 (D. K.).

Notes. - The host termites were determined to be Cubitermes sankur-

ensis Wasmann by R. M. C. Williams. Specimens are in the collection of

the British Museum (Natural History), London, and the Kistner collection.

The species is named after Mr. Rudyerd Boulton, Director of the Atlantica

Ecological Research Station for all his help in locating termites.

Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 18-27, -- Pronota: 18, Termitusa atlanticus; 19, T. boultoni; 20, T. cameroni;

21, T. cloveri; 22, T. coatoni; 23, 7. “deligner 24, J. dyoasi, 29, I. escalarae, 20,

hexasetosa; 27, T. hystrix.

Kistner: A revision of the Subtribe Termitusina 13

Termitusa cameroni Reichensperger

Figures 20, 49, 73

Termitusa cameroni Reichensperger 1929: 136 (Tanganyika, Ngeren-

gere, with Cubitermes glebae (Sjoestedt); Seevers 1957: 212 (redescription).

Distinguished from T. atlanticus, to which it is most closely related,

by the chaetotaxy of the abdominal tergite VIII and the shape of the male

genitalia.

Color light reddish brown throughout with the legs and antennae some-

what lighter than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with numerous fine yellow setae interspersed

among the macrosetae. Macrochaetotaxy of pronotum as in Fig. 20; setae

counts are 5,3,2,3,2. Macrochaetotaxy of elytra as in Fig. 49; setae counts

are 2,3,3,2. Macrochaetotaxy of abdominal tergites H-VIII as follows: 2, 4,

4,4,4,4,4. There are numerous light yellow setae scattered in amongst the

macrochaetae of all parts of this species. Macrochaetae on tergites VII and

VIII are thinner and lighter in color than those on the other tergites. Ter-

gite VIII with many yellow setae on the apical half, which tend to obscure the

larger setae. No macrochaetae on paratergites. Macrochaetotaxy of abdom-

inal sternites III-VIII as follows: 8,8,8,8,6,6. Median lobe of male gen-

italia shaped as in Fig. 73. Spermatheca unknown.

Measurements. - Pronotum length, 0. 43; elytra length, 0.35. Number

measured, 1.

Material examined. - 1 male, Tanzania, Ngerengere, (F. M. N. H.).

Notes. - The host termites were determined to be Cubitermes glebae

Sjoestedt. While we excavated numerous C. glebae nests while we were in

Tanzania none of our nests yielded specimens of this species.

Termitusa cloveri, n. sp.

Figures 21, 46, 50, 78, 96

Distinguished from T. muneris to which it is most closely related by

the chaetotaxy of the elytra and the shape of the male genitalia and female

spermatheca.

Color dark reddish brown throughout with the abdomen somewhat light-

er than the head, pronotum, and elytra. Dorsal surface of the head, pro-

notum, elytra, and abdomen smooth and shiny with chaetotaxy as follows.

Pronotum with chaetotaxy as shown in Fig. 21; with row counts from the med-

ian to the outside as follows: 5,3,1,3,3. Elytra with chaetotaxy as shown in

Fig. 50 and row counts as follows: 2,3,2,1,2. There are no short fine setae

between the macrochaetae. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2,4,4,4,4,4,4. The macrosetae on tergites VII and VIII are much

thinner and shorter than the setae on the other segments. Segment VIII has

additionally a large number of yellow setae coming out of the apical third;

these tend to blend in with the row of setae across the anterior end of this.

Paratergites without macrochaetae. Macrochaetotaxy of abdominal sternites

III- VIII as follows: 8,8,8,8,8,6. Macrochaetotaxy of abdominal segment IX

with 3 on each of the median portions and 2 on each lateral portion. Male

14 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 28-37. -- Pronota: 28, T. katangensis; 29, T. leleupi; 30, T. lujae; 31, T. minor;

32, T. mixaseta; 33, T. mundus; 34, T. muneris; 35, T. pasteelsi; 36, T. quadricollis;

S121 Desina., ! M

genitalia shaped as in Fig. 78. Female spermatheca shaped as in Fig. 96.

Measurements. - Pronotum length, 0. 49-0. 52; elytra length, 0. 38-

0.41. Number measured, 10.

Kistner: A revision of the Subtribe Termitusina 15

Holotype. - Male, no. 14602, Zambia, 10 mi ex Mbala-Mpulungu, 22

May 1970, from the vicinity of the queen of nest T-338, Coll. J. R. Clover,

D. H. and A. C. Kistner, no. 2175. In the collection of D. H. Kistner.

Paratypes. - Zambia: 36, same data as holotype, (D. K.); 54, same

locality and nest as holotype, from other parts of the nest, Coll. J. R.

Clover, D. H. and A. C. Kistner, no. 2174, (D. K.). Zaire: 2, Elisabeth-

ville, 3 Jan. 1949, Coll. N. Leleup, no. 8, (M. R. A. C., D. K.); 1,

Fig. 38-46, -- Pronota: 38, T. ruellei; 39, T. sheasbyi; 40, T. sjoestedti; 41, T. super-

hirsutus; 42, T. ugandae; 43, T. wasmanni; 44, T. williamsi. Lateral lobes of male

genitalia: 45, T. hexasetosa; 46, T. cloveri.

16 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Katanga, Tshila-Tembo, 20 Feb 1949, Coll. N. Leleup, no. 1, (M. R. A. C.);

2, Kyapamoka, Source dela Mukusu, 11 Feb. 1949, Coll. N. Leleup, (M. R.

A. Co) K_),

Notes. - The host termites were determined to be Cubitermes inclitus

(Silvestri) by R. M. C. Williams. Specimens are in the collections of the

British Museum (Natural History), London, and Kistner. The species is

named after Mr. James Clover, one of the collectors.

Termitusa coatonin. sp.

Figures 1, 22, 51, 74

Closely related to Termitusa quadricollis from which it is distinguish-

ed by the chaetotaxy of abdominal segment VIII and the presence of many

secondary setae between the macrosetae of the pronotum.

Color dark reddish brown throughout with legs and antennae some-

what lighter than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with a covering of fine yellow setae interspersed

among the macrochaetae. Macrochaetotaxy of pronotum as in Fig. 22; setae

counts are 4,3,3,3. Macrochaetotaxy of elytra as in Fig. 51; setae counts

are 2,3,3,1,3. Macrochaetotaxy of abdominal tergites II-VIII as follows: 2,

4,6,6,6,6,2. There are numerous fine yellow setae interspersed among the

macrosetae of the abdomen. Macrosetae on tergites VII and VII much

smaller and thinner and lighter in color than the macrosetae on the other

tergites. Paratergites without macrosetae. Macrochaetotaxy of abdominal

sternites III-VIII as follows: 12,12,12,10,10, 6-8. The macrosetae on

sternite VIII are arranged irregularly in 2 rows. Median lobe of male gen-

italia shaped as in Fig. 74. Female unknown. Note that the chaetae shown

in Fig. 1 are somewhat impressionistic.

Measurements. - Pronotum length, 0. 46; elytra length, 0.35. Number

measured, 1. ,

Holotype. - Male, no. 14597, Zambia, Mbala (Abercorn), 15 May 1970,

ex nest T-327, Coll. J. R. Clover, D. H., A. C., A. H., and K, M. C.

Kistner, no. 2119.

Paratypes. - 1 male, same data as holotype, (D. K.).

Notes. - These specimens came from a mixed colony of Thoracotermes

lusingensis Harris and Cubitermes oblectatus Williams. Because no Term-

itusa were recovered from any pure colonies of Thoracotermes it is highly

likely that the actual host of the species is the Cubitermes. The host ter-

mites were determined by R. M. C. Williams, specimens of which are in the

British Museum (Natural History) and Kistner. The species is named after

me W. G. H. Coaton in honor of his studies of the termites of Southern

rica.

Termitusa delignein. sp.

Figures 23, 52, 79, 97

Distinguished from T. pasteelsi, to which it is most closely related,

Kistner: A revision of the Subtribe Termitusina 17

by the shape of the carinate male genitalia, slight differences in the female

spermatheca, and its smaller size.

Color dark reddish-brown throughout with the appendages and antennae

somewhat lighter thanthe rest of thebody. Dorsal surface of the head, pro-

notum, and elytra smooth and shiny with very few secondary setae between

the macrochaetae. Macrochaetae relatively short in this species. Macro-

chaetotaxy of pronotum as in Fig. 23; setae counts are 4,2,3,3. Macro-

chaetotaxy of elytra as in Fig. 52; setae counts are 2,3,1,2. Macrochaeto-

taxy of abdominal tergites II-VIII as follows: 2,4,4,4,4,4,2. The macro-

chaetae on abdominal tergites VII and VIII are shorter and thinner than those

on preceeding segments. Macrochaetae on tergites II and the middle macro-

chaetae of tergite III are also short and thin. Many secondary setae on ter-

gites VII and VIII where they are particularly conspicuous toward the pos-

terior third of tergite VII. Paratergites without macrosetae. Macrochaeto-

taxy of abdominal sternites III-VIII as follows: 10,8,6,6,6,6. Six of the 8

macrochaetae on abdominal sternite III are shorter and thinner than all the

rest of the macrochaetae. Median lobe of male genitalia carinate, shaped

as in Fig. 79. Spermatheca shaped as in Fig. 97.

Measurements, - Pronotum length, 0. 41-0. 48; elytra length, 0. 34-

0.39. Number measured, 10.

Holotype. - Male number 14933, Gabon, Mekob, (Ivindo), 14 km. up-

stream from Makokou, 8 May 1962, Coll. J. M. Pasteels and J. Deligne, no.

1072. In the collection of Deligne and Pasteels, Brussels.

Paratypes. - 17, same data as holotype, (J. M. P., D. K.).

Notes. - The host termites were determined to be Cubitermes heghi

Sjoestedt, by J. M. Pasteels, and are in the collection of Deligne and

Pasteels, Brussels.

Termitusa dybasin. sp.

Figures 24, 53, 98

Most closely related to T. leleupi from which it is distinguished by the

chaetotaxy of the pronotum and elytra and the shape of the female sperm-

atheca.

Color reddish-brown throughout with the antennae and legs somewhat

lighter than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny without light yellow setae scattered in amongst the

heavier macrosetae. Macrochaetotaxy of pronotum as in Fig. 24; counts are

6,3,3,3, Macrochaetotaxy of elytra as in Fig. 53; setae counts are 2,1,3,1,

2. Macrochaetotaxy of abdominal tergites II-VIII as follows: 2,4, 4, 4,4, 4,0.

There are some light yellow setae scattered in amongst the darker setae of

the abdomen. Macrosetae on segment VII are lighter in color and thinner

than the macrosetae of the other abdominal tergites. Paratergites without

macrosetae. Macrochaetotaxy of abdominal stergites III-VIII as follows: 8,

8,8,6,6,0. Spermatheca shaped as in Fig. 98. Male genitalia unknown.

Measurements. - Pronotum length, 0. 48; elytra length, 0.34. Number

measured, 1.

Holotype. - Female, 14907, Cameroun, Tiko, October 1933, Coll. H.

Piening. In the collection of the Field Museum of Natural History, Chicago.

Notes. - The host of this species is unknown although there is a

18 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

notation on the label that it was taken from a termite nest. The species is

named for Henry S. Dybas, Curator of Insects in the Field Museum of Nat-

ural History, Chicago.

Termitusa escalarae Fauvel

Figures 25, 54, 75, 99

Termitusa escalarae Fauvel 1906: 289 - Institut Royal des Sciences

naturelles de Belgique, Brussels- (Biafra, Cabo San Juan, Rio Muni,with

Cubitermes sp.); Wasmann 1911: 163 (Gabon); Seevers 1957: 211 (rede-

scription).

Related to T. mundus from which it is distinguished by the chaetotaxy

of the abdominal sternite VIII, the presence of secondary setae between the

macrochaetae of the pronotum, and the shape of the male genitalia and sperm-

atheca.

Color reddish brown throughout with the pronotum and head somewhat

darker than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny with very few secondary setae between the macro-

chaetae. Macrochaetotaxy of pronotum as in Fig, 25; setae counts are 4,2,

3,3. Macrochaetotaxy of elytra as in Fig. 54; setae counts are 2,1,3,2.

Macrochaetotaxy of abdominal tergites ]-VIII as follows: 2,2,4,4,4, 4, 2.

Macrosetae on tergite VII and VIII are smaller, thinner, and somewhat light-

er in color than the macrochaetae on the rest of the tergites. Apical 4th of

tergite VIII covered with many light colored secondary setae. Paratergites

without macrosetae. Macrochaetotaxy of abdominal sternites III-VIII as fol-

lows: 8,8,8,6,6,6. Median lobe of male genitalia shaped as in Fig. 75.

Spermatheca shaped as in Fig. 99.

Measurements. - Pronotum length, 0. 42-0. 45; elytra length, 0. 31-

0.34. Number measured, 10.

Material examined. - Type, Biafra, Cabo San Juan, Aug. 1901, Coll.

Escalera, (I. R. S. N. B.); 18, Cameroons, Douala, Sept. 1912, Coll.

Rothkirch, (N. H. M., D. K.).

Notes. - The termites are as yet unidentified.

Termitusa hexasetosa n. Sp.

Figures 26, 45, 55, 80, 100

Closely related to T. quadricollis from which it is distinguished by the

chaetotaxy of the pronotum and elytra, and the shape of the median lobe of the

male genitalia and spermatheca.

Color dark reddish brown throughout with the antenna and legs a little

lighter than the rest of the body. Dorsal surface of the head, pronotum,

elytra, and abdomen smooth and shiny with essentially no secondary setae

scattered in among the primary macrosetae. Macrochaetotaxy of pronotum

as in Fig. 26; setae counts are 4,3,2,3,4. Macrochaetotaxy of elytra as in

Fig. 55; setae counts are 2,2,2,3,3. Macrochaetotaxy of abdominal tergites

II-VIII as follows: 2,4,6,6,6,6,2. There are very few shorter and thinner

Kistner: A revision of the Subtribe Termitusina 19

Fig. 47-56. -- Elytra: 47, Termitusa atlanticus; 48, T. boultoni; 49, T. cameroni; 50,

T. cloveri; 51, T. coatoni; 52, T. delignei; 53, T. dybasi; 54, T. escalarae; 55, T.

hexasetosa; 56, T. hystrix. it ba Sg

20 Contr, Amer. Ent. Inst., vol.10, no.4, 1974

Pig. 57-66. -= Elytra: 67, -T.. kKatangensis; 98, 7. leleupi; 59, T. lujae; 60, T. minor,

61, T. mixaseta; 62, T. mundus; 63, T. pasteeisi; 64, T. quadricollis; 65, T. regina,

66, T. sheasbyi.

Kistner: A revision of the Subtribe Termitusina 21

setae in among the macrosetae. Macrosetae on tergites VII and VIII are

shorter and thinner than the macrosetae on the other tergites. Paratergites

without macrosetae. Macrochaetotaxy of sternites III-VIII as follows: 12,

12,12,10,10,4-10. Median lobe of male genitalia shaped as in Fig. 80,

Spermatheca shaped as in Fig. 100.

Measurements. - Pronotum length, 0.51-0.55; elytra length, 0. 38-

0.40. Number measured, 7.

Holotype. - Male, no. 14615, Zaire, Katanga, Tshila-Tembo, 20 Feb.

1949, Coll. N. Leleup, ex termite nest no. 1. In the collection of the

Musée Royal de l'Afrique Centrale, Tervuren.

Paratypes. - Zaire, Katanga: 2, same data as holotype, (M. R. A. C.,

D. K.); 2, Massif des Kundelungu, 15 Jan. 1949, Coll. N. Leleup, ex ter-

mite nest no. 7, (M. R. A. C.,D. K.); 2, Kundelungu, 1750 m. 23 Apr. 1950,

Coll. N. Leleup, ex nest M. C. no. 7, (M. R. A. C., D. K.).

Notes. - The host termites have not been identified.

Termitusa hystrix Wasmann

Figures 27, 56, 76, 101

Termitusa hystrix Wasmann 1911: 159, 163, pl. 5, Fig. 10 (Belgian

Congo, Sankuru, with Cubitermes fungifaber (Sjoestedt); Hegh 1922: 613,

Fig. 421; Seevers 1957: 211 (redescription)

Closely related to T. ugandae from which it is distinguished by the

chaetotaxy of the pronotum and elytra and the shape of the male genitalia

and spermatheca.

Color reddish brown throughout with the abdomen somewhat lighter than

the head, pronotum, and elytra. Dorsal surface of the head, pronotum, ely-

tra, and abdomen, smooth and shiny with very few punctures scattered ran-

domly about. Chaetotaxy of pronotum as shown in Fig. 27 with the setae

counts from the middle to the outside as follows: 4,3,3,3. Macrochaetotaxy

of elytra as in Fig. 56 with the row counts as follows: 2,3,1,2. The 2

rows on the outside of elytra are much finer and shorter than the 2 rows on

the interior of the elytra. There are no fine short setae interspersed be-

tween the major setae on either the pronotum or the elytra. Macrochaeto-

taxy of abdominal tergites II-VIII as follows: 4,4,4,4,4,4,2. Those on

Segments VII and VIII are much thinner and shorter than the setae on the rest

of the segments, Paratergites without macrochaetae, although the most lat-

eral setae on the sternites are so close to the paratergites as to appear to

come from them. Macrochaetotaxy of abdominal sternites III-VIII as follows:

8,8,6,6,6,6. Macrochaetae on sternite III are shorter and thinner than the

macrochaetae on the rest of the segments. Macrochaetae on sternites VI-

VIII appear to be set on special thickening of the exoskeleton somewhat re-

sembling tubercles. Segment IX with 3 macrochaetae on each of the center

parts and 1 macrochaeta on each of the lateral parts. Median lobe of male

genitalia shaped as in Fig. 76. Spermatheca shaped as in Fig. 101.

Measurements. - Pronotum length, 0. 45-0. 47; elytra length, 0. 33-

0.35. Number measured, 2.

Material examined. - Type and cotype, Zaire, Kasai, Sankuru, Nov.

1907, Coll. E. Luja, (N. H. M.); 1 male, 1 female, Zaire, Yangambi, 4

22 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

July 1960, Berlese, nest no. 297, Coll. J. Décelle, no. 298, (D. K.).

Notes. - The host for the specimens taken at Yangambi were deter -

mined to be Cubitermes fungifaber (Sjoestedt) by Kumar Krishna. Speci-

mens of the hosts are in the collection of the American Museum of Natural

History, New York.

Termitusa katangensis n. sp.

Figures 28, 57, 81, 102

Closely related to T. hexasetosa from which it is distinguished by the

chaetotaxy of the pronotum, the chaetotaxy of the elytra, and the shape of the

male genitalia and the female spermatheca.

Color dark reddish brown throughout with the legs and antenna some-

what lighter than the rest of the body. Dorsal surface of the head, pronotum,

elytra, and abdomen smooth and shiny with numerous fine yellow setae

scattered in among the dark macrochaetae. Macrochaetotaxy of pronotum as

in Fig. 28; setae counts are 5,3,1,3,3. Macrochaetotaxy of elytra as in Fig.

57; setae counts are 2,3,3,2,3. Macrochaetotaxy of abdominal tergites II-

VIII as follows: 2,4,6,6,6,6,2. There are many fine yellow setae scattered

in among the darker macrochaetae on the abdomen, expecially on tergite VII

and the apical portion of tergite VII. Paratergites without macrochaetae.

Macrochaetotaxy of abdominal sternites III-VIII as follows: 12,12,12,10,10,

4-8, Median lobe of male genitalia shaped as in Fig. 81. Spermatheca

shaped as in Fig. 102.

Measurements. - Pronotum length, 0.45-0.49; elytra length, 0. 35-

0.38. Number measured, 10.

Holotype. - Male, no. 14616, Zaire, Elisabethville, 15 Mar. 1949,

Coll. N. Leleup, ex nest no. 54. In the collection of the Musée Royal de

l'Afrique Centrale, Tervuren.

Paratypes. - 13, same data as holotype, (M. R. A. C., D. K. a

Notes. - The host termites have not been identified.

Termitusa leleupin. sp.

Figures 29, 58, 82, 103

Distinguished from T. muneris to which it is most closely related by

its larger size, the chaetotaxy of the elytra, and the shape of the male gen-

italia and spermatheca.

Color dark reddish brown throughout with the abdomen and legs some-

what lighter than the rest of the body. Dorsal surface of the head, pro-

notum, and elytra smooth and shiny with many fine yellow setae scattered in

among the macrosetae. Macrochaetotaxy of the pronotum as in Fig. 29;

setae count is 5,3,1,3,3. Macrochaetotaxy of elytra as in Fig. 58; setae

count is 2,3,2,1,3. Macrochaetotaxy of abdominal tergites II-VIII as fol-

lows: 2,4,4,4,4,4,2. There are many fine yellow setae scattered in among

the macrosetae. Macrosetae on tergites VII and VIII are finer and thinner

than the macrosetae on the rest of the abdomen. Paratergites without

macrosetae. Macrochaetotaxy of abdominal sternites IJJ-VIII as follows:

Kistner: A revision of the Subtribe Termitusina 23

2-8, 2-8,8,8,6,6; the 2 on sternites III and IV are lateral. Male gen-

italia shaped as in Fig. 82. Female spermatheca shaped as in Fig. 103.

Measurements, - Pronotum length, 0.52-0.55; elytra length, 0. 36-

0.39. Number measured, 10.

Holotype. - Male, no. 14539, Zaire, Katanga, Tshila-Tembo, 20 Feb.

1949, Coll. N. Leleup, in termite Nest No. 1. In the collection of the Musée

Royal de l'Afrique Centrale, Tervuren.

Paratypes. - 10, same data as holotype, (M. R. A. C., F. M. N. H.,

D. K.); 3, Zaire, Katanga, Tshila-Tembo, 20 Feb. 1949, Coll. N. Leleup,

from an ant nest, (M. R. A. C., D. K.); 6, Zaire, Katanga, Lukuni, 30 Dec.

1948, Coll. N. Leleup, ex termite nest no. 9, (M. R. A. C., D. K.).

Notes. - The host termites have not been identified. The species is

named for the collector, Dr. N. Leleup.

Termitusa lightningae n. sp.

Figures IA -C

Related to T. escalarae and T. mundus but easily distinguishable from

them by the chaetotaxy of the abdomen, the pronotum, the elytra, and the

shape of the female spermatheca.,

Color dark reddish brown throughout with the abdomen somewhat

lighter than the rest of the body; legs somewhat lighter than this. Dorsal

surface of the head, pronotum, and elytra smooth and shiny with a sparse

vestiture of microsetae. Macrochaetotaxy of pronotum as in Fig. I C;

counts from the midline are: 4, 3, 1, 3, 4. Macrochaetotaxy of elytra as

follows: 3, 3, 1, 3, 4; as in Fig. I A. Macrochaetotaxy of abdominal ter-

gites II-VIII as follows: 2, 2, 4, 4, 4, 4, 2. All of the macrosetae are inter-

spersed with numerous pale yellow setae. Setae on tergites VII and VIII are

shorter and thinner than those on the rest of the tergites. Paratergites with-

out macrosetae. Macrochaetotaxy of abdominal sternites III-VIII as follows:

10, 10, 19, 8, 8, 6. All of the macrosetae are interspersed with numerous

pale yellow setae. Male unknown. Spermatheca shaped as in Fig. I B.

Measurements. - Pronotum length, 0.42; elytra length, 0.30. Number

measured, 1.

Holotype. - Female, number 14946, Guinea, Kamsar, 9 October 1972,

ex nest T-427, Coll. D. H., A. C., A. H., and K. M. C. Kistner, No. 2489.

In the collection of D, H. Kistner.

Notes. - The host termites were determined to be Cubitermes sp. by

Kistner, specimens of which are in the collection of the British Museum (Na-

tural History), London and the collection of D. H. Kistner. The termites

were not similar to any available in the Kistner collection. The species is

named for the Farrell Lines ship S.S. African Lightning on whose decks the

nests from Kamsar, Guinea were necessarily sorted. This name not only

honors a great merchant vessel but also her captain, E. Shuttleworth, and

crew who were So helpful to us in our collecting efforts.

Termitusa lujae Wasmann

Figures 30, 59, 88

24 Contr, Amer. Ent. Inst. ,.vol..10; no.4, 1974

Termitusa lujae Wasmann 1911: 156, 159, 164 - Naturhistorisch

Museum, Maastricht - (Belgian Congo, Sankuru, with Cubitermes sankur-

ensis Wasmann; Seevers 1957: 211 (redescription)

Most closely related to T. boultoni from which it is distinguished by

the chaetotaxy of the pronotum, elytra, and the shape of the male genitalia.

Color dark reddish brown throughout with the abdomen somewhat light-

er than the rest of the body, legs somewhat lighter than this. Dorsal sur-

face of the head, pronotum, and elytra smooth and shiny beneath the very

dense vestiture of setae. Macrochaetotaxy of pronotum as in Fig. 30; counts

from the midline are: 4,4,1,3,3. Macrochaetotaxy of elytra as follows:

3,3,3,4,3; as in Fig. 59. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2,6,6,6,6,6,2. All of the macrosetae are interspersed with num-

erous pale yellow setae. Setae on tergites VII and VIII are shorter and thin-

ner than those on the rest of the tergites. Paratergites without macrosetae.

Macrochaetotaxy of abdominal sternites III-VIII as follows: 10,10,8,8,8,8.

Median lobe of male genitalia shaped as in Fig. 88. Spermatheca unknown.

Measurements. - Pronotum length, 0. 41-0. 43; elytra length, 0. 36-

0.38. Number measured, 2.

Material examined. - Type plus 3 co-types, Zaire, Kasai, Sankuru,

Jan 1905, Coll. E. Luja, with Cubitermes sankurensis Wasmann, no. 19,

(NL Hy M:, M. R. A. C.);1, Kondué; Coll. B. Luja, (fF. M,N. H.).

Fig. 67-71. -- Elytra: 67, Termitusa sjoestedt; 68, T. superhirsutus; 69, T. ugandae;

70, T. wasmanni; 71, T. williamsi. a ss

Kistner: A revision of the Subtribe Termitusina 95

Termitusa malawiensis n. sp.

Figures I D-F

Closely related to T. lujae and T. bandai from which it is distinguished

by the chaetotaxy of the pronotum, elytra, and the shape of the female sperm-

atheca.

Color dark reddish brown throughout with the legs and antennae some-

what lighter than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with numerous fine yellow setae interspersed

among the macrochaetae. Macrochaetotaxy of the pronotum as in Fig. I F;

setae counts are 3, 3, 2, 3. Macrochaetotaxy of elytra as in Fig. I D;

setae counts are 2, 2, 3, 2. Macrochaetotaxy of abdominal tergites II-VIII

as follows: 2, 6, 6, 6, 6, 6, 6. There are numerous light yellow setae

scattered in among the macrochaetae on the abdomen. Macrochaetae on ter-

gites VII and VIII are thinner and lighter in color than those on the other ter-

gites, Tergite VIII with many yellow setae on the apical half. Dorsal mar-

gin of tergite VIII indented. No macrochaetae on paratergites. Macrochae-

totaxy of abdominal sternites III-VIII as follows: 8, 8, 10, 8, 8, 6-8. Male

unknown. Female spermatheca shaped as in Fig. I E.

Measurements, - Pronotum length, 0.39; elytra length, 0.32. Number

measured, 1.

Holotype. - Female, 14965, Malawi, 10 mi. ex Kusungu-Kusungu Na-

tional Park, 14 April 1973, ex nest T-589, Coll. D. H. and A. C. Kistner,

No. 3047. In the collection of D. H. Kistner.

Notes. - The host termites with which this species was associated were

determined to be Cubitermes tenuiceps (Sjoestedt) by Kistner. The host ter-

mites are in the collection of the British Museum (Natural History), London

and the Kistner Collection. The labrum of the termites were only slightly

shorter than labrums of C. tenuiceps from coastal Tanzania determined by

Mr. R. M. C. Williams.

Termitusa minor n. sp.

Figures 31, 60, 104

Related to T. regina, from which it is distinguished by the chaetotaxy

of the pronotum and the shape of the spermatheca.

Color dark reddish throughout with the antenna and the appendages

somewhat lighter than the rest of the body. Dorsal surface of the head, pro-

notum, and elytra smooth and shiny with virtually no secondary setae be-

tween the macrochaetae. Macrochaetotaxy of pronotum as in Fig. 31; setae

counts are 5,3,3,3. Macrochaetotaxy of elytra as in Fig. 60; setae counts

are 2,2,2,2,2. Macrochaetotaxy of abdominal tergites II-VIII as follows: 2

4,4,4,4,4,2. There are very few light yellow setae interspersed among the

macrosetae. Posterior third of abdominal tergite VIII has numerous sec-

ondary setae. Paratergites without macrosetae. Macrochaetotaxy of abdom-

inal sternites III-VIII as follows: 2-8,6,6,8,6,6; the 2 on sternite III are

lateral. Spermatheca shaped as in Fig. 104. Male unknown.

Measurements. - Pronotum length, 0.45; elytra length, 0. 35-0. 36.

Number measured, 2.

26 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Holotype. - Female, no. 14619, Zaire, Kundelungu, 1750 m, Apr.

1950, Coll. N. Leleup, ex termite nest MC 8. In the collection of the Musée

Royal de l'Afrique Centrale, Tervuren.

Paratype. - 1 female, same data as holotype, (D. K.).

Notes. - The identity of the host termites is not known.

Termitusa mixaseta n. sp

Figures 32, 61, 83

Closely related to T. sheasbyi from which it is distinguished by the

chaetotaxy of the abdomen, the pronotum, the elytra, and the shape of the

male genitalia and spermatheca.

Color dark reddish brown throughout with the appendages and antenna

somewhat lighter than the rest of the body. Dorsal surface of the head,

pronotum, and elytra smooth and shiny with no secondary setae scattered

about. Macrochaetotaxy of pronotum as in Fig. 32; setae counts are 4,3,1,

3,5. Macrochaetotaxy of elytra as in Fig. 61; setae counts are 3,3,2,3,3.

Macrochaetotaxy of abdominal tergites II-VIII as follows: 2,4,4,6,6,4, 2.

There are numerous thin light setae scattered in amongst the setae of tergite

VII and VII particularly the posterior half of tergite VII. Paratergites with-

out macrosetae. Macrochaetotaxy of abdominal sternites II-VI as follows:

12,12,12,12,10,6-10. The 2 rows of setae on sternite VIII are not absolutely

distinct. Male genitalia shaped as in Fig. 83. Female unknown.

Measurements. - Pronotum length, 0.51-0.52; elytra length, 0.39-

0.40. Number measured, 2.

Holotype. - Male, no. 14618, Zaire, Katanga, Massif des Kundelungu,

15 Jan. 1949, Coll. N. Leleup, ex termite nest no. 7. In the collection of

the Musée Royal de l'Afrique Centrale, Tervuren. ,

Paratype. - Male, same data as holotype, (D. K.).

Notes. - The host termites are not identified.

Termitusa mundus (Bernhauer)

Figures 33, 62, 105

Glaphyrinus mundus Bernhauer and Paulian 1942: 375 (Field Museum of

Natural History, Chicago, (Cameroons, Edéa, in the upper galleries of a

termitary).

Termitusa mundus Seevers 1957: 212 (redescription and changed

generic name. )

Distinguished from T. escalarae by the chaetotaxy of the abdominal

sternite VIII, the elytra, and the shape of the spermatheca.

Color dark reddish brown throughout with the appendages and antennae

somewhat lighter than the rest of the body. Dorsal surface of the head,

pronotum, and elytra smooth and shiny with no secondary setae scattered

about. Macrochaetotaxy of pronotum as in Fig. 33; setae counts are 4,2,3,3.

Macrochaetotaxy of elytra as in Fig. 62; setae counts are 2,3,1,2. Macro-

Kistner: A revision of the Subtribe Termitusina 27

chaetotaxy of abdominal tergites II-VIII as follows: 2,2,4,4,4,4,2. The

setae on tergites VII and VIII are shorter and thinner than those on the rest

of the segments. Paratergites without macrosetae. Macrochaetotaxy of

abdominal stergites III-VIII as follows: 8,8,8,6,6,4. Male genitalia un-

known. Spermatheca shaped as in Fig. 105.

Measurements. - Pronotum length, 0.45; elytra length, 0.32. Number

measured, 1.

Material examined. - Female Holotype, Cameroon, Edéa, Coll. P.

Lepesme, R. Paulian, and A. Villiers, (F. M. N. H.).

Notes. - The host of this species is unknown. While the paper has 2

authors, the species are credited to Bernhauer in the text.

Termitusa muneris, n. Sp.

Figures 3, 4, 6-8, 10, 13, 15, 34, 89, 106

Most closely related to T. cloveri from which it is distinguished by the

chaetotaxy of the elytra and the shape of the male genitalia and spermatheca.

Color reddish brown throughout with the head and pronotum a little

darker than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny except for the presence of numerous setae which are

characteristic of the species. Macrochaetotaxy of the pronotum as in Fig.

34; counts are 5,3,1,3,3 with many smaller, thinner, and lighter colored

setae scattered between them. Macrochaetotaxy of elytra with counts as

2,3,1,3,3 with many smaller, thinner, and lighter setae scattered between

them especially on the inner half and the basal half. Macrochaetotaxy of

abdominal tergites II-VIII as follows: 2,4,4,4,4,4,2. The macrosetae on

segments VII and VIII are smaller, thinner, and somewhat lighter in color

than those on the rest of the segments. All segments with smaller, thinner,

and lighter setae on the surface between and anterior to the apical, darker

setae. Paratergites without macrosetae. Macrochaetotaxy of abdominal

sternites III-VIII as follows: 8,8,8,8,8,6. Segment VIII with an additional

brush of lighter and thinner setae on the posterior third. Macrochaetotaxy

of abdominal segment IX as follows: 3 each on the median pieces and 2 on

each lateral piece. Median lobe of male genitalia shaped as in Fig, 89.

Spermatheca shaped as in Fig. 106.

Measurements, - Pronotum length, 0.40-0.45; elytra length 0. 30-0. 38.

Number measured, 10.

Holotype. - Male, no. 14614, Zambia, Mbala (Abercorn), 15 May 1970,

ex nest T-329, Coll. J. R. Clover, D. H., A. C., A. H., and K. M. C.

Kistner, no. 2126. In the collection of D. H. Kistner.

Paratypes. - Zambia, Mbala (Abercorn): 27, same data as holotype,

(D. K.); 3 males, 1 female, 10 May 1970, ex nest T-317, same collectors,

no. 2088, (D. K.); 4, 14 May 1970, ex nest T-324, same collectors, no.

2111, (D. K.); 16, 16 May 1970, ex nest T-330, same collectors, no. 2130

(D. K.); 57, 16 May 1970, ex nest T-331, same collectors, no. 2132 (D. K.);

62, 16 May 1970, ex nest T-335, same collectors, no. 2145, (D. K.); 12,

Zambia, 10 mi. ex Mbala-Mpulungu, 22 May 1970, ex nest T-339, Coll. J.R.

Clover, ‘D,. Hs. aa Cy, Asc, and Bh OC. Kistwersmoe: 2177, (D.K.); 12

same data as previous, 22 May 1970, ex nest T-340, same collectors, no.

219, {Dy Ye

28 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Notes. - All of the host termites were determined to be Cubitermes

muneris Sjoestedt by R. M. C. Williams. Specimens of the hosts are in the

collection of the British Museum (Natural History), London, and D. Kistner.

Fig. 72-81. -- Median lobes of male genitalia: 72, Termitusa atlanticus; 73, T.

cameroni; 74, T. coatoni; 75, T. escalarae; 76, x. _hystrix, nt, a. boultoni: 78, i

cloveri; 79, T. ~delignei,; 80, kT hexasetosa; 81, T. Kkatangensis.

Kistner: A revision of the Subtribe Termitusina 29

Termitusa pasteelsi n. sp.

Figures 35, 63, 84, 107

Termitusa sp. Pasteels 1969: 18, gland; 5, behavior.

Distinguished from T. delignei, to which it is most closely related, by

the chaetotaxy of the pronotum, elytra, and abdomen, and the shape of the

male genitalia and female spermatheca.

Color dark reddish brown throughout with the appendages and antennae

somewhat lighter than the rest of the body. Dorsal surface of the head, pro-

notum, and elytra smooth and shiny with very few secondary setae between

the macrochaetae. Even the macrochaetae are relatively short in this spe-

ies. Macrochaetotaxy of pronotum as in Fig. 35; setae counts are 4,2,3,3.

Macrochaetotaxy of elytra as in Fig. 63; setae counts are 2,3,1,2. Macro-

chaetotaxy of abdominal tergites II-VI] as follows: 2,4,4,4,4,4,2. The

macrochaetae on abdominal tergites VII and VIII are shorter and thinner than

those on preceeding segments. Macrochaetae on tergite II and the middle

macrochaetae of tergite III are also short and thin. Many secondary setae

on tergites VII and VIII where they occupy the posterior third of tergite VII.

Paratergites without macrosetae. Macrochaetotaxy of abdominal sternites

III- VIII as follows: 8,8,6,6,6,6. Median lobe of male genitalia shaped

as in Fig. 84. Spermatheca shaped as in Fig. 107.

Measurements. - Pronotum length, 0.55-0.62; elytra length, 0. 41-

0.43. Number measured, 10.

Holotype. - Male, no. 14923, Gabon, Mekop (Ivindo), 14 km upstream

from Makokou, 8 May 1962, Coll. J. Deligne and J. M. Pasteels no. 1040.

In the collection of Deligne and Pasteels, Brussels.

Paratypes. - Gabon: 10, same data as holotype, (J. M. P., D. K.); 2,

between Makokou and Mekembo, 8 June 1962, Coll. J. M. Pasteels, no.

1069,, (D.. K.).

Notes. - The host termites were determined to be Cubitermes heghi

Sjoestedt by R. M. C. Williams.

Termitusa quadricollis Cameron

Figures 36, 64, 90, 108

Termitusa quadricollis Cameron 1952: 331 (Musée Royal de l'Afrique

Centrale, Tervuren) Belgian Congo: Massif des Kundelungu, no host;

Seevers 1957: 213 (redescription)

Distinguished from T. coatoni and T. hexasetosa by the chaetotaxy of

the pronotum and elytra and the shape of the male genitalia and spermatheca.

Color reddish brown throughout with the apex of the abdomen and the

legs and antenna somewhat lighter than the rest of the body. Dorsal surface

of the head, pronotum, and elytra smooth and shiny, without secondary setae

between the large setae on the head and pronotum. Macrochaetotaxy of pro-

notum as in Fig. 36; setae counts are 4,3,3,3. Macrochaetotaxy of elytra

as in Fig. 64; setae counts are 2,1,2,1,3. Macrochaetotaxy of abdominal

tergites II-VIII as follows: 2,4,6,6,6,6,2. There are many yellow setae

30 Contr. Amer. Ent. Enst., vol.10, no.4, 1974

between the macrochaetae between tergites VII and VIII. Macrochaetae on

tergites VII and VIII smaller and thinner than the macrochaetae on the other

abdominal tergites. Tergite VIII with a thick vestiture of fine yellow setae

on the apical half. Paratergites without any macrosetae. Macrochaetotaxy

of abdominal sternites III-VIII as follows: 12,12,12,10,10,6-8. Male gen-

italia shaped as in Fig. 90. Female spermatheca shaped as in Fig. 108.

Measurements. - Pronotum length, 0. 44-0. 46; elytra length, 0. 35-

0.37. Number measured, 5.

Material examined. - Type and 5 paratypes, Zaire, Katanga, Massif

des Kundelungu, 15 Jan. 1949, Coll. N. Leleup, with termite no. 7, (M. R.

A. C.); 3, same data as type series, (M. R. A. C., D. K.); 2 males, Zaire,

Katanga, Elisabethville, 27 Dec. 1948, Coll. N. Leleup, in small termite

nest, (M. R. A. C., D. K.).

Notes. - The host termites have not been identified.

Termitusa regina n. Sp.

Figures 37, 65, 109

Closely related to T. cloveri from which it is distinguished by the

shape of the spermatheca and the chaetotaxy of the elytra.

Color dark reddish brown throughout with the antenna and legs some-

what lighter than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with numerous fine yellow setae scattered even-

ly between the macrosetae. Macrochaetotaxy of pronotum as in Fig. 37;

setae counts are 5,3,1,3,3. Macrochaetotaxy of elytra as in Fig. 65; setae

counts are 3,3,2,2,2. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2,4,4,4,4,4,4. There are numerous fine setae in between the

macrosetae on the tergites. Macrosetae on tergites VII and VII smaller,

thinner, lighter in color than those on the rest of the tergites. No macro-

setae on the paratergites. Macrochaetotaxy of abdominal sternites III-VII

as follows: 12,10,10,10,8,6-8. Spermatheca shaped as in Fig. 109. Male

unknown.

Measurements. - Pronotum length, 0.49; elytra length, 0.39. Number

measured, 1.

Holotype. - Female, no. 14605, Zaire, Elisabethville, 27 Dec. 1948,

Coll. N. Leleup, ex termite nest no. 11. In the collection of the Musée

Royal de l'Afrique Centrale, Tervuren.

Notes. - The termite host has not been identified.

Termitusa ruellei n. sp.

Figures 38, 85, 110

Most closely related to T. sheasbyi from which it is distinguished

by the chaetotaxy of the pronotum, the shape of the spermatheca, and the

slightly smaller male genitalia. Closely related to T. lujae from which

it is distinguished by the chaetotaxy of the pronotum and elytra.

Color dark reddish brown throughout with the abdomen somewhat light-

er than the rest of the body; legs somewhat lighter than this. Dorsal sur-

Kistner: A revision of the Subtribe Termitusina 31

Fig. 82-93. -- Median lobes of male genitalia: 82, Termitusa leleupi; 83, T. mixaseta;

84, T. pasteelsi; 85,-T. ruellei; 86, T. sjoestedti; 877, T. ugandae; 88, T. lujae; 89, T.

muneris; 90, T. quadricollis; 91, T. sheasbyi; 92, T. superhirsutus; 93, T. williamsi.

32 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

face of the head, pronotum, and elytra smooth and shiny with a very dense

vestiture of setae. Macrochaetotaxy of pronotum as in Fig. 38 counts from

the midline are 4,3,3,3. The upright macrosetae are scarcely distinguish-

able from the very numerous lighter and more recumbent microsetae.

Macrochaetotaxy of elytra as in T. sheasbyi; counts are: 2,2,2,4. Macro-

chaetotaxy of abdominal tergites II-VIII as follows: 2,4,4,4,4,4,2. All of

the macrosetae are interspersed with numerous pale yellow setae. Setae on

tergites VII and VIII are shorter and thinner than those on the rest of the

tergites. Paratergites without macrosetae. Macrochaetotaxy of abdominal

sternites III-VIII as follows: 14,12,12,12,10,4-10. Median lobe of male

genitalia shaped as in Fig. 85. Spermatheca shaped as in Fig. 110.

Measurements. - Pronotum length, 0. 42-0. 46; elytra length, 0. 32-

0.34, Number measured, 9.

Holotype. - No. 14631, Republic of South Africa, Transvaal, Water-

berg District, Rankin's Pass, 22 Nov. 1971, ex nest TM25657, Coll. J. L.

Sheasby, T-851. In the National Collection of Insects, Pretoria.

Paratypes. - 8, same data as holotype, (N. C. I., D. K.).

Fig. 94-116. -- Spermathecae: 94, Termitusa atlanticus; 95, T. boultoni; 96, T. cloveri;

97, T. delgnei; 98, T. dybasi; 99, T. escalarae; 100, T. nexaseicas 19. 7. hystrix;

102, cr katangensis; 165,57, leleupi: 104, T. minor; 105, T. mundus; 106, ~'T. muneris;

107, T. pasteelsi, 108, T. -quadricollis; 109, T. regina; 110, T. ruellei: Lit, T.

shea sbyi: 1 ay ae sjoestedti: i: TES. es superhirsutus; be i ee ie ‘ugandai: Ti6, ‘vs wasmanni.

Kistner: A revision of the Subtribe Termitusina 33

Special plate I. -- Termitusa lightningae: A, Right elytron; B, Spermatheca; C, Pronotum,

T. malawiensis: D, Left elytron; E, Spermatheca; F, Pronotum. T. viegasi: G, Median

lobe of male genitalia; H, Spermatheca; 1, Elytron; J, Laceral lobe of male genitalia; K,

Pronotum.,

34 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Notes. - The host termites were identified as Cubitermes pretorianus

Silvestri by R. M. C. Williams and are in the National Isoptera Collection,

Pretoria, and British Museum of Natural History, London, The species is

named after Dr. J. E. Ruelle.

Termitusa sheasbyi n. sp.

Fioures 39, 66, 91, 111

Closely related to T. ruellei from which it is distinguished by the

chaetotaxy of the pronotum, the shape of the spermatheca, and the some-

what larger male genitalia. Also related to T. lujae from which it is

distinguished by all of the above characteristics.

Color dark reddish brown throughout with the abdomen somwhat light-

er than the rest of the body; legs somewhat lighter than this. Dorsal surface

of the head, pronotum, and elytra smooth and shiny beneath the very dense

vestiture of setae. Macrochaetotaxy of pronotum as in Fig. 39; counts

from the midline are: 4,3,3,4. Macrochaetotaxy of elytra as follows: 2,2,

3,4; as in Fig. 66. Macrochaetotaxy of abdominal tergites Il-VIII as follows:

2,4,4,4,4,4,2. All of the macrosetae are interspersed with numerous pale

yellow setae. Setae on tergites VII and VIII are shorter and thinner than

those on the rest of the tergites. Paratergites without macrosetae. Macro-

chaetotaxy of abdominal sternites TII-VIII as follows: 14,12,12,12,12,10,

All of the macrosetae are interspersed with numerous pale yellow setae.

Median lobe of male genitalia shaped as in Fig. 91. Spermatheca shaped

as in Fig. 111.

Measurements. - Pronotum length, 0. 44-0.47; elytra length, 0. 32-

0.35. Number measured, 10. :

Holotype. - Male no. 14231, Republic of South Africa, Transvaal, 16

km ex Sentrum-Matlabas, 20 Nov. 1971, ex nest TM25650, Coll. J. L.

Sheasby, T-849. In the National Collection of Insects, Pretoria.

Paratypes. - 63, same data as holotype, (N. C. I., D. K.); 18, same

locality, date, and collector, ex nest TM25651, no. T-849, (N. C. I., D. K.).

Notes. - Both nests of host termites were determined to be Cubitermes

transvaalensis Fuller by R. M. C. Williams and are in the National Isoptera

Collection, Pretoria and British Museum (Natural History), London. The

species is named after the collector, Mr. J. L. Sheasby.

Termitusa sjoestedti Wasmann

Figures 40, 67, 86, 112

Termitusa sjoestedti Wasmann 1905: 200 - Naturhistorisch Museum,

Maastrict, (Cameroons, Cap Debundscha, with Cubitermes fungifaber); 1911:

163, Pl. 5, Fig.9; Hegh 1922: Fig. 421; Seevers 1957:210, (redescription,

Cameroons, Longji)

Most closely related to T. pasteelsi, from which it is distinguished by

the chaetotaxy of the abdomen and the shape of the spermatheca.

Color reddish brown throughout with the elytra and head somewhat

Kistner: A revision of the Subtzibe Termitusina 35

darker than the pronotum and abdomen and the legs and antennae somewhat

lighter than that. Dorsal surface of the head, pronotum, and elytra smooth

and Shiny with chaetotaxy as follows: Pronotum with the chaetae counts of

4,2,3,3; arranged as in Fig. 40. Elytra with chaetae counts of 2,3,1,2;

arranged as in Fig. 62. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2,4,4,4,4,4,2. The very prominent macrosetae on tergites III-VI

are set in prominent elongated projections of the chiton which look like tub-

ercles. Macrosetae on tergites VII-VIII are smaller, thinner, and lighter

in color than those on the preceeding segments. Macrosetae on tergite VIII

are interspersed in a dense brush of long fine pale setae. In general the

macrosetae of the pronotum and elytra are not interspersed with numerous

fine yellow setae, although there seems to be an apical row of yellow setae

on each of the abdominal tergites. Paratergites without macrosetae. Macro-

chaetotaxy of abdominal sternites III-VUI as follows: 8,8,8,8,8,8. The

macrosetae on abdominal sternite III are thinner and shorter in length than

those on the subsequent segments. Median lobe of male genitalia shaped as

in Fig. 86. Spermatheca shaped as in Fig. 112.

Measurements. - Pronotum length, 0.49-0.52; elytra length, 0. 35-

0.36. Number measured, 2.

Material examined. - Type and cotype, Cameroons, Cap Debundscha,

with Cubitermes fungifaber, (N. H. M.); 4, Cameroons, Longji, (F. M. N.

Hi, )s

Termitusa superhirsutus n. sp.

Figures 41, 68, 92, 113

Most closely related to T. sjoestedti from which it is distinguished by

the chaetotaxy of the pronotum, elytra, and the shape of the spermatheca and

the male genitalia.

Color dark reddish brown throughout with the abdomen somewhat light-

er than the rest of the body, legs somewhat lighter than this. Dorsal sur-

face of the head, pronotum, and elytra smooth and shiny beneath the very

dense vestiture of setae. Macrochaetotaxy of pronotum as in Fig. 41; counts

are meaningless because microsetae are as large as macrosetae. Macro-

chaetotaxy of elytra as in Fig. 68; counts are meaningless because micro-

setae cannot be destinguished from macrosetae. Macrochaetotaxy of abdom-

inal tergites II-VIII as follows: 2,4,4,4,4,4,2. All of the macrosetae are

interspersed with numerous pale yellow setae. Setae on tergites VII and VIII

are shorter and thinner than those on the rest of the tergites. Paratergites

without macrosetae. Macrochaetotaxy of abdominal sternites II-VIII as

follows: 8,8,8,8,8,8. All of the macrosetae are interspersed with numerous

pale yellow setae. Median lobe of male genitalia shaped as in Fig. 92.

Spermatheca shaped as in Fig. 113.

Measurements. - Pronotum length, 0.50-0.51; elytra length, 0. 29-

0.30. Number measured, 2.

Holotype. - 1 female, no. 14594, Zambia, Mbala (Abercorn), 23 May

1970,..ex nest T-341, Coll, J.. R.: Clover,..D; H.,.and A,,C..Kistner. no.

2190. Inthe collection of D. H. Kistner.

Paratype. - 1 male, same data as holotype, (D. K.).

Notes. - The termite hosts were determined to be Cubitermes

36 Contr. Amer. Ent. Inst., vol. 10; no.4;, 1974

sankurensis Wasmann by R. M. C. Williams. Host termites are in the col-

lections of the British Museum (Natural History), London, and the Kistner

collection.

Termitusa ugandae n. sp.

Figures 42, 69, 87, 115

Closely related to T. hystrix from which it is distinguished by the

larger size, slight differences in the chaetotaxy of the pronotum and elytra

and the shape of the median lobe of the male genitalia and the spermatheca.

Color dark reddish brown throughout with the abdomen a little lighter

than the head, pronotum, and elytra and the legs and antennae still lighter

than that. Dorsal surface of the head, pronotum, elytra, and abdomen

smooth and shiny with macrochaetae arranged in distinctive patterns which

are not interspersed with fine yellow setae. Macrochaetotaxy of pronotum

as in Fig. 42; with setae counts of 5,3,2,3. Macrochaetotaxy of elytra as

shown in Fig. 69 with the following counts, 3,1,4,3,2, starting from the

suture. Macrochaetotaxy of abdominal tergites II-VIII as follows: 4,4, 4,4,

4,4,2. Macrosetae on tergite VII and VIII are much smaller and thinner than

the macrosetae on the other tergites. Macrosetae on tergites III-VI appear

to be set on small tubercles. Paratergites without macrosetae. Macro-

chaetotaxy of abdominal sternites III-VIII as follows: 8,8,8,6,6,6. The

setae on sternites III are smaller and thinner than the setae on the other

sternites. Macrochaetotaxy of abdominal segment IX as follows: 2 median

parts with 4 macrosetae each, 2 outers with 2 macrosetae each. Sperm-

atheca shaped as in Fig. 115. Median lobe of male genitalia shaped as in

rie or

Measurements. - Pronotum length, 0.59-0. 62; elytra length, 0. 39-

0.41. Number measured, 9.

| Holotype. - Male, no. 14612, Uganda, Zika Forest, 9 mi ex Entebbe-

Kampala, 11 July 1970, ex nest T-390, Coll. D. H., A. C., A. H., and

K. M. C. Kistner, no. 2377. In the collection of D. H. Kistner.

Paratypes. - 8, same data as holotype, (D. K.).

Notes. - The host termites were determined to be Cubitermes spec-

iosus (Sjoestedt) by R. M. C. Williams. Specimens of the host termites are

in the collection of the British Museum (Natural History), London, and

Kistner.

Termitusa viegasin. Sp.

FiguresI G-K

Most closely related to T. atlanticus from which it is distinguished by

the chaetotaxy of the elytra, the shape of the male genitalia, and the shape of

the spermatheca.

Color light reddish brown throughout with the elytra and head some-

what darker than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with numerous fine yellow setae interspersed

among the macrosetae. Macrochaetotaxy of pronotum as in Fig. I K; setae

Kistner: A revision of the Subtribe Termitusina 37

counts are 5, 3, 1, 3, 3. Macrochaetotaxy of elytra as in Fig. I I; setae

counts are 2, 3, 2, 2. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2, 4, 4, 4, 4, 4, 2. Macrosetae on tergites VII and VIII are

shorter and thinner than the rest of the tergites. There are numerous light

yellow setae on the apical third of abdominal segment VIII which is only

slightly indented on its posterior border. Paratergites without macrosetae.

Macrochaetotaxy of abdominal sternites III-VIII as follows: 8, 5, 8, 8, 2-6, 6;

the 2 on sternite VII are lateral, Median lobe of male genitalia shaped as in

Fig. I G. Lateral lobe of male genitalia shaped as in Fig. I J. Spermatheca

shaped as in Fig. I H.

Measurements, - Pronotum length, 0.36 -0. 40; elytra length, 0. 26-

9.39. Numer measured, 5.

Holotype. - Female, No. 15000, Angola, Rio Mussanguege, nr. Calon-

do, 26 December 1972, ex nest T-505, Coll. D. H. Kistner, S. A. Peles,

and R. J. Swift, No. 2774. In the collection of the Museo do Dundo.

Paratypes. - 4, same data as holotype, (M. D., D. K.).

Notes. - The host termites with which this species was associated

were determined to be Cubitermes sp. by Kistner. The host termites are

smaller than both C. pallidiceps and C. glebae. In fact, the heads were

even shorter than those of C. sankurensis. The host termites are in the col-

lection of the British Museum (Natural History), London, the Museo do Dun-

do, and the Kistner collection. The species is named for Sr. J orge Viegas,

Resident Director of DIAMANG, Dundo through whose courtesy we were ex-

tended many privileges.

Termitusa wasmanni n. sp.

Figures 43, 70, 116

Most closely related to T. boultoni from which it is distinguished by

the chaetotaxy of the pronotum and elytra and the shape of the female sperm-

atheca.

Color reddish brown throughout with the antenna and legs somewhat

lighter than the rest of the body. Dorsal surface of the head, pronotum,

and elytra smooth and shiny with many light yellow setae scattered in among

the darker and longer macrosetae. Macrochaetotaxy of pronotum as in Fig.

43; setae counts are 4,3,3,3. Macrochaetotaxy of elytra as in Fig. 70; setae

counts are 2,3,3,3. Macrochaetotaxy of abdominal tergites I[-VIII as fol-

lows: 2,4,6,6,6,6,2. There are many light yellow setae scattered in among

the darker setae of the abdomen. The macrosetae on segment VII and VIII

are lighter in color and thinner than the macrosetae on the other segments.

Paratergites without macrosetae. Macrochaetotaxy of abdominal sternites

INI-VIII as follows: 12,10,10,10,10,12. The row on sternite VIII appear to

be broken into 2 rows but these are not clearly defined. Spermatheca shaped

as in Fig. 116. Male genitalia unknown.

Measurements. - Pronotum length, 0.50; elytra length, 0.35. Number

measured, 1.

Holotype. - Female, 14598, Rhodesia, Mt. Selinda, 13 Apr. 1970, ex

nest T-292, Coll. J. R. Clover, D. H. and A. C. Kistner, and R. Boulton,

no.1994, In the collection of D. H. Kistner.

38 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Special plate II. -- Termitusa bandai: A, Right elytron; B, Pronotum; C, Spermatheca;

D, Lateral lobe of male genitalia; E, Median lobe of male genitalia. Thoracotermes

carrissoensis: F, Left elytron, G, Pronotum; H, Spermatheca; I, Median lobe of male ge-

nitalia; J, Lateral lobe of male genitalia. Termitusodes lativentris: K, Spermatheca.

Kistner: A revision of the Subtribe Termitusina 39

Notes. - The host termites were determined to be Cubitermes trunca-

tus (Holmgren) by R. M. C. Williams. Specimens of the host termites are

in the collections of the British Museum (Natural History), London, and

Kistner. The species is named after Rev. Eric Wasmann who did much of

the pioneering work in the study of termitophiles.

Termitusa williamsi n. sp

Figures 44, 71, 93

Most closely related to T. superhirsutus from which it is distinquished

by the chaetotaxy of the pronotum, elytra, and abdomen as well as the shape

of the male genitalia and spermatheca.

Color reddish brown throughout with the legs and antennae somewhat

lighter than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny with no thin yellow setae between the macrosetae.

Macrochaetotaxy of pronotum as in Fig. 44; with the following setae counts,

0,3,1,3,3, counting from the median row. Macrochaetotaxy of elytra as

shown in Fig. 71; setae counts are 2,3,3,3; the most anterior seta on the

lateral border and the 1 most frequently broken off. Macrochaetotaxy of

abdominal tergites II-VIII as follows: 2,4,4,4,4,4,2. Macrochaetae on ter-

gites VII and VIII are shorter and thinner than the macrochaetae on the other

tergites. No macrochaetae on the paratergites. Macrochaetotaxy of abdom-

inal sternites IJI-VIII as follows: 8,8,8,8,6,6. Setae on sternite III are

thinner and somewhat shorter than the setae on the rest of the sternites.

Spermatheca shaped as in T. muneris (Fig. 106). Median lobe of male gen-

italia shaped as in Fig. 93.

Measurements. - Pronotum length, 0. 38-0. 40; elytra length, 0. 28-

0.30. Number measured, 10.

Holotype. - Male, no. 14613, Zambia, Mbala (Abercorn), 10 May 1970,

ex nest T-316, Coll, J, R. Clover, D: H., A. C., A. H., and K. My, Cc,

Kistner, no. 2086. In the collection of D. H. Kistner.

Paratypes. - 68, same data as holotype, (D. K.).

Notes. - The host species were determined to be Noditermes sp. by

R. M. C. Williams. Specimens of the termites are in the British Museum

(Natural History), London, and Kistner. The species is named after Mr. R.

M. C. Williams who provided many of the host determinations in this paper

and has done such a splendid revision of the African species of the genus

Cubitermes.

Genus Thoracotusa new genus

Most closely related to Termitusa from which it is distinguished by the

position of the emarginate mesocoxal acetabulae which only proceed about

half way down to the posterior edge of the metasternum whereas in Termitusa

and Termitusodes these proceed almost all the way to the posterior border

of the metasternum (Fig. 136, 147, and 8).

Overall shape limuloid, as in Termitusa and Termitusodes. Head

40 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 117-124, -- Thoracotusa generic characters: T. alzadae: 117, abdominal segments

II-V; 118, mesoleg; 120, abdominal segment VII; 121, abdominal segment IX of male; 122,

proleg; 124, abdominal segment VII]. T. kymrae: 119, metaleg; 123, abdominal segment

VIO. he

capsule more or less oval in shape, hypognathus in life. Dorsal surface of

head without further sculpture, without a neck, and without a distinct nuchal

ridge. Eyes present, reniform, with many facets directed both anteriorly

and laterally. Gula fairly wide, mentum fused to gula and extremely short.

Submentum fairly long, shaped as in Fig. 129, not fused to the mentum. Max-

illary acetabulae not distinctly margined. Antennae 11-segmented, shaped

as in Fig. 125. Maxillae shaped as in Fig. 128, palpi 4-segmented; apical

segment attached to most apical part of proceeding segment. Mandibles only

slightly asymmetrical, shaped as in Fig. 126 and 127. Labium shaped as

in Fig. 129, palpi 2-segmented with the 1st segment about twice as long as

the 2nd segment. There is a thin seta on the tip of the 2nd segment which

may represent an extremely regressed 3rd segment but it does not even show

up on most slides. Ligula distinct. Labrum shaped as in Fig. 130, very

thinly sclerotized where it attaches to the clypeus.

Kistner: A revision of the Subtribe Termitusina 41

Pronotum shaped as in Fig. 142, with the sides reflexed ventrally as

shown. Pronotum without further significant sculpture; chaetotaxy variable

by species. Prosternum shaped as in Fig. 131, with relatively short art-

iculating apodemes, acarinate between the legs but with emarginate articul-

ating surfaces for the procoxa. Coxal cavities closed behind by the very

thinly sclerotized strap-like mesothoracic peritremes which are embedded

in membrane around the coxal cavity. Elytra shaped as in Fig. 143 and 144,

note the ventral reflection of the hypomera, such that these clamp onto the

lower side of the meso- and metasternum. Meso- and metanotum shaped as

in Fig. 132. Meso- and metasternum shaped as in Fig. 135 and 136; meso-

thoracic coxal cavities widely separated with distinctly margined acetabulae.

These margins only extend about half the length of the metasternum (Fig.

136). Wings present, of normal size, with the usual staphylinid venation.

The species is able to fly in life. Pro-, meso-, and metalegs shaped as in

Fig. 122, 118, and 119, respectively; tarsal formula 4-4-5.

Abdomen somewhat flattened dorsally, but otherwise subconical in

form. Abdominal segment I fused to the metanotum (Fig. 132). Abdominal

II represented by the tergite alone. Segments III-VI represented by a ter-

gite, 2 pairs of paratergites, and 1 sternite each; the inner and outer para-

tergites approximately equal in width. Segment VII represented by a tergite

and sternite and 1 long pair of paratergites; the inner paratergites are re-

duced to a small sclerite in each anterior corner. Segment VIII represented

by the tergite and sternite only. Segment IX 4-valved, shaped as in Fig.

134, The male segment IX has extremely long anteriorly directed apodemes

and it also has a small ventral sclerite probably representing a remnant of

sternite IX. Male genitalia bulbous, variable by species. Spermatheca of

female sclerotized, shape slightly variable by species.

Type species. - Thoracotusa alzadae n. sp.

KEY TO SPECIES OF THORACOTUSA

1. Abdominal tergites V-VI with 8 macrochaetae .....T. alzadae (p. 41 :

Abdominal tergites V-VI with 4 macrochaetae . inc te aaa ek pacers

2. Elytra with 2 setae on lateral edge......ccccecccecs T. kymrae - (p. 44 ;

Elytra with 4 setae on lateral edge.........¢. T. carrissoensis (p. 43 )

Thoracotusa alzadae n. sp.

Figures 117, 118, 120-122, 124-135, 137-139, 142, 143

Distinguished from T. kymrae by the shape of the spermatheca, the

chaetotaxy of the pronotum, and the chaetotaxy of the abdomen.

Color dark reddish brown throughout, with the head, pronotum, and

elytra a little darker than the abdomen. Dorsal surface of the head, pro-

notum, elytra and abdomen smooth and shiny with a distinctive chaetotaxy.

Chaetotaxy of the pronotum as shown in Fig. 142; counts are 4,3,3,4 or 5

(rarely). In slide preparations, these setae show up very well even if they

are broken off as the setae bases leave a small hole in the pronotum which

42 Cont. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 125-134, -- Thoracotusa generic characters: T. alzadae: 125, antennae; 126, right

mandible; 127, left mandible; 128, maxillae; 129, labium and submentum; 130, labrum;

131, prosternum; 132, meso-and metanotum with attached abdominal segment I; 133,

abdominal segment VII; 134, abdominal segment IX of female.

is easily visible. Some of those in the photograph were broken off. They

have been redrawn in from the study of other specimens. The specimen

shown exhibits the unusual situation where there is an odd seta put in the

wrong place on the right hand side of the pronotum. The left hand side of

the pronotum shows the more usual chaetotaxy. Chaetotaxy of elytra as

shown in Fig. 143; counts are 2,3,3,3. When these setae break off the holes

are not as immediately apparent in slide preparation. Macrochaetotaxy of

abdominal tergites II-VIII as follows: 2,6,8,8,8,4,0. The 2 on tergite II

are extremely small and on the lateral corners. The 2 middle setae of ter-

Kistner: A revision of the Subtribe Termitusina 43

gite III are much shorter and smaller and thinner than the other 4 which are

2 on each side. The 4 ontergite VII are extremely small and thin. On ter-

gite VIII while we have listed it as 0, it has many very fine, thin yellow

setae on its posterior third. Paratergites without setae. There is always

the odd specimen that has an odd seta out of place on the tergite. This setais

usually much thinner and smaller than the other setae whichare given in the

counts. Abdominal sternite III with 2 macrosetae at each lateral edge and

with an apical row of 10 very short, thin black setae between them. Sternite

IV with 10 macrosetae. Sternite V with 10 macrosetae. Sternite VI with 10

macrosetae, VII with 8, VIII with 8. Median and lateral lobes of male gen-

italia shaped as in Fig. 137 and 138. Spermatheca shaped as in Fig. 139.

Measurements. - Pronotum length, 0.51-0.54; elytra length, 0. 35-

0.38. Number measured, 10.

Holotype. - Male, no. 14536, Zambia, Mbala (Abercorn), 12 May 1970,

ex nest T-320, Coll. J. R. Clover, D. H., A. C., A. H., and K. M.-C.

Kistner, no. 2099, (D. K.).

Paratypes. - Zambia, Mbala (Abercorn): 358, same data as holotype,

(D. K,); 5, 15 May 1970; ex nest T-327, Coll. J. R. Clover, D. H:, A. C.,

A. H., and K. M. C. Kistner, no. 2119, (D. K.); 9, 23 May 1970, ex nest

T-342, Coll. J. R. Clover, D. H., A. C., A. H., and K. M. C. Kistner,

no. 2195, (D. K.). Zaire: 2, Katanga, Tshila-Tembo, 20 Feb. 1949, Coll.

N. Leleup, no. 1, (M. R. A. C., D. K.). Angola: 331, km 10, Estrada de

Chingufo, nr. Dundo, 25 November 1972, ex nest T-472, Coll. D. H. and

A. H. Kistner, and R. J. Swift, (M. D., D. K.); 3, Rio Mussanguege, nr.

Calondo, 24 December 1972, ex nest T-503, Coll. D. H. Kistner, S. A.

Peles, and R. J. Switt, (M. D., D: K.).

Notes. - The host termites from Zambia were identified as Thoraco-

termes lusingensis Harris by R. M. C. Williams, and are in the collection

of the British Museum (Natural History), London, and the Kistner collection.

Nest no. T-327 was a mixed colony containing T. lusingensis, Cubitermes

oblectatus Harris and Crenetermes sp. However, since T. alzadae was the

only species taken with pure colonies of T. lusingensis we do not regard the

other termites as probable hosts. Host termites from Angola were deter-

mined to be T. lusingensis by A. de Barros Machado (M. D., B. M. N. H.,

and D. K.).

Thoracotusa carrissoensis n. Sp.

Figures II F - I

Distinguished from T. kymrae by the chaetotaxy of the pronotum, the

chaetotaxy of the elytra, and the shape of the spermatheca. When males are

known of T. kymrae there will undoubtedly be additional characters there.

Color dark reddish brown throughout, approaching black. Dorsal sur-

face of the head, pronotum, elytra, and abdomen smooth and shiny with no

significant microsculpture. Pronotum with chaetotaxy as shown in Fig. II G;

counts are 4, 3, 1, 3, 3. Elytra with chaetotaxy as shown in Fig. II F;

counts are 1, 2, 3, 4. Macrochaetotaxy of abdominal tergites II-VIII as

follows: 2, 2, 4, 4, 4, 4, 2. The 2 setae on the abdominal tergite II are at

the lateral borders and are very small. The 2 setae on abdominal tergite III

are at the lateral borders. The 4 setae on tergite VII are very small and

4a Contr. Amer, Ent. Inst., vol.10, no.4, 1974

thin; and all of the many setae on tergite VIII are thin, 1 on each side stands

out as darker and these might be macrosetae. Paratergites without macro-

setae. Macrochaetotaxy of abdominal sternites III-VIII as follows: 8, 8, 8,

6, 6, 6. Spermatheca shaped as in Fig. II H. Median lobe of male genitalia

shaped as in Fig. II I. Lateral lobes of male genitalia shaped as in Fig. II J.

Measurements. - Pronotum length, 0. 48-0.52; elytra length, 0. 35-

0.38. Number measured, 10.

Holotype. - Male, number 14999, Angola, Pare Dr. Carrisso, nr.

Dundo, 5 November 1972, ex nest T-442, Coll. D. H. Kistner, and R. J.

Swift, No. 2545. In the collection of the Museo do Dundo.

Paratypes. - Angola, Parc Dr. Carrisso, nr. Dundo: 16, same data

as holotype, (D. K.); 6, 8 November 1972, ex nest T-449, Coll. D. H. Kist-

ner, andR. J. Swift, No. 2569, (M. D., D. K.); 8, 4 November 1972, ex

nest T-440, Coll. D. H. and A. H. Kistner, and R. J. Swift, No. 2537, (D.

K., M. D.); 23, 4 November 1972, ex nest T-441, Coll. D. H. and A. H.

Kistner and R. J. Swift, No. 2539, (M. D., D. K.); 2, 8 November 1972,

ex nest T-448, Coll. D. H. Kistner and R. J. Swift, No. 2568, (D. K.).

Notes. - All colonies of the host termites were determined to be

Thoracotermes macrothorax (Sjoestedt) by A. de Barros Machado, speci-

mens of which are in the collection of Museo do Dundo and the collection of

the author. These specimens of T. macrothorax have shorter heads than T.

macrothorax from Liberia.

Thoracotusa kymrae n. sp.

Figure 119, 123, 136, 140, 141, 144

Distinguished from T. alzadae by the chaetotaxy of the pronotum, the

chaetotaxy of the abdomen, and the shape of the spermatheca.

Color dark reddish brown throughout, approaching black. Dorsal sur-

face of the head, pronotum, elytra, and abdomen smooth and shiny with no

Significant microsculpture. Pronotum with chaetotaxy as shown in Fig. 141;

counts are 4,3,3,3. Elytra with chaetotaxy as shown in Fig. 143; counts are

1,2,3,1,2. Macrochaetotaxy of abdominal tergites II-VIII as follows: 2,4, 4,

4,4,4,4. The 2 setae on abdominal tergite II are at the lateral borders and

are very small. The 2 center setae on abdominal tergite III are small. The

4 setae on tergite VII are very small and thin and all of the many setae on

tergite VIII are thin, 1 on each side sort of stand out as darker and might be

macrosetae. Paratergites without macrosetae. Macrochaetotaxy of abdom-

inal sternites HI-VIII as follows: 8,8,8,6,6,6, Spermatheca shaped as in

Fig. 140. Male unknown.

Measurements. - Pronotum length, 0.52-0.55; elytra length, 0. 38-

0.41. Number Measured, 4.

Holotype. - Female, no. 14537, Zaire, Yangambi, 5 July 1960, ex

nest, Coll. D. H. and A. C. Kistner and R. Banfill, no. 304. In the collect-

ion of D. H. Kistner.

Paratypes. - Zaire, Yangambi: 1 female, same data as holotype, (D.

K.); 2 females, 6 July 1960, ex nest, Coll. D. H. and A. C. Kistner, and

R. Banfill, no. 312, (D. K.).

Notes. - Both colonies of host termites were determined to be Thora-

cotermes macrothorax (Sjoestedt) by Kumar Krishna, specimens of which

Kistner: A revision of the Subtribe Termitusina 45

Fig. 135-144. -- Thoracotusa alzadae: 135, meso- and metasternum; 137, median lobe of

male genitalia; 138, lateral lobe of male genitalia; 139, spermatheca; 142, pronotum; 143,

elytron. T. kymrae: 136, meso- and metasternum; 140, spermatheca; 141, pronotum;

144, elytron.

are in the collection of the American Museum of Natural History, New York.

Genus Termitusodes Pasteels

Termitusodes Pasteels 1967: 62, type species, T. lativentris Seevers

by original designation.

46 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

147 | 148

Fig. 145-149. -- Termitusodes generic characters: T. lativentris: 145, antennae with

attached part of head capsule; 146, labium and maxillae; 147, meso- and metasternum;

148, right mandible, 149, left mandible.

Kistner: A revision of the Subtribe Termitusina 47

Most closely related to Termitusa from which it is distinguished by the

shape of the abdomen and the shape of the labrum. There are other subtle

differences between the segment IX complex wherein there are fewer setae

in Termitusodes but these are not characters we usually associate with the

generic ranking.

Overall shape as in Termitusa. Head capsule oval in shape. Dorsal

surface of the head without further sculpture, the head is without a distinct

neck and without a nuchal ridge. Eyes present, reniform, with many facets

directed both anteriorly and laterally. Ventral surface of the head without

unusual sculpture. Gula with sides relatively straight, mentum extremely

short and fused to the gula. Mentum almost completely membranous but dis-

tinct from the submentum. Antennae shaped as in Fig. 145, 11-segmented,

with the terminal segment asymmetrical. Labrum elongate, shaped as in

Fig. 153. Mandibles slightly asymmetrical, very slender, shaped as in

Fig. 148 and 149. Maxillae shaped as in Fig. 146 with elongate lacinia and

galea; palpi 4-segmented, terminal segment immobilized to the side of the

3rd segment. Maxillary acetabulae not distinctly margined. Labium shaped

as in Fig. 146; palpi 2-segmented; ligula distinct.

Pronotum shaped as in Fig. 163 and 165. Chaetotaxy variable by

Species. Extent of the ventral reflection of all borders of the pronotum

about the same in all genera. Prosternum fairly large, shaped as in Fig.

165 with relatively short anterolateral articulation processes. Prosternum

evenly rounded between the legs. Prothoracic coxal cavities closed behind

by membrane in which are located the very thin straplike mesothoracic peri-

tremes. Elytra shaped as in Fig. 159 and 160, chaetotaxy variable by spec-

ies. Note that the lateral borders are deflected ventrally but are not folded

back on the ventral side as in Thoracotusa. Meso- and metanotum shaped as

in Fig. 150. Note the extreme reduction of abdominal tergite I which is

fused to the metanotum. Meso- and metasternum shaped as in Fig. 147.

Mesothoracic coxae widely separated with an acarinate, evenly rounded

mesosternal process produced between them. Mesocoxal cavities distinctly

marginate with the posterior margin proceeding all the way to the posterior

border of the metanotum. Wings present, of normal size and with the usual

staphylinid venation. Pro-, meso-, and metalegs shaped as in Fig. 155,

157, and 156 respectively; tarsal formula 4-4-5,

Abdomen subconical in form (Fig. 152), tapering from anterior to pos-

terior, such that the segment VII is 18.2% less wide than segment III. This

differs from Termitusa and Thoracotusa in that their abdomens taper 27.2%

from segment III to segment VII. Abdominal segment I fused to the meta-

notum. Abdominal segment II represented by the tergite alone. Abdominal

segments II-VI represented by a tergite, 2 pairs of paratergites, anda

sternite each. Abdominal segment VII represented by a tergite and sternite,

1 pair of paratergites which are full length and the inner paratergites which

are reduced to a small tab at the upper inner corner of each outer parater-

gite. Segment VIII represented by the tergite and sternite only. The para-

tergites of all segments are almost completely fused together. The line of

fusion shows on the paratergites on segments 3 and 4 but very faintly. The

other segments show no line of fusion whatsoever. One opening to the tergal

gland is present on each side of the midline on the anterior margin of tergite

Vil. Segment IX 4-valved (shaped as in Fig. 151) with the male having ex-

tremely long anterior apodemes and a very thin ventral piece which probably

represents sternite IX, Male genitalia bulbous, shape variable by species.

48 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Female spermatheca sclerotized, presumed variable by species. Lateral

lobe of male genitalia shaped as in Fig. 161 and 162.

Type species. - Termitusodes lativentris (Seevers)

KEY TO SPECIES OF TERMITUSODES

i Tereive Ti with. 4 macrosetac.. 6. 62s. 6 aise edie os Sew ee Sore 2

Tergite III with 6 macrosetae; pronotum with many secondary

setae Delween the. macrosetac..... .-2)4-. 6 6 6 eo: T. leleupi (p.52)

Doth) Vera be ik with 4 Mac rOSeia Gs. voces pe oe ese ess ek Peed sw es 2

Tercite 1V auth 6 macroseta 6... ns esc ee .T. summa (p.52)

3, (2) Pronotum with 4 rows of macrosetae; with 3 macrosetae

ie PO AAO Ie EGS ase). es eee oe ee hc ee T. congolesis (p. 48)

Pronotum with 4 rows of macrosetae; with 4 macrosetae

TM PC CO OWS cess ees ewe ol ak we ws cee ee Gade oe 4

4. (3) Paratergites narrow, paratergite IV 1/10 the width of the

corresponding tergite (ex Pasteels 1967). .... T. media (p.52 )

Paratergite wider, paratergite IV 1/12 the width of the

eorrespondine lengthy © 6 Se bb erai tess Coates T. lativentris (p.51 )

Termitusodes congolensis n. sp.

Figures 160, 162, 165, 166

Distinguished from all other species of Termitusodes by the chaeto-

taxy of the pronotum, chaetotaxy of the abdomen, and the shape of the male

genitalia.

Color light reddish brown throughout with the head and pronotum and

elytra a little darker than the rest of the body. Dorsal surface of the head,

pronotum, and elytra smooth and shiny with chaetotaxy as follows. Pro-

notum with chaetae as shown in Fig. 165; counts are 3,2,1,3, from the mid-

dle to the outside. Macrochaetotaxy of elytra as shown in Fig. 160, counts

are 2,3,2. Macrochaetotaxy of abdominal tergites II-VIII: 0,4,4, 4,4, 4,2.

All of the setae are extremely short though they are longer on tergite II

than all the rest. Paratergites without setae. Sternites with only apical

rows of 6 setae each; 2 near each lateral border and 2 in the middle of ster-

nites III-VI and 6 evenly spaced on the posterior border of sternites VII and

VIII. Median lobe of male genitalia shaped as in Fig. 166. Lateral lobe of

male genitalia shaped as in Fig. 162. Female unknown.

Measurements. - Pronotum length, 0.40; elytra length, 0.36. Number

measured, 1.

Holotype. - Male, no. 14307, partly on slides, Zaire, Yangambi, 4

July 1960, Berlese: nest 297, Coll. J. Decelle, no. 298. In the collection

of D. H. Kistner

Notes. - The host termites were determined to be Cubitermes fung-

ifaber (Sjoestedt) by Kumar Krishna. Specimens are in the collection of the

American Museum of Natural History, N. Y.

Kistner: A revision of the Subtribe Termitusina

Fig. 150-157. -- Termitusodes generic characters: T. lativentris: 150, meso- and

metanotum with attached abdominal segment I; 151, abdominal segments VIII and IX

of male; 152, abdominal segments II-VII; 153, labrum; 154, abdominal sternites VIII;

155, proleg; 156, metaleg; 157, mesoleg.

50 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Fig. 158-166. -- Termitusodes leleupi: 158, elytron; 163, pronotum. T. lativentris:

159, elytron; 161, lateral lobe of male genitalia; 164, pronotum. T. congolensis: 160,

elytron; 162, lateral lobe of male genitalia; 165, pronotum; 166, median lobe of male

genitalia.

Kistner: A revision of the Subtribe Termitusina 51

Termitusodes lativentris (Seevers)

Figures 145-157, 159, 161, 164, 167, II K

Termitusa lativentris Seevers 1957: 213, Fig. 32, Field Museum of

Natural History, Chicago, (French Congo, Brazzaville, 1 Apr. 1948, Coll.

A. E. Emerson with Noditermes cristifrons (Wasmann)).

Termitusodes lativentris, Pasteels 1967: 62, (Gabon, nr. Makokou,

with Pericapritermes magnificus Silvestri),

Distinguished from all other species by the chaetotaxy of the pronotum,

the abdomen, and the shape of the male genitalia.

Color reddish brown throughout with the abdomen and legs somewhat

lighter than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny with chaetotaxy which is species specific. Pronotum

with chaetotaxy as shown in Fig. 164; counts are 4,2,2,and 3 counting from

the middle of the pronotum outward, without short yellow setae interspersed.

Chaetotaxy of elytra as shown in Fig. 159, counts are 2,3,2. Elytra also

with a number of smaller yellow setae interspersed between these. Macro-

chaetotaxy of abdominal tergites II-VHI as follows: 0,4,4,4,4,4,2. Tergite

VIII also with many long yellow setae covering the apical third. Paratergites

without macrosetae. Sternite III with very few scattered setae other than

the apical row, sternites IV, V, VI, and VII with an apical row and another

row about the middle of the sternite with a variable number of setae. Ster-

nite VIII with an apical row and 2 anteapical setae near each lateral margin.

Median lobe of male genitalia shaped as in Fig. 167. Lateral lobe of male

genitalia shaped as in Fig. 161. Spermatheca shaped as in Fig. II K.

Measurements. - Pronotum length, 0.49-0.50; elytra length, 0. 29-

0.30. Number measured, 2.

Material examined. - The type series, Congo Republic, Brazzaville,

1 Apr] 1948, Coll. Alfred E. Emerson, (F. M. N. H., D. K.). Angola: 1,

Pare Dr. Carrisso, nr. Dundo, 7 November 1972, ex nest T-445, Coll. D.

H. Kistner, and R. J. Swift, No. 2559, (D. K.); 4, Mussangue Forest, nr.

Dundo, 16 November 1972, ex nest T-461, Coll. D. H. Kistner, and R. J.

Swift, No. 2603, (M. D., D. K.).

Fig. 167-168. -- Termitusodes lativentris: 167, median lobe of a male genitalia. T.

leleupi: 168, spermatheca.

92 Contr. Amer. Ent. Inst., vol.10, no.4, 1974

Notes. - The original type series of the species was taken from a

mixed colony containing termites determined as Noditermes cristifrons

(Wasmann) and Pericapritermes magnificus (Silvestri) according to Alfred

Emerson (personal communication). Specimens of the host termites are in

the Emerson collection of the American Museum of Natural History, New

York. The host termites from Angola were determined to be P. magnificus

(Silvestri) by Kistner, specimens of which are in the collections of the

British Museum (Natural History), London, and Kistner, The P. magnificus

were in mounds built by Cubitermes fungifaber (also determined by Kistner) ;

see behavior section for experiments with the hosts,

Termitusodes leleupi n. sp.

Figures 158, 163, 168

Distinguished from all other species by the chaetotaxy of the pronotum

and abdomen and the shape of the spermatheca.

Color reddish brown throughout with the abdomen and legs somewhat

lighter than the rest of the body. Dorsal surface of the head, pronotum, and

elytra smooth and shiny with a thick vestiture of secondary setae between the

rather short macrosetae. Macrochaetotaxy of pronotum as shown in Fig.

163; setae count is 1,1,2,3. Macrochaetotaxy of elytra as in Fig. 158; setae

count is 2,1,2,2. Macrochaetotaxy of abdominal tergites II-VHI as follows:

2,6,6,6,6,4,2. Tergite VIII also with a number of long yellow setae cover-

ing the apical third. Paratergites without macrosetae. Macrochaetotaxy of

abdominal sternites ITJ-VIII as follows: There seems to be an apical row of

a variable number of thin setae which are scarcely distinguishable from the

secondary setae which cover the posterior half of each sternite. Sperm-

atheca shaped as in Fig. 168. Male unknown.

Measurements. - Pronotum length, 0.42; elytra length, 0.35. Number

measured, 1.

Holotype. - Female, no. 14611, Zaire, Katanga, Massif des Kun-

dulungu, 1 Jan. 1949, Coll. N. Leleup, ex small termite nest. In the col-

lection of the Musée Royal de l'Afrique Centrale, Tervuren.

Notes. - The identity of the host termites is unknown.

Insertae sedis

The genus contains 2 other species; Termitusodes media Pasteels and

T. summa Pasteels. These 2 species are adequately described by Pasteels

(1967) and are only included in the key to species herein.

Kistner:

A revision of the Subtribe Termitusina 53

Fig. 169-170, --Cubitermes muneris nests: 169, typical undisturbed nest; 170, regener-

ated nest, 2 weeks after we chopped the top off of it.

54 Cont. Amer. Ent. Inst. vol.10, no.4, 1974

BEHAVIOR AND GLANDS

The behavior of Termitusa pasteelsi was first described by Pasteels

(1969, p. 5), who stated that while the species was very common in the nests

of Cubitermes heghi, it did not have a very close relationship with their

hosts. Very quick and agile, they clamber under and between the termites

who often ignor them. They often carry their abdomen in a vertical position.

While the termites sometimes move their mouthparts on the termitophile and

vice versa, these interactions do not last as long as they do in the Termito-

gastrina. Pasteels saw exchanges of food between Termitusa and their hosts

which had the character of robbery as the termites were disturbed and made

attempts to disengage the termitophile and even to chase it. Termitusa was

observed taking food from one termite in which case it went through the

begging behavior with its legs. In another exchange, the Termitusa took food

as it was passed from 1 worker to another.

My observations parallel those of Pasteels. We made detailed studies of

Termitusa atlanticus, T. boultoni, T. muneris, and T. viegasi. T. muneris

is found in the conspicuous termite nests of Cubitermes muneris, 2 varieties

of which are shown in Fig. 170 and 171. Many of the species can be quite

common in the nests and population counts of 60-80 are not uncommon. We,

too have observed that Termitusa frequently walks with its abdomen in a ver-

tical position (Fig. 173) and that they can walk over, under (Fig. 172), and be-

side of (Fig. 174) the termite soldiers and workers without any interference

whatsoever from soldiers, workers, or nymphs. These beetles would fre-

quently walk up to the side of a termite (Fig. 175) and walk along side of itfor

awhile, rubbing its abdomen against the thorax or abdomen of the termite be-

side it. When it raised the abdomen while walking alongside the termite, the

termite frequently responded by shifting its head toward the abdomen as if to

palpate it but it did not linger there as if to do some serious licking. We, too,

observed very quick food grabbing by the termitophile.

In spite of the apparent lack of antagonism between the termites and

Termitusa, the Termitusa did not thrive well in close contact with the ter-

mites. Our observations were made in Petri dishes and we normally crowd

many termitophiles and termites together to force more interactions. The

termites lived well in these chambers and we could keep small termite

samples alive for a week or more this way. However, the Termitusa seldom

lasted more than 1 or 2 days. Ido not know how to interpret this but I want

to point out that well-integrated forms suchas Termitomimus usually live as

long as the termites do under identical conditions.

Thoracotusa are even less integrated in the Thoracotermes nests than

Termitusa are in the Cubitermes nests. Thoracotusa alzadae carries its

abdomen down further on its back than Termitusa does. They stay away from

the active areas of termite activity and hide themselves in small folds of the

filter paper where they could get at the termites, but the termites were un-

likely to locate them. When a termite got close to Thoracotusa, it presented

the very tip of its abdomen. There would then be a momentary stop of the

termites' action which would allow the Thoracotusa to get away. There were

no movements of the termite's mouthparts to indicate that something was

exchanged. Thoracotusa also steals food from the termites. The behavior of

T. kymrae and T. carrissoensis was the same as that observed for T. alza-

dae.

Kistner: A revision of the Subtrive Termitusina 55

Fig. 171-172, --Termitusa muneris: 171, crawling beneath a soldier of Cubitermes

muneris

termite.

; 172, walking with its abdomen nearly vertical while being palpated by a worker

56 Cont. Amer. Ent. Inst. vol.10, no.4, 1974

Fig. 173-174. --Termitusa muneris: 173, walking to the side of a worker; 174, walking

alongside of the worker termite while rubbing its head and pronotum against the abdomen.

Kistner: A revision of the Subtribe Termitusina 57

The specimens of Termitusodes lativentris that we studied in Angola

were found in mixed colonies of Cubitermes fungifaber, Pericapritermes ma-

enificus, and another species of a still undetermined Termitid genus. While

these termites have separate galleries in nature, these were always mixed

when we dissected the nests. We then placed the Termitusodes in company

with each termite successively. Cubitermes and the unidentified genus were

antagonistic and attempted to bite the Termitusodes, while Pericapritermes

never snapped their mandibles at them nor attempted to bite them. This sup-

ports the contention of Pasteels that Pericapritermes is the true host of the

species.

Our studies of sections of Termitusa muneris, T. boultoni, and Thora-

cotusa alzadae showed no important differences from the findings of Pasteels

(1969). Like him, we found no unique glands. We found reduced tergal gland

reservoirs which are less reduced in Thoracotusa where the measurements

were: length, 105-110 4, width, 150-160, and depth of 72-86“. The post-

pleural glands of abdominal segment VIII are reduced in both genera. We

found type 1 cells scarcely distributed at the articulations of legs, antennae,

paratergites, and anterior edges of tergites and sternites but their position

from specimen to specimen is not constant. Maxillary glands are present in

both genera.

Fig. 175. --Termitusa cloveri, seta base on the posterior edge of abdominal tergite VI.

X 2400.

58 Cont. Amer. Ent. Inst. vol.10, no.4, 1974

ULTRASTRUCTURE OF SETAE

The macrosetae given in the counts on the abdomens sometimes have

very large sockets. To see the precise structure of these sockets and to see

if there were any glandular pores associated with them we placed a specimen

of Termitusa cloveri and Thoracotusa alzadae under the scanning electron

microscope. Fig. 176 shows the socket from 1 angle while Fig. 177 shows

a Similar setae from the reverse angle. No pores are apparent.

A characteristic which has not been used before but will probably be of

value at the generic level is the fringe of microtrichia along the posterior edge

of tergite VII. These are shown in Fig. 178, 179, and 180. They are inter-

esting microtrichia in that they apparently have no sockets and look like out-

growths of the edge itself. No other tergite has these nor does the sternite of

VII (Fig. 181). They are found in all genera of Aleocharinae which we have

examined and differ in the number of rows and in the form the microtrichia

take. They are barely visible in normal cleared slide preparations after you

know they should be there. They are probably to be found in many subfamilies

of Staphylinidae as Hammond (1972, Fig. 1) recently showed their presence on

Conosoma phloxum Olliff in addition to the peg-like setae of the males. They

have also been shown to be present in Microbledius and Bledius by Herman

(1972).

Fig. 176. --Thoracotusa alzadae, seta base on the posterior edge of abdominal tergite IV.

X 2400.

Kistner: A revision of the Subtribe Termitusina 59

Fig. 177, --Termitusa cloveri, junction of abdominal tergites VII and VIII showing fringing

microtrichia on the posterior edge of tergite VII. X 600.

INTERPRETATION

The behavior and gland distribution both support the interpretation that

the Termitusina are termitophiles which are adapted to the lives of the ter-

mites but are not integrated into the social life of the termites. They make

their living by stealing food from the termites in the adult stage. Where the

larvae are reared is still unknown. All species are host specific, so far as

we know, so it would appear that the colonial life of each species is some-

thing to which the beetles must somehow adapt in an evolutionary sense. Far

more likely is that the larvae, presently unknown, are even more strikingly

60 Cont. Amer. Ent. Inst. vol.10, no.4, 1974

dependent upon a given species of termite.

Fig. 178. --Termitusa cloveri, abdominal tergites VII and VIII emphasizing the setae of

tergite VOI. Note the presence of spatulate setae. This is the part that is presented to

the termite during close encounters.

Kistner: A revision of the Subtribe Termitusina 61

Fig. 179. --Thoracotusa alzadae, posterior edge of tergite VII, viewed from the side

emphasizing the microtrichia. X 2400.

62 Cont. Amer. Ent. Inst. vol.10, no.4, 1974

Fig. 180. --Thoracotusa alzadae, posterior edge of abdominal sternite VI; note absence

of microtrichia.

LITERATURE CITED

Bernhauer, Max and P. Paulian 1942. Coléoptéres Staphylinides du Camer-

oun (Mission P. Lepesme, R. Paulian, et A. Villiers, 1939). Rev.

Zool. Bot. Afr. 35 (1941): 344-375.

Blackwelder, Richard E. 1952. The generic names of the beetle family

Staphylinidae. U. S. Nat. Mus. Bull. 200: IV+484 p.

Cameron, Malcolm 1952. New Staphylinidae from the Belgian Congo. Rev.

Zool. Bot. Afr. 46: 323-332.

Fauvel, Albert 1906. Staphylinides de la Guinée Espagnole. Mém. Soc.

Espan. Hist. Nat. 1: 285-292.

Fenyes, Adalbert 1918. Coleoptera: Family. Staphylinidae, subfamily

Aleocharinae. Genera Insectorum 173A: 1-110.

Hammond, P. M. 1972. The microstructure, distribution, and possible

function of peg-like setae in male Coleoptera. Entomol. Scand. 3: 40-

04,

Kistner: A revision of the Subtribe Termitusina 63

Hegh, E. 1922, Les Termites, Bruxelles, 756 p.

Herman, Lee H. 1972. Revision of Bledius and related genera. PartI. The

aequatorialis, mandibularis, and semiferrugineus groups and two new

genera (Coleoptera, Staphylinidae, Oxytelinae). Bull. Amer. Mus. Nat.

Hist. 149(2): 111-254.

Kistner David H. 1968. A taxonomic revision of the termitophilous tribe

Termitopaedini, with notes on behavior, systematics, and post-

imaginal growth (Coleoptera: Staphylinidae). Misc. Publ. Entomol.

Soc. Amer. 6(3): 142-179.

1970a. Revision of the termitophilous tribe Termitohos-

pitini (Coleoptera: Staphylinidae) I. The genus Hetairotermes with a

numerical analysis of the relationships of Australian species. Pacific

Insects 12 (3): 467-484.

1970b. Taxonomic revision of the Termitophilous tribe

Termitohospitini (Coleoptera: Staphylinidae). II. The genus Ter-

mitoecia Bernhauer. Pacific Insects 12(3): 485-491.

Pasteels, Jacques M. 1967 Contribution a l'étude systématique des staphylins

termitophiles du Gabon (Coleoptera Staphylinidae, Aleocharinae). Rev.

Zool. Bot. Afr. 74(1-2): 43-69.

1969. Les glandes tégumentaire des staphylins ter-

mitophiles III. Les Aleocharinae des genres Termitopullus (Coro-

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Termitusa (Termitohospitini, Termitusina). Insectes Soc. 16(1): 1-26.

Reichensperger, August 1929. Neue Amisengaste und ein neuer Termiten-

east. (Pauss. Hist. Staph. ). Entomol. Blat. 25: 132-137.

Seevers, Charles H. 1957. A monograph on the termitophilous Staphylinidae

(Coleoptera). Feldiana: Zool. 40: 1-334.

Sokal, R. R. and P. H. A. Sneath 1963. Principles of numerical taxonomy.

Freeman and Co., San Francisco, 360 p.

Wasmann, Erich 1905. Termitusa. Nouveau genre d'Aleochariens termito-

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Soca

ent | es ev

4 “ ery aigeaand! abe

Fe PER x.

Contributions

of the

American Entomological Institute

Volume 10, Number 5, 1974

9 So

A REVISION OF THE GENUS

SYNECHES WALKER

(DIPTERA: EMPIDIDAE)

FOR NORTH AMERICA AND THE ANTILLES

D. Dee Wilder

TABLE OF CONTENTS

PSC a a ee ae ae | ee ee ee eG a 1

RCTNO WE EDGE EN a nw oe ee es ee oe pe ee Bes ee 1

INTRODUCTION. coca Scs ES ae oe Sak & ee ac, OE er ara Z

Ds Ca etice pre Auig iee Shed Sen By git ae 3

WER ee ee Re ee ae 3

KEY TO THE SPECIES OF SYNECHES FOR NORTH

AMERICA AND THE ANTILLES iis ok ihe ae Sh eee ee as 4

Pe er SST CN A AOL rata Stake hy a as Bek 8h ag eee gow hg 4a es wo 6

Syrecies Hyams COGUINGL. fo 80 2.2 FR eae 2 ee aE Pe b

OU ROR ONS A Fe ee pe ae We eles eGR ode Bald ae 8

Be OW aie Ea EER AE ens See 9

Sinecties on G1 peneis MEIARGEL ooo go hei ows PEs ea eae wees Laas 11

ROE ey i oe oe eens Said oS SEEN Gey Gane ee 12

ES FCCC SAY) ok id Ge ER wee sles Rd Che Oe Oe ee eB ee 12

DUCES VETS UR NYA ps ee 1 ek cope bo a aeRO oe ORD oa ee os 14

EG C8 BO eR ks len be Oe eal ee ale te Sa SAO 15

SVHECRES TRACTOCHACIONUE NT : SPiekiel ois pi 8 Fe eeeia sleet se oe ers ele ds ee ye 13

Ween eS CC OGGIE i. a ye ny OS ye pe wipe Rey oe cated oh 16

ee es es a ee iin hs deanna ae eae ee oe a 17

Syneches quaaraneularis Wheeler & Melander .. 6.6... eee ee 18

Boe es OG era Sas Ska A palscne mace bape Se, eels 19

Bee ek OCI Gao hee Alans te hes Sle ale Sa wk ce eS 10

OP CMS RO. CD ah ah ha PL A Wb EEO Ce ee ae a he 21

eo Fee as hw hehehe Mek ek die we oh Ga 22

BVOCs ViICUS COINGI Ti. SSDs) | GRR, ees Ce eet 23

BU COR CSUR AIR OL CCHISIZ ASS 6. 9. any ge SE rs eee ds ny ee ve WU 23

POT ee es ere tics a BiG hn bs Sieminca wa wted! vidas Ra leeaie acs ieie ale 25

FIGURES 2 oc) ds NCES RI tm gf Benea Se RR Rr Uo Ua De enn ean gs ae 26

PDE PO SCLIN EPIC NA ieee ee he simp ea wie Rig alo ee eke ee 31

A REVISION OF THE GENUS

SYNECHES WALKER

(DIPTERA: EMPIDIDAE)

FOR NORTH AMERICA AND THE ANTILLES!

D. Dee Wilder2

ABSTRACT

The genus Syneches Walker is reviewed. A dignostic key and

illustrations of wings, male genitalia and legs are presented for the

identification of the seventeen species of Syneches known from North

America and the Antilles. Twelve species are redescribed and the

following species described as new: S. frosti from Florida, S. pallidus from

Jamaica, S. macrochaetosus from Puerto Rico, S. vittatus from the south-

western United States, S. vineus vineus from Dominica, W. I., S. vineus

medinai from Puerto Rico, and S$. vineus jamaicensis from Jamaica.

Gloma phthia and S. testaceus are for the first time listed as synonyms of

S. thoracicus.

ACKNOWLEDGMENTS

I would like to thank the following institutions for loans of specimens: Academy of

Natural Sciences, Philadelphia (ANSP); California Academy of Sciences (CASC);

Carnegie Museum, Philadelphia (ICCM); Canadian National Collection of Insects

(CNCD); Cornell University Insect Collection (CUIC); Frost Entomological Museum,

Penn State University (PSUC); Field Museum of Natural History (FMNH); Florida

State Collection of Arthropods (FSCA); Illinois State Natural History Survey (INHS);

Iowa State University (ISUI); Los Angeles Co. Museum of Natural History (LACM);

Museum of Comparative Zoology, Harvard University (MCZC); Ohio State University

(OSUC); Snow Entomological Museum, University of Kansas (KSUC); Washington

State University (WSUC); University of Arizona (UAIC); University of Minnesota

(DEFW); Michigan State University (MSUC); Museum of Zoology, University of

IA thesis submitted to the Department of Entomology, eo & State University, in partial

fulfillment of the degree of Master of Science.

2Present address: Department of Entomology, Pennsylvania State University, University Park,

PA 16802.

2 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Michigan (UMMZ); University of Nebraska (DEUN); U.S. National Museum

(USNM); University of Wisconsin (UWEM); British Museum (Natural History)

(BMNH). |

My special thanks go to Dr. L. V. Knutson and Mr. George Steyskal, both of the

U.S. National Museum, for their invaluable assistance throughout this project. Mr.

Steyskal very kindly examined the types of rufus Loew, albonotatus Loew and pusillus

Loew and made some helpful notes on these specimens. I would also like to thank

Mr. Kenneth G. V. Smith of the British Museum (Natural History) for his prompt

attention to my problems with certain species. Special thanks also go to my major

professor, Dr. Roland L. Fischer for his many of hours of patient help.

INTRODUCTION

The genus Syneches, included in the subfamily Hybotinae, was erected by Walker

in 1852 for the North American species simplex Walker. The genus is now known to

be world wide in distribution, occurring in North America primarily east of the Rocky

Mountains. The group has been divided into subgenera, but according to Smith

(1969) the division does not hold up on a world basis. Syneches, sens. str. is the only

subgenus known in the Western Hemisphere.

Of the available specimens belonging to this genus, many could not be identified

because of poor keys and inadequate descriptions. There were also numerous

species which were undescribed. This study was undertaken with the objective of

producing a key and a set of descriptions to simplify the identification of all known

North American and West Indian Syneches.

Genus Syneches Walker

Damalis Fabricius, 1805, Syst. Antliat., p. 147. Type species, curvipes Fabricius

(Westwood, 1835 Ann. Soc. Ent. France 4: 684).

Syneches Walker, 1852, Ins. Saund. Dipt. 1: 165. Type species, simplex Walker.

Pterospilus Rondani, 1856, Dipt. Ital. 1: 152. Type species, Hybos muscarius

Fabricius.

Syneches, subgenus Harpamerus Bigot, 1859, Rev. Mag. Zool. 11: 306 (as genus).

Types species signatus Bigot.

Syneches, subgenus Epiceia Walker, 1860 Proc. Linn. Soc. Lond. 4: 149. Type

species, ferruginea Walker.

The genus Damalis Fabricius originally included four species; two of these were

empidids and two asilids. Wiedemann (1823) placed the two asilids in Damalis and

the two empidids in the genus Hybos. Westwood (1835) designated D. curvipes, one of

the empidids, the type species of the genus.

Hull (1962) proposed continuance of the name, Damalis, in the Asilidae, ‘‘because

the mass of literature has, with only one or two exceptions, continued to treat these

Wilder: North American Syneches 3

asilids under the name Damalis, and because students of the Empididae have

uniformly rejected the name Damalis.”’

Flies belonging to the genus Syneches are quite distinctive and can be separated

from all other empidids by the following characteristics:

Head hemispherical, occiput flattened; eyes meeting from antennae to

prominent ocellar triangle; antennae two-segmented, basal segments

fused, outer segment oval with apical or subapical hair-like arista.

Thorax very highly arched, scutellum margined with many hair-like

bristles.

Legs hairy; hind femora usually somewhat enlarged and spinose beneath,

always with several flexor bristles.

Wings with anal angle usually strongly rectangular, pedicel of second and

third veins long.

BIOLOGY

There is no recorded information on the immature stages of any hybotine empidids.

It is assumed, as with most other empidids, that the larvae are predaceous, living in

soil, leaf litter or rotting wood.

Schwartz (1891) has described the feeding behavior of an undetermined species of

Syneches. This predaceous behavior has since been observed by the writer and is

described in detail under thoracious (Say).

Species of the genus Syneches have been considered to be insects of grassland

areas. While this is undoubtedly true of some species such as simplex Walker, which

has been collected by sweeping in grassland areas, it does not hold for others. The

writer has collected thoracious, pusillus Loew and rufus Loew only in lush forest

vegetation. Ecological data on labels indicate that vineus n. sp. and macrochaetosus

n. sp. are also forest inhabiting species.

It might be added that Syneches as well as many other empidids are easily collected

at lights, to which they often come in very large numbers.

MORPHOLOGY

Earlier descriptions and keys to species of the genus Syneches have relied largely on

color as the main character. This is unfortunate since many of the species are

especially variable in color. There are several more stable characters which are useful

in separating the species; a short explanation of some of these follows.

The length of the hairlike arista is quite consistent within a species. The arista is

usually curved, and as a result is difficult to measure. For this reason, the arista

length is used in the descriptions only when necessary, and then only as the ratio of

arista to third antennal segment length. The apical antennal segment is called the

third antennal segment even though the two basal joints are fused.

4 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

On the occiput is a v-shaped depression in many species which extends from the

ocellar prominence to the cervix, herein termed the posterior ocellar depression.

Occasionally there is a polished dorsoventral band in the same area, termed the

posterior ocellar polished band.

A very consistent character is the extent of polished areas on the pleuron. When

colors or patterns of notal pollen are used, the insect should be examined from several

different angles.

Terminology for naming wing veins and cells is conventional (Figure 3).

The legs in most of the species considered in this paper are quite hairy. Therefore

when looking for certain bristles on the legs, one must be careful to distinguish

between spines, bristles and hairs, bristles being stiffer and longer than hairs.

The fore tibiae of all Syneches bear a depression on the anterodorsal basal third,

the fore tibial depression. Smith (1969) calls this a “‘gland-like leg structure.”’

All Syneches considered in this revision have long hairs on the abdomen. These

hairs are shorter dorsally and longer laterally. They are also longer on the anterior

and posterior margins of abdominal segments, and are absent on the posterior half of

the first abdominal segment.

Although the male genitalic characters in this group are somewhat variable, they

are still fairly diagnostic. Unfortunately, the genitalia must be removed and

macerated before the important features are sufficiently visible. The most constant

structures within each species are the apical margin of the epandrium and the general

shape of the hypandrium; however species with wide geographical ranges may show

considerable variation in these characters, especially at the extremes of their ranges.

KEY TO THE SPECIES OF SYNECHES FOR

NORTH AMERICA AND THE ANTILLES

1. Wings with dark spot beyond tip of second vein

Rice 5, ah eG 2 Se ON Ba oe simplex Walker

Wings without dark spot beyond tip of second vein. .............. 0.000. 2

2. Hind femora swollen compared with middle femora, with numerous

bristles below and usually with short, stout spines; body usually

MORO SE AG 4 We ky ee oe Ge ke a es ee Oe 10

Hind femora not swollen, similar to middle femora, at most 5 strong bristles

bellows Uody usually less than 4 mim. in leneth.--.. wk ee 3

3. VV LOS Wit 2 Wel Ge MiCG SOMA. 28) cu lcnenicoiauue arieeaans aan on 4

Wigs witient Wellecetinied SQW aii fous hgh epg a iy telat ales an om S

4. Thorax black; stigma grey, filling marginal cell

to apex walling Ghia meek ta pee. Sl es vineus vineus N. sp.

Thorax brown; stigma dark brown, quadrate, not filling marginal

Cell t@apexscis wots: teed isk ie quadrangularis Wheeler and Melander

5. Abdomen dusted brown, apical band on each segment grey pollinose; hind

femora dark brown; Arizona, New Mexico................. vittatus Nn. sp.

Abdomen brown, apical bands, if present, very narrow ..............0000e. 6

Wilder: North American Syneches 5

6. Thorax yellow to light brown; hind legs yellowish with dark femoral-

tibial joints, a dark band at middle of tibiae, and dark basitarsi and

GIStita rst (Rpt 0) ih a Oh etisalat debilis Coquillett

Thorax dark brown to black; hind legs not conspicuously banded ........... Zz

7. Second vein slightly curved at apex of first vein; ratio of marginal cell to

submarginal cell at apex of first vein approximately 1/1.5 ............. 8

Second vein straight at apex of first vein; ratio of marginal cell to

submarginal cell at apex of first vein approximately 1/1 ............... 10

8. Hind femora dark brown to black, Dominica .............. vineus vineus N. sp.

Hind femora-yellow to Teht bPOWi oo as to sal ere ye eee ae). 9

9. Hypandrium without median apical notch, female unknown,

PUCHIOURICG: 0. occ oie ee ee a ey Ones vineus medinai N. ssp.

Hypandrium with median apical notch,

FEE a PORE CU RENEE ny pee eee See vineus Jamaicensis N. SSp.

10. Arista less than five times length of third antennal segment; wings

datk brown Paguie 00) 66) oa ins oo). a acemdes pees eee ater Melander

Arista five or more times length of third antennal segment; wings light

brown,to hyaline (Figure 56) enwe. cgae. ee eee ees pusillus Loew

11>;Onenotopleural, one supraalar bristle present 2. ee PoC Tee 12

Two notopleural no supraaliar bristles present’. 6-2). 0 ipod Ve when nel a5

12. Stigma short; Guadtate or oval, distmet, /75%°) 2k Ore ORE Pd Oe 13

Stigma elongate, indistinct, nearly filling apex of marginal

COMING Sigs ek ee ae eee inversus Curran

13. First posterior cell markedly wider at wing margin than at

APLePIOMCLOSSYEeIU, ees NUL DANGCG | aga es oy sea eens 14

First posterior cell not widened at wing margin (Figure 62),

ICS OATS PIG) oe ee ee eure oor aa pallidus n. sp.

14. Thorax yellow with golden pollen; middle tibiae without long ventral

bristle. at asd) tid: Jamisied oe ie oe i ee ee ew bacis (Walker)

Thorax brown with brown pollen and silvery pollinose longitudinal

mesoscutal stripes; middle tibiae with long ventral bristle

at basal third (Figure.14), Puerto Rico... 455. 55 ee es macrochaetosus N. sp.

13. Stigma distinet, muchdarker than wing caullins Lear uh eee o 16

Stigma diffuse or net much darker than wing’ so. uy SARS Aw ek 18

16. Stigma quadrate; legs yellow, hind femora, tibiae and basitarsi

BFOWN apically. ee UIT. ase quadrangularis Wheeler and Melander

Stigma round or oval; color of legs variable, hind femora, tibiae and

tarsi never. distinetly darkeranieally <. cau. funn oe. ook. forces 17

17. Stigma elongate oval; ratio of marginal cell to submarginal cell at

apex of first vein. 13/1 or tess (Figure 32) ue. kv. ee arcs os thoracious (Say)

Stigma round to oval; ratio of marginal cell to submarginal cell at apex of

Tipst VEIN. 2.0/7 | OF MOIS LP IOUBO OD hla he ha winds ey A rufus Loew

6 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

18. Notum dark brown dusted with grey and brown stripes, legs

Pirech) your fee Cet ree Ore re ey COCO Tk ere we oe UG 20

Notum, if dark, usually without distinct grey and brown stripes;

Pee Mee dre POW. oe cee re eer See ee cee eee. 19

19. Hind. tibiae tuberculate ventrally; white pollinose humeral

tees OSC oe re ee et ee ee es longipennis Melander

Hind tibiae not tuberculate; white pollinose humeral patches

Olesen Te To cee oe ee eo ae albonotatus Loew

20. Sternopleura not polished; femoral-tibial joints dark brown (Figure 11);

fore tibial depression dark brown...) 2025. 20s 2 OS frosti n. sp.

Sternopleura partly polished; femoral tibial joints only slightly darker

than rest of leg; fore tibial depression yellow........... hyalinus Coquillett

Syneches simplex Walker

Figures 10, 30, 37, 59

Syneches simplex Walker, 1852, Ins. Saund., Dipt. 1: 16S.

Syneches punctipennis van der Wulp, 1867, Tijdschr. v. Ent. 10: 139.

Head reddish brown to black, grey pollinose; posterior ocellar depression rec-

tangular, posterior ocellar polished band present. Two long and two to four shorter

ocellar bristles. Antennae yellow to black, third segment usually black at base, yellow

at apex, arista bare.

Thorax yellow to black; northern specimens grey or brown pollinose, sometimes

with greyish pollinose mesoscutal stripes, southern specimens gold and silvery

pollinose, variously patterned. Pollen at base of dorsocentral bristles darker than that

of remainder of thorax. Integument of humeri of dark specimens yellow. Numerous,

more than six, humeral bristles; two subequal notopleurals; one long and two minute

postalars. Pleurae yellow to brown pollinose, except for small polished spot on

sternopleuron. Halteres brown to black, stem usually lighter.

Wings 3-4 mm. long, subhyaline to brown; stigma small, brown, located at end of

first vein; another small brown spot at end of second vein. Second vein straight below

end of first vein; third and fourth veins parallel beyond discal cell. Marginal cell less

than twice as wide as submarginal cell at end of first vein.

Legs yellow to black, tibiae and tarsi often lighter. Middle tibiae with one an-

terodorsal and two apical bristles. Hind femora without anterodorsal bristles, below

usually with six to seven stiff hair-like bristles. Hind tibiae with one prominent an-

terodorsal and numerous subapicals.

Abdomen yellow to black, brown to gold pollinose; abdominal hairs yellow to

brown. Abdomen of females frequently lighter than thorax. Male genitalia as figured

(Figures 30, 37).

Type locality and type: ‘‘United States.’’ Type deposited in British Museum, ap-

parently lost.

Wilder: North American Syneches 7

Distribution: This is perhaps the most commonly collected species of Syneches.

Over 750 specimens have been examined from eastern North America, ranging from

Nova Scotia to Florida west to Galveston, Texas and north to Elbow, Saskatchewan;

two specimens from Logan, Utah.

Flight period: January to December in southern parts of range, May to October in

northern parts of range.

Remarks: This species, the type species of the genus, bears no close relationship to

any other North American species. It is easily distinguished by the wing spot beyond

the end of the second vein.

Melander (1928) states that, according to Dr. S. W. Williston, simplex was

described in 1849 as Gloma phthia Walker. Mr. Kenneth G. V. Smith, who has

examined the type of phthia in the British Museum (Natural History), has stated

(1962, 1971) that the two insects are distinct. Examination of a wing photograph of

the type of phthia has shown it to be conspecific with thoracicus, described by Say in

1823. Based on this information, phthia, the name used in place of simplex since

1928, is now placed in synonomy with thoracicus.

This species has been collected by the writer by sweeping grassy areas near forests.

It also may be commonly collected at lights. Hundreds of specimens were collected

from 26 June to 1 September 1971 at greenhouse lights situated near a forest and a

large grassy field in Michigan. It is interesting to note that exhaustive sweeping and

searching yielded no specimens during the daylight hours at this locality. |

Syneches hyalinus Coquillett

Figures 27, 36, 63

Syneches hyalinus Coquillett, 1895, Proc. U.S. Natl. Museum 18: 437.

Head dark reddish brown, grey pollinose; posterior ocellar depression present. Two

long and six short pale ocellar bristles. Antennae yellow to rufous, third segment with

short grey pubescence; arista subdorsal, more than five times length of third antennal

segment, pale, finely pubescent except for extreme apex.

Thorax light to dark brown, greyish pollinose except for three longitudinal

mesoscutal stripes and a lateral spot with brown pollen. Pollen on prescutellar

triangle flattened, the spot contrasting with rest of notum when viewed from different

angles. Integument of postalar calli lighter than that of notum, pollinosity similar.

Three weak humeral bristles; two notopleurals, posterior slightly longer; three

postalars, one long and two minute. Pleurae grey pollinose except for large polished

spot on sternopleuron. Halteres brownish, stem lighter.

Wings 4-4.5 mm in length, hyaline to light brown; stigma indistinct, pale brown,

filling marginal cell from before end of first vein to end of second vein. Second vein

nearly straight beneath end of first vein; third and fourth veins parallel beyond discal

cell. Marginal cell less than twice as wide as submarginal cell at end of first vein.

Legs yellow to brownish yellow, yellow haired; coxae darker, dusted greyish. An-

terodorsal bristle of middle tibiae brown, shorter than or subequal to posterior

notopleural; apical posteroventral slightly shorter, light brown to yellow. Hind

femora enlarged with two or three anterodorsal bristles; below usually with two or

8 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

three, rarely four or five, spines on basal half. Hind tibiae with two subequal an-

terodorsal bristles, both longer than femoral bristles; one long, usually pale,

posterodorsal at level of second anterodorsal.

_ Abdomen brown, brown pollinose, except first segment with grey pollen, ab-

dominal hairs pale. Genitalia as figured (Figures 27, 36).

Type locality and type: Holotype.—°, Maryland, Type No. 3240 in U.S. National

Museum.

Records: MARYLAND. 1°, Riverdale, Prince Georges Co., 11 June 1916 (F.R.

Cole). 1c) Hyattsville, Prince Georges Co., 2 Aug. 1908 (A.L. Melander). 1¢{ College

Park, Prince Georges Co., 25 July 1937 (C. T. Greene). NEW YORK. 70%’, Babylon,

L.I., Suffolk Co., 13 July 1937 (F. S. Blanton); 20°, 26 July 1933 (Blanton and

Borders). 12, Jones Beach, L. I., Suffolk Co., Aug. 1963 (W. Sikora). NEW JERSEY.

1 of 229 Malaga, Gloucester Co., 4 Aug. 1907 (C. T. Greene). 6 JG’ Avalon, Cape May

Co., 22 July 1894 (C. W. Johnson). 1°, Westville, Gloucester Co., 21 July 1896 (C. W.

Johnson). 1%, Pemberton, Burlington Co., 11 July 1909 (C. T. Greene). 1%, Cape May

Co., 3 Aug. (C. W. Johnson). VIRGINIA. 20° Cape Henry, Princess Anne Co., 28

June 1939 (A. L. Melander). 1°, 1 mile S.E. of Lanoka, New Kent Co., 1-6 July 1956

(N. D. Richmond). NORTH CAROLINA. 1c Smithfield, Isle of Wight Co., 7 July

1946 (M. W. Wing). GEORGIA. 1% , Whitmarsh Island, Chatham Co., 28 June

1958, (C. F. Hartley). 1 o, Ft. Stewart Area, 25 June 1956 (C. F. Hartley). FLORIDA.

1 o', Ft. Myers, Lee Co., 27 Feb. 1918 (Blatchley).1 co, Torreya St. Park, Liberty Co.,

20 May 1966 (H. V. Weems, Jr.); 1 9, ea 1965. 2 oo", Gainesville, Alachua Co., 9

Sept. 1964 (R.P. Esser).

Remarks: This species is very closely related to S. frosti n. sp., from which it is

easily distinguished by yellow femoral-tibial joints, smaller upper eye facets and the

polished spot on the sternopleuron. Some specimens of longipennis Melander and

albonotatus Loew may resemble this species, but hyalinus may be separated from the

former by the non-tuberculate hind tibiae and from the latter by possessing yellow,

instead of brown legs.

Data on labels indicate that this species is attracted to lights.

Syneches frosti n. sp.

Figures 11, 19, 46, 65

Head dark reddish brown, grey pollinose; posterior ocellar depression present. Two

long and six short pale ocellar bristles present. Antennae rufous to brown, third

segment with long grey pubescence; arista apical, less than five times length of third

antennal segment, pale, densely pubescent except for dark base and apical fifth.

Thorax light brown to black, grey pollinose, except for three longitudinal

mesoscutal stripes and two lateral, nearly confluent spots with brown pollen. Pollen

on prescutellar triangle flattened, the spot contrasting with rest of notum when

viewed from different angles. Integument of postalar calli lighter than that of notum,

pollinosity similar. Three weak humeral bristles; two dark notopleurals, posterior

slightly longer; three postalars, one subequal to anterior notopleural and two minute.

Pleurae grey pollinose. Halteres brownish, stem lighter.

Wilder: North American Syneches 9

Wings 4.5-5 mm in length, hyaline; stigma, if visible, pale yellow. Second vein

nearly straight beneath end of first vein; third and fourth veins parallel beyond discal

cell. Marginal cell at end of first vein less than twice as wide as submarginal cell at

same point.

Legs yellow to brownish yellow, yellow haired; coxae darker, dusted greyish; apices

of femora, bases of tibiae and fore tibial depression dark brown. Anterodorsal bristle

of middle tibiae brown, subequal to or slightly longer than posterior notopleural;

apical posteroventral bristle slightly shorter, light brown to yellow. Hind femora

enlarged with three to five anterodorsal bristles, below with two to five spines on basal

half. Hind tibiae with two anterodorsal bristles, both subequal to or slightly longer

than longest femoral bristle; one long pale posterodorsal at level of second an-

terodorsal.

Abdomen brown, brown pollinose, first segment grey pollinose; abdominal hairs

pale. Genitalia as figured (Figures 19, 46).

Type locality and type: Holotype. —°, Florida, Archbold Biological Station,

Highlands Co., 10 Feb. 1962 (S. W. Frost). Type No. 72141 in U.S. National Museum.

Allotype. —¥ same data as holotype (USNM).

Paratypes. Same data as holotype with following dates: 300, 17, 10 Feb. 1962; 19,

11 Jan. 1962; 165 14 Jan. 1962; 10.17 Jan. 1962; 19 18 Jan. 1962; 1c’, 23 Jan. 1962;

10 26 Jan. 1962; 10% 2 Feb. 1962; Ic", 4 Feb. 1962; ', 5 Feb. 1962; 40; 19, 8 Feb.

1962; 20%, 24 Feb. 1963; 10%, 27 Feb. 1963; 200} 4 March 1963; 1c, 6 March 1962; 1g

10 March 1963; 1c% 11 March 1963; 1c; 12 March 1963; 1cf 13 March 1963; 1%, 23

March 1962; 10", 19, 24 March 1963; 299 30 March 1963; 60% 19, 1 April 1963; 14 6

April 1963; 1%, 8 April 1962; 1c’, 13 April 1963. 1c’, 1%, Archbold Biological Station,

10 May 1965 (L. L. Pechumen). 1 9 Archbold Biological Station, 23 April 1967 (B. V.

Peterson). 19, Gainesville, Alachua Co., 28 May 1959 (H. V. Weems, Jr.). Paratype

distribution: 2c", 2 9(CNCI); 2 co 29¢ (MSUC); 2¢'c' 299 (FSCA); 1c’, 1° (CUICQ);

1c", 12 (BMNH); remainder (USNM).

Remarks: This species is closely related to hyalinus Coquillett, but is distinct in

having upper eye facets larger, sternopleuron entirely pollinose and apices of femora

and bases of tibiae dark. Some specimens may resemble longipennis and albonotatus,

but differ from the former by the non-tuberculate hind tibiae and from the latter by

virtue of the largely yellow legs.

Syneches frosti n. sp. has been collected exclusively at lights.

This species is named in honor of Dr. Stuart W. Frost, who collected the majority of

the speciemns examined, and whose enthusiasm has been a continuing inspiration to

the writer.

Syneches albonotatus Loew

Figutes 13,33)57

Syneches albonotatus Loew, 1862, Berl. Ent. Zeitsch. 6: 195.

Head dark reddish brown, grey pollinose; posterior ocellar depression present. Two

long and two (rarely four) shorter ocellar bristles. Antennae rufous to brown, third

segment with short grey pubescence; arista subdorsal, five times the length of third

10 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

antennal segment or longer, brownish with fine golden pubescence except for extreme

tip.

Thorax dark brown to black, rarely light brown, greyish pollinose, usually with two

to three longitudinal mesoscutal stripes and a large side spot with brown pollen.

Humeri, two large posthumeral patches and prescutellar triangle with pollen flat-

tened, these areas contrasting with rest of notum when viewed from different angles.

Integument of postalar calli lighter than that of notum, pollinosity similar. Four,

rarely three, weak humeral bristles; two dark notopleurals, posterior longer; three

postalars, one long and two minute, Pleurae grey pollinose except for polished spot on

sternopleuron. Halteres brown to dark brown, stem slightly lighter.

Wings 4.5-S mm long, brown; stigma varying from pale to dark brown, with in-

distinct posterior borders, nearly filling apex of marginal cell. Second vein nearly

straight beneath end of first vein; third and fourth veins parallel beyond end of discal

cell. Marginal cell acute at distal end; slightly wider at end of first vein than sub-

marginal cell at same point.

Legs dark reddish brown to nearly black, rarely light brown; brown haired; tibiae

and tarsi occasionally light brown, coxae dusted greyish. Anterodorsal bristle of

middle tibiae brown to black, variable in length; posteroventral dark, also variable.

Hind femora enlarged with two to three anterodorsal bristles on apical half, below

with abundant long spines, usually six or more on basal half. Hind tibiae with two

anterodorsal bristles both shorter than longer femoral bristles; subapical

posterodorsal longer than anterodorsals.

Abdomen brown to black, brown pollinose, first segment grey pollinose; abdominal

hairs pale. Genitalia as figured (Figures 18, 33).

Type locality and type: Holotype. —°, District of Colombia, Type No. 1112 in

Museum of Comparative Zoology, Harvard University.

Records: NEW JERSEY. 1°, Basking Ridge, 9 June 1936 (W. R. Douglas).

PENNSYLVANIA. 1°, Westmoreland Co., July. VIRGINIA. 19, Falls Church,

Fairfax Co., 24 June 1912 (C. T. Greene). GEORGIA. 1c’, 19, Austell, Cob Co., 27

Aug. 1910 (C. W. Johnson). FLORIDA. 1c’, Gainesville, Alachua Co., 28 May 1959

(H. V. Weems, Jr.) OKLAHOMA. 60% 19, Pawhuska, Pawnee Co., 10 June 1960 (G.

C. Eickwort and W. T. VanVelzen). TEXAS. 1c’, 2.5 miles S. W. of Forestburg,

Montague Co. 28 April 1951, (CNMH Paleont. Exp., Patterson leg.). 1c Dallas,

Dallas Co., 3 May (Osten Sacken).

~ Remarks: This species can be separated from hyalinus and frosti by the dark hind

femora, and from longipennis by the nontuberculate hind tibiae. It is also

distinguishable by the white pollinose humeral patches, which can be seen only when

the insect is held at certain angles.

Despite the disjunct distribution, these specimens unquestionably belong to the

same species. The specimens from Texas and Oklahoma are darker in color than the

others, and have a slightly different shape to the hypandrium.

This species has been collected at lights as well as on Elm and the blossoms of

Rhus.

Wilder: North American Syneches 11

Syneches longipennis Melander

Figures 20, 45, 49

Syneches longipennis Melander, 1902, Am. Ent. Soc. Trans. 28: 346.

Head brown to dark reddish brown, grey pollinose; posterior ocellar depression

present. Two long and two short ocellar bristles. Antennae yellow to brown with short

greyish pubescence; arista subdorsal, usually six times length of third antennal

segment, pubescent, sometimes finely plumose, except for base and hair-like apex.

Thorax light brown to dark reddish brown, pale pollinose, some specimens with

three to four golden pollinose mesoscutal stripes, others with a dark median in-

tegumental stripe. Pollen on prescutellar triangle flattened, the spot contrasting with

remainder of notum when viewed from different angles. Three humeral bristles; two

notopleurals, posterior markedly longer; four postalars, one long and three very

small. Pleuron pale pollinose except for large polished area on sternopleuron which

extends to ventral edge of mesopleuron. Halteres brownish, stem usually lighter.

Wings 6 mm. in length, light to dark brown; stigma usually indistinct, pale brown,

filling marginal cell from before end of first vein to end of second vein. Second vein

but slightly curved beneath end of first vein; third and fourth veins parallel beyond

discal cell. Marginal cell at end of first vein slightly wider than submarginal cell at

same point.

Legs brownish yellow to brown, coxae pale pollinose. Anterodorsal bristle of middle

tibiae brown to black and longer than notopleurals; apical posteroventral subequal to

posterior notopleural, brown. Hind femora of males greatly swollen, usually as wide

as abdomen; of females, not quite as swollen; with four to five strong anterodorsal

bristles; below with numerous long, stout spines, especially in males. Hind tibiae with

small tubercles on the ventral surface; two subequal anterodorsal bristles, both

considerably shorter and more slender than longer femoral bristles; and

posterodorsal slightly longer than anterodorsals.

Abdomen light to dark brown, pollen varying in color from golden to grey; ab-

dominal hairs pale to dark. Genitalia as figured (Figures 20, 45).

Type locality and type: Holotype. —c’, North Carolina, Hertford Co., 9 June 1895

(C. W. Johnson). Type in U.S. National Museum.

Records: GEORGIA. 1c{ Burton, 21 May 1911, 1800 ft. alt. (D. C. Bradley).

ALABAMA. 1c’, Thomasville, Clarke Co., 20 April 1910 (W. D. Pierce). TEN-

NESSEE. 1c Allardt, Fentress Co., c. 1200 ft., 13 June 1924 (T. H. Hubbell).

MISSISSIPPI. 1c’ 19, Tishomingo State Park, Tishomingo Co. 31 May 1958 (T. E.

Moore).

Remarks: This species can be distinguished from all other North American

Syneches by the presence of small tubercles on the ventral surface of the hind tibiae

and by the greatly swollen, spinose hind femora.

Although most specimens have three humeral bristles, the holotype appears to have

only two. This species exhibits a large range of variation in shape of the hypandrium

and epandrium.

12 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Syneches rufus Loew

Figures 8, 17, 34, 52

Syneches rufus Loew, 1861, Berl. Ent. Zeitsch. 5: 320.

Head light to dark reddish brown, grey pollinose; posterior ocellar depression

present. Two long and two shorter ocellar bristles. Antennae yellow to reddish brown,

arista yellow, pubescent except for apex.

Thorax yellow to dark brown, yellowish to brown pollinose, longitudinal mesoscutal

integumental stripe sometimes present. Three to five small humeral bristles; two,

rarely three, notopleurals of varying lengths; one long and one or two minute

postalars. Pleurae yellow to brown pollinose except for polished sternopleuron and

mesopleuron. Halteres yellow to brown, stem sometimes lighter.

Wings 4.5-5 mm in length, hyaline to light brown; stigma round to oval, pale yellow

to brown, never filling apex of marginal cell. Second vein curved below end of first -

vein; third and fourth veins nearly parallel beyond discal cell. Marginal cell greater

than 2.5 times as wide as submarginal cell at end of first vein.

Legs yellow to dark brown, pro- and mesotibiae and all tarsi usually lighter than

remainder of leg. Middle tibiae with at least one anterodorsal and two subapical

bristles. Hind femora with two anterodorsal bristles, below usually with four or fewer

prominent spines on basal half. Hind tibiae with two anterodorsal bristles and one

longer subapical anteroventral.

Abdomen yellow to black, often darker than thorax, yellow to brown pollinose;

abdominal hairs yellow to brown. Genitalia as figured (Figures 17, 34).

Type locality and type: New York. Type No. 1114 in Museum of Comparative

Zoology, Harvard University. Two female specimens examined by Mr. George

Steyskal.

Distribution: Ontario, Quebec, Minnesota, Wisconsin, Michigan, Ohio, Penn-

sylvania, Massachusetts, New York, Maine, and Northern [linois and Virginia.

Flight period: 25 June to 16 October.

Remarks: This species is closely related to thoracicus, but may be separated by the

wider marginal cell, more rounded stigma and fewer ventral bristles on the basal half

of the hind femora.

This species has been taken in forest vegetation, in a tamarack bog, and at lights.

185 specimens were examined.

Syneches thoracicus (Say)

Figures 1, 4, 9, 51

Hybos thoracicus Say, 1823, Jour. Nat. Sci. Phil. 3: 76.

Gloma phthia Walker, 1849, List Dipt. Brit. Mus. 3: 492. (NEW SYNONYM)

Syneches testaceus Melander, 1928, Genera Insectorum. Fasc. 185: 44. (NEW

SYNONYM)

Head reddish brown to black, grey pollinose; posterior ocellar depression present.

Two long and two or four shorter ocellar bristles. Antennae yellow to brown, arista

densely pubescent except for apex.

Wilder: North American Syneches 13

Thorax yellow to black; lighter specimens may have one to three dark longitudinal

mesoscutal integumental stripes; uniformly golden to brown pollinose. Three, rarely

five, small humeral bristles; two notopleurals, posterior longer; one long and two

minute postalars. Pleurae gold to brown pollinose except for large polished spot on

sternopleuron and mesopleuron. Halteres yellow to black, stem sometimes lighter.

Wings 4-5 mm. in length, subhyaline to dark grey or brown, stigma elongate oval,

pale yellow to black, sometimes nearly filling apex of marginal cell. Second vein

gently curved before end of first vein; third and fourth veins nearly parallel beyond

discal cell. Marginal cell one to two times as wide as submarginal cell at end of first

vein.

Legs from yellow to dark brown; pro- and mesotibiae and all tarsi usually lighter

than remainder of leg. Middle tibiae with at least one long anterodorsal and two

subapical bristles. Hind femora with two to four anterodorsal bristles, below usually

with more than five prominent spines on basal half. Hind tibiae with two anterodorsal

bristles and one or two subapical anteroventrals.

Abdomen yellowish to black, often darker than thorax, brown to gold pollinose;

abdominal hairs yellow to brown. Genitalia as figured (Figures 1, 4).

Type locality’ and type: Pennsylvania. Holotype lost.

Distribution: Ontario, Quebec and Maine south to Arkansas, Alabama and

Georgia and west to Kansas and Nebraska. Flight period: 14 May to 14 August.

Remarks: This species may be distinguished from rufus, the most closely related

species, by the presence of an elongate oval stigma, a narrower marginal cell, and

more ventral spines on the basal half of the hind femora.

643 specimens of this species were examined.

The name phthia has, since 1928, been used incorrectly in place of simplex.

Examination of a photograph of the wing of the type of phthia has proven that this

insect is actually thoracicus (see detailed note under simplex).

S. thoracicus is an extremely variable species, especially in wing characters which

are usually very stable in species of Syneches (stigma, shape and color, shape of

marginal cell). Melander described testaceus from a single female, using characters

such as the stigma and the marginal cell. Examination of the holotype rom Kansas

City, Missouri, and several male specimens from the same general area which fit the

description has shown testaceus to be within the range of variation of thoracicus.

The feeding habits of this species were observed in Stewart Co., Tennessee, 9-10

June, 1971, by the writer. They are very similar to the predaceous behavior of an

unidentified species of this genus described by Schwartz in 1891.

Numerous S. thoracicus were observed resting on the undersides of leaves during

the day in the forest. Towards dusk the insects began to fly singly in rectangular

patterns near clumps of vegetation at varying heights. Occasionally one of the flies

was observed to dart into vegetation and capture a minute insect with its powerful

hind legs. It then flew, oriented in a vertical position, prey still trapped in the hind

legs, to a nearby leaf and without alighting caught on to its edge with the tarsal claws

of one of the fore legs. The fly, still suspended by one leg, then manipulated the prey

with the remaining legs, inserted its mouthparts and sucked out the fluids.

When disturbed or when finished with the prey (usually one to three minutes), the

fly dropped the insect and again flew to rest on the underside of a leaf. Both sexes

14 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

were observed feeding in this manner, but despite the density of the insects, no mating

behavior was observed. Schwartz also reported a high density of the species he ob-

served.

Schwartz makes the statement, ““The long and very sharp claws are well adapted

for taking a firm hold on the surface of the leaf, and the long pulvilli assist in the

operation by pressing on the edge or the underside of the leaf.’’ One might assume

that since all Syneches have long curved tarsal claws and large pulvilli (Figure 12),

they all have this type of feeding behavior.

Syneches inversus Curran

Figures 3, 28, 48

Syneches inversus Curran, 1928, Ent. Bull. Dept. Agric. Jamaica 4: 35.

Head blackish, grey pollinose; posterior ocellar depression present. Two long and

four short ocellar bristles.

Thorax brown pollinose on the anterior two thirds, except the broad sides,

elsewhere above and on the pleura, greyish; black from dorsal view. Pleurae grey

pollinose, posterior half of sternopleura and mesopleura polished below. Four weak

humeral bristles; one notopleural, one supraalar; one long and three short postalars.

Halteres yellow.

Wings 4.5 mm. in length, light brown, darker apically, stigma elongate, dark

brown. Second vein not curved below end of first vein; third and fourth veins parallel

beyond discal cell. Marginal cell at end of first vein subequal to submarginal cell at

same point. )

Legs: Front and middle legs yellowish, coxae and trochanters brown; hind legs

reddish. Anterior tibiae with several bristles above in two rows; middle tibiae with a

long posterodorsal bristle at the basal fourth and three to four ventral bristles on the

apical half; hind femora with two long, strong anteroventral bristles near the middle

and several shorter bristles toward the base; on the apical half below with about eight

short, stout apically inclined spines; posterior tarsi with a row of four anteroventral

bristles on the swollen first segment; hind tibiae enlarged apically.

Abdomen brown, first segment slightly darker, pale pollinose; abdominal hairs

pale. Genitalia as figured (Figures 28, 48).

Type locality and type: Jamaica, Cinchona. Deposited in British Museum (Natural

History).

Records: JAMAICA.1°%,Cinchona, 25 August 1923. 20%} 6 miles N. Mavis Bank, 2

June 1966 (Byrd K. Dozier).

Remarks: This species can be distinguished from all other West Indian Syneches

by the presence of an elongate, indistinct stigma and a swollen, spinose basitarsus.

One of the two specimens of this species examined was compared with the holotype

in the British Museum (Natural History) by Mr. Kenneth G. V. Smith.

Wilder: North American Syneches 15

Syneches pallidus n. sp.

Figures 13, 22, 47, 62

Head black, grey pollinose; posterior ocellar depression present. Two long and four

short ocellar bristles. Antennae yellow, third segment with fine golden pubescence;

arista more than six times length of third antennal segment, dark, finely pubescent

except for apex.

Thorax with anterior half of notum brown with three longitudinal yellow in-

tegumental stripes; posterior half yellow to white, very pale pollen throughout. Six

weak humeral bristles; one notopleural, one supraalar; one long and several short

postalars. Pleurae pale pollinose except for large polished area of sternopleuron and

mesopleuron. Halteres pale brown, stem whitish.

Wings 5-5.5 mm. long, pale brown, stigma dark brown, rectangular. Second vein

curved slightly before end of first vein; third and fourth veins parallel beyond discal

cell. Marginal cell at end of first vein subequal to submarginal cell at same point.

Legs from nearly white to pale tan except for brown median band and apex of hind

femora, median band of middle and hind tibiae and hind tarsi. Front tibiae with four

to six prominent dorsal bristles. Anterodorsal bristle of middle tibiae slightly shorter

than length of basitarsus, ventral apical bristle markedly shorter. Hind femora with

five to six anterodorsal bristles along entire length; below with numerous strong

spines, two anteroventrals evidently longer than others. Several apical bristles slightly

differentiated. Hind basitarsus slightly swollen and spinose on anterior edge.

Abdomen pale brown, pale pollinose; abdominal hairs pale. Genitalia as figured

(Figures 22, 47).

Types: Holotype. —c, Jamaica, Hardwar Gap, 4000 ft., 13 July 1966 (Howden and

Becker). Type No. 12692 in Canadian National Collection.

Allotype. —9, same data as holotype, (CNCI).

Paratypes. — Same data as holotype with following dates: 1c’, 19, 13 July 1966; 19,

6 July 1966; 399, 25 July 1966. 12, Try. Good Hope, 8 August 1966 (Howden and

Becker). Paratype distribution: 10, 12, (USNM); 12, (MSUC); 19, (IJSM); remainder

in CNCI.

Remarks: This species is readily distinguished from other West Indian species by

the combination of large size, pale color and conspicuously banded legs.

All specimens examined had apparently been treated by some method, prior to

mounting, which resulted in fading. Certain artifacts of this fading were ignored in

the description, such as the partially whitened stigma, which is almost certainly

absent in specimens which have not been processed in this manner. The colors of the

faded fly, however, are those given in the description.

Syneches macrochaetosus n. sp.

Figures 12, 31, 39, 53

Head dark brown, grey pollinose; posterior ocellar depression present. Two long

and two short ocellar bristles. Antennae dark brown, third segment with in-

conspicuous pale pubescence; arista more than five times length of third antennal

segment, dark, finely pubescent except for apex.

16 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Thorax light brown to brown; anterior half of mesonotum brown pollinose except

for two longitudinal stripes and three lateral spots which are grey pollinose; posterior

half grey pollinose with brown pollen at base of bristles. Pollen on prescutellar

triangle flattened, the spot contrasting with rest of notum when viewed from different

aspects. Integument of postalar calli lighter than that of notum, pollinosity similar.

Five weak humeral bristles; one notopleural, one supraalar; four postalars, one longer

than supraalar and three very weak. Pleurae pale pollinose except for large polished

area on sternopleuron and mesopleuron. Halteres brown, stem white to yellow.

Wings 3.5-4mm. in length, light brown, stigma dark brown, rectangular, with a

hyaline area at its apex. Second vein nearly straight beneath end of first vein; third

and fourth veins parallel beyond discal cell. Marginal cell one and one half to two

times as wide as submarginal cell at end of first vein.

Legs: Front and middle legs yellow to light brown except for dark distitarsus and

rarely dark middle tibia; hind legs dark brown except for yellowish trochanters and

femoral and tibial bases. Front tibiae with three to four prominent dorsal bristles;

anterodorsal and ventral bristles of middle tibiae both subequal to length of tibia;

apical posteroventral bristle markedly shorter. Hind femora with two to three short

anterodorsal bristles, below with abundant short and two to four long posterodorsal

spines. Hind tibiae tuberculate ventrally with two anterodorsal bristles of varying

lengths, apical posterodorsal stout but very short. Hind basitarsus slightly swollen;

spinose on anterior edge.

Abdomen brown, occasionally first three segments yellow; brown to pale pollinose;

abdominal hairs brown. Genitalia as figured(Figures 31, 39).

Types; Holotype. —c, Puerto Rico, Maricao LT, July 1960 (J. Maldonado C.) Type

No. 72142 in USNM.

Allotype. —?, same data as holotype. (USNM)

Paratypes. — 61%, 1199, same data as holotype. 10°, Caribbean N. For., Toro

Negro Unit Road No. 564, K.4.8., 2870 ft. alt., Dona Juana Forest on vegetation on

forest floor, 17 August 1965, 18°9’N, 66°29'W. (S. Medina Gaud); 19, K.5.7., 2880

ft. alt., 17 August 1965 (S. Medina Gaud). 1c", Barranquitas, Orocovis Road No. 143,

K.2.7., on mixed vegetation by roadside, 17 August 1965 (S. Medina Gaud). 19,

Luquillo Forest, El Yunque Biol. Sta., Molindero Road, elevation 2100 ft., at black

light, 2 Jan. 1963 (Paul and Phyllis Spangler). Paratype distribution: 5c, 5%,

(CNCD; 39°", 3 S¥(MSUCQ); 39°, (ISUD; 300", 3 9%, (IJSM); 10°, 1 9, (BMNH);

remainder in USNM.

Remarks: This species is easily distinguished from all other West Indian Syneches

by the long setae on the middle tibiae.

Syneches debilis Coquillett

Figures 6, 25, 42, 58

Syneches debilis Coquillett, 1895, Proc. U.S. Natl. Mus. 18: 436.

Head dark reddish brown, grey pollinose; posterior ocellar depression and

posterior ocellar polished band present. Two long and two short ocellar bristles.

Antennae dark brown to black, arista usually black, apex sometimes yellow, ap-

parently bare, about ten times length of third antennal segment.

Wilder: North American Syneches IF

Thorax light brown to yellow, golden pollinose. Thorax may be marked with dark

integumental median longitudinal stripe, notopleural spot, prescutellar longitudinal

stripe and humeri. Five, rarely six humeral bristles; two stout subequal notopleurals;

three postalars, one long and two minute. Pleurae yellow to dark brown with sparse

golden pollen, most of sternopleura and mesopleura polished. Halteres dark brown to

black, stem yellow.

Wings 3 mm in length, pale brown, stigma, if visible, light brown, indistinct.

Second vein curved beneath end of first vein; third and fourth veins parallel to slightly

converging beyond discal cell. Marginal cell at end of first vein as wide as sub-

marginal cell at same point. |

Legs light brown to yellow; apex of hind femora, middle of hind tibiae, hind

basitarsi and middle and hind distitarsi dark brown to black. Middle tibia with

strong black anterodorsal at basal fourth; subapical posteroventral shorter than

anterodorsal, yellow. Hind femora with four to five anteroventral bristles; hind tibia

with two dark anterodorsal bristles, at least one of which is longer than anteroventral

femoral bristles. Hind tibia slightly swollen at apex.

Abdomen light brown to black, brown pollinose, first segment with greyish pollen;

abdominal hairs golden to brown. Genitalia as figured (Figures 25, 42).

Type locality and type: Holotype. —%, Washington, D.C. Type No. 3239 in U.S.

National Museum.

Distribution: New Jersey, Maryland, Washington, D.C., Virginia, Florida, Georgia.

Remarks: This species can be distinguished from most North American species by

its small size, and from other small species (pusillus, vittatus, vineus) by the light

thoracic color. It differs from bacis, which keys to debilis in Melander (1902), in not

having a well-defined oval stigma.

70 specimens of this species were examined.

Syneches bacis (Walker)

Figure 64

Platypalpus bacis Walker, 1849, List. Dipt. Ins. in Brit. Mus. 3: 510.

Head dark brown, grey pollinose; posterior ocellar depression present. Two long

and two shorter ocellar bristles. Antennae yellow, apical segment and arista brown.

Thorax yellow, golden pollinose. Three humeral bristles; one notopleural, one

supraalar; one long and two shorter postalars. Pleurae pale pollinose, subshining;

parts of sternopleuron and mesopleuron polished.

Wings 3.25 mm. in length, hyaline with a well-defined oval stigma. Second vein

nearly straight beneath end of first vein; third and fourth veins parallel.

Legs yellow, hind tibiae brownish at tip. Front tibiae with several prominent

bristles; middle tibiae with long anterodorsal at basal third, numerous long ventral

and anteroventral bristles on apical half. Hind femora with apparently three an-

terodorsal bristles, two small bristles visible on posteroventral surface; several weak

subapical bristles present on hind tibiae.

Type locality and type: Holotype. — Sex uncertain, Jamaica, (P. H. Gosse). Type in

British Museum (Natural History).

18 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Remarks: This species can be distinguished from other West Indian Syneches by

the combination of yellow thorax; entirely yellow legs, except for tip of hind tibiae;

and the presence of a well-defined oval stigma.

The type of this species is damaged, the abdomen is missing as are three of the legs

and the wings. The only hind leg is glued to part of the mount in such a manner that

the anterior and ventral surfaces are obscured. Figure 64 is a photograph of the wing

of the holotype, according to Mr. K. G. V. Smith. Mr. Smith most kindly allowed the

writer to examine the type.

Also present on the pin is a Syrphid, Mesograpta arcifera Loew, presumably taken

as prey. This seems unlikely, however, since the prey is much larger than that usually

taken by Syneches of even larger size than bacis. :

Syneches quadrangularis Wheeler and Melander

Syneches quadrangularis Wheeler and Melander, 1901, Biol. Cent. Am., Dipt.

Suppl., p. 374.

Head dark reddish brown, grey pollinose; posterior ocellar polished band present.

Two long and two shorter ocellar bristles. Antennae brown, arista six times length of

third antennal segment.

Thorax brown with two light brown longitudinal mesoscutal stripes on anterior

third; yellow pollinose; integument of humeri and postalar calli yellow. Three

humeral bristles; two subequal notopleurals; one long and two minute postalars.

Pleurae brown, nearly black at bases of legs, grey pollinose. Halteres brown, stem

yellow.

Wings hyaline, stigma dark brown, quadrate. Marginal cell not widened at the

stigma; third vein not rigidly straight.

Legs yellow, hind femora, tibiae and basitarsi brown apically. Middle tibiae with

brown dorsal bristle on basal third and one yellow apical bristle.

Abdomen dark brown, brown pollinose; abdominal hairs brown.

Type locality and type: Holotype. —<, Tabasco, Mexico. Type in British Museum

(Natural History).

Remarks: This species can be easily distinguished from all other North American

Syneches by the combination of brown thoracic color and the presence of a distinct

quadrate stigma.

This description was made after examination of the holotype, generously loaned to

the writer by Mr. K. G. V. Smith. The specimen, however, was damaged, having

neither hind legs nor wings. Several other important features were also obscured or

missing. Melander’s description (1902) served as a basis for describing missing or

obscured characters.

Wilder: North American Syneches 19

Syneches pusillus Loew

Figures 2, 26, 40, 56

Syneches pusillus Loew, 1861, Berl. Ent. Zeitsch. 5: 320.

Head dark reddish brown, grey pollinose; posterior ocellar polished band present.

Two long and two shorter ocellar bristles. Antennae yellow to black, third segment

pubescent, arista five to ten times length of third antennal segment.

Thorax brown to black; brown or brown and grey pollinose; integument of humeri

and postalar calli sometimes lighter than that of rest of notum, pollinosity similar

throughout. Usually more than four humeral bristles; two notopleurals, posterior

usually longer; three postalars; one long and two minute. Pleurae pale brown and/or

grey pollinose, sternopleura and anterior strip of mesopleura polished. Halteres dark

brown to black, stem yellowish.

Wings 2.5-3.5 mm. in length, hyaline to light brown, stigmal area of marginal cell

rarely darker than rest of wing. Second vein straight beneath end of first vein; third

and fourth veins nearly parallel beyond discal cell. Width of marginal cell at end of

first vein subequal to width of submarginal cell at same point.

Legs varying in color from a uniform brown to black with yellow pro- and meso-

tarsi, bases and apices of metatibiae, and middle three segments of metatarsi.

Middle tibiae with dorsal bristle on basal third. Three, rarely more, ventral hair-like

bristles on apical two thirds of hind femora; tibia with anterodorsal bristle at one-half

its length; one long pale subapical and numerous small apical bristles.

Abdomen brown to black; brown or grey and brown pollinose; abdominal hairs

dark. Genitalia as figured (Figures 26, 40).

Type locality and type: Type No. 1113 in Museum of Comparative Zoology,

Harvard University. Loew stated he had both sexes, but according to Mr. George

Steyskal, who kindly examined the types, no abdomens remain on the two specimens.

One label reads ‘‘Chicago, III., Loew’’; and the other, ’’N. W., Loew.”’

Distribution: Ontario and Maine south to Florida, west to Louisiana, Arkansas,

Kansas, Nebraska and South Dakota. Flight period: southern parts of range, January

to July; northern parts of range, May to September.

Remarks: This species is very close to ater Melander, differing mainly by the

smaller size, lighter wings and longer arista. It can be separated from vineus by the

straight second vein and from vittatus n. sp. by the absence of banding on the ab-

domen.

173 specimens of pusillus were examined. A majority of the western specimens

differ from those farther east by the color of the legs. Western specimens have femora

and tibiae nearly black and all tarsi entirely yellow; eastern specimens almost always

have brown femora and hind basitarsi.

20 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Syneches ater Melander

Figures 7, 21, 35, 60

Syneches ater Melander, 1928, Genera Insectorum 185: 40.

Head dark reddish brown to black, grey pollinose; posterior ocellar polished band

present. Two long and two shorter ocellar bristles. Antennae dark brown to black,

third segment pubescent, arista bare, not more than five times length of third an-

tennal segment.

Thorax dark brown to black; brown or brown and grey pollinose; integument of

humeri and postalar calli sometimes lighter than that of rest of notum, pollinosity

similar throughout. Three to five, rarely more, humeral bristles present; two

notopleurals; posterior usually longer; three postalars, one long and two minute.

Pleurae pale brown and/or grey pollinose, sternopleuron partly polished. Halteres

dark brown to black, stem yellowish.

Wings 3.5-4 mm. in length, pale to dark brown, stigmal area of marginal cell

darker than rest of wing. Second vein straight beneath end of first vein; third and

fourth veins nearly parallel beyond discal cell. Marginal cell usually slightly wider

than submarginal cell at end of first vein.

Legs brown, hind legs darker; tarsi, bases and apices of tibiae, middle segments of

tarsi usually lighter, brown to yellow. Middle tibiae with dorsal bristle on basal third.

Two or three, rarely more, ventral hair-like bristles on apical two thirds of hind

femora; tibiae with anterodorsal bristle on apical half, one pale subapical and

numerous small apical bristles.

Abdomen dark reddish brown to dark brown, brown or grey and brown pollinose;

abdominal hairs dark. Genitalia as figured (Figures 21, 35).

Type locality and type: Holotype. —c, Pennsylvania, Hazleton, Luzerne Co., 28

June 1912 (Dr. Deitz). Type in U.S. National Museum.

Records: NOVA SCOTIA. &’, Lockeport, 28 July 1958 (J. R. Vockeroth).

NEWFOUNDLAND. 1c Terra Nova National Park, 7 July 1961 (C. P. Alexander).

ONTARIO. 1c’, Mer Bleu, 5 mi E. Ottawa, 19 June 1966, Malaise trap (D. D.

Munroe); same locality with following dates: 3005 499 25 June 1966; 105 49 9 27 June

1966; 1% 1 July 1966; 1c’, 229, 5 July 1966; 299, 7 July 1966; 229, 11 July 1966; 19, 19

July 1966. 20% 299, Ottawa, 4 July 1964 (JJ. R. Vockeroth). MAINE. 2c 2?9 Mt.

Desert Island, Hancock Co., 16 July 1963 (JJ. S. Naeger). 1% Fort Kent, Aroostook

Co., 19 Aug. (A. L. Melander). MASSACHUSETTS. 19, Athol, Worcester Co., 13

July 1933 (A. L. Melander). CONNECTICUT. 299, Storres, Tolland Co., 6 July 1931

(A. L. Melander). NEW YORK. 10, Lynbrook, Nassau Co., 24 July 1959 (G. C. Eick-

wort). 12 Zurich Bog, Wayne Co., 25 July 1964 (L. L. Pechuman). 19, Orient, Long

Island, Suffolk Co., 5 July 1952 (R. Latham). 1c, 19, Babylon, Suffolk Co., 16 June

1934 (Blanton and Borders). 13, Ellis, Slaterville, 13 June 1904. NEW JERSEY. 20%;

Lakehurst, Ocean Co., 26 June 1931 (A. L. Melander). MARYLAND. Ic’, College

Park, Prince Georges Co., 28 July 1935 (C. T. Greene). MINNESOTA. 1c’, Q-S WRC,

T46N R10W Sec. 9, Basswood Lake, Lake Co., 18 July 1952 (E. McConnell).

Wilder: North American Syneches Zl

Remarks: This species is very close to S. pusillus and can be distinguished only by

the larger size, slightly darker wings and shorter arista. All of these characters overlap

with pusillus, and in transitional areas, Michigan and Pennsylvania, the two species

are nearly indistinguishable.

At the beginning of this study, it was hypothesized that pusillus and ater were

simply different variations of the same species, and that Melander, looking at one

very dark insect, had erroneously considered it distinct.

The length of the arista was the most concrete and easily measurable character.

The antennae of 17°C’ and 17 °¢of ater and an equal number of pusillus were

removed and mounted on slides. The lengths of the third antennal segment and of the

arista were measured for each specimen and recorded along with a measurement of

wing length. All measurements overlapped between the two species.

A statistical test could not be run on arista length alone because specimens of ater

were, in general, larger in body size than those of pusillus, and arista length varied to

some degree with body size. A ratio of arista length to third antennal segment length

was chosen because the length of the third antennal segment varies directly with size

of the insect.

The ratio varied from 3.5 to 5.0 in ater and from 5.0 to 10.0 in pusillus. A Student’s

T-test was run on the sets of ratios and a t-value of 10.01 obtained. This is significant

at the “1. percent ‘level:

On the basis of these data, it was concluded that ater and pusillus are distinct, and

nearly always separable on the basis of a combination of color, size and ratio of the

arista to the third antennal segment length.

Syneches vittatus n. sp.

Figures 16, 32, 38, 55, 61

Head black, grey pollinose; posterior ocellar depression and posterior ocellar

polished band present. Two long and two shorter pale ocellar bristles. Antennae dark

brown to black, arista bare.

Thorax brown to black, greyish pollinose, variously marked with brown pollinose

patches; brown pollen always present at bases of dorsocentral bristles. Humeral

bristles variable, usually more than six; two notopleurals, posterior longer; one long

and two shorter postalar bristles. Pleura greyish pollinose except for polished spot on

sternopleuron. Halteres brown to black, stem usually lighter.

Wings 3 mm. in length, subhyaline to hyaline, stigma absent. Second vein curved

beneath end of first vein; third and fourth veins parallel beyond discal cell. Marginal

cell at end of first vein about half as wide as submarginal cell at same point.

Legs brown; coxae, trochanters, femora and sometimes tibiae quite dark. Dorsal

bristle on basal third of middle tibiae small, subapicals pale and subequal to dorsal

bristle. Hind femora with pale hairs below; anterodorsal on basal half of hind tibiae

barely differentiated; two subapicals present.

Abdomen dark brown to black, brown pollinose, posterior band on each abdominal

segment grey pollinose; abdominal hairs pale. Genitalia as figured (Figures 32, 38).

» Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Types: Holotype. —S& Arizona, Cochise Stronghold, Dragoon Mts., Cochise Co. 29

June to 2 July, 1970, U.V. Light trap (R. J. Shaw). Type deposited in University of

Arizona Insect Collection.

Allotype. —?, same data as holotype. (UAIC).

Paratypes. —19, same data as holotype. Others with following dates: 19, 29 June

to 2 July 1970; 2 99 , rain during nights 20, 21, 18-21 July 1970; 19 , July 31-Aug. 2,

1970. 200", 19, Pena Blanca, Santa Cruz Co., 10 July 1961, U.V. Light trap

(Werner, Nutting). 300", 4 99, Patagonia Mountains, Santa Cruz Co., 5500 ft., 10

Cochise Co., 4 July 1940 (D. E. Hardy and R. H. Beamer). 1° , 12 , Sunnyside

Canyon, Huachuca Mts., Cochise Co., 9 July 1940 (D. E. Hardy and R. H. Beamer).

Paratype distribution: 20° , 3 cb", (KSUC); 10° ,12 , (MSUC); 4 ov, 222 , (CNCI);

200 | 222, (USNM); remainder at UAIC.

Remarks: This species can be distinguished from all other North American

Syneches by the banded abdomen and the narrow marginal cell.

Syneches vineus n. sp.

Figures 5, 29, 43, 50

Head black, greyish pollinose; posterior ocellar depression present; posterior

ocellar polished band wide. Two long and two shorter ocellar bristles. Antennae

black; arista bare.

Thorax: Notum black, shining, sparsely covered with long brown pollen. Two to

four humeral bristles; two notopleurals, posterior distinctly longer; one long and two

shorter postalars. Pleurae dark brown, brown pollinose, sternopleuron and

mesopleuron largely polished. Integument of postalar calli and a small posterior

humeral spot light brown to yellow. Halteres black, stem lighter.

Wings 3 mm. in length, grey to hyaline; stigma absent to dark grey. Second vein

with a long gentle curve below end of first vein; third and fourth veins slightly con-

vergent beyond discal cell. Marginal cell at end of first vein distinctly narrower than

submarginal cell at same point.

Legs dark brown to black; front and middle tibiae and tarsi yellow; bases and

apices of hind tibiae and middle three tarsal segments yellow; hind basitarsi and all

distitarsi brown. Dorsal bristle on basal third of middle tibia longer than

notopleurals; one pale ventral subapical, subequal to dorsal bristle. Hind femora not

swollen, below with one hair-like bristle and one or two long hairs. Anterodorsal

bristle on basal half of hind tibiae subequal to ventral femoral bristle; two subapicals,

one pale, both slightly shorter than anterodorsal. Hind basitarsus with two subapical

hair-like bristles.

Abdomen dark brown to black, shining; pollen brown, sparse; abdominal hairs

black. Genitalia as figured (Figures 29, 43).

Types: Holotype. —CG Dominica, W. I., Fond Figues R., Rain forest, 9 Feb. 1965

(W. W. Wirth). Type No. 72170 in U.S. National Museum.

Allotype. —%, same data as holotype.

Wilder: North American Syneches 23

Paratypes. — Same data as holotype with following dates: 40’, 2 2¢ 9 Feb. 1965, 3

Feb. 1965; 19, 25 Jan. 1965; 299 9-12 March 1965; 299 Fond Figues R. 400’, 29

April 1966 (R. J. Gagne); 1c, 12 April 1966. Pt. Mulatre, 15 Sept. 1966 (D. L.

Jackson). 1%, Pont Case, 20 Sept. 1965 (D. L. Jackson). 1c5 Clarke Hall, Cocoa Trail,

18 Jan. 1965 (W. W. Wirth). 1°, trail, 1 mi. n. junc. rds. to Rosalie and Castle Bruce,

1300’, 29 March 1966 (R. J. Gagne). 12, 5 mi. e. Pont Casse, 11 April 1966 (R. J.

Gagne). 19, 2 mi. e. Pont Casse, 6 May 1966 (R. J. Gagne). Paratype distribution: 1c’,

12, (CNCD); 10’, 12, (MSUC); 10%, 19, (JSM); remainder (USNM).

Remarks: This species can be distinguished from other West Indian Syneches by

the small size and dark body color. It can be separated from vineus jamaicensis Nn. ssp.

and from vineus medinai n. ssp. by the presence of a distinct grey stigma and long

subapical bristles on hind basitarsus.

Syneches vineus medinai n. ssp.

Figures 23, 44, 61

Morphologically very similar to S. vineus vineus, differing in the following

characters: Wings hyaline, stigma absent. Thoracic pollen sometimes greyish on

posterior third of notum; posterior notopleural bristle only slightly longer than an-

terior. Legs light brown, front and middle tibiae and tarsi yellow; hind legs yellow

except for brown coxae, trochanters, and apical third of hind femora; all distitarsi

brown. Hind basitarsus without subapical bristles. Apical margin of hypandrium

convex (Figure 44).

Types: Holotype. —% Puerto Rico, Caribbean N. For. Toro Negro Unit Rd. No.

564, K.5.7., 2880 ft. alt., Dona Juana Forest at entrance to recreation area, 18°8’N,

66°30’W, 17 Aug. 1965 (S. Medina Gaud).

Paratypes. —1%, same data as holotype; 1%, Caribbean N. For. Toro Negro Unit

Rd. No. 564, K.4.8., 2870 ft. alt., Dona Juana Forest on vegetation on forest floor,

18%’N, 66°29’W, 17 August 1965 (S. Medina Gaud); 1c} K.12.6 from Carey on rd.

No. 15 to Guayama, swept from grass, 1892’30’’N, 66°7’45”’W, 1 Sept. 1964 (S.

Medina Gaud and E. Medina). 10", Maricao LT., July 1960 (J. Maldonado C.). Ic’,

K.22.7 from Carey on road No. 15 to Guayama, on vegetation, mainly Bidens pilosa,

80°6’N, 66 9’W (S. Medina Gaud and E. Medina). Paratype distribution: 1c’ im each

of the following museums: USNM, CNCI, MSUC, ISUI, and IJSM.

Syneches vineus jamaicensis n. ssp.

Figures 15, 24, 41,54 —

Morphologicaily very similar to S. vineus vineus, differing mainly in the following

characteristics: Wings hyaline, stigma milky white, probably hyaline. Thoracic pollen

greyish on posterior third of notum. Legs yellow to white, probably light brown,

distitarsi darker; hind basitarsi without subapical bristles. Hypandrium with a

- median apical notch (Figure 41).

Types: Holotype. —, Jamaica, Hardwar Gap, 4000 ft., 25 July 1966 (Howden and

Becker). Type No. 12693 in Canadian National Collection.

24 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Allotype. —?, same data as holotype. (CNCI).

Paratypes. — Same data as holotype with following dates: 400" ,6?9, 25 July 1966;

1%, 229, 6 July 1966; 299 10 July 1966; 4cb, 229, 13 July 1966; 305 29 July 1966.

Paratype distribution: 200’, 2 2(USNM); b&b’, 19, (MSUC); Ic’, 19, (IJSM); remainder

in CNCI.

Remarks: The preceeding three insects are morphologically quite similar in many

respects. The characters used to separate them, however, might normally warrant

placement in separate species. For example, the median apical notch on the epan-

drium of jamaicensis and the absence of this notch on medinai appears to be a very

constant character; but in vineus vineus both the notched and unnotched condition

exist. Therefore, without examination of a larger number of specimens, one cannot be

certain that the character is constant in jamaicensis and medinat.

Of the specimens examined, only those of vineuws vineus were in satisfactory con-

dition. Only five males of medinai were examined. Although the series of jamaicensis

was fairly large, all specimens had been stored or prepared in a manner which caused

fading. It is unfortunate that such a process was used on all of the specimens because

it caused fading of many body parts and the collapse of certain portions of the thorax.

Color patterns could only be inferred by comparison with similarly treated specimens

of ater.

Because of the larger degree of variation in other Syneches, the unsatisfactory

condition of the relatively small numbers of specimens and the definite geographical

isolation, it seems more reasonable, at this time, to place the three groups at a sub-

species level. It is very possible that the three are distinct species, but this cannot be

determined until a larger number of specimens in satisfactory condition from these

and other islands have been examined to determine the range of variation in each

group.

Wilder: North American Syneches 23

LITERATURE CITED

Hull, F.M. 1962. Robberflies of the World. The genera of the family Asilidae. Part I.

U.S. National Mus. Bul. 224: 53.

Melander, A.L. 1902. A Monograph of the North American Empididae. Amer. Ent.

Soc. Trans., 28: 195-367.

1928. Diptera. Fam. Empididae. Fasc. 185, 434 pp., 8 pls. In

Wytsman, P., ed., Genera Insectorum. Bruxelles.

Schwartz, E.A. 1891. Feeding habit of a species of Empididae. Proc. Ent. Soc. Wash.

2: 146.

Smith, K. G. V. 1962. Studies on the Brazilian Empididae. Trans. Roy. Ent. Soc.

Lond. 114: 195-266.

1969. The Empididae of Southern Africa. Ann. Natal Mus. 19: 1-

347.

1971. A revision of Francis Walker’s Types of North American

Empididae (Diptera). Bull. Br. Mus. Nat. Hist. Entomology 26: 347-370.

Westwood, J. O. 1835. Insectorum nonnullorum novorum (ex ordine dipterorum)

descriptiones. Soc. Ent. de France, Ann. (ser. 1), 4: 681-685.

Wiedemann, C. R. W. 1823. Aussereuropaische zweiflugelige Insekten. Vol. 1, xxxii

+ 608 pp., 7 pls. Hamm.

26 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Cows

Fig. 1 Epandrium of Syneches thoracicus (Say).

Fig. 2. Meta leg of S. pusillus Loew.

Fig. 3. Wing of S. inversus Curran.

Fig. 4. Hypandrium of S. thoracicus (Say).

Figs. 5-7. Metal leg of: S. vineus vineus n. ssp. 5; S. debilis Coquillett 6; S. ater Melander 7.

Wilder: North American Syneches 27

Figs. 8-13. Meta leg of: Syneches rufus Loew 8; S. thoracicus (Say) 9; S. simplex Walker 10; S. frosti n. sp.

11; S. macrochaetosus n. sp. 12; S. pallidus n. sp. 13.

Fig. 14. Meso leg of S. macrochaetosus n. sp.

Fig. 15. Meta leg of S. vineus jamaicensis n. ssp.

Fig. 16. Meta leg of S. vittatus n. sp.

28 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

Figs. 17-32. Epandrium of: Syneches rufus Loew 17; S. albonotatus Loew 18; S. frosti n. sp. 19; S.

longipennis Melander 20; S. ater Melander 21; S. pallidus n. sp. 22; S. vineus medinai n. ssp. 23; S.

vineus jamaicensis n. ssp. 24; S. debilis Coquillett 25; S. pusillus Loew 26; S. hyalinus Coquillett 27; S.

inversus Curran 28; S. vineus vineus n. ssp. 29; S. simplex Walker 30; S. macrochaetosus n. sp. 31; S.

vittatus n. sp. 32.

Wilder: North American Syneches 29

9 IO

Figs. 33-48. Hypandrium of: Syneches albonotatus Loew 33; S. rufus Loew 34; S. ater Melander 35; S.

hyalinus Coquillett 36; S. simplex Walker 37; S. vittatus n. sp. 38; S. macrochaetosus n. sp. 39; S.

pusillus Loew 40; S. vineus jamaicensis n. ssp. 41; S. debilis Coquillett 42; S. vineus vineus n. ssp. 43; S.

vineus medinai n. ssp. 44; S. longipennis Melander 45; S. frosti n. sp. 46; S. pallidus n. sp. 47; S. in-

versus Curran 48.

SO Pe

30 Contrib. Amer. Ent. Inst., vol. 10, no. 5, 1974

coe

RES

Figs. 49-65. Wing of: Syneches longipennis Melander 49; S. vineus vineus n. ssp. 50; S. thoracicus (Say)

51; S. rufus Loew 52; S. macrochaetosus n. sp. 53; S. vineus jamaicensis n. ssp. 54; S. vittatus n. sp. 55;

S. pusillus Loew 56; S. albonotatus Loew 57; S. debilis Coquillett 58; S. simplex Walker 59; S. ater

Melander 60; S. vineus medinai n. ssp. 61; S. pallidus n. sp. 62; S. hyalinus Coquillett 63; holotype of S.

bacis (Walker), (not to same scale) 64; S. frosti n. sp. 65.

Contributions

of the

American Entomological Institute

Volume 10, Number 6, 1974

MORPHOLOGY AND TAXONOMY OF THE KNOWN PUPAE

OF COCCINELLIDAE (COLEOPTERA) OF

NORTH AMERICA, WITHA

DISCUSSION OF PHYLOGENETIC RELATIONSHIPS

Duang Trung Phuoc

and

Frederick W. Stehr

TABLE OF CONTENTS

Page

INTRODUCTION]: 6 Feet rR Ene Es 93 5 SEP EE a VE P hs PPE 2

CLASSIFICATION OP COCCINBLEIDAR 2. ch. aia ore i a aes mene we & + 3

METHODS OF STUDY AND SOURCES OF MATERIAL ................0.. 3

COMPARATIVE MORPHOLOGY oe. o eeu ek as age EP ce aes ce 6

KEY TO SUBFAMILIES AND TRIBES OF THE AVAILABLE PUPAE

OF COCCINELEIDAR Stes Os Hii ph ee a oS SP a I a 11

SUBFAMILY SIICHOLOUINAD Sole ioe tee ee we Crd ge ee 14

TRIBEGERANGUIING 0 8 jiwacwwswitss » eahe dete woteud ape ek 65 14

Gene S eS CASO on ess oe ap easels 5 ny Was bade ok es le eg ee 14

DAR asta pusiiis( ecOIe) oS i tae hc ee eee eo eee 14

TRIBE STICHOLO INT err vk ce CO a Od ee ae 15

Genus MICROWEPEISEA Cdekerell. 00: seeaeste. aarwachien ele 15

Microweiséa ovalis (econte) . igsiek O23 WR teh So 8 Ye 15

SLBFAMILY LOCCTOUIIN A Beco ie cst an ca acne uw oye ce uae aera neues camer 15

TRIBE COC CIDULINE Oris aie oes 5 RN ea PY 16

Key to Genera of the Available Pupae of Coccidulini:. 044 04 eso ke 16

Genus GIN DORTS-C asey Cr ee, CF BA i A os es all 16

LindorusigpriwtaeBiaisdelly: soe oa 5, 6 BIER AMIS OI ee ek 16

Genus RHIZOBMIS Stepnens own 6 6 PURER, WO, WSU t7

Rhizobius ventralue(EMeNson) 66 shea eee a nee ee ee eee 17

PRR SC ys isl cogisy nitou qciiabence ae cca as We ecm cane ee NR anual 18

Genus ZAGLOBA.C ae go ey gg roe 4 EM 8 Re Gl oe hd I 18

Zasioba ormata (Hoye 23 05-4 REA ) EGR TA Ne 18

TREBE OVINE 5205 eR TEAC, OF Ea is wae a 4/5 uk eokid son AW 18

Genus RODOR Ait set Pe A, ke cola 5 tive a Saw eae won eae Ae 18

PR OMOLEA CONGIIICIIS WAUSAU) 6 65.6 55 6 scaled See hbase kn Sek We A aS kN 19

SUBFAMILY BPG as wei viaia Vie oie 4b paie ssa ebeders men ie 20

TRIBE EPLGACHINI NE ois wala iw bag weak aceite ee NO Rac dle ee 20

Genus EPILACEHINA Cheval sie oe cee eee CR OTT nn PS as 20

Key to Species of the Available Pupae of Epilachna.:.. ..0 000 e 20

Epilachna bored (Pa oieiey! 5G CORR, We ARO PE OL Re a 21

Epilachna.vanvestis asa: : oa 3x SARI SA ERIE ce 21

FE ULGCHIGE BP 6 ee ok ek EAS hae OND FARGO i ba yes

NB ADIL SCT MINNA i eee eee Sve ee ees 22

RE a a es Wives aces 22

Key to Genera of the Available Pupae of eS Gace Depo ee enn Sane § 23

ents YY Pir A erro i ee ea Po Fein oe ao

Key to Species of the Available Pupae of aoe Be ey eet ees ais aes 24

pers Oroidia Say ee er eS 25

Hy peraspis lateralis Maisany S65 2.0500 Fo ek 5 de ee a) ieee tes 26

Teas DON EEC OIG os ee ew So Rea ee 26

Ay peraspis ecu anecunce Casey... ess es Pe ES. Zt

Hypores pis eieta Lecunie ii ei Fa PSE He eR as 28

Hy peraspis quadrioculata (Motschulsky) <2... 2... ee oe ee es 29

SIPC iis OCI Sey lg FS ewes ee OR be Baw 29

PPV CTOR IN GROUT IVIOtG LECOINE o.oo ica pee eb cly Re oy Ee pate ecees 30

ors THALASSA Mulsamt . 2. kc ces mieten ay Gs canst Macaeed Hy 31

tT hajlassa moniczuniae Mulsant.... on be. ols LYELL 31

RE a a ss eee ee ak ge ee 32

Key to Genera of the Available Pupae of Scymnini ................... eee

Crete SOY MINUS Rielgnt oc ee ec GPU A a4

Sepronus creperus Maulsant » 23 Sea PS a 8

Genus CR YPPOLAEMUS Mulsant. .. ie See Oo aI any 33

Crvproiaemus montrouzier: Mulsalt . oo. ok ce eRe ec ee bes oe

Rt OE Or a ay Oe ee lee thea ea ees 34

Cons SP POR LS Weise ns ek, RCP. 34

Key to Species of the Available Pupae of Siethorus. 2.361). ws pinaaesws ds. 5

rer ory aie Cae oe ree DL LM Tse s 35

DOCInOrIS PieLes <ASey . Se ig PO nar eudtes ceed ds. es 36

Sie boris puncruey Lecomte 3.0 occ. afliade. 2 LEE 4 Sei. 37

TRIPE HILO CRI be al ie FP BOI Cees 37

Key. to Genera of the Available Pupae of Chilocorint..3) ninco. peitau dc... 38

re A eC a i ie au nn Wl eee ace de 39

Ciigcbrs Pe ineras Wise a er a ce Sey Le ele eo 39

RCO CLS Vib Ga ee ne eae eae Ve eh ney view 40

OReUR Cet Oe oe cc ees eee Een os sa ke 40

Re ee a ee ee oes heey her ae ee ue at 41

Key to Species of tne Avaliavle Pupae Of AXION . vias A 41

ANTON Dieter TOMIVIET oe Cink hk sc aaie oo aA EG eho RSE Pe 42

Axion tripustulain (DeEGeen os 6 aere dA PE eee es 42

Genus. EXOCHOMUS Redteniacket i} sidcti cut os aninge easisoW sca. 43

Key to Species of the Available Pupae of Bxochomus . oie check 43

EXOcHOMmus Hoew@erl VIOENAI kas no eee A Ete ie hd Ee Raley As Ce es 43

EMOCEOMUS CUDERSIS IIUMINIORK: oh i 5 os 46 ay ke 6 4 9s EES doe 44

Ces Sie MOTDES Chao socials Vai oie caw lose ee a SS a4

Wri nodes Sutiuralis (PAOriClsy ss «ooh vo we ees Gk ds RA ee AA

SUBPAMILY COC Pe ey ae nae eb eb is oe a gees 45

TRIBE COCCINED LUND 633 vss dee) sano macevriecs healers +04 ss 45

Key to Genera of the Available Pupae of Coccinellini ..................... 45

Genus COCCI MEE Dinars ons noe ce ee dy a A te ee ee 49

Key to Species of the Available Pupae of Coccinella... 1.0.0... ccc eee. 50

Coccinella irifascinta Tiare a ASP Me oi Re i ee St pis

Coccinella transversoputtata ¥ aderimann. ooo. ee ew ee 8 ee ws 53

Cocetrena WOveminGtad ICIS he ia a ol a ease ett pe aeaa ee 54

Coccinella MrOnLiCohs WU IRat ioe ieee Ve iw aG denen a te bade ge ee a

Coccinelia seplem punctate Lannaeus 2.576 c5 Se ice ce ee he 9 56

(pets FA ea i ce a hd Wa Ei ek he 56

Acialin Papsieraia (raed) co sc os ik i a kn ee ew 56

ens CYC es a eee sien oes HINES CE ae es 57

Key to Species of the Available Pupae of Cycloneda..............00 0 cues 58

CVCLONCG WHE ON 5 eo ules paved Res teh Cy eee wd Oh Rd 59

Cyeionedd San mie (LANA GOGS) 6 css cde wy ve dt ee Ve SR iain db iced a0

Ceres MAU SAP IIA. Fs ee ihe vc eae eh ee wee eee 60

I leet CTO) se i A es in Pee Se 60

Mulsaniiia had eien Cage) ive ong AeA re CYS ee a ey Se 61

Crores COU 1 0 eee si Pe ie ee EE US ee es et ee ee 61

Olla abdominalis (Say) ......... Ts Pe aaa wis aaa seer Ma 61

Genus PROPY AEA Wiiisate ie es ee ee ee Se eek eee ey en 63

Propylaea quatuordecimpunctata (Linnaeus) ......... 0.0.06 c eee ee ees 63

Genus META I Re ee a ee ca eo ak eA oO N Saas see Be 64

Neonarnionia venustd (MeISneIMe!) 6.6 be ea can wey eee nee ne 64

COTS FE Ey et arte ea eee Cec nn bak ei eo serene ws 65

Key to Species of the Available Pupae of Anatis........ ped ueeeslh Asit: eriye ake oye erties 66

Anat OCCNO UNGOS ke cS ca ee ewe eis Guar heb ee eee 66

ANGNS GIIACCOURDUNCIAIA OUNCE fh ce ak ew 8 eco ole 8 bine es ge 8s 67

Gates ASO ee CP ee ad ee eo te a hom alge remo uNe 67

Anisocalvia quatvorqecimputiata (Linnaeus) oi ee ns ea es 67

Sorts SY NON Y CHA WIGISeND po Ne vn ce ee rite ae ae Naa a eens 68

SOnOnNVCHE Crandis (TV AUNOCI Es. tore ia sc) 6 eet eas Owe ay oe ee 68

Coens PII AL ICA Cri os Pelee een cn eben Ste tees baa eS 69

Key to Species of the Available Pupae of Hippodamia..................6. 70

Hippodamia tredecimpanctata (Linndeus) 0. i es ae ck eta ess 71

PTI PORUNING DARENINGTIEN OAR) oon elas Cl cs nat eh eee goes SA ae £9

AI POROHT a Glare ars) ke ee eh ee etek eee tiny ks cae eae is

Pi ppodaias COR VORPCnS UETIU.. oo 6 cas cae hs ce an ye OC ewe ae a 74

Fi npodaiid BUTANE OAAEG CRIED) oi iee sed nes vy ce BAL a ae he 74

Genus COLEOMEGILLA TINIDErIgke So vind c awa cece hee e eaee aed tees fis.

ColgOnmmenila Mat ate (OOtIOEl) o.oo eee ae bas Ue kK eS 7

Cents A eee Ne tes ok cs oe ok ee ea a Lew awa ses 76

Neentia seviare (WISI). i oc cae ka ek ki ca ee hee 76

Certs HIPs ete a ee ee ed eA ak Cu ee ed ea ve

PPOs COE CA 5 oc ch ach Gk owe PRE ee Ae ek Aw 8 a

Genas PSYLLORORACKevrolal 9.2 a es 78

Psyllobora-vipmiimaculata (Say). ee A EI, 78

DISCUSSION ANG CONCEUSIONS 20 igs oe Ge chert Lee 79

EITBRATURECTIED: = er ee Sr ee, 83

PUGS 10). kk ae 85

MORPHOLOGY AND TAXONOMY OF THE KNOWN PUPAE

OF COCCINELLIDAE (COLEOPTERA) OF

NORTH AMERICA, WITH A

DISCUSSION OF PHYLOGENETIC RELATIONSHIPS

Duang Trung Phuoc

and

Frederick W. Stehr

ABSTRACT

The structure of the pupae of 32 known genera and 49 species from 11

tribes of coccinellid pupae (mainly of North America) which are positively

associated with adults either by collecting in the field or by rearing in the

laboratory have been studied and described. Keys to known subfamilies,

tribes, genera and species are presented. Based on the pupal characters,

the tribal relationships within the family are discussed.

The COCCINELLINAE (Coccinellini and Psylloborini) are highly

advanced with very little connection with the remainder of the family. The

remainder of the family is grouped in a separate stem with

EPILACHNINAE splitting off very early and acquiring highly modified

structures to fit their exclusively phytophagous habit.

The COCCIDULINAE (Coccidulini and Scymnillini) and _ the

STICHOLOTINAE (Sticholotini and Serangiini) are regarded as being

the most primitive groups, and are closely related. Thus, they are grouped

on the same stem of the phylogenetic tree. The Coccidulini (=Rhizobiini)

are more closely related to the EPILACHNINAE than any other groups of

COCCINELLIDAE.

The CHILOCORINAE (Chilocorini) show some relationships with the

STICHOLOTINAE and the SCYMNINAE, but independently evolved on

a distinct stem.

The SCYMNINAE (Stethorini, Scymnini and MHyperaspini) have

recently evolved. The Hyperaspini appear to be more primitive compared

with Scymnini and Stethorini. The Stethorini are considered to be the

most highly evolved within the SCYMNINAE, and have the closest affinity

to the Chilocorini.

2 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

ACKNOWLEDGEMENTS

We wish to thank the following individuals and institutions for the loan or donation

of materials: Donald M. Anderson, Systematic Entomology Laboratory, USDA, U.S.

National Museum, Washington D.C.; Kenneth S. Hagen, University of California,

Berkeley, California; Robert E. Waites, College of Agriculture, University of Florida,

Gainesville Florida; Roland W. Portman, Department of Entomology, University of

Idaho, Moscow Idaho; Raymond D. Eikenbary, Department of Entomology,

Oklahoma State University, Stillwater Oklahoma and Donald C. Cress, Department

of Entomology, Michigan State University, East Lansing, Michigan.

We wish to thank Roland L. Fischer for his help with the morphology and various

other aspects, and to thank Lyle Matteson and Bill Ravlin for their care in testing the

Keys:

INTRODUCTION

The morphology of immature insects, especially of holometabolous groups, is an

additional source of taxonomic information which unfortunately has not been used

for many groups. According to Van Emden (1957) immature characters may be

significant in the separation of sibling species, and may help to confirm or to

reorganize the classification and phylogeny of groups of insects which have previously

been classified only upon adult characters.

The classification of the family Coccinellidae has been based largely on the mor-

phology of adults and larvae. The pupae of Coccinellidae have been little studied.

There have been a few short notes or brief descriptions on the morphology, but often

only for a single species. This morphological study of the pupae has provided another

source of data for more precisely understanding and interpreting the relationships

and the natural classification within the family.

Coquillet (1889), Gorham (1892), Pinto da Fonseca and Autuore (1931), Kanervo

(1941), Kapur (1943), Kesten (1969), and Marthur and Srivastava (1966) have briefly

described the pupae of Vedalia cardinalis (Mulsant), Orcus coeruleus (Mulsant),

Calvia 15-punctata Fabricius, Solanophila candestina (Mulsant), Thea bisoctonotata

Mulsant, Anatis ocellata Linnaeus, Stethorus gilvifrons Mulsant, and Hyperaspis

vinciguerrae Capra.

Palmer (1914) studied a group of common coccinellid species and provided brief

descriptions of the pupae of Hippodamia convergens Guerin, Hippodamia sinuata

Mulsant, Hippodamia parenthesis (Say), Coccinella 5-notata Kirby, Coccinella

monticola Mulsant, Coccinella sanguinea Linnaeus, and Olla abdominalis (Say).

Binaghi (1941) has carefully studied the pupae of the Chilocorini. He offered a key

to genera and the species Chilocorus bipustulatus Linnaeus, Exochomus 4-pustulatus

Linnaeus, Exochomus auritus Scriba, Exochomus nigripennis Erichson, and

Exochomus sjodstedti Weise.

Phuoc and Stehr: Pupae of Coccinellidae 3

Recently, Savoiskaya (1962a) has redescribed the pupae of the tribe Chilocorini.

Three genera were taxonomically separated (Chilocorus L., Brumus Muls., and

Exochomus Redt.) and the following species were described: Brumus 8-signatus

Gebl., Chilocorus bipustulatus Linnaeus, Exochomus flavipes Thunberg, Exochomus

melanocephalus Zubk., and Exochomus semenovi Weise.

Savoiskaya (1962c) has also described and studied the larvae and pupae of the

genus Coccinula Dobzhansky. He prepared a key separating the pupae of the genus

Coccinula, including the following species: C. redimita Weise, C. 14-pustulata

Linnaeus, C. sinuato-marginata Faldermann, and C. elegantula Weise.

CLASSIFICATION OF COCCINELLIDAE

Modified from Sasaji (1971)

(*) indicates tribe with available pupae

STICHOLOTINAE SCYMNINAE

Sukunahikonini Hyperaspini (*)

Serangiini (*) Scymnini (*)

Sticholotini (*) Stethorini (*)

Shirozuellini Cranophorini

| Aspidimerini

COCCIDULINAE Ortaliini

Lithophilini

Coceidulini (*) GEO COMNAE

Scymnillini (*) Telsimiini

Exoplectrini Platynaspini

Noviini (*) Chilocorini (*)

EPILACHNINAE COCCINELLINAE

Epilachnini (*) Psylloborini (*)

Coccinellini (*)

Discotomiini

METHODS OF STUDY AND SOURCES OF MATERIAL

The specimens examined were mainly from the U.S. National Museum and from

the rearing program carried on throughout the study. Pupae obtained by rearing are

preferred since identifications are positive. Pupae collected in association with adults

4 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

in the field are not as positively identified as reared specimens; however, it is an

excellent way to get a lot of material with minimal effort. The latter method can best

be used if the adults are allowed to emerge, and the pupal exuviae are associated with

the newly emerged adult. Not only can the specimens be correctly identified this way,

but the exuviae can be examined almost as well as the pupa itself.

In addition to the above material, several important pupae have been obtained

from Kenneth S. Hagen, University of California, Berkeley, California; Robert E.

Waites, University of Florida, Gainesville, Florida; Roland W. Portman, University of

Idaho, Moscow, Idaho and Raymond D. Eikenbary, Oklahoma State University,

Stillwater, Oklahoma.

The following pupae were available for this study:

STICHOLOTINAE 3

Delphastus pusillus (Leconte) (Serangiini)

Microweisea ovalis (Leconte) (Sticholotini)

COCCIDULINAE

Rhizobius ventralis (Erichson) (Coccidulini)

Lindorus lophantae Blaisdell (Coccidulini)

Rodolia cardinalis Mulsant (Noviini)

Zagloba ornata (Horn) (Scymnillini)

EPILACHNINAE

Epilachna varivestis Mulsant (Epilachnini)

Epilachna borealis (Fabricius) (Epilachnini)

Epilachna sp. (Epilachnini)

SCYMNINAE

Scymnus creperus Mulsant (Scymnini)

Cryptolaemus montrouzieri Mulsant (Scymnini)

Hyperaspis binotata (Say) (Scymnini)

Hyperaspis cincta Leconte (Scymnini)

Hyperaspis lateralis Leconte (Scymnini)

Hyperaspis octavia Casey (Scymnini)

Hyperaspis oculaticauda Casey (Scymnini)

Hyperaspis postica Leconte (Scymnini)

Hyperaspis psyche Casey (Scymnini)

Hyperaspis quadrioculata (Motschulsky) (Scymnini)

Hyperaspis quadrivittata Leconte (Scymnini)

Thalassa montezumae Mulsant (Hyperaspini)

Stethorus atomus Casey (Stethorini)

Stethorus picipes Casey (Stethorini)

Stethorus punctum LeConte (Stethorini)

CHILOCORINAE

Chilocorus bivulnerus Mulsant (Chilocorini)

Orcus chalybeus (Boisd.) (Chilocorini)

Phuoc and Stehr: Pupae of Coccinellidae 5

Axion plagiatum (Olivier) (Chilocorini)

Axion tripustulatum (DeGeer) (Chilocorini)

Exochomus hoegei Gorham (Chilocorini)

_Exochomus cubensis Dimmock (Chilocorini)

Brumoides suturalis (Fabricius) (Chilocorini)

COCCINELLINAE

Coccinella trifasciata Linnaeus (Coccinellini)

Coccinella transversoguttata Faldermann (Coccinellini)

Coccinella novemnotata Herbst (Coccinellini)

Coccinella septempunctata Linnaeus (Coccinellini)

Coccinella monticola Mulsant (Coccinellini)

Adalia bipunctata (Linnaeus) (Coccinellini)

Cycloneda munda (Say) (Coccinellini)

Cycloneda sanguinea (Linnaeus) (Coccinellini)

Mulsantina picta (Randall) (Coccinellini)

Mulsantina hudsonica (Casey) (Coccinellini)

Olla abdominalis Say (Coccinellini)

Neoharmonia venusta (Melsheimer) (Coccinellini)

Propylaea quatuordecimpunctata (Linnaeus) (Coccinellini)

Anatis ocellata Linnaeus (Coccinellini)

Anatis quindecimpunctata Olivier (Coccinellini)

Anisocalvia quatuordecimguttata Linnaeus (Coccinellini)

Synonycha grandis (Thunberg) (Coccinellini)

Hippodamia parenthesis (Say) (Coccinellini)

Hippodamia tredecimpunctata (Linnaeus) (Coccinellini)

Hippodamia convergens Guerin (Coccinellini)

Hippodamia glacialis (Fabricius) (Coccinellini)

Hippodamia quinquesignata (Kirby) (Coccinellini)

Coleomegilla maculata DeGeer (Coccinellini)

Naemia seriata (Melsheimer) (Coccinellini)

Eriopis connexa ((Germar) (Coccinellini)

Psyllobora vigintimaculata Say (Psylloborini)

Pupae were examined with a stereo-microscope with magnification from 6X to

200X. Pupae were submerged in a small dish containing alcohol with sandy subtrate

on which the specimens can be oriented easily. The background may be adjusted from

light to dark by using various colored sands which helps to reveal inconspicuous

characters such as setae. Illumination was provided by an adjustable light source

which could be varied from very dim to very bright. In addition, different colored light

filters were helpful, a deep blue filter being most useful. The dimensions of the pupae

were measured with an ocular micrometer.

6 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

COMPARATIVE MORPHOLOGY

General

Pupae of Coccinellidae are exarate with all appendages free. The larval exuvium

usually folds back and covers the anal end of the pupa (Figs. 1, 244). However, in

CHILOCORINI and NOVIINI the pupa is enclosed in an almost intact last larval

exuvium, broken only by a dorsolongitudinal slit through which the pupa can be seen

(Fig. 239); and in HYPERASPINI (Hyperaspis and Thalassa montezumae) the last

larval exuvium entirely covers the ventral surface of the body including the head.

The size may range from minute (1mm in length) as in Microweisea ovalis, to

moderately large (12mm or more in length) as in Anatis quindecimpunctata, Anatis

ocellata and Synonycha grandis. The body is usually slightly elongate oval. However,

in Coccinella, Olla, and Axion the body is very rounded-oval and strongly convex

dorsally; and in Hippodamia, Coleomegilla, Paranaemia, and Eriopis, the body is

elongate-oval.

The maculation pattern also varies a great deal among coccinellid pupae. However,

it is possible to distinguish two main groups. The first group consists of all members

of COCCINELLINAE (Coccinellini and Psylloborini) in which the dorsal surface of

the body including the elytra is usually distinctly marked with dark spots (brown,

dark brown or black) or with pale spots (yellowish). The second group consists of the

remaining species of the family in which the body is usually entirely pale, yellowish or

brownish or brown, with spots rarely present.

Setae and external texture of the body wall

All members of subfamily COCCINELLINAE have setae that are very short and

fine, with setal length never exceeding the thickness of the lateral margin of the

pronotum. The remaining members of the family are usually very densely and

coarsely setiferous, with the setal length greatly exceeding the thickness of the lateral

margin of the pronotum.

Setae may be very coarse and bristle-like (macrosetae) (Fig. 108) as in most

SCYMNINAE, CHILOCORINAE (Chilocorus) and EPILACHNINAE. Macrosetae

may be born on a flat cuticular ring (Figs. 108, 206, 241) as in Epilachna, Hyperaspis,

Cryptolaemus and Thalassa, or on a tubercle (Fig. 22, 110) as in Stethorus.

Microsetae, which are distinguished from macrosetae by being either more slender

or very short and fine, are present on most species. In the COCCINELLINAE

microsetae are usually borne on a slightly elevated cuticular ring with the tip either

slightly enlarged and truncated (Fig. 7, 8) as in Coccinella, Adalia, Hippodamia and

Psyllobora, or pointed as in Cycloneda, Anatis, Mulsantina, and Synonycha, or borne

on a small tubercle or chalaza with the distal end strongly curved as in Propylaea and

Anisocalvia (Figs. 172, 177). Very long and fine hair-like setae have only been found

on Orcus chalybeus (Fig. 29).

Ornamentation of the body surface (excluding ‘‘wrinkles’’) varies from being

entirely smooth as in all members of the STICHOLOTINAE, EPILACHNINAE, and

SCYMNINAE, and some members of the COCCINELLINI (Anatis and Synonycha),

to being slightly rough with the surface clothed with very slightly rounded bumps (Fig.

7) as in Cycloneda and Mulsantina. The body surface may also be covered with very

Phuoc and Stehr: Pupae of Coccinellidae rf

fine sharp spines as on the elytral surface of Coccinella, and Hippodamia, and on the

hind wing apex of Coccinella and Adalia (Figs. 8, 179-184). Spines may become more

conspicuous and larger as on Axion and Exochomus (Fig. 25).

Head

The head capsule of coccinellid pupae is of the opisthognathous type in which the

mouth parts are deflected in a posteroventral position. Epicranial sutures are nearly

obsolete or wanting but may be discerned as deep depressions in Coccinella and

Hippodamia. The frontal sutures are distinctly visible in Stethorus and Rodolia (Figs.

17-22, 50, 51). The cranial capsule is often well and homogeneously sclerotized except

in STETHORINI where the frontal area remains membranous and_ usually

protuberant (Figs. 20, 22).

The eyes do not vary much among coccinellid pupae. The inner lateral portion of

the ocular suture is well defined, but the eye facets are not well defined.

The antennae, in contrast, vary a great deal from one group to another. In most

cases, the antenna is long and may exceed the distance between the eye and the widest

lateral margin of the pronotum as in Propylaea quatuordecimpunctata, or Psyllobora

vigintimaculata (Figs. 44, 46). In others, it only extends to the widest lateral margin of

the pronotum (Figs. 9, 10, 32) as in Coccinella, Hippodamia, Anatis, Cycloneda,

Adalia, Microweisea and Epilachna. In a few cases the antenna is shorter, extending

less than half the distance between the eye and the widest lateral margin of the

pronotum (Figs. 12, 15, 19) as in Synonycha grandis, Hyperaspis, Zagloba ornata and

all members of the CHILOCORINI. The scape, flagellum and club of the antenna are

distinct in most cases, but segmentation is poorly developed.

The scape may be enlarged and expanded caudally with the anterior surtace

flattened as in Coleomegilla maculata, Naemia seriata, or strongly convex as in

Coccinella, Hippodamia and Olla. Only Cryptolaemus montrouziert (SCYMNINI)

and Epilachna sp. (EPILACHNINI) have a scape which is setiferous on the anterior

surface (Figs. 10, 31, 59).

The flagellum is usually slender, slightly elbowed as in Coccinella, Hippodamia,

and Anatis, or C-shaped as in Psyllobora and Propylaea. The proximal end of the

flagellum is greatly enlarged and somewhat “‘bisegmented”’ in Hyperaspis, Scymnus

nad Cryptolaemus, but is normal in all other species of the family. The distal end of

the antenna is usually recognizable as a club which is often broadly enlarged and

usually has four rings of well-developed papillae (Figs. 52, 53). The club may be said

to be ‘‘distinct’’ when the diameter is much greater than that of the flagellum (Figs.

52, 53) or “‘indistinct’”’ when the diameter of the club and the flagellum are subequal

(Figs. 55-57). In the latter case, the ‘‘club’”’ can be recognized by the presence of

papillae as in Anatis, Synonycha and Propylaea, but in Zagloba, Hyperaspis and the

CHILOCORINI the flagellum is short and tapered distally, with no papillae present

(Figs. 61-63).

The Clypeus and labrum are usually united into a clypeolabrum in which the

clypeolabral suture may be seen as a transverse arched depression. The clypeolabrum

is usually large, subquadrate, subhexagonal or trapezoidal, and usually as long as

wide (Figs. 9, 14, 32) except in Rhizobius ventralis, Rodolia cardinalis nad Psyllobora

vigintimaculata where it is twice as wide as long (Figs. 11, 46, 50). The apical margin

a. | Cénuib Amer Ent dase Volo N6 61974

may be truncated (Fig. 46) as in Psyllobora vigintimaculata, slightly convex as in

Anatis and Synonycha (Fig. 32), slightly to deeply concave as in Coccinella,

Cycloneda, Olla and Adalia, or deeply notched as in Eriopis connexa, and most

Hippodamia (Figs. 41, 49). The clypeus is greatly dilated laterally and conceals the

antennal bases only in the CHILOCORINI (Figs. 23-29).

The mandible is usually bifid and pointed at the tips as in COCCINELLINAE,

STICHOLOTINAE and SCYMNINAE, (the posterior tip of the mandible of the

latter is greatly reduced in size). In Epilachna the mandible is well developed for the

phytophagous habit, with the chewing tips angulated, truncated or pointed as seen

from the apex, and the mesal area broadly concave as in Figs. 72-78. In members of

the tribe CHILOCORINI, in contrast, the mandible is simple at the tip (Figs. 64-67).

Throughout the family the mola is usually rather well developed.

The maxilla is bulbous in appearance. The palpus is large, trapezoidal, or lan-

ceolate, and is glabrous except in Zagloba ornata and Cryptolaemus montrouzieri

where it is monosetose at the apex (Figs. 84, 85), and in EPILACHNINAE where the

palpus is densely setiferous (Figs. 9, 10, 83).

The galea is also bulbous, as viewed from the top, appearing subquadrate or

subpentagonal with a small hook-shaped lobe at the inner anterior angle caused by

the impression of the mandibular tip (Figs. 79-82). The galea is small, but greatly

enlarged in Psyllobora and Epilachna where the greatest width of the galea is

subequal to the base of the maxillary palpus (Figs. 83, 92). The galeal surface is

smooth, but may be armed with compact and sharp spinules in Anatis, Mulsantina,

Cycloneda, and Adalia (Figs. 79-82).

The lacinia is located beneath the galea, small and not as well defined.

The labium is bulbous with a distinct mentum. The labial palpi are large, stout,

and short, with the tip rounded except in Chilocorus and Epilachna. In Delphastus

the labial palpi appear more slender.

The hypopharynx is enlarged anteriorly as seen from the apex and is appressed

laterally by the large and bulbous maxillary galeae (Figs. 95, 100). The labium is

entirely glabrous, except in Epilachna where the labial palpi are monosetose at the

tip, bisetose at the lateral margins of the base, and monosetose at the base of the

ligula (Figs. 99, 100). In Hyperaspis (Figs. 96, 97) the anterior face of the ligula is

densely setiferous.

The thorax

The pronotum is large and immarginate as in Coccinella and STICHOLOTINAE,

or strongly marginate apically as in most members of the COCCINELLINAE except

Coccinella, Adalia, Hippodamia and Eriopis. In CHILOCORINI and Thalassa

montezumae (HYPERASPINI) the pronotum is greatly expanded laterally (par-

ticularly along the posterior margin), pushing the lateral margin of the pronotum

downward and forward, and in some cases (as in Axion) the lateral side may descend

considerably beyond the lower margin of the eye (Fig. 24), giving the pronotum as

seen from the top a crescent shape (Figs. 106, 107). In the COCCINELLINAE, the

pronotum is usually subquadrate with the anterior margin very slightly concave or

almost straight. In contrast, the remaining species in all groups (including Epilachna)

usually have the anterior margin of the pronotum deeply concave (Fig. 108). The

Phuoc and Stehr: Pupae of Coccinellidae 9

posterior margin of the pronotum is often rounded and convex (Figs. 105, 112, 115)

except in Stethorus where it is broadly tuberculated (Fig. 110), and in Scymnus where

it is expanded caudally into a rounded lobe (Fig. 109). The lateral margins of the

pronotum are straight or slightly concave in all cases, with the edge often rounded

except in Hippodamia where it is greatly expanded into a sharp knife-like edge (Fig.

133). A medio-longitudinal pale line is usually seen in COCCINELLINAE (Figs. 111-

140).

The mesonotum is usually trapezoidal in shape, with the anterior margin much

longer than the posterior one, and often spotted in the COCCINELLINAE (Fig. 1).

The scutellar area may be recognized as an elevated area at mid-base.

The metanotum is more or less trapezoidal with the posterior margin usually

straight except in HY PERASPINI where the margin is slightly sinuate (Fig. 236). The

metanotum is also often spotted in COCCINELLINAE.

The elytra are elongate oval or rectangular, bending ventrally to cover most of the

hind legs and abdominal sterna. The elytral area of coccinellid pupae may be divided

as in Figs. 5 and 6, in which the apex possesses the most significant characters. In

most cases the lateral angle is obtusely rounded and almost continuous with the

lateral margin (Fig. 5), but especially in Hippodamia (Figs. 6, 164-170), the lateral

angle sharply expands anteriorly into a well defined rounded lobe. The epipleuron is

usually wide and slightly concave. In COCCINELLINAE, the lateral margin of the

elytron is strongly marginate (Fig. 161) but in the rest of the family it is immarginate

(Fig. 162). The elytral surface is smooth except in some members of the COC-

CINELLINI where the surface is covered with microscopic rounded bumps

(micronodulated, Fig. 7), and in the CHILOCORINAE (especially in Axion,

Exochomus and Brumoides) where the elytral surface is obviously spinose (Fig. 25).

The hind wing is usually semi-sclerotized or membranous, bulbous and tapered

apically. The surface is always smooth and glabrous except in Coccinella and Adalia

where the apex is covered with very fine and sharp spines, (microspinulated, Figs.

179-184), and in the STICHOLOTINAE and COCCIDULINAE where the apex of

hind wing is finely setiferous (Figs. 232-234, 243).

The structure of the legs of coccinellid pupae is homogeneous throughout the family.

The articulations are poorly developed, with the femur and tibia apparently ‘‘fused”’

together along their margins. The tarsus is weakly segmented, but the terminal

segment is rather clearly defined by being more slender and cylindrical. Claws are

often undefined and obtuse except in Psyllobora vigintimaculata where the claws are

more distinct and pointed. In Exochomus the terminal segment is more slender, with

the diameter about one-fourth the dorsal length of the tarsus (similar to Fig. 104).

Only in Stethorus and Scymnus are the coxae more widely separated posteriorly, with

the distance between the procoxae being one-half as wide as that between the

metacoxae (Fig. 238).

The abdomen

Abdominal terga: In general, coccinellid pupae may be placed in two groups, one

group with broad intersegmental conjunctivae clearly visible between segments 4 and

5, 5 and 6, and 6 and 7 (Figs. 1, 244, 245). These intersegmental conjunctivae appear

to be as strongly sclerotized as the terga with the exception of a transverse line which

10 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

allows the conjunctiva to fold upon itself. This morphological adaptation is ex-

clusively in the COCCINELLINAE and allows the abdomen to move more freely. The

conjunctivae are finely setiferous except in Hippodamia where they are glabrous.

The second group consists of the remaining species of the family in which all terga

are closely opposed to one another without exposed conjunctivae (Figs. 240-243),

resulting in greatly reduced flexibility of the abdomen.

As mentioned above, the dorsum of the abdomen of COCCINELLINAE is

distinctly maculate, while the remaining species of the family are usually immaculate

and pale yellowish or brownish.

Urogomphi: Abdominal tergum 9 is usually modified into one pair of lateral cerci-

like processes or urogomphi which serve as grasping appendages to permit the pupa

to attach to the substrate within the last larval exuvium. Tergum 9 also bears a dorsal

median lobe or pygidium which is glabrous in COCCINELLINAE and

STETHORINI, and_ finely setiferous in CHILOCORINI, SCYMNINI,

HYPERASPINI, COCCIDULINI, SCYMNILLINI, SERANGIINI,

STICHOLOTINI, and EPILACHNINI. The shape of the urogomphi is strikingly

different from one group to other, except for the STICHOLOTINAE which have no

urogomphi (Figs. 195-198). The urogomphi may be long, slender and subcylindrical,

with the tip simple and obtusely rounded as in Epilachna and Scymnus (Figs. 199,

201). In Stethorus, the urogomphal tip is slightly flared into a flat subcircular

distal disk (Figs. 210, 211). In Lindorus lophantae the tip is tapered to a point (Fig.

202). In another case restricted to the COCCINELLINAE, the urogomphi appear less

slender, with the distal end twisted and folded ventroanterad into a bilobed distal disk

(Figs. 218-229) as seen ventrally. The inner lateral side of the urogomphus is usually

straight except in Anatis, Synonycha, Anisocalvia and Neoharmonia where a spine-

like process (Fig. 223) is present. In HYPERASPINI the urogomphi are quite short,

with the distal disk very well developed, sclerotized and double parenthesis-shaped as

seen ventrally (Figs. 207,208). In Cryptolaemus montrouzieri, the urogomphi branch

at the distal one-third into a large sausage-like process (Figs. 205, 206), which is quite

different from Scymnus where the urogomphi are unbranched. The urogomphi of

CHILOCORINI are very characteristic, with either single or double branches and

with the distal end greatly enlarged into rounded mushroom-shaped tips (Figs. 215-

217).

Abdominal pleura: The structure of the abdominal pleura is very homogeneous in

coccinellid pupae. They often are greatly expanded laterally into subquadrate or

rhomboidal plates, with the lateral margin usually simple, straight, or slightly

convex (Figs. 171, 172, 174) except in Evriopis connexa, where the lateral margin of

pleura 3, 4, and S is slightly angulate (Fig. 175), and in Anisocalvia, Anatis and

Synonycha where the posteriolateral angle projects laterocaudad into a conical or

long, cylindrical spine-like process (Figs. 176-178). In Neoharmonia venusta the

posteriolateral angle of the pleura expands slightly caudally on segments 3, 4 and 5

(Fig. 171). This is intermediate between the ‘‘Anatis type’ and the undifferentiated

type. Pleura 1 and 2 are always hidden under the elytra and pleuron 9 is often fused

with the base of the urogomphus. In HYPERASPINI, unlike all other groups, the

abdominal pleura strongly curve ventrally, and thus are invisible from above (Fig.

236).

Phuoc and Stehr: Pupae of Coccinellidae 11

Abdominal sterna: The structure of the abdominal sterna is also very homogeneous

among coccinellid pupae. There are usually nine, however, the first two are greatly

reduced in size and hidden beneath the hind coxae except in a few cases where they

tirely flat and small, whereas in the ° it is bipartite (Figs. 215, 218), and mammillate

with the tip of the mammilla usually more sclerotized than the base. In SERANGIINI

and STICHOLOTINI the tip is very large (Figs. 195-198).

The Spiracles

The prothoracic spiracles are elongate oval in most cases, or rounded as in Adalia

bipunctata and Eriopis connexa.

The abdominal spiracles are located on the anterolateral angle of the abdominal

tergum and usually differ from group to group, except for members of tribe

HYPERASPINI, in which the spiracles are wanting (Fig. 236). The different types of

spiracles can be distinguished as follows:

— In EPILACHNINI all abdominal spiracles are slightly and subequally

pedunculate, with the peduncles semisclerotized or membranous (Fig. 235). This

type of spiracle is also present in the COCCIDULINI and Chilocorus; however, the

peduncle progressively decreases in length as the spiracles approach the posterior end

of the abdomen (Figs. 234, 242).

— In CHILOCORINI and STETHORINI, only the first abdominal spiracles are

pedunculate, with the peduncle very long, conical and strongly sclerotized in

CHILOCORINI (Figs. 185-189, 240, 242), or very slender, usually cylindrical, and

semisclerotized or membranous in STETHORINI (Figs. 190-192). The remaining

abdominal spiracles of these two tribes are without peduncles (Fig. 240).

— Also, in the CHILOCORINI, (Axion, Exochomus, Brumoides), in addition to

the peduculate spiracles, one pair of pit-like gland openings is present between the

anterior margin of the first abdominal tergum and the posterior margin of the

metanotum (Figs. 187, 239, 240). These are absent in Chilocorus and Orcus.

— Finally, the most common type of spiracle is observed in SCYMNINI,

SCYMNILLINI, SERANGIINI, STICHOLOTINI, COCCINELLINI and

PSYLLOBORINI where the spiracular peduncle is wanting, the opening is usually

elongate oval or circular,and usually well sclerotized (Figs. 1, 244, 245). The pair of

spiracles on the first abdominal segment is much larger than the rest, and is mostly

hidden beneath the elytron (Figs. 1, 244, 245).

KEY TO SUBFAMILIES AND TRIBES OF THE

AVAILABLE PUPAE OF COCCINELLIDAE

1 Abdomen flexible with intersegmental conjunctivae of abdominal

3 to 6 exposed (Figs. 244, 245); body finely setiferous, setae shorter

than the thickness of the lateral margin of the pronotum; dorsal

surface distinctly maculate (Figs. 1, 244, 245),

CC Ac kG Ce EN EHR OT CE RAG ow ROME eS 2

2(1)

3(1’)

3’

4(3’)

S(4’)

eS?

6(S)

6’

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Abdomen compact, without intersegmental conjunctivae exposed

(Figs. 240-243); body coarsely setiferous with various types of setae,

the longest setae considerably longer than the thickness of the

lateral margin of the pronotum (Figs. 9-22); dorsal surface

ically WimachInse Ss re eee as 3

Clypeolabrum usually as long as wide (Figs. 32-37); galea small, with

the greatest width one-half as wide as the base of the maxillary

palpus (Figs. 90, 91); size medium to large, over 5 mm in length

Pe a Pe ee en COCCINELLINI

Clypeolabrum much wider than long (Fig. 46); galea greatly

enlarged, with the greatest width as wide as the base of the

maxillary palpus (Fig. 92); size small, less than Smm

iM length ee eS ea a PSYLLOBORINI

Mandibles greatly enlarged with chewing tip broadly concave

mesally (Figs. 72-78); maxillary palpi densely setiferous (Figs. 9,

10, 83); abdominal spiracles subequally pedunculate (Fig. 235)

PPILACHNINAE Jo) Sher re Oe EPILACHNINI

Mandibles not enlarged, tip pointed, simple or bifid, maxillary

palpi glabrous or monosetose (Figs. 12, 31, 84, 85); abdominal

spiracles variable but never subequally pedunculate............... 4

Clypeus broadly expanded laterally, concealing antennal bases

(Figs. 23-29); mandible simple at tip (Figs. 64-67); pupa usually

enclosed in the last larval exuvium; and visible through a dorso-

longitudinal slit (Fig. 239),

CHIPOGCORINAB ee a Oe CHILOCORINI

Clypeus normal, antennal bases visible (Figs. 11-22); mandibles

usually bifid at tip (Figs. 68-71); pupa usually free from last larval

SxUviat Fig, 234) ON ee ee Le a 5

Hind wing apex finely setiferous dorsally; setae fine

SNC THM 2G eee. ee ee Se ee ee 6

Hind wing apex glabrous; setae coarse or bristle-like,

SCYMBINAR o250 8) os a a eee i eas 10

Urogomphi wanting (Figs. 195-198); clypeolabrum as long as or

longer than wide (Figs. 13, 14), STICHOLOTINAE .............. ,

Urogomphi well developed (Figs. 11, 12, 50); clypeolabrum wider

fam lone, COCCIDULINAE. ka ea ees ee ek ee) 8

7(6)

8(6’)

9(8)

10(S’)

10’

11(10’)

11’

Phuoc and Stehr: Pupae of Coccinellidae 13

Pronotum with a dorsolongitudinal depression (Fig. 13); abdominal

pleura glabrous; posterior margin of pygidium deeply concave

(Figs. 195-196), Delphastus pusillus (Leconte)

wei te. PEG, Sa ees a ee ees bee SERANGIINI

Pronotum without such a dorsolongitudinal depression; abdominal

pleura setiferous; posterior margin of pygidium subparallel with

anterior margin (Figs. 197-198), Microweisea ovalis (Leconte) ........

ik ick ee ne ee STICHOLOTINI

Abdominal pleura 3 to 5 greatly expanded laterally with lateral

margin strongly convex (Fig. 234); dorsum of abdominal tergum

mon-tuberculate (Pig’ 204) he ee 9

Abdominal pleura 3 to 5 not expanded laterally, subquadrate

(Fig. 243); dorsum of abdominal terga each with one transverse

pair of tubercles (Fig. 243), Rodolia cardinalis Mulsant

Ol ON Be eee ae ee ee ee ee ee NOVIINI

Antennae tapered distally (Figs. 12, 61) and short, extending toa

point about midway between the eye and the widest lateral

margin of the pronotum (Fig. 12); first four abdominal spiracles

normal; maxillary palpus monosetose (Figs. 12, 84),

ZOgIGRa OfHATA TION) oe ee es es SCYMNILLINI

Antennae enlarged distally into a distinct club with well developed

papillae (Fig. 11), and extending to the widest lateral margin of

the pronotum (Fig. 11); first four abdominal spiracles slightly

pedunculate (Fig. 234); maxillary palpus glabrous...

FO Mii eka heli os a ee ca COCCIDULINI

Head with frontal area membranous and usually protuberant

(Figs. 20, 22); first abdominal spiracles pedunculate (Figs. 190-

192 237) femoata SlabvOusy sy cei oe eas een STETHORINI

Head with frontal area as sclerotized as the other areas; first

abdominal spiracles not pedunculate or obsolete;

fEIMOPA SOPMOHOUS. Ghiticcck dices SUN ect ha wh Ce ih oe eae 11

Abdominal pleura visible from above, abdominal spiracles

present; antennal club distinct (Figs. 31, 58, 59)

Fy wet a a. CRE Gt ALE Gat ee a nC ee ene ry SCYMNINI

Abdominal pleura not visible from above (Fig..236); abdominal

spiracles absent; antennal club indistinct

PIGS OTN ck a ee pees vere HYPERASPINI

14 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

SUBFAMILY STICHOLOTINAE

Diagnosis

This is the only known group of pupae of Coccinellidae which lack urogomphi.

However, they appear to have a strong affinity to Coccidulinae by having the hind

wing apex finely setiferous dorsally on the lateral angle (Figs. 232-234). Moreover, the

simple mandibular tip allies the Sticholotinae with members of Chilocorinae

(Chilocorini) which have also retained this primitive type of mandible.

TRIBE SERANGIINI

Genus DELPHASTUS Casey

Specimens examined

The study was based on three pupae of Delphastus pusillus from the U.S. National

Museum, collected in Havana, Cuba, 29 September 1928 by Brinner.

Diagnosis

Serangiini and Sticholotini are the only two tribes for which pupae were available in

the subfamily Sticholotinae. They are usually minute in size (about 1mm to 1.50mm

in length). Serangiini can be recognized by the dorsolongitudinal depressed line on

the pronotum (Figs. 13, 233), the abdominal pleura and terga 3 to 5 are glabrous,

and the posterior margin of the pygidium is deeply notched. In the Sticholotini the

dorsum of the pronotum is convex, all abdominal terga and pleura are uniformly

setiferous, and the anterior and posterior margins of the pygidium are subparallel.

Delphastus pusillus (Leconte)

(Figs. 13, 60, 87, 101, 195, 196, 233)

Description

Length: 1.5mm; width: Imm. Body small, rounded oval, yellowish, immaculate,

and densely setiferous dorsally. Head pale, setiferous. Antennae long with club

greatly enlarged and slightly compressed dorsoventrally, with 3 rings of poorly

developed papillae; antennal scape and the last 2 ‘‘segments’’ of flagellum somewhat

papillated and enlarged (Fig. 13). Clypeolabrum narrow, with lateral sides sub-

parallel, greatly narrowed apically, and apical margin slightly concave (Fig. 13).

Mandibles simple at tip. Maxillary palpi long, slender, somewhat cylindrical (about 3

times or more longer than wide), and slightly curved inward at base (Fig. 13).

Pronotum elongate oval as seen from above, with a wide dorsolongitudinal

depression line (Figs. 13, 233). Metanotum large and as long as the first two ab-

dominal terga combined (Fig. 242). Elytra pale yellowish, immaculate, and densely

setiferous. Hind wings semi-membranous with dorsal surface of lateral apical angle

densely setiferous. Legs short and rather robust.

Phuoc and Stehr: Pupae of Coccinellidae 15

Abdominal terga yellowish, immaculate, closed tightly together, with the first two

terga and terga 6 to 9 setiferous, whereas terga 3 to 5 are very finely setiferous or

apparently glabrous. Pygidium small with posterior margin deeply notched.

Urogomphi wanting (Figs. 195, 196). Abdominal pleura subquadrate, usually in a

verticle position and glabrous. Abdominal sterum 9 of the ° characteristically

modified with a pair of greatly elongate and conical “‘gonopods’’ (Fig. 196). In the o ,

the sternum remains flat and small (Fig. 195).

TRIBE STICHOLOTINI

Genus MICROWEISEA Cockerell

Specimens examined

The study was based on 2 pupae of Microweisea ovalis from the U.S. National

Museum, collected in New Orleans, Louisiana, on 13 July 1923.

Diagnosis

See Delphastus pusillus for the separation of these two tribes.

Microweisea ovalis (Leconte)

(Figs. 14, 197, 198, 232)

Description

Length: 1.25mm; width: 0.75mm. Body pale brownish, immaculate and apparently

setiferous. Very similar to Delphastus pusillus.

Head longer than wide. Antennae long, extending to the widest lateral margin of

the pronotum; club enlarged and with well developed papillae; flagellum cylindrical

and slender. Clypeolabrum subtriangular, with apex very narrow and truncate (Fig.

14). Mandibles simple at tip. Maxillary palpi slender, very long and cylindrical (Fig.

14).

Pronotum nonmarginate and densely setiferous. Elytron brownish, and con-

spicuously and densely setiferous. Dorsal side of abdomen uniformly and densely

setiferous. Pygidium small and subrectangular. Abdominal pleura subquadrate, with

the 3rd and 4th pleura conspicuously and densely setiferous. Venter of the abdomen

entirely pale and sparsely setiferous. In the 2 , the ‘“‘gonopods’’ (9th abdominal

sternum) are conical and greatly elongate (Fig. 198). Sternum 9 of the o” is short and

inconspicuous (Fig. 197).

SUBFAMILY COCCIDULINAE

Diagnosis

Coccidulinae are separated from Sticholotinae by the presence of one pair of

prominent urogomphi and by abdominal pleura 3 to 6 being greatly expanded

16° Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

laterally, giving the lateral margin of the pleuron a strongly convex appearance (Fig.

234). These two subfamilies in turn are distinguished from the remaining groups of

Coccinellidae by the characteristically setiferous hind wing apex (Figs. 232-234).

TRIBE COCCIDULINI

Diagnosis

The Coccidulini is the only group of pupae with at least the first four pairs of

abdominal spiracles slightly pedunculate (Fig. 234). This character is also observed in

Epilachnini, but in the Epilachnini all abdominal spiracles except those on segment 9

are slightly and subequally peduculate. The above character separates Coccidulini

from Scymnillini whose abdominal spiracles are circular or nearly so (= not

pedunculate). Furthermore, the long antennae with a distinct papillated club, and the

slender urogomphi with the tip unevenly pointed, are very characteristic of this tribe.

In the Scymnillini, the antennae appear short with an indistinct club, papillae are

entirely absent, and the urogomphi have enlarged tips (Figs. 12, 203, 204).

Key to Genera of the Available Pupae of Coccidulini

1 Body very densely setiferous, dorsum nearly clothed with

short, thin, semi-erect setae whose tips strongly

curve back to the body surface; macrosetae (long

setae) sparsely distributed; pygidium shorter than

aodominaltersum (06... 2. Rhizobius ventralis (Erichson)

1s Body less densely setiferous, setae erect with tip not

curved back to body surface; pygidium as long as

qodominal teroudi To ee Lindorus lophantae Blaisdell

Genus LINDORUS Casey

Lindorus lophantae (Blaisdell)

(Figs. 11, 202, 234)

Specimens examined

The study was based on four pupae of Lindorus lophantae from the U.S. National

Museum, reared by the Bermuda Department of Agriculture, 13 June 1952.

Diagnosis

This species, in general, is very easily separated from Delphastus pusillus and

Microweisea ovalis by possessing very prominent and slender urogomphi (Fig. 202),

and by the clypeolabrum being much wider than long (Fig. 11). In Delphastus and

Microweisea the urogomphi are wanting, and the clypeolabrum is longer than wide

(Figs. 13, 14, 195-198).

Phuoc and Stehr: Pupae of Coccinellidae 17

Description

Length: 2mm; width: 1mm. Body elongate oval, densely setiferous, pale and im-

maculate. Head wide, clypeolabrum wider than long with apex slightly convex (Fig.

13). Antennae long, extending to the widest lateral margin of the pronotum; club

distinct and large, elongate cylindrical, with 4 rings of well developed papillae.

Mandibles unequally bifid at tip.

Pronotum pale or brownish, densely setiferous both dorsally and ventrally, and

immarginate with the anterior margin concave. Posterior margin of metanotum

slightly sinuate (Fig. 234). Elytron pale and setiferous (except brownish in one

specimen); lateral margin immarginate. Hind wings membranous, gradually tapering

toward apex where dorsal surface is setiferous (Fig. 234). Legs short and robust with

femora densely setiferous.

Abdominal terga subequal in length, each with 2 pairs of conspicuous groups of

setae located at spiracular and dorsal areas. Spiracular group consisting of four

conspicuous setae, whereas dorsal group consists of three (Fig. 234). Pygidium large,

trapezoidal, setiferous and as long as the 7th abdominal tergum. Urogomphi strongly

sclerotized, extremely slender and cylindrical, tapering, and pointed apically (Fig.

202). First 4 abdominal spiracles slightly pedunculate (Fig. 234). Abdominal pleura 1

and 2 hidden under the elytron, pleura 3 to 6 greatly expanded laterally, with lateral

margin strongly convex and conspicuously setiferous (Fig. 234). Abdominal sterna

pale, densely setiferous along posteriomarginal areas. Sternum 8 small, about one-

half as long as sternum 7.

Genus RHIZOBIUS Stephens

Rhizobius ventralis (Erichson)

Specimens examined

The study was based on two pupae of Rhizobius ventralis from the U.S. National

Museum collected in California on 15 June 1896.

Description and diagnosis

Very similar to Lindorus lophantae in many respects except for the following

characters:

Length: 3.5-4mm; width: 1.5-1.8mm. Body more densely setiferous, with setae

semi-erect and with the distal ends strongly curved back to the body surface.

However, because of the poor condition of the specimens, it is very difficult to

distinguish this species from Lindorus lophantae in terms of chaetotaxy. Pygidium

much shorter than abdominal tergum 7.

18 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

TRIBE SCYMNILLINI

Genus ZAGLOBA Casey

Specimens examined

The study was based on a single pupa of Zagloba ornata from the U.S. National

Museum reared by the Bermuda Department of Agriculture from California stock, 13

June 1952.

Diagnosis

See diagnosis of Coccidulini for the separation of these two tribes.

Zagloba ornata (Horn)

(Figs. 12, 61, 84, 102, 203, 204)

Description

Length: 2.5mm; width: 1.5mm. Body pale brownish, very densely setiferous,

similar to Lindorus lophantae except for the following characters:

Head pale yellowish, slightly longer than wide (Fig. 12). Antennae short, extending

to about one-fourth the distance between the eye and the widest lateral side of the

pronotum; scape and the first two ‘‘segments”’ of flagellum greatly enlarged (much

larger than distal portion which is somewhat cylindrical and rounded at tip) (Fig. 12,

61). Clypeolabrum subquadrate and sparsely setiferous, apical margin slightly

concave (Fig. 12). Mandible unequally bifid at tip. Maxillary palpi long and cylin-

drical, with the apex rounded and monosetose (Figs. 12, 84).

Pronotum and mesonotum slightly darker (brownish) than any other areas.

Abdomen appearing more densely setiferous than Lindorus. Abdominal spiracles

subcircular and normal (without peduncle). Abdominal pleura 3, 4, and 5 greatly

expanded laterally, rounded and densely setiferous (Fig. 234). Urogomphi short (as

long as pygidium), with distal end slightly enlarged and curved ventrally (Figs. 203,

204).

TRIBE NOVIINI

Genus RODOLIA Mulsant

Specimens examined

The study was based on 8 pupae of Rodolia cardinalis collected in Gainesville,

Florida, on 14 July 1972 by R. E. Waites. Two pupae are deposited in the Entomology

Museum of Michigan State University, the remainder in the Department of En-

tomology and Nematology, University of Florida.

Phuoc and Stehr: Pupae of Coccinellidae 19

Diagnosis

Noviini can be separated from Coccidulini (=Rhizobiini) and Scymnillini by the

subvertical position of the abdominal subquadrate pleura whose lateral margins are

usually straight; and by the two-branched uorgomphi. In Coccidulini and Scym-

nillini, the abdominal pleura are usually greatly expanded laterally, are horizontal in

position with strongly convex lateral margins. The urogomphal apices are pointed or

slightly enlarged. The Noviini are placed in the subfamily Coccidulinae with the

Coccidulini and Scymnillini because they possess the distinctive finely setiferous hind

wing apex.

Rodolia cardinalis Mulsant

(Figs. 50, 51, 230, 231, 243)

Description

Length: 4-4.5mm; width: 2.5-3mm. Body elongate oval. The abdomen is sub-

rounded in cross section, retaining much of the larval aspect and is twice as long as

the thorax. Dorsum yellowish to brownish, with undefined brown spots on mid-dorsal

area of abdominal terga where a slight depression is located. In general, the body is

densely setiferous with the setae on the pronotum and the dorsum of the abdomen

appearing coarser than on the other areas of the body surface. The body is almost

enclosed in the last larval exuvia.

Head broad, as wide as long. Antennae short, extending slightly beyond the outer

lateral margin of the eye, without papillae. Club indistinct from the flagellum. Scape

enlarged and rounded (Fig. 50). Clypeolabrum large, much wider than long, with the

apical margin slightly concave (Fig. 50). Mandibles unequally bifid at the apex, with

the posterior tip greatly reduced in size. Maxillary palpi with the apex enlarged.

Pronotum immarginate, mostly pale except for one pair of pooly defined brownish

discal spots. Surface of the pronotum densely setiferous, and setae adjacent to discal

spots strongly pigmented. Meso- and metanotum brown except scutellar area on

mesonotum pale yellowish. Elytron elongate oval, pale whitish except for brownish

basal and scutellar areas. Lateral margin immarginate. Epipleura sparsely setiferous

and slightly concave. Hind wing apex finely setiferous dorsally, especially along

lateral margin. Legs short and robust with femora densely and finely setiferous. |

Abdominal terga yellowish to brownish. Each tergum usually with a dorsally joined

pair of depressions where the coloration is darker than other areas, and one pair of

transverse-banded tubercles running the tergal width, on which setae are very dense.

Tubercles much reduced in size or wanting in terga 6 to 9. Urogomphi bipartite, the

outer branch with apex simple, the inner branch shorter but with apex modified into

a parenthesis-shaped distal disk (Figs. 230, 231). This character indicates some

relationships with Hyperaspini (SCYMNINAE) and Chilocorini (CHILOCORINAE)

whose urogomphi are usually bipartite. Abdominal spiracles small, inconspicuous

and normal (nonpedunculate). Abdominal pleura subquadrate and in a subvertical

position. Abdominal sterna entirely pale.

20 Contribs Nabe. "Bat It. Vols 10/No.6, £974

SUBFAMILY EPILACHNINAE

TRIBE EPILACHNINI

Diagnosis

This is the only group of coccinellid pupae in which the chewing tip of the man-

dibles is distinctly and broadly concave mesally (Figs. 72-78). The maxillary palpi are

densely setiferous (Figs. 9, 10, 83), and the abdominal spiracles are membranous and

slightly pedunculate (Fig. 235).

Genus EPILACHNA Chevrolat

Description

Length: 7.5mm-8.00mm; width: 4.5-Smm. Body moderately large, pale yellowish,

usually immaculate and densely setiferous.

Head pale, as long as wide. Antennae (Fig. 9) long, extending to the widest lateral

margin of the pronotum. Club monosetose distally, and indistinct, with three or four

rings of papillae (Fig. 57). Flagellum long, subquadrate in cross section and slightly

elbowed or C-shaped. Clypeolabrum narrow, subtrapezoidal with the apical margin

concave (Fig. 9). Mandible large, visible without removing the clypeolabrum; the

chewing tip bifid as seen from the top and distinctly and broadly concave mesally

(Figs. 72-78). Maxillary palpi densely setiferous (Fig. 83).

Pronotum nearly twice as wide as long, the posterior margin arcuate and the an-

terior margin deeply concave. Macrosetae dense but confined to the margins of the

pronotum (Fig. 108). Elytron pale, immaculate, and three times as long as wide.

Epipleura glabrous and wide. Hind wing pale and entirely glabrous.

Abdomen compact, usually immaculate and pale, with macrosetae somewhat

sparsely distributed on dorsal surface. In contrast, the microsetae are more densely

distributed over the rest of the body. Pygidium small and finely setiferous.

Urogomphi slender, subcylindrical and straight at apex (Fig. 199). All abdominal

spiracles distinctly pedunculate (with the peduncle usually as long as the diameter of

the spiracular opening).

Key to Species of the Available Pupae of Epilachna

1 Anterior surface of the antennal scape setiferous (Fig. 10);

both anterior and posterior mandibular tips pointed as

Seer Wom tie ADEN Pig. TI ey ee Epilachna sp.

iss Anterior surface of the antennal scape glabrous

(Figs. 9,57); anterior mandibular tip truncate as seen

frourthe apex (Fics: 74, 78) ee ee ee eS 2

Phuoc and Stehr: Pupae of Coccinellidae zy

2(1’) Posterior mandibular tip sharply pointed as seen from the

apex (Fig. 78); one pair of distinct eye-like brown spots

present On. AodOiMaltetea tho oe ae ee

pias Posterior mandibular tip angulate or truncate as seen from

the apex (Fig. 74); abdominal terga 3 to 5 immaculate

aa SC eye Oe Epilachna varivestis Mulsant

Epilachna borealis (Fabricius)

(Figs. 9,57; 75, 78)

Specimens examined

The study was based on a single pupa associated with adults, from the U.S.

National Museum collected in Quincy, Florida 9 August 1944.

Diagnosis

Epilachna borealis has one pair of eye-like brownish spots per segment on ab-

dominal terga 3 to 5 and only the posterior tip of the mandible is pointed as seen from

the apex. In Epilachna sp. both the posterior and anterior mandibular tips are

pointed, and in Epilachna varivestis, the posterior mandibular tip is truncate or

angulate as seen from the apex. The last two species are usually entirely pale dorsally.

Description

Length: 7mm; width: 5mm. Body pale yellowish, broadly oval with macrosetae

sparsely distributed on dorsal surface.

Head entirely pale yellowish. Antennae long, with flagellum C-shaped and scape

distinct and glabrous. Mandible with the posterior tip pointed and the anterior tip

truncate as seen from the apex (Fig. 78). Dorsum of abdomen _ sparsely

macrosetiferous. Abdominal terga pale and immaculate except terga 1 and 2 which

have a brownish posterior margin, and terga 3 to S which have one pair of brown eye-

like spots.

Epilachna varivestis Mulsant

(Figs. 72-74, 83, 99, 100, 108, 173, 199, 200, 235)

Specimens examined

The study was based on 8 pupae from the U.S. National Museum, | pupa collected

in Tippecanoe Co., Indiana, on 4 September 1970 by L. Matteson and 2 pupae

collected in Michigan in August 1972 by D. C. Cress. The last two pupae are

deposited in Entomology Museum at Michigan State University, East Lansing,

Michigan.

22 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Diagnosis

Epilachna varivestis is characterized by the truncate or angular posterior man-

dibular tip as seen from the apex (Fig. 74). This differs from the other two available

Epilachna pupae (Epilachna borealis and Epilachna sp.) whose posterior mandibular

tip is sharply pointed (Figs. 77, 78).

Description

Length: 6.5-7mm; width: 4.5mm. In general this species is very similar to E.

borealis except that the body of E. varivestis is usually pale yellowish, and abdominal

terga 3 to 5 are entirely immaculate. In addition, the number of spiracular

macrosetae on abdominal segments 1-3 varied from 4 to 6, and the structure of the tip

of the mandibles is strikingly different from E. borealis by having an angulate

posterior mandibular tip as viewed from the apex (Fig. 74).

Epilachna sp.*

(Figs. 10, 76, 77)

Specimens examined

The study was based on a single pupa from the U.S. National Museum, apparently

from Brazil.

Diagnosis

This is the only known species among the available Epilachna pupae whose an-

tennal scape is setiferous. In E. varivestis and E. borealis the antennal scape is en-

tirely glabrous.

Description

Length: 7mm; width: Smm.

Body densely macrosetiferous, immaculate and shiny brownish. Antennae long,

with four rather well-developed papillae. Antennal scape bisetose or trisetose on

anterior surface (Fig. 10). Clypeolabrum trapezoidal with the apical margin truncate

or slightly concave and densely setiferous. Mandible unequally bifid with anterior and

posterior tips pointed as seen from the apex; the posterior tip twice as large as the

anterior one (Fig. 77). Maxillary palpi more densely setiferous toward outer lateral

margin. Macrosetae dense and confined cephalad of abdominal tergum 5.

SUBFAMILY SCYMNINAE

TRIBE HYPERASPINI

Diagnosis

The members of this tribe possess very short antennae with the distal end tapering,

pointed and lacking papillae (Fig. 62). Moreover, the Hyperaspini may additionally be

separated from the Scymnini by the lack of abdominal spiracles and the short

urogomphi, with the distal disk strongly sclerotized and double parenthesis-shaped as

seen ventrally (Figs. 207-208).

* A new species being described by Robert Gordon, Systematic Entomology Laboratory, USDA, c/o U.S.

National Museum, Washington, D.C. 20560 in a USDA Tech. Bull., 1975.

Phuoc and Stehr: Pupae of Coccinellidae 23

Key to Genera of the Available Pupae of Hyperaspini

1 Body large, over 6mm; Pronotum greatly expanded laterally,

pushing the posteriolateral margin downward beyond lower margins

of the eyes (Fig. 16); Macrosetae only fringing marginal

areas of the pronotum, elytra and abdomen (Fig. 236).

(Thalassa montezumae Mulsant) ................ Thalassa Mulsant

e Body small, under 5mm. in length; Pronotum not expanded

laterally, with posteriolateral margin above the lower

margins of the eyes (Fig. 15); Macrosetae more

densely and uniformly distributed over entire dorsal surface

of the pupa, except for a few species with

discal area of the pronotum finely setiferous

RRS) FEA SSE Ote ee roe SE Bee Hyperaspis Chevrolat

Genus HYPERASPIS Chevrolat

Diagnosis

This genus is distinctly separated from Thalassa montezumae by its smaller size

(under Smm.), and by the densely and uniformly macrosetiferous dorsal surface of the

body, especially on the elytra and the abdomen. In Thalassa the body appears

glabrous except for the macrosetiferous marginal areas. Thalassa are over Smm. in

length.

Description

Length 2.5-4mm; width: 1.2-2mm. Body broadly rounded-oval, dorsum pale

yellowish to brownish and densely setiferous with three types of setae. The first type of

seta is the bristle-like macroseta. The dorsum of the pupa is usually densely and

uniformily macrosetiferous except for a few species whose discal area of the pronotum

has sparser macrosetae. The second and third types of setae are called microsetae.

The second type may be as long as the first, but more slender and finer. The third type

is very short, with the diameter slightly greater than the second type, and about one-

tenth as long.

Head usually glabrous and pale except for the macrosetiferous vertex (Figs. 246-

248). However, in Hyperaspis binotata and Hyperaspis lateralis, the entire head is

densely macrosetiferous (Figs. 15, 249). Antennae short, extending to about one-third

the distance between the eye and the widest lateral margin of the pronotum, with

distal half subcylindrical and tapering toward apex with no papillae present (Fig. 62).

Clypeolabrum gradually narrowed apically with apical margin slightly concave.

Mandibles unequally bifid at tip with the posterior tip considerably reduced in size

(Figs. 68, 69).

Pronotum yellowish, immarginate; macrosetae may be uniformly and densely

distributed over the dorsal surface of the pronotum as in the case of H. lateralis, H.

24 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

binotata and H. quadrivittata (Fig. 263), or they may appear sparser or absent as in

H. oculaticauda and H. quadrioculata. Posterior margin of metanotum slightly

sinuate. Lateral margin of the elytron immarginate; epipleura wide and glabrous.

Legs robust and short, with the anterior surface of pro- and mesofemora usually

microsetiferous except in H. lateralis and H. binotata whose pro- and mesofemora are

macro- and miicrosetiferous (Figs. 15, 249). All abdominal terga densely

macrosetiferous. Urogomphi very short, flattened at base; distal disk strongly

sclerotized, consisting of a large kidney- to crescent-shaped outer lobe and a small,

elongate oval- to comma-shaped inner lobe as seen ventrally (Figs. 207, 250-256). In

H. quadrivittata, the inner lobe is wanting and the outer lobe is greatly reduced in size

and rounded (Figs. 257-258). Abdominal spiracles absent, abdominal pleura

subquadrate and in a vertical position, thus invisible from above. Abdominal sterna

pale and finely setiferous.

Key to Species of the Available Pupae of Hyperaspis

1 Head densely macrosetiferous including clypeolabrum

(Fig. 249); macrosetae present on anterior surface of

pro; ad Mes@iemora 2 6 ee eae. 8

1’ Head apparently glabrous except for the macrosetiferous

vertex (Figs. 246-248); macrosetae absent on anterior surface

Op OO aNd MesoremOla 9 ee ee ee 2

2(1’) Pyeiciunl MaCrOsetilerOUS 2. ee Coe re 3

z Pyerdsiin apparenily ClabrOuUs oo ee oe ve ee eh A

3(2) Macrosetae absent on discal area of the pronotum (Figs. 264-265);

lateroapical margin of the elytron macrosetiferous (Fig. 259);

inner lobe of urogomphus elongate oval, outer lobe large

and subrectanoular (rig. 251)... yes Hyperaspis octavia Casey

a Macrosetae uniformly distributed on the pronotum (Fig. 263);

lateroapical margin of the elytron finely setiferous (Figs. 260-262);

inner lobe of urogomphus absent, outer lobe small

and subrounded or kidney-shaped (Figs. 257-258)

ieciy gages ee a OU ie ate t es Hyperaspis quadrivittata Leconte

4(2’) Macrosetae slender and longer than the length of

BDGCMMN A Per CN LS oe oe ee a ae es acm cee can seieeia 2): )

4’ Macrosetae stout and shorter than the length of

POCORN TOROU I Lo Bae a as Pe eee ee eas ea a fi

S(4) Posterior margin of the pronotum apparently glabrous

A DD) ue a a as Hyperaspis quadrioculata (Motschulsky)

Phuoc and Stehr: Pupae of Coccinellidae 25

s Posterior margin of the pronotum macrosetiferous

(iy 204) ee. Oe eee ee 6

6(S’) Macrosetae apparently arranged in a single transverse

row on the vertex (Fig. 246); macrosetae on elytron

gradually becoming more slender toward the sutural, scutellar

and basal areas (Fig, 260) 5 no. caw. Hyperaspis oculaticauda Casey

6’ Macrosetae apparently arranged in two transverse rows on the

vertex (Fig. 247); macrosetae on elytron subequal in size

(Pig: 262) ac. ol Belews ws eee Sarl Hyperaspis postica Leconte

7(4’) Excluding apical area, macrosetae uniformly distributed over

the sutiace of the eWiton. . ick wucade ss. Hyperaspis cincta Leconte

7 Excluding apical area, macrosetae absent or sparse on

median area at the basal one-fifth of the elytron

MI 200) es ees oy oe oe a Hyperaspis psyche Casey

8(1) Frontal area densely macrosetiferous (Fig. 15); macrosetae

covering entire surface of abdominal pleura 4 to 7

Jk ees ieee a ee Hyperaspis binotata (Say)

8’ Frontal area finely setiferous or apparently glabrous (Fig. 249);

macrosetae sparse and confined to the outer lateral

marginal area of abdominal pleura 4 to 7

Ned ce auegetl yh aici cease apie aie tgae Hyperaspis lateralis Mulsant

Hyperaspis binotata (Say)

(Figs. 15, 62, 68, 69, 86, 96, 97, 207)

Specimens examined

The study was based on 30 pupae collected on cottony maple scale in association

with adults in E. Lansing, Michigan 5 July 1971 by Dang T. Phuoc. The specimens

are deposited in the Entomology Museum at Michigan State University.

Diagnosis

This species and H. lateralis represent a group of available pupae which have a

densely macrosetiferous head and pro- and mesofemora. Pupae of the remaining

available Hyperaspis species have the head (except for the macrosetiferous vertex) and

all femora finely setiferous. See the diagnosis for H. lateralis for characters to

separate it from H. binotata.

Description

Length: 2.5-4mm; width:1.2-2mm. Body broadly rounded-oval. Dorsum entirely

brownish and uniformly and very densely macrosetiferous.

26 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Head densely macrosetiferous including the frontal area (Fig. 15). Labium with

setiferous ligula (Figs. 96, 97).

Pronotum yellowish, immarginate, densely and uniformly macrosetiferous.

All abdominal terga uniformly and densely macrosetiferous, including the pygid-

ium. Abdominal sterna entirely pale and finely setiferous.

Hyperaspis lateralis Mulsant

(Fig. 249)

Specimens examined

The study was based on 12 pupae from K. S. Hagen’s collection, collected in

California in March 1971. Two pupae are deposited in the Entomology Museum at

Michigan State University; the remainder are returned to K. S. Hagen, University of

California, Berkeley, California.

Diagnosis

See diagnosis. of H. binotata for separation of these two species from other

Hyperas pis.

Description

Hyperaspis lateralis is very similar to H. binotata except that the available

specimens of lateralis vary from 4mm to 5mm in length and 3mm to 3.5mm in width.

In addition, the frontal area of Jateralis is finely setiferous and the abdominal pleura

(especially from 4 to 7) are sparsely macrosetiferous, with the macrosetae confined to

the outer lateral marginal area. In H. binotata, the head capsule is uniformly

macrosetiferous (including the frontal area), and abdominal pleura 4 to 7 are densely

macrosetiferous with the macrosetae covering the entire surface of the pleuron.

Hyperaspis postica Leconte

(Figs. 247, 255, 256)

Specimens examined

The study was based on two pupae from K. S. Hagen’s collection, collected in -

Placerville, California. One pupa is deposited in Entomology Museum at Michigan

State University, the other is returned to K. S. Hagen, University of California,

Berkeley, California.

Diagnosis

This species represents the pupae of a group of available species including H.

quadrioculata, H. oculaticauda, H. cincta and H. psyche whose pygidium is ap-

parently glabrous. H. postica, H. oculaticauda and H. quadrioculata are separated

from H. cincta and H. psyche by the slender macrosetae whose length may exceed the

length of abdominal tergum 1. In H. cincta and H. psyche, the macrosetae appear

stout and much shorter than the length of abdominal tergum 1. Furthermore, H.

postica can be distinguished from H. oculaticauda by the double subparallel tran-

sverse row of macrosetae on the vertex, compared with only a single transverse row of

Phuoc and Stehr: Pupae of Coccinellidae a

macrosetae on H. oculaticauda. H. postica can be distinguished from JZ.

quadrioculata by the posterior margin of the pronotum being densely

macrosetiferous, while in H. quadrioculata the pronotum appears macrosetiferous

along both the anterior and lateral margins, but posterior margin is not (Fig. 265).

Description

Length: 2.5-3mm; width: 1.8-2mm. Dorsum brownish, sparsely macrosetiferous.

Macrosetae slender, longer than the length of abdominal tergum 1.

Head apparently glabrous and pale, except for two distinct transverse rows of setae

confined to the vertex (Fi ig. 247). Clypeolabrum narrowed apically with apical margin

concave (Fig. 247).

Pronotum with macrosetae densely fringing the marginal areas, with the discal area

sparsely macrosetiferous (as in Fig. 264). Elytron with only the apical area (including

lateroapical margin) finely setiferous. The remaining surface of the elytron is

macrosetiferous with all macrosetae subequal in size.

Anterior surface of all femora finely setiferous. Abdominal terga subequal in

length. Macrosetae apparently confined to the posterior half of terga 1 to 8. Pygidium

apparently glabrous.

Urogomphi as seen wise with the outer lobe varying from subrounded to

elongate kidney-shaped (Figs. 255-256), and with the inner lobe greatly reduced in

size.

Hyperaspis oculaticauda Casey

(Figs. 246, 254, 260, 264)

Specimens examined

The study was based on five pupae from K. S. Hagen’s collection, collected in

Plumas Co., California. Two pupae are deposited in the Entomology Museum at

Michigan State University. The remaining specimens are returned to K. S. Hagen,

University of California, Berkeley, California.

Diagnosis

See diagnosis of H. postica for separation of these two species.

Description

Length: 2.5mm; width: 1.5mm. This species is very similar to H. postica in many

respects except that the vertex is armed with only a single transverse row of

macrosetae (Fig. 246), and the macrosetae on the elytron are sparser and become

gradually more slender toward the sutural, scutellar and basal areas (F ig. 260). Outer

lobe of the urogomphus as seen ventrally appearing elongate kidney-shaped (Fig.

254).

28 — Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Hyperaspis cincta Leconte

(Fig. 252)

Specimens examined

The study was based on ten pupae from K. S. Hagen’s collection, collected in

Fresno Co., California. Two pupae are deposited in the Entomology Museum at

Michigan State University. The remaining specimens are returned to K. S. Hagen,

University of California, Berkeley, California.

Diagnosis

Hyperaspis cincta and H. psyche are the only available pupae whose macrosetae on

the dorsum appear rather stout, with the setal length never exceeding the length of

abdominal tergum 1. This differs from the pupae of the remaining available

Hyperaspis species whose macrosetae are more slender, and whose setal length

usually exceeds the length of abdominal tergum 1. Moreover, this species can be

separated from H. psyche by the macrosetae being uniformly distributed over the

elytral surface (excluding the apical area). In H. psyche the macrosetae at the basal

one-fifth of the median area appear sparser than on the surrounding areas.

Description

Length: 3-3.5mm; width: 1.5-2mm. Dorsum shining brown and densely

macrosetiferous. Macrosetae stout, with the setal length shorter than the length of

abdominal tergum 1.

Head apparently glabrous except the vertex is densely macrosetiferous.

Pronotum with macrosetae fringing the marginal areas, and the discal area ap-

parently without macrosetae.

Elytron with apical area including lateroapical margin finely setiferous; the

remaining area is densely and uniformly macrosetiferous.

Anterior surface of all femora apparently glabrous. Abdominal terga densely and

uniformly macrosetiferous except for the apparently glabrous pygidium. Abdominal

pleura glabrous except for pleura 3 to 7 which usually have macrosetae confined to

the outer lateral marginal area.

Urogomphus as seen ventrally with the outer lobe large, slender and sickle-shaped.

Inner lobe greatly reduced in size (Fig. 252).

Hyperaspis psyche Casey

(Figs. 250, 261)

Specimens examined

The study was based on 10 pupae from K. S. Hagen’s collection. Two pupae are

deposited in the Entomology Museum at Michigan State University, the remaining

specimens are returned to K. S. Hagen, University of California, Berkeley, California.

Diagnosis

See diagnosis of H. cincta for the separation of these two species.

Phuoc and Stehr: Pupae of Coccinellidae 29

Description

Length: 2.5-3mm; width: 1.2-1.5mm. This species is very simlar to H. cincta in

many respects except that the macrosetae appear sparser on the dorsum, especially on

the abdominal terga where they tend to be denser on the posterior half of each

tergum. The macrosetae on the elytron are not as uniformly distributed over the

elytral surface as H. cincta, excluding the apical area where microsetae replace the

macrosetae. Macrosetae appear distinctly sparser in the basal one-fifth of the median

area than elsewhere on the elytron of H. psyche pupae.

Hyperaspis quadrioculata (Motschulsky)

(Figs. 253, 265)

Specimens examined

The study was based on a single pupa from K. S. Hagen’s collection, collected in

San Mateo Co., California. The pupa is returned to K. S. Hagen, University of

California, Berkeley, California.

Diagnosis

This is the only available pupa that has the pronotum apparently glabrous except

for the macrosetiferous anterior and lateral margins. This differs from the pupae of

the remaining available species in which the pronotum is either densely setiferous

over the dorsal surface or the macrosetae fringe the marginal areas.

Description

Length: 3.5mm; width: 2.2mm. Dorsum pale yellowish and rather sparsely

macrosetiferous.

Head pale and apparently glabrous except for 2 groups of macrosetae (8 setae in

each group) on the single available pupa.

Pronotum almost entirely glabrous, especially on discal area and posterior margin,

while macrosetae fringe the anterior and lateral marginal areas (Fig. 265). Meso- and

metanotum also appear almost entirely glabrous except for the presence of a few

macrosetae along posterior margin.

Elytron sparsely macrosetiferous except for apical area and lateroapical margin

being finely setiferous.

Anterior surface of all femora finely setiferous.

Macrosetae on abdominal terga rather sparse, but tending to be denser in posterior

half of each tergum. Pygidium apparently glabrous.

Urogomphus as seen ventrally with the outer lobe broadly elongate oval (Fig. 253),

and with the inner lobe greatly reduced in size.

H 'yperaspis octavia Casey

(Figs. 248, 251, 259)

Specimens examined

The study was based on two pupae from K. S. Hagen’s collection, collected in

Michigan. The pupae are returned to K. S. Hagen, University of California, Berkeley,

California.

30 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Diagnosis

This species represents a group of available pupae including H. quadrivittata

whose pygidium is macrosetiferous. This also occurs in H. binotata and H. lateralis,

however, these two species differ considerably from H. octavia and H. quadrivittata

by their densely macrosetiferous head, pro- and mesofemora. In H. octavia and H.

quadrivittata the head (excluding the vertex) and all femora are apparently glabrous.

Moreover, H. octavia can be separated from H. quadrivittata by the macrosetiferous

apical area and lateroapical margin of the elytron, whereas H. quadrivittata has the

apical area of the elytron finely setiferous.

Description

Length: 2.5-3.5mm; width: 1.5-2mm. Dorsum brownish and_ densely

macrosetiferous. Setae appear slender, with the setal length about as long as the

length of abdominal tergum 1.

Head pale and apparently glabrous excluding the densely macrosetiferous vertex

(Fig. 248).

Pronotum finely setiferous discally, whereas the marginal areas are fringed with

macrosetae.

Elytron densely macrosetiferous, including the apical area and lateroapical margin

(Fig. 259).

Anterior surface of all femora finely setiferous. Abdomen densely macrosetiferous

dorsally including pygidium. Urogomphus as seen ventrally with outer lobe

subrectangular, with the inner anterior angle slightly pointed and curved mesally, the

inner lobe small, rounded or elongate-oval (Fig. 251).

Hyperaspis quadrivittata Leconte

(Figs. 257, 258, 262, 263)

Specimens examined

The study was based on three pupae from K. S. Hagen’s collection collected in

Plumas Co., California. One pupa is deposited in the Entomology Museum at

Michigan State University, the remaining pupae are returned to K. S. Hagen,

University of California, Berkeley, California.

Diagnosis

This is the only available species which has the urogomphi greatly reduced in size

with the inner lobe wanting. This separates this species from the remaining available

Hyperaspis pupae which always have the inner urogomphal lobe present. See the

diagnosis of H. octavia for the separation of these two species.

Description

Length: 2.5-3mm; width: 1.2-1.5mm. Dorsum_ yellowish and_ regularly

macrosetiferous. Setal length as long as or longer than the length of abdominal

tergum 1.

Phuoc and Stehr: Pupae of Coccinellidae 31

Head finely setiferous except for the densely macrosetiferous vertex (as in Fig. 248).

Pronotum densely macrosetiferous with macrosetae distributed uniformly over the

dorsal surface (Fig. 263).

Elytron densely macrosetiferous excluding the finely setiferous apical area and

lateroapical margin (Fig. 262).

Anterior surface of all femora finely setiferous. Abdomen densely and regularly

macrosetiferous dorsally, including the pygidium.

Urogomphus as seen ventrally appearing simple with inner lobe absent. Outer lobe

small, rounded to subrounded and sometimes subkidney-shaped (Figs. 257, 258).

Genus THALASSA Mulsant

Thalassa montezumae Mulsant

(Figs. 16, 208, 209, 236)

Specimens examined

The study was based on one pupa and one exuvia of Thalassa montezumae from

the U.S. National Museum, collected from Harlingen, Texas, 13 September 1932.

Diagnosis

The apparently glabrous dorsum of the body, except for the marginal areas which

are densely and coarsely setiferous, distinctly separates this species from Hyperaspis

whose body is densely, coarsely and homogeneously setiferous.

Description

Length: 6mm; width: 4-4.5mm. Body large, broadly-rounded oval.

Head brown, with bristle-like setae confined to vertex and upper portion of frontal

area; the rest of the head covered with fine and slender or short conical setae (Fig. 16).

Pronotum with anterior margin deeply concave and greatly expanded laterally,

with the posterior lateral side descended beyond the lower margin of the eye. Body

surface brown and covered with fine, slender setae, except for the yellowish marginal

areas which are fringed with long, bristle-like setae (Figs. 16, 236). Meso- and

metanotum apparently glabrous and brownish except for the yellowish scutellar area.

Elytron brown, densely and finely setiferous, except for sutural and lateral margins

which are fringed with long, bristle-like, stout setae. Legs robust with finely setiferous

femora.

Abdominal terga brownish (except the first 2 yellowish), broadly tuberculate

laterally, where dense, bristle-like setae are confined (Fig. 236). Abdominal pleura

subquadrate, invisible from above, and fringed with bristle-like setae on the lateral

margins (Fig. 236). Abdominal spiracles absent.

32 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

TRIBE SCYMNINI

Diagnosis

This tribe, in a general way, is very similar to the Hyperaspini, but the antennae

(Figs. 58, 59) have a distinct club and the abdominal spiracles are normally present.

Moreover, the homogeneous sclerotization of the head capsule, with the frontal suture

wanting, and with dense setae on the labrum, femora and pygidium separates this

tribe from Stethorini which characteristically possess the membranous frontal area,

the glabrous labrum, legs, and pygidium.

Key to Genera of the Available Pupae of Scymnini

1 Body uniformly and densely setiferous; anterior surface of

antenna scape glabrous (Fig. 58); pronotum projected caudad at

mediobasal area into a turbercle-like rounded lobe

(Fig. 109), and apical margin continuous with the vertex

of head (Fig. 30); maxillary palpi glabrous

poe o cee. ee ae Ge ee ees SCYMNUS Kugelann

1’ Body apparently densely and coarsely setiferous only

along marginal areas (Fig. 241); anterior surface of

antennal scape setiferous (Figs. 31, 59); pronotum with basal

margin arcuate, mediobasal area normal and apical margin not

continuous with the vertex of the head; maxillary palpi

monosetose (Figs. 31,85) .......0.5..... CRYPTOLAEMUS Mulsant

Genus SCYMNUS Kugelann

Scymnus creperus Mulsant

(Figs. 30, 58, 109, 162, 201)

Specimens examined

The study was based on a single pupa of Scymnus creperus from the U.S. National

Museum, collected in Amherst, Massachusetts, on 18 July 1941 by M. E. Smith and

identified by E. A. Chapin.

Diagnosis

Species of Scymnus are quite different from Stethorus for the head is slightly and

homogeneously sclerotized and all abdominal spiracles are normal, whereas in

Stethorus the frontal area is membranous and usually protuberant and the first

abdominal spiracles are prominently pedunculate.

Description

Length: 3mm; width: 1.5mm. Body generally pale or yellowish, slightly elongate

oval, and very densely setiferous.

Phuoc and Stehr: Pupae of Coccinellidae 33

Head pale, homogeneously sclerotized and setiferous, vertex largely visible from

above. Antennae short, extending to about one-third of the distance between the eye

and the widest lateral margin of the pronotum. Antennal scape projected caudally

into a short spine-like process (Fig. 58). The first 2 segments of the flagellum

distinctly enlarged, and the club with papillae rather well developed. Clypeolabrum

shorter than wide, with lateral sides rounded and apical margin truncate. Mandibles

bifid. Maxillary palpi slightly enlarged apically.

Pronotum immarginate, with apical margin continuous with vertex of the head

(Fig. 30). Mediobasal area expanded caudally into a large tubercle-like rounded lobe

(Fig. 109). Elytron homogeneously and densely setiferous, including the apical and

sutural margins. Hind wings glabrous. Legs short and robust, with anterior surface of

femora densely setiferous. Metacoxae widely separated, the distance twice as great as

between the procoxae (Fig. 238).

Dorsum of abdomen pale or yellowish and densely setiferous, with 2 types of setae.

The macrosetae are twice the length of microsetae and consistent in location. They

can be distinguished on each tergum as dorsal setae (usually a group of 3 somewhat

transversely aligned setae), and subspiracular setae (usually a group of 2 setae).

Pygidium small and very densely setiferous. Urogomphi cylindrical and very slender

(as long as or slightly longer than the 8th abdominal tergum), with the distal end very

slightly curved ventrally (Fig. 201). Abdominal spiracles subcircular and normal.

Abdominal pleura subquadrate or rhomboidal and very densely setiferous, especially

along outer lateral half. Ventral side of abdomen pale and sparsely setiferous.

Genus CRYPTOLAEMUS Mulsant

Cryptolaemus montrouzieri Mulsant

(Figs. 31, 59, 85, 205, 206, 241)

Specimens examined

The study was based on 4 pupae of Cryptolaemus montrouzieri from the U.S.

National Museum, collected in Honolulu, Hawaii, 1 September 1894.

Diagnosis

This is the only known species of the Scymninae which has the antennal scape

setiferous on the anterior surface (Figs. 31, 59) and with the maxillary palpus

monosetose at apex (Figs. 31, 85). Cryptolaemus can also be separated from Scymnus

by the appearance of dense and coarse setae confined along the marginal areas of the

body as seen dorsally. Furthermore, the urogomphi of Scymnus are slender and

straight, whereas in Cryptolaemus, the urogomphi prominently branch mesally into a

large sausage-like process (Figs. 205, 206).

Description

Length: 4-Smm; width: 2.5-3.5mm. Similar to Scymnus in many respects except

the following.

Body moderately large in size; in general, very densely and coarsely setiferous along

marginal areas of the body.

34, Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Head pale, finely setiferous except the labrum and antennal scape which have long

and slender setae. Antennae short, extending only half the distance between the eye

and the widest lateral side of the pronotum; club with 4 rings of well developed

papillae; flagellum short and subquadrate in cross section; scape somewhat rounded

and setiferous on anterior surface (Fig. 59). Clypeolabrum wider than long with apical

margin very narrow and slightly concave. Labral surface (including margins) densely

setiferous. Maxillary palpi monosetose at apex (Fig. 31).

Pronotum brownish, immarginate, densely and coarsely setiferous along lateral

and basal margins. Meso- and metanotum brownish and apparently glabrous.

Elytron brownish but pale at apex and epipleura; surface apparently glabrous except

for sutural, apical and lateral margins which are densely and coarsely setiferous (Fig.

241). Hind wings glabrous. Legs coarsely setiferous on anterior side of distal ends of

femora (Fig. 31). Metacoxae narrowly separated.

Abdomen brownish and apparently glabrous dorsally, except that the lateral

margins of each abdominal tergum are broadly tuberculate and armed with dense

and coarse setae. Pygidium finely setiferous dorsally. Urogomphi short and stout; as

much sclerotized as the immediately previous tergum, and projected medially into a

long sausage-like process (distal disk) (Fig. 205). Abdominal pleura semi-sclerotized,

tuberculate dorsally and laterally and armed with dense and coarse setae along lateral

margins (Fig. 241) except pleura 1, 8 and 9 which are glabrous. Abdominal sterna

pale and finely setiferous.

TRIBE STETHORINI

Genus STETHORUS Weise

Diagnosis

This is the only known tribe of Coccinellidae in which the frontal area of the pupae

is membranous and usually protuberant (Figs. 17-22)..This character separates

Stethorini from all other groups of the family where the pupal head capsule is

homogeneously sclerotized. Stethorus is also characterized by having one pair of

prominent tubercles at the base of the pronotum (Fig. 110). This character indicates

some degree of affinity to Scymnus which has a single broad, rounded lobe directed

caudally at the base of the pronotum (Fig. 109).

Description

Length: 1.50mm-1.60mm; width: 0.9mm-lmm. Body shiny brownish, somewhat

flattened dorsoventrally, elongate oval, tuberculated, and coarsely setiferous. Head

pale, eyes very large and sub-rounded. Frontal suture sharply visible (Figs. 17, 19, 21).

Frontal area entirely membranous, often protuberant (Figs. 20, 22). Antennae long,

with very well-developed papillae arranged in 3 rings, but with the last two rings

incomplete by lacking the dorsal and lateral papillae. Clypeolabrum glabrous and

subrectangular, much shorter than wide, with apical margin deeply concave (Figs. 17,

19, 21). Mandible unequally bifid at tips. Maxillary palpi large and cylindrical. Labial

palpi slender and cylindrical.

Phuoc and Stehr: Pupae of Coccinellidae 35

Pronotum shiny brown, immarginate and broadly bituberculate mediobasally, with

posteriolateral angle angulate (Fig. 110). Elytron elongate oval, brownish and coarsely

setiferous, with the epipleura wide. Hind wings brownish at apex and glabrous. Legs

short, robust and glabrous; metacoxae widely separated, twice as widely separated as

procoxae (Fig. 238).

Abdomen compact, segments subequal in length and somewhat densely tuber-

culate dorsally. Pygidium small, whitish and glabrous. First pair of abdominal

spiracles prominently pedunculate (Figs. 190-192), the rest normal and circular (Fig.

237). Urogomphi rather long, straight, and slightly flared at distal end into an oval

flat disk (Figs. 210, 211). Abdominal pleura conspicuously setiferous, subrhomboidal

and curved ventrally. Ventral side of abdomen entirely pale, each sternum with one

pair of median prominent setae except sterna 1, 2, 8 and 9.

Key to Species of the Available Pupae of Stethorus

1 Body sparsely setiferous, tubercles on mesonotum and

subscutellar area of elytron prominent (Fig. 237);

first abdominal spiracle somewhat flattened laterally and short (Fig.

192); 2nd abdominal pleuron glabrous

a ei as ic BELLS EE AUS GaPS ES Page ane veal de tee aera Stethorus atomus Casey

¥ Body densely setiferous, tubercles on mesonotum and

subscutellar area of elytron obsolete; first abdominal spiracle

very slender and cylindrical (Figs. 190, 191); 2nd abdominal

picivonactioreus sh.). ofed Sie wollen fee ase sie: tures 2

2(1’) Body dark and shiny brown, the membranous area

on the frons large, broadly oval, seldom protuberant,

with surface slightly concave; abdominal pleuron with more

tah TO setaetiiiils tua Ed ee Stethorus punctum Leconte

z Body pale brownish, the membranous area on the frons

small, circular and usually protruded into a short

cylindrical process (Fig. 20); abdominal pleuron with less

than Heetae tic, bas Loubanik nd wa oebed-. Stethorus picipes Casey

Stethorus atomus Casey

(Figs. 21, 22, 110, 192, 210, 211, 237, 238)

Specimens examined

The study was based on 7 pupae from U.S. National Museum collected in

association with adults in Brownsville, Texas, 3 September 1957 by U. L. Stegman

and determined by E. A. Chapin.

36 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Diagnosis

This is the only known species of Stethorus which has the first abdominal spiracles

with a short and somewhat flattened but conspicuous peduncle (Fig. 192). All other

known species have the first abdominal spiracles slender and cylindrical (Figs. 190,

191).

Description

Length: 1.5mm; width: 0.9mm. Body bronze brownish; prominently tuberculated

dorsally.

Head strongly sclerotized, especially on vertex except for the membranous and

protuberant oval frontal area. Antennae short and well papillated. Scape and the first

segment of flagellum largely expanded caudally along lower side (Fig. 21).

Clypeolabrum glabrous, wider than long, narrowed apically and deeply concave at

apical margin (Fig. 21).

Mediobasal tubercles on the pronotum large, with 2 to 4 setae on each (Figs. 22,

110). Mesonotum with one large and prominent dorsal tubercle at the scutellar area

bearing 2 pairs of setae (actually a pair of tubercles entirely fused together) (Fig. 237).

Metanotum depressed dorsally, with tubercles greatly reduced in size and bearing 3

pairs of dorsal setae and one pair lateral setae (Fig. 237). Elytron very elongate oval,

setae on discal area more sparse than on lateral margin; sutural and apical margins

glabrous, but subscutellar area prominently tuberculate (Fig. 237). Hind wing

somewhat strongly sclerotized at apical area and glabrous. Legs brownish at tibio-

femoral joints.

Dorsum of the abdomen brownish except for the pale last segment. First ab-

dominal spiracles flattened laterally, with base wider than one-half the length of Ist

abdominal tergum (Figs. 192, 237). Each abdominal tergum usually with one pair of

dorsal tubercles bearing 3 setae on each, and one pair of subspiracular tubercles

bearing 2 setae, except tergum 1 in which the subspiracular tubercles are wanting,

and terga 8 and 9 which are entirely glabrous. Pygidium pale translucent white;

urogomphi cylindrical, as long as the 8th sternum, slightly flared distally and ending

in a flat subcircular disk more or less perpendicular to the urogomphal axis (Figs.

210, 211). Abdominal pleura rhomboidal and curved ventrally; each pleuron with 4

setae occupying the outer half except for pleura 1, 2, 8 and 9 which are glabrous.

Ventral abdomen pale; sterna 4, 5, 6, and 7 each with one pair of median setae;

sternum 3 largest with 2 pairs of anteriomedial and posteriomedial setae.

Stethorus picipes Casey

(Figs. 19, 20)

Specimens examined

The study was based on 9 pupae from the U.S. National Museum, collected in

Yakima, Washington, 4 September 1925. |

Phuoc and Stehr: Pupae of Coccinellidae oT

Diagnosis

This species can be separated from Stethorus atomus by being more densely

setiferous and paler, with the 1st abdominal spiracle very slender and cylindrical (as

long as or longer than adjacent setae) (Fig. 190), and the lack of a large scutellar

tubercle on the metanotum.

Description

Length: 1.6mm; width: Imm. Similar to Stethorus atomus except for the following

characters: In general, body paler and more densely setiferous, however, setae much

more slender and slightly curved distally.

Head pale with membranous area on the frons circular and sometimes protruding

into a short cylindrical process (Fig. 20). Antennal papillae not well developed and

greatly reduced in number. Mediobasal tubercles on the pronotum more densely

setiferous and each with more than four setae. Scutellar tubercles on the mesonotum

much reduced in size or wanting. Discal area of elytron as densely setiferous as lateral

margin.

First abdominal spiracles cylindrical, slender and as long as the neighboring setae

(Fig. 190), the diameter of spiracular opening equal to that of the spiracles on the

second segment. Abdominal pleura densely setiferous (6-8 setae on each) except that

pleura 1, 8 and 9 are glabrous and pleuron 2 bears two to three setae.

Stethorus punctum Leconte

(Figs. 17, 18, 190, 191)

Specimens examined

The study was based on 1 pupa and 3 exuviae from the U.S. National Museum,

collected in a greenhouse in Amherst, Massachusetts, 22 July 1942, by M. E. Smith.

Description

Very similar to Stethorus picipes, except body darker and more shiny. The

membranous area on the frons is large and broadly oval, with the surface slightly

concave (Figs. 17, 18). First abdominal spiracles with peduncle brownish, somewhat

conical, and shorter than neighboring setae (Fig. 191).

SUBFAMILY CHILOCORINAE

TRIBE CHILOCORINI

Diagnosis

Members of Chilocorini are very easily recognized by the greatly dilated epistoma

which is expanded laterally to conceal the antennal bases (Fig. 23).

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Similar to the Stethorini in that one pair of well pedunculated spiracles is also

present in all members of the Chilocorini, but here the peduncle is more strongly

sclerotized and more prominent. |

Furthermore, the pupae of Chilocorini are often easy to recognize by having the

pupa enclosed in the last larval exvuvium except for a dorsolongitudinal slit which

exposes the pupa.

1’

2(1)

3(2)

3’

Key to Genera of the Available Pupae of Chilocorini

First abdominal segment with one pair of pit-like gland

openings located between anterior and posterior margins

of the first abdominal tergum and metanotum

respectively (ics 730 eet Joraiod ra RO. Bree) 2 2

First abdominal segment without such a pair of pit-like

SMA PeMING Se. aah oe amore AN aew ar ate adv d aE OA 4

Pronotum crescent-shaped with anterior angle subacute

(less than 90°) (Figs. 106, 107); legs somewhat slender,

with long tarsi, the width of terminal segment

is one-fourth as long as dorsal length

of the tarsus (Fig. 104); size under 4mm

Se ey eG ek Oe ay oN VAS oho eG A 3

Pronotum U-shaped with anterior angle near 90°

(Fig. 105); legs short and robust, tarsi

not as long, with the width of terminal

segment more than one-fourth as long

as dorsal length of the tarsus (Fig. 103);

size latge, over pmuiiin length was, elisiike UarGd hs AXION Mulsant

Spiracular peduncle long, with the outer lateral side

viewed from metanotum more than three times as

long as the diameter of spiracular opening

(Fig. 187); dorsal surface inconspicuously spinose,

with spines less than one-half of setal length

PE PLIOLE ES SUL UP EIR 9 ion decal kl Cae eK RA OS BRUMOIDES Chapin

Spiracular peduncle short, with the outer lateral

side viewed from metanotum less than three

times as long as the diameter of spiracular

opening (Figs. 185, 186); dorsal surface conspicuously

spinose, with spines one-half as long as setae

meee ceig ics SAG ak Fag meee RM eM Cece EXOCHOMUS Redtenbacher

Phuoc and Stehr: Pupae of Coccinellidae 39

4(1’) Size large, over Smm in length, with large bristle-like

setae on dorsal surface (Figs. 23, 242); base of

labrum much narrower than apex of clypeus

(Fig. 23): caiwwahlow .trenlp caus 2 bebee claire - CHILOCORUS Leach

4’ Size small, under 5mm in length, with dense

and long hair-like setae giving the body a fuzzy

appearance (Fig. 29); never with bristle-like setae

on dorsal surface; base of labrum subequal to

apex of clypeus (Fig. 29), Orcus chalybeus

oo .igen eal weniedes heb rige nae! denies ORCUS Mulsant

Genus CHILOCORUS Leach

Chilocorus bivulnerus Mulsant

(Figs. 23, 66, 67, 88, 93, 212, 213, 242)

Specimens examined

The study was based on 5 pupae of Chilocorus bivulnerus from the U.S. National

Museum, collected in Florida.

Diagnosis

This species represents one group of species of Chilocorini including Orcus

chalybeus that have no gland openings on the dorsum of first abdominal segment.

The pupae of the remaining available genera (Axion, Brumoides and Exochomus)

possess one pair of dorsal pit-like gland openings located between the anterior margin

of the 1st abdominal tergum and the posterior margin of the metanotum (Figs. 187,

239). Thus, Chilocorus and Orcus appear to have strong affinities. Chilocorus can be

separated from Orcus by the much shorter and coarse setae, and the labral base being

much narrower than the clypeal apex (Fig. 23). In Orcus the setae appear very long

and slender, giving the body a fuzzy appearance, and the labral base is as wide as the

clypeal apex (Fig. 29).

Description

Length: 4.5-Smm; width: 2.5-2.8mm. Body elongate oval, entirely enclosed in the

last larval exuvia, brownish, slightly rugose and finely setiferous. Epistoma largely

dilated laterally and concealing antennal bases (Fig. 23). Antennae short and

somewhat annulated, tapering apically and pointed at apex (Fig. 23). Labrum densely

setiferous, small and much narrower than clypeal apex whose margin is slightly

concave (Fig. 23). Mandible simple at tip (Figs. 66, 67). Maxillary palpi glabrous,

large and subcylindrical (Fig. 88). Galea bulbous, glabrous and hook-shaped as seen

from the top. Labial palpi cylindrical, slender, and rounded apically (Fig. 93).

Pronotum large, immarginate, ‘‘U”’ shaped, brownish and densely setiferous, with

bristle-like setae on discal area. Mediolongitudinal line yellowish and very distinct,

with anterior angles pale and finely setiferous. Meso- and metanotum brownish with 2

groups of discal bristle-like setae. Elytron immarginate along lateral margin,

40 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

brownish at base, then gradually lighter toward apex, and finely setiferous except for

the subscutellar area which bears bristle-like setae (Fig. 242). Lateroapical angle

obtuse and rounded; epipleura wide. Hind wings semi-sclerotized and glabrous,

tapering apically, with apex rounded. Legs short, yellowish to light brown; femora

robust (Fig. 23).

Abdominal terga brownish except the spiracular area of the Ist segment pale;

posterior margin of terga 3 to 6 finely dentulate. Large bristle-like setae confined to

one pair of dorsal tubercles on each tergum except terga 7, 8 and 9 where dorsal

tubercles are nearly obsolete, and tergum 1, while large bristle-like setae are present

on the subspiracular area. Pygidium minute in relation to the 8th tergum and finely

setiferous. Urogomphi long, subcylindrical, tapering toward apex, and ending in a

kidney-shaped disk (Figs. 212, 213). Abdominal spiracles circular and slightly

pedunculate, with the peduncle of the first abdominal spiracle large and conspicuous

(Fig. 242). Abdominal pleura partially visible trom above, semi-sclerotized and

subquadrate. Abdominal sterna pale yellowish, the 7th sternum very large and twice

as long as the 8th. In the 9, sternum 9 is distinctly bimammillate (Fig. 212).

Genus ORCUS Mulsant

Orcus chalybeus (Boisduval)

(Figs. 29, 214)

Specimens examined

The study was based on 30 pupae of Orcus chalybeus collected from Sydney,

Australia, and deposited in the U.S. National Museum. However, because of poor

condition of the specimens detailed studies of this species have been omitted.

Diagnosis

See Chilocorus diagnosis for the separation of these 2 species.

Description

Length: 3mm; width: 2mm. Body apparently setiferous, very fuzzy with long hair-

like setae, and brownish on dorsal surface. Head wide, hexagonal, with base of

labrum subequal to clypeal apex (Fig. 29), and apical margin of the labrum convex.

Pronotum brown with lateral margins greatly descended beyond the lower side of

the eyes (Fig. 29). Elytron brown at the base and at the sutural area where long, hair-

like setae are very dense...then gradually lighter toward apex and lateral margin.

Abdominal terga brownish and very fuzzy, with long hair-like setae; the first ab-

dominal segment with only one pair of short pedunculate lateral spiracles.

Phuoc and Stehr: Pupae of Coccinellidae 41

Genus AXION Mulsant

Diagnosis

This species is representative of one group of species of the Chilocorini with the

dorsum of the first abdominal segment possessing one pair of gland openings located

between the anterior margin of the first abdominal tergum and the metanotum.

Axion differs from Exochomus and Brumoides by its very large size (over 5mm in

length) and horseshoe-shaped pronotum (Fig. 105). In Exochomus and Brumoides the

size is much smaller (under 5mm in length) and the pronotum is crescent-shaped

(Figs. 106, 107).

Description

Length: 5.5mm; width: 4-4.5mm. Body broadly rounded-oval, brownish to very

dark brown with yellowish spots; dorsal surface densely spinose (Fig. 25) and finely

setiferous.

Head pentagonal (Fig. 24), brownish, with frontal area spinose. Antennae short,

mostly hidden under pronotum and epistoma, the latter dilated laterally (Fig. 24),

flagellum subquadrate in cross section, with distal end truncated and club indistinct

(Fig. 63). Clypeolabrum as wide as distal end of epistoma and greatly narrowed

apically (Fig. 24). Mandibles simple at tip and mola well developed (Figs. 64-65).

Maxillary palpi trapezoidal, elongate oval in cross section, with sides subparallel

(Figs. 24, 89). Labial palpi short, subcylindrical, with apex rounded (Fig. 94).

Pronotum large, ‘‘U’’-shaped with apical margin deeply concave and surface

densely spinose (Fig. 105). Prothoracic spiracle oval. Elytron elongate oval,

lateroapical angle obtuse and rounded, lateral side immarginate. Epipleura wide and

slightly concave, and elytral surface finely nodulated and densely spinose. Hind wing

semi-sclerotized with apex rounded. Legs short and robust, with femorotibial joints

brownish.

Abdominal terga without dorsal tubercles and densely spinose, each with 4 con-

spicuous groups of fine setae arranged in circles consisting of 2 dorsal and 2 lateral or

spiracular setae. The area inside the ring of setae is often smooth. Urogomphi

biramose, with distal ends mushroom-shaped and greatly enlarged (Fig. 215).

Pygidium minute in relation to the 8th tergum and glabrous. Abdominal segment 1

with 1 pair of lateral, prominently pedunculate spiracles, and 1 pair of characteristic

pit-like gland openings located between the anterior margin of the first abdominal

tergum and the metanotum. Abdominal sterna smooth and pale, with sternum 7

twice as long as sternum 6.

Key to Species of the Available Pupae of Axion

1 Yellow spiracular spots on the first abdominal

tergum present; the opening plane of the

first abdominal spiracles perpendicular to peduncular

axis (Fig. 188); peduncle pale yellowish

s iehes opi ues tuk Pate oa Aaa Delain Axion plagiatum (Olivier)

42 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

1’ Spiracular spots on the first abdominal tergum

absent; the opening plane of the first

abdominal spiracles oblique to the peduncular

axis (Fig. 189); peduncle dark brown

We bv oe oe ee a re ee Axion tripustulatum (DeGeer)

Axion plagiatum (Olivier)

(Figs. 24, 25, 63-65, 89, 94, 95, 103, 105, 188, 215, 239, 240)

Specimens examined

The study was based on 4 pupae associated with adults from the U.S. National

Museum collected in Arizona, 19 June 1901, by Prescott, and from Los Angeles,

California, by Coquillet.

Diagnosis

This species is separated from Axion tripustulatum by the presence of a large pair

of subquadrate spiracular yellow spots on abdominal tergum 1, and the opening of

the first abdominal spiracle being perpendicular to the peduncular axis, with the

peduncle pale yellowish. In tripustulatum, the spiracular opening of the first ab-

dominal segment is oblique to the peduncular axis, and the peduncle is dark brown or

black.

Description

Length: 5.5mm; width: 3.5mm. Body rounded-oval and moderately convex dor-

sally.

Head light brown with spines confined in the frontal areas (Fig. 24). Pronotum dark

brown, with two large discal yellow spots. Metanotum dark brown with mid-basal

area yellow. Elytron dark brown at basosutural area, then gradually lighter toward

apex and lateral margin. Abdominal terga dark brown except for medio-dorsal yellow

areas; terga 6, 7, 8, 9 light brownish and the first abdominal tergum with 2 large

subquadrate spiracular yellowish spots. Opening of the spiracles on abdominal

tergum | perpendicular to the peduncular axis. Peduncle pale yellowish (Fig. 188).

Axion tripustulatum (DeGeer)

(Fig. 189)

Specimens examined

The study was based on three pupal exuviae from the U.S. National Museum,

collected in association with adults in College Park, Maryland, on 14 July 1873, and

on 4 July 1940, by C. V. Riley and W. H. Anderson respectively. Adults were iden-

tified by E. A. Chapin.

Description and diagnosis

Very similar to A. plagiatum except the spiracular area on the first abdominal

tergum is brownish, and the opening of the spiracles on the first abdominal tergum is

oblique to the peduncular axis, with the peduncle dark brown or black (Fig. 189).

Phuoc and Stehr: Pupae of Coccinellidae 43

Genus EXOCHOMUS Redtenbacher

Description

Very similar to Axion except size much smaller (3-3.5mm in length). Dorsum light

brown, with yellowish spots. Pronotum crescent-shaped with apical angles subacute

(Fig. 106). Legs somewhat slender, with long tarsi in which the diameter of terminal

segment is about one-fourth as long as dorsal length of the tarsus (similar to Fig. 104).

Body spines one half as long as setae.

Key to Species of the Available Pupae of Exochomus

1 Elytron and frons never spinose; labral apex

truncate or slightly concave (Fig. 26);

peduncular spiracles on the first abdominal

segment cylindrical near the distal

ee i ue ee eats Exochomus hoegei Gorham

1’ Elytron and frons with scattered spines; labral apex

subacute or convex (Fig. 27); peduncular

spiracles on the first abdominal segment

conical, not quite cylindrical near the

Pista ott (Wie TOO) . 1.4 hvac esa 5 2 Exochomus cubensis Dimmock

Exochomus hoegei Gorham

(Figs. 26, 106, 189)

Specimens examined

The study was based on six specimens from the U.S. National Museum, collected in

Douglas, Arizona, 2 October 1956, by J. H. Russel.

Diagnosis

This species can be easily separated from Exochomus cubensis by the concave

clypeolabrum, with the spiracular peduncle cylindrical near the distal end. In E.

cubensis, the spiracular peduncle is conical and the clypeolabrum is convex.

Description

Length: 3.5mm; width: 2.5mm. Head pentagonal (or hexagonal because of labral

apex truncated or slightly concave). Antennae short, slightly elbowed, with the bases

hidden under dilated epistoma (Fig. 26). Pronotum U-shaped, with basal margin

arcuate and largely extended forward, pushing the lateral margins forward where the

posteriolateral angles are very obtuse (Fig. 106). Dorsal surface of pronotum densely

spinose medially, then gradually sparser laterally and with the margins never covered

by spines. Mediodorsal surface of meso- and metanotum glabrous and smooth.

Elytron finely setiferous and micronodulated but never spinose, brownish at base and

along sutural area, and gradually lighter toward apex.

44 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Mediodorsal areas of abdominal terga glabrous and smooth. First five abdominal

terga brown except mediodorsal and spiracular areas yellowish; the rest immaculate

and pale yellowish; tergum 8 entirely smooth. Peduncular spiracles of the first ab-

dominal segment somewhat slender and cylindrical near the distal end (Fig. 185), and

with the diameter of spiracular opening much less than that of the pit-like gland

opening. Abdominal sterna pale, surface smooth.

Exochomus cubensis Dimmock

(Figs. 27, 186)

Specimens examined

The study was based on 2 pupae from the U.S. National Museum collected in Cuba

by Dimmock.

Diagnosis and description

Similar to Exochomus hoegei in many respects except for the following.

Size from 3-3.5mm in length and 2-2.5mm in width. Frontal area and elytral

surface spinose. Labral apex subacute or convex (Fig. 27). Peduncular spiracles of the

first abdominal segment conical (Fig. 186) (in E. hoegei, the peduncular spiracles are

cylindrical near distal end), the spiracular opening with the diameter as wide as or

wider than that of the pit-like gland opening. Dorsally pale or very light brown, the

first abdominal tergum pale yellowish with two large brownish subquadrate in-

terspiracular spots located between gland opening and peduncular spiracle.

Genus BRUMOIDES Chapin

Brumoides suturalis (Fabricius)

(Figs. 28, 104, 107, 187, 216, 217)

Specimens examined

The study was based on 16 pupae of Brumoides suturalis reared by A. G. Selhime,

from Florida, 1955, and deposited in the U.S. National Museum.

Diagnosis and description

Similar to Exochomus in many respects, except for the following.

Length: 3-3.5mm; width: 1.5-2mm. Body fusiform, finely setiferous and very in-

conspicuously spinose on dorsal surface where the spines are about one-third to one-

fifth as long as the setae.

Head and dorsal surface pale yellowish. Pronotum crescent-shaped, with anterior

angle subacute (Fig. 107), usually pale yellowish or very light brown, especially on

mediodorsal area. Peduncular spiracles on the first abdominal segment conical and

long; peduncle height may exceed the distance along posterior margin of the

metanotum between the pit-like laterodorsal gland opening and the elytral sutural

impression line (Fig. 187).

Phuoc and Stehr: Pupae of Coccinellidae 45

SUBFAMILY COCCINELLINAE

Diagnosis

This is the largest group of the family, consisting of the tribes Coccinellini,

Psylloborini and Discotomiini. Pupae of the latter were not available. Pupae of

Coccinellinae are characterized by the exposed intersegmental conjunctivae between

the abdominal terga 3 and 4, 4 and S, 5 and 6, and 6 and 7, allowing for a strongly

flexible abdomen, and by having very fine setae which give the body an apparent

glabrous aspect. In addition, the dorsum of the pupa of the Coccinellinae is distinctly

maculate (even though adults of some species may be immaculate), while all other

members of the family are usually immaculate and pale or brownish throughout (even

though adults of some species may be maculate).

TRIBE COCCINELLINI

Diagnosis

This tribe according to several authors is divided into several different tribes such

as Coccinellini, Hippodamini, Anisostictini, and Synonychini. But, as Boving (1917),

Savoiskaya (1960) and Sasaji (1968a,b, 1971) have pointed out, there are many in-

termediate characters among these “‘tribes,’’ and the group should be regarded as a

single tribe, the Coccinellini, as we do here.

Pupae of the Coccinellini and Psylloborini are almost identical morphologically,

except that the mouthparts are somewhat modified to suit the fungiphagous habit in

the Psylloborini. The clypeolabrum is much wider than long, with the apical margin

truncate, the maxillary palpi have the apex broadly expanded (twice as wide as the

base), and the galea is greatly enlarged, with the greatest width as wide as the base of

the maxillary palpus.

In contrast, in the Coccinellini, the clypeolabrum is as long as, or slightly longer

than wide, with the apical margin usually concave, the maxillary palpi have the width

of the apex subequal to that of the base, and the galea is usually small, with the

greatest width one half as wide as the base of the maxillary palpus.

Key to Genera of the Available Pupae of Coccinellini

1 Legs long, with front femora extended considerably

beyond the widest lateral margins of the pronotum

(Figg. 41,43; ABest I 12

Legs short, with front femora not extended beyond the

| widest lateral margins of the pronotum

(TG Soe MO) civ atin ele aenun Ga Meee aye ies i. 2

At) Hind wing apex finely and densely spinulate

ay ae OG ee ee ae 3

3(2)

6(S’)

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Hind wing apex simooth or micrOnod Wate 56... hole We eben es 4

Elytral surface densely spinulate (Fig. 8); only

ventromarginal areas of hind wing apex finely

and densely spinulate (Figs. 181-183);

prothoracic spiracle elongate oval

ee ert ier are res ee COCCINELLA Linnaeus

Elytral surface smooth, both dorso- and ventromarginal

areas of hind wing apex finely and densely spinulate

(Figs. 179-180); prothoracic spiracle rounded

(Fig. 193) Adalia bipunctata (Linnaeus)........... ADALIA Mulsant

Urogomphi branched mesally into a spine-like process

ai tise Wasa) One-fiird (Mies 270-226) i ee ee oc ee ees S

Urogomphi not branched mesally (Figs. 227, 228) ................... 9

Abdominal pleura 3 to 5 with posterior lateral angle

projected into a long spine-like process

(Figs. 176-178); antennal club indistinct, with

the diameter subequal to that of the flagellum

(FP ige O49 5)) uct See ee ease tei AUS. cdlest. aie 7

Abdominal pleura 3 to 5 subquadrate, with posterior

lateral angle only slightly expanded laterocaudally

(Fig. 171); antennal club distinct and subspherical,

with the diameter much larger than that

i Ue Tapellinr Wigs < sa Ser Sorts). os cawirnwe SRE SARs 2 6

Antennal scape strongly convex anteriorly and dis-

tinctly separated from the flagellum (Fig. 34);

body rounded, apparently glabrous; posterior

lateral angle of abdominal pleura 3 to 5 slightly

risose, Old epdomindlis (S89) vos bo ee i eos eke ee OLLA Casey

Antennal scape flattened anteriorly and hardly separated

from the flagellum (Fig. 40); body slightly

elongate oval and finely setiferous; posterior

lateral angle of abdominal pleura 3 to 5

deeply rugose and punctate (Fig. 171), Neoharmonia

venusta (Melsheimet).« isan coated incall NEOHARMONIA Casey

(5)

8(7)

Q’

Phuoc and Stehr: Pupae of Coccinellidae 47

Abdominal pleura with the spine-like process projected

laterocaudad and short, not exceeding the greatest

width of the corresponding pleuron (Figs. 177-178);

antennae long, extending over two-thirds of the

of the pronotium and the‘eye (Fives. 62,45). (28. 2 F700 2 8

Abdominal pleura with the spine-like process projected

lateroanterad and long, exceeding the greatest

width of the corresponding pleuron (Fig. 176);

antennae short, extending to about half the

distance between the widest lateral margin of

the pronotum and the eye, Synonycha grandis

CUM ON coe oe bee ee ee ee SYNONYCHA Mulsant

Apical margin of clypeolabrum slightly concave (Fig. 45);

spots large, almost covering the entire segment,

but poorly defined; setae coarse, borne on a

conspicuous tubercle with distal end strongly curved

(Fig. 177) Anisocalvia quatuordecimguttata Linnaeus

PUR Ey Bee ae Se ig eee Aig We hearers ANISOCALVIA Crotch

Apical margin of clypeolabrum sinuate or slightly convex

(Fig. 32); spots smaller, with a well defined

border; setae finer, borne on a flat cuticular

ring on an inconspicuous tubercle 67 60 ANATIS Mulsant

Antennae very long, if straight, then extending beyond

the widest lateral margins of the pronotum

(Figs. 44, 56); flagellum C-shaped; antenna with six rings

of papillae; club not so distinct, with the

diameter subequal to that of the flagellum (Fig. 56);

body rather coarsely setiferous, with chalaza-type

setae, especially along lateral margin of abdominal

pleura 3 to 5 (Fig. 172), Propylaea

quatuordecimpunctata (Linnaeus)........... PROPYLAEA Mulsant

Antennae shorter, if straight, not extending beyond

the widest lateral margins of the pronotum;

flagellum slightly elbowed; antenna with less

than six rings of papillae; club distinct,

with the diameter greater than that of the

flagellum (Figs. 52, 53); body apparently glabrous or

finely setiferous, with setae borne on a flat

Ube ealin rire ie Sigs PERI Mar ee tad aah le MAGEE nese Sw ore gad 10

10(9’)

10’

11(10)

1V’

1200

12)

13(12’)

1413’)

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Antennal club subspherical (Fig. 52); surface of the

galea with less than five conspicuous fine

beet hits, 6182) oe ee te Ha ee ee 11

Antennal club cylindrical-elongate or subrectangular

block-shaped (Fig. 53); surface of the galea

with at least five conspicuous fine teeth

(Pie SO) c. pene a2 oases ecu locuenle. MULSANTINA Weise

Antennae actually with three rings of papillae;

scape greatly swollen (Fig. 34); body apparently glabrous and rounded,

pale silky white dorsally,

Olla abdominalis (Say) ....... Os hoy ete ae ok) OLLA Casey

Antennae actually with four rings of papillae

(Figs. 38, 39); scape flat and not so

distinct from flagellum; body finely setiferous,

yellowish, and slightly elongate-oval

Sy eS a ee ee CYCLONEDA Crotch

Exposed conjunctivae on abdominal segments 4 to 7 glabrous;

elytron with lateral angle broadly expanded anteriorly

into a distinct rounded lobe (Figs. 158, 159)

eS ala. Gee Get ers ob a HIPPODAMIA Dejean

Exposed conjunctivae on abdominal segments 4 to 7 finely

setiferous; elytron with lateral angle obtuse and

rounded, almost continuous with the lateral margin;

never expanded anteriorly into a distinct lobe

(PiO5 155, $56) Gb oe ee ea eta OR a kk 13

Apical margin of clypeolabrum deeply notched (Fig. 49);

anterior margin of pronotum non-marginate;

lateral margin of abdominal pleura 3-4 slightly angulate

(Fig. 175), Eriopis connexa (Germat) wis 03 4). 2. 2 ERIOPIS Mulsant

Apical margin of clypeolabrum slightly concave (Figs. 42, 43);

anterior margin of pronotum strongly marginate

(Figs. 124-126); lateral margin of abdominal pleura 3-4

rounded Convex (Pig 174A S25 te ee nw oe ee os 14

Lower side of anterior edge of the pronotum

descending to two-thirds of the eye length

(Fig. 42); first abdominal spiracles entirely hidden

under the elytra, Naemia seriata (Melsheimer)

Re ee ie NAEMIA Mulsant

Phuoc and Stehr: Pupae of Coccinellidae 49

14 Lower side of anterior edge of the pronotum never

descending to two-thirds of the eye length (Fig. 43);

first abdominal spiracles mostly hidden under the

elytra, Coleomegilla maculata DeGeer

OOP ae COLEOMEGILLA Timberlake

Genus COCCINELLA Linnaeus

Diagnosis

Coccinella represents one group in the subfamily Coccinellinae in which the body is

usually rounded-oval, moderately convex dorsally, and with short legs in which the

front femora never extend beyond the widest lateral margins of the pronotum as

viewed ventrally. By contrast, in the other group as represented by Hippodamia,

Coleomegilla, Eriopis, Paranaemia and Naemia, the body is elongate-oval, and has

long legs in which the front femora extend considerably beyond the widest lateral

margins of the pronotum. Adalia is very similar to Coccinella. The elytral surface is

clothed with fine sharp spines in Coccinella, whereas in Adalia, the elytral surface is

smooth. In addition, in Adalia, both dorsal and ventral surfaces of the hind wing apex

are clothed with fine sharp spines (Figs. 179, 180), whereas in Coccinella, only the

ventral surface of hind wing apex is sharply spinulated (Figs. 181-183). These two

genera are quite different from the remainder of Coccinellini whose hind wing apex is

usually smooth.

Description

Length: 5.5-8mm; width: 4-Smm. Body slightly elongate oval, strongly convex

dorsally and finely setiferous.

Head somewhat\hexagonal, slightly longer than wide, surface deeply rugose, brown

to black, frontal area usually pale whitish or light yellow. Mediolongitudinal line well

defined throughout the head length. Eye dark brown. Antennae long and slender,

exceeding half the distance between the widest lateral margin of the pronotum and

the eye (Fig. 35). Antennal club subspherical and distinct from flagellum, and with

four rings of papillae, the 3rd and the 4th incomplete (lacking some lateral papillae),

with the ventral papillae of the 4th flattened but visible. Flagellum slender and

subquadrate in cross section; antennal scape suboval and convex anteriorly.

Clypeolabrum usually dark brown, with lateral edge slightly sinuate. Labrum almost

black, surface with deep longitudinal and subparallel wrinkles. Apical margin of the

labrum concave or sinuate-concave (Fig. 35) and very narrow, about one half as wide

as the base, with the lateral margin rounded convex. Mandibles bifid at tip, with mola

well developed (Figs. 70, 71). Maxillary palpus large, trapezoidal, usually dark brown

or black and rugose, with lateral margin slightly convex (Fig. 35). Galea smooth and

brownish. Lacinea immediately beneath the galea with tip wedge-shaped. Labial

palpus with deep longitudinal subparallel wrinkles.

Pronotum immarginate, usually yellowish with dark brown spots along marginal

areas; mediolongitudinal line always present and the anterior margin deeply rugose.

Mesonotum trapezoidal, usually yellowish or brownish with one pair of circular spots,

50 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

and the scutellar area elevated in the middle. Metanotum usually with two large oval

spots at base where deep subparallel wrinkles are clearly visible. Prothoracic spiracle

elongate oval (Fig. 194).

Elytron large, somewhat rectangular, with wide and slightly concave epipleuron:

the lateroapical angle slightly expanded anteriorly, forming a somewhat rounded lobe

(Figs. 141-152). Elytral surface densely covered with very fine sharp spines. Color

pattern may change from light yellow with dark spots or transverse bands to very dark

brown or black with small yellowish areas. Hind wing apex slightly sclerotized, finely

and densely spinulated along ventrolateral and ventrosutural margins (Figs. 181-183).

Abdomen usually yellowish with dark brown spots on dorsal surface. Terga 4 to 6

slightly narrowed medially, and “detachable” from one another to show the in-

tersegmental conjunctivae which are as strongly sclerotized as the terga and finely

setiferous. Posterior margins of terga 3 to 6 finely and sharply dentulate. One pair of

dorsal tubercles may be present on terga 2 to 8. Abdominal spiracles circular or

nearly so, and gradually diminishing in size toward posterior of the abdomen; usually

the first five spiracles well sclerotized. Abdominal pleura subquadrate with pleura

one and two hidden and partly hidden under elytron respectively; pleura 8 and 9

somewhat fused together. Urogomphal disk bilobed, with inner lateral side of base

straight (Fig. 219). Abdominal sterna immaculate, yellowish and finely setiferous.

Third sternum the largest, with anterior margin deeply sinuate, and with 2 sublateral

is flat and subrectangular, whereas, in the 9 , it is bibulbous, with the nipple-like tip

well sclerotized (Figs. 2, 219).

Key to Species of the Available Pupae of Coccinella

1 Hind wing apex finely spinulated along ventrolateral

and ventrosutural margins (Fig. 183); elytron with

at least two transverse bands (Figs. 141-142)

See RUE Area beoly aed odie Coccinella trifasciata Linnaeus

iE Hind wing apex finely spinulated along ventrolateral

margin only (Figs. 181-182); elytron without

er AlioveIae Dalida Saat hae Poe. Ges Borde Se te eae oat fat 2

2(1’) Elytral surface more than half dark brown or black,

WOUSPOURRIe Poe eee he henil ted SEE gon bie. aoe: 3

2 Elytral surface half or less than half dark brown

or binge may nance Gistiticnanot(sstiests ts) o0Seui dh beersiavrob is i)

3(2) Elytron with dark area(s) occupying over three-

Tourths of the totabeinioce ities oadud ae bidieswdl. soak athe, wal: 4

i Elytron with dark area(s) never occupying three-

fourths ofthe total suvlace vex atuoail'ss vllaitiew Jeet he BROAN) i

A(3)

3(2’)

6(S’)

7(3’)

Phuoc and Stehr: Pupae of Coccinellidae 51

Elytron with subhumeral and basal areas pale, and

with a narrow pale stripe extending from

outer apical angle to the distal end of lateral

margin (Fig. 148); length of spinulated area

on hind wing apex one-half as long

as elytral apex (Fig. 181); pronotum yellowish with

dark brown marginal areas and no spots

Kel iG ee eRe sh Be Coccinella transversoguttata Faldermann

Elytron with subhumeral and basal areas pale (Fig. 152);

length of spinulated area on hind wing

apex shorter than one-half of elytral

apex (Fig. 182); pronotum brownish with darker

margins and 2 pairs of circular and

elongate oval pale spots

ed Es GE et Oe TEs EON ES Coccinella monticola Mulsant

Elytron with median spot at basal two-fifths twice as

wide as long (Fig. 145); subscutellar spot

triangular and usually present (Figs. 146, 147)

PON a aN G PRE Coccinella transversoguttata Faldermann

Elytron with median spot at basal two-fifths never twice

as wide as long (Fig. 149); subscutellar

INT Se TO aca ce a Mer Ri alae tae 6

Elytral spots unequal; median spot at basal

two-fifths subquadrate, larger or as large

as metanotal spot (Fig. 149); disto-lateral

SPOk MOSAIC EH AA Pa Coccinella novemnotata Herbst

Elytral spots subequal; median spot at basal

_ two-fifths rounded, oval and much smaller

than metanotal spot (Fig. 143); disto-lateral

spot also rounded, oval, and located

at distal end of lateral margin (Fig. 143)

Py ELE PG Ey aE Coccinella septempunctata Linnaeus

Subscutellar and humeral dark areas somewhat truncate

anterad, with a well-defined contour (Fig. 144);

a subscutellar spot may be distinct;

mesonotal spot half as large as

metanatal ery a EM ee Coccinella septempunctata Linnaeus

52 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

(ot Subscutellar and humeral dark areas somewhat

pointed anterad, and with not so

well-defined contour (Figs. 150-151); subscutellar spot

never present; mesonotal spot much less

than half as large as metanotal spot

ee ee Coccinella novemnotata Herbst

Coccinella trifasciata Linnaeus

(Figs. 112, 141, 142, 183)

Specimens examined

The study was based on 25 pupae which were reared in the laboratory from adults

collected in Clinton Co., Michigan, on 30 May 1971, by Dang T. Phuoc. The

specimens are deposited in the Entomology Museum at Michigan State University

and in the U.S. National Museum.

Diagnosis

This is only known species of Coccinella with a very well defined maculation

pattern on dorsal surface of the body, with at least 2 transverse dark brown bands on

each elytron, and frequently a third, and with the hind wing apex finely spinulated

along both ventrolateral and ventrosutural margins. This is distinctly different from

the remaining known species of Coccinella whose hind wing apex is only spinulated

along the ventrolateral margin.

Description

Length: 6-6.5mm; width: 4-4.5mm. Body slightly elongate oval, finely setiferous

and yellowish, with distinct dark brown spots and transverse bands on dorsum of the

body.

Head with most of frontal area yellowish. Eyes and antennae dark brown. Antennal

scape distinct, convex anteriorly. Club distinctly subspherical. Clypeus usually pale or

light brown at base. Labrum with the apical margin concave and dark brown except

for a mediolongitudinal whitish line. Mandibles unequally bifid at apex.

Pronotum with anterior margin dark brown except for the median area, one pair of

large claw-shaped spots at posterior angles, and one pair of large subcircular spots at

the base (Fig. 112). In dark forms, one pair of median light brown spots may be

present. Meso- and metanotum yellowish, each with one pair of small circular and

one pair of large oval spots. Elytron (Figs. 141, 142) always with sutural area and apex

dark brown. Subscutellar spot triangular, separated or fused with a transverse band

extending to humeral angle. The second and the third transverse bands extending

from the median to lateral margin at basal two-fifths and three-fourths respectively.

The third band may be reduced to a circular spot in light forms. Hind wing apex

slightly sclerotized and very finely spinulated along ventrolateral and ventrosutural

margins (Fig. 183).

Abdominal terga 2 to 6 each with one pair of large spots, the rest immaculate or

with very light spots. Tergum 2 with undefined spots consisting of one pair of

spiracular and one pair of small dorsal spots. Tergum 3 similar to 2 but with large,

Phuoc and Stehr: Pupae of Coccinellidae 53

subquadrate and subrectangular, more well-defined spots; the spiracular and dorsal

spots may fuse together. Terga 4, 5 and 6 each with only one pair of subquadrate

dorsal spots. Posterior margins of terga 3 to 6 dark brown or black at portions

corresponding to spots, and very finely and densely spinulated (including the lower

surface). Abdominal pleura immaculate and yellowish, except that pleura 2 and 3 are

almost covered by a large subquadrate dark brown spot. All abdominal sterna

yellowish and immaculate; sternum 9 flat in the cand bibulbous in the Qwith the

nipple-like tip well sclerotized.

Coccinella transversoguttata Faldermann

(Figs. 113, 114, 145-148, 181, 194, 218-220)

Specimens examined

The study was based on 25 pupae and 20 exuviae which were reared in the

laboratory from adults collected in Saginaw Co., Michigan, on 5 August 1971 by

Dang T. Phuoc. The specimens are deposited in the Entomology Museum at

Michigan State University and in the U.S. National Museum.

Diagnosis

This species differs from Coccinella trifasciata by the hind wing apex being

spinulated only along the ventrolateral margin (Figs. 181, 182). It can be separated

from Coccinella novemnotata by the presence of tergal spots on abdominal segment 1,

and by the median spot on the elytron at the basal one-third always being twice as

wide as long (in C. novemnotata it is subquadrate). In addition, in dark forms, the

elytron is dark brown or black over three-fourths of the surface, whereas in C.

novemnotata, the dark area never reaches three-fourths of the elytral surface.

Description

Length: 6.5-8mm; width: 4.5-5mm. Body slightly elongate oval, yellowish, with

well-defined, dark brown spots on dorsum and finely setiferous.

Head dark brown, with frontal area usually pale yellowish. In dark forms, the head

is completely black except for one pair of circular yellowish spots on the frons.

Mediolongitudinal pale whitish line always present. Eyes and antennae dark brown.

Clypeolabrum dark brown except for mediolongitudinal pale line. Labral apex

narrowed, with apical margin concave or sinuate concave. Surface of labrum deeply

rugose, with deep longitudinal and subparallel wrinkles. Mandibles subequally bifid

at apex.

Pronotum pale yellowish, in most cases with one pair of medio-apical subquadrate

spots and two pairs of large claw-like spots, (one at anterior and one at posterior

angles) (Fig. 113). In dark forms all of these spots may fuse together except medially,

thus framing the anterior and lateral areas of the pronotum in dark brown (Fig. 114).

In addition, one pair of mediobasal spots may be present, and one pair of very light

brown, rectangular discal spots may also be present in some forms (Fig. 114). Meso-

and metanotum yellowish to brown, each with one pair of distinct large spots. Elytron

varies from very light to very dark. In most cases, elytral suture, lateral margin, and

apex light to dark brown. In light forms, a median subrectangular spot (twice as wide

54 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

as long) at basal 1/3 is always present, a subcircular median spot at basal 2/3 is

present in most cases, and these two spots occasionally narrowly fuse (Fig. 147). A

subscutellar triangular spot is also very frequently present (Figs. 146, 147). In dark

forms, the elytron is excessively black or dark brown except for the yellowish

subhumeral and basal areas and a narrow yellowish stripe extending from the suturo-

apical angle to the posterior end of lateral margin. The apex of hind wing slightly

sclerotized and finely spinulated along the ventrolateral side (the length of this

spinulated area is one-half as long the lateroapical margin of the elytron). In light

forms, at least one pair of dark brown or black spots is present on each abdominal

tergum except that terga 7 to 9 are usually immaculate. The small spots on terga 1

and 2 are subcircular and oval respectively. The spots on terga 4 to 6 are large,

subquadrate, or subrectangular. Tergum 3 usually has 2 pairs of large dorsal spots

and 2 pairs of large spiracular spots, with the anterior margin of the spots deeply

sinuate. Abdominal pleura usually immaculate and yellowish, except for the third

which is almost entirely dark brown. In dark forms, all abdominal terga are brownish

to dark brown with yellow spots, except that the Ist tergum is yellowish with two

subcircular dorsal brown spots, and the 9th tergum is pale and immaculate. Terga 2

and 3 each with a median keyhole-like yellow spot. Terga 4 to 7 each with one pair of

spiracular yellow spots diminishing in size from 4 to 7, and with a median vase-

shaped yellowish spot. Tergum 8 brownish and immaculate.

Abdominal pleura dark brown with a yellow spot on each, except that pleuron 1 is

pale whitish, pleuron 2 brownish, pleuron 3 completely dark brown, pleuron 4 entirely

yellowish and pleura 8 and 9 pale to very light brown.

Coccinella novemnotata Herbst

(Figs. 1, 2, 8, 35, 52, 70, 71, 149-151)

Specimens examined

The study was based on 18 pupae and two pupal exuviae which were reared from

adults collected at the Gull Lake Biological Station, Kalamazoo Co., Michigan, on 20

May 1971, by Dang T. Phuoc. The specimens are deposited in the Entomology

Museum at Michigan State University and in the U.S. National Museum.

Diagnosis

This species is very similar to Coccinella transversoguttata except that the tergal

spots are absent on abdominal segment 1. On the elytron, the median spot at the

basal one-third is subquadrate, whereas in C. transversoguttata it is subrectangular,

twice as wide as long. In dark forms, the dark area of the elytron never reaches three-

fourths of the surface, but does so in C. transversoguttata.

Description

Length: 6-7mm; width: 4-Smm. In general, very similar to Coccinella tran-

sversoguttata.

Head dark brown with two yellowish rounded spots which are medially fused on the

frons. Labrum strongly convex dorsally, with longitudinal subparallel wrinkles

Phuoc and Stehr: Pupae of Coccinellidae 3D

usually shallower than those of the clypeus. In a few cases, the surface of the labrum is

smooth without such wrinkles. In most cases, anterior marginal spots of pronotum

fused to each other except in the median area, and the posterior angular spot may join

the apical angular spot along the lateral margin. Meso- and metanotum yellowish,

with one pair of small circular and one pair of large oval spots on each. Elytron always

dark brown along sutural area, lateral margin, and laterapical angle. In light forms,

at least two median spots are present on each elytron ... a large subrectangular spot

on the basal two-fifths, and a small and elongate longitudinal spot parallel to the

sutural margin at the basal three-fifths. These two spots are often fused together

(Figs. 149-150). In dark forms, these median spots are completely fused with the dark

sutural area, but this dark area never exceeds three-fourths of the elytral surface.

Both the subscutellar and humeral dark area are tapered and pointed but without a

well-defined border. The border is well-defined in C. septempunctata. First ab-

dominal tergum immaculate. Urogomphi brownish or darker than abdominal sterna.

Coccinella monticola Mulsant

(Figs. 115, 116, 152, 182)

Specimens examined

The study was based on 11 dry pupae loaned from the USS.

National Museum, collected and identified by Dimmock.

Diagnosis

This species is very similar to Coccinella transversoguttata and C. novemnotata but

the spinulated area on the hind wing apex appears shorter than one-half the length of

the lateroapical side of the elytron. In C. transversoguttata and C. novemnotata it is

as long as the length of the lateroapical side of the elytron.

Description

Length: 5.5-6mm; width: 3.5-4mm. Head completely dark brown without frontal

yellow spots except the mediolongitudinal yellow line running the head length;

surface deeply rugose.

Pronotum in most cases brownish, except apical and lateral margins are dark

brown, with one circular sublateral and one elongate oval dorsal yellowish spot. These

two spots may fuse medially in some forms (Figs. 115, 116). The longitudinal median

pale line clearly present. Meso- and metanotum brownish except for elevated and pale

scutellar area. Elytron as in dark forms of Coccinella transversoguttata, but lacking

the yellowish stripe extending from suturoapical angle to distal end of lateral margin

(Fig. 152). The spinulated surface along ventrolateral margin of hind wing apex

shorter than one half of elytral apex. All legs are brown or brownish at coxae,

femorotibial joints and tarsi.

Abdomen also as in dark forms of Coccinella transversoguttata except that median

yellowish spots on abdominal terga 4 to 7 are large-mouthed-vase-shaped.

56 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Coccinella septempunctata Linnaeus

(Figs. 143, 144)

Specimens examined

The study was based on 3 pupae from the U.S. National Museum collected in

Denmark on 19 July 1893 by E. Rosenbero.

Diagnosis

This is the only species examined with subequal and well-defined oval spots on the

elytron in light forms. In dark forms the elytral spot pattern is somewhat similar to

the dark forms of Coccinella novemnotata; however, the humeral and scutellar ends

of the dark area appear rounded or truncated with a well-defined contour. In C.

novemnotata the above areas are poorly defined and usually tapered and pointed.

Description

Length: 6mm; width: 4mm. Head as C. transversoguttata, but more rugose and

usually entirely dark brown.

Pronotum usually pale along lateral margins. Circular spot on mesonotum large,

about one-half as large as that on metanotum.

Elytron usually brown along sutural margin, lateroapical angle and part of the

lateral margin. In light forms, three well-defined subequal oval spots (much smaller

than metanotal spots) are present. The subscutellar spot and the mediosutural spot at

the basal two-fifths may be more or less fused with the dark sutural area. The lateral

spot at the distal end of the lateral margin is free. Sometimes a humeral spot may be

present, but it is often fused with the lateral marginal dark area (Fig. 143). In dark

forms, the spots are not so distinct, but they are slightly darker than the surrounding

areas. This dark form is very similar to the dark forms of Coccinella novemnotata, but

the scutellar and humeral ends of the dark area are rounded or somewhat truncated

with a well-defined contour (Fig. 144) in C. septempunctata.

Genus ADALIA Mulsant

Adalia bipunctata (Linnaeus)

(Fios. 36, 37, 2, 91, 117-119, 179, 180,192)

Specimens examined

The study was based on 12 pupae which were reared from adults collected in

Kalamazoo Co., Michigan, 8 April 1972 by Dang T. Phuoc. The specimens studied

are deposited in the Entomology Museum at Michigan State University and in the

U.S. National Museum.

Diagnosis

This genus is very similar to Coccinella. As previously mentioned these are the only

2 genera of Coccinellidae examined which have the hind wing apex finely spinulated.

Phuoc and Stehr: Pupae of Coccinellidae 37

Adalia is separated by the presence of spinules on the dorsal and ventral surface of

the hind wing apex, whereas in Coccinella the hind wing apex is spinulated on the

ventral surface only.

Description

Length: 4.5-S5mm; width: 3.5-4mm. Body slightly elongate oval, convex dorsally,

yellowish with large dark brown spots on dorsal surface and finely setiferous.

Head light to dark brown, nearly smooth but with a few wrinkles.

Mediolongitudinal pale line always present. Eyes dark brown. Antennae long, ex-

tending over half the distance between the widest lateral margin of the pronotum and

the eye; club subspherical with three rings of well developed papillae, the third ring

incomplete by lacking some dorsal papillae; flagellum subquadrate in cross section;

scape flat, subtriangular and slightly expanded caudally. Clypeolabrum hexagonal

(Figs. 36, 37), slightly rugose with longitudinal wrinkles visible but shallow. Clypeus

narrow at base. Labrum tapered apically and with apical margin deeply concave (Fig.

37). Maxillary palpi large, trapezoidal, with outer lateral margin convex (Fig. 36).

Surface of galea armed with a few fine teeth at anterior inner angle (Fig. 82). Labial

palpus rugose and short.

Pronotum immarginate to slightly marginate along apical margin; surface rugose

and irregularly brownish to dark brown. In most cases the apical and posterior

angles, mediobasal area and mediolongitudinal line are pale yellowish. Prothoracic

spiracles rounded (Fig. 193). Mesonotum brownish to dark brown with two sub-

triangular pale spots at posterior angles. Scutellar area elevated in the middle with a

triangular pale spot. Mediolongitudinal line pale and greatly enlarged anteriorly.

Apical angles of mesonotum elevated. Metanotum slightly elevated along median

line, with two large dorsal dark brown spots fused together in some cases and oc-

cupying most of the surface.

Elytron immaculate dark brown at median and sutural area, then gradually lighter

toward apex and lateral margin. A median light brown, poorly defined spot may be

present in some forms at about basal two-fifths. Lateral margin usually pale yellowish

or light brown. Hind wing apex brownish, slightly sclerotized, finely and densely

spinulated along ventrosutural and both ventro- and dorsolateral margins (Figs. 179-

180). Legs short, brownish except for pale yellowish femoral bases and tibiotarsal

joints. Surface of the leg slightly rugose with a few transverse wrinkles.

Abdomen as in Coccinella, but slightly darker, each abdominal tergum with one

pair of small subquadrate spiracular and one pair of large subrectangular dorsal dark

brown spots. Spots on the first tergum greatly reduced in size; tergum eight brownish

and immaculate. Abdominal pleura brownish with pale yellowish margins except for

pleura 8 and 9 which are completely pale. Urogomphi as in Coccinella.

Genus CYCLONEDA Crotch

Diagnosis

This genus is representative of the remaining species of Coccinellini which possess a

smooth hind wing apex. Cycloneda shows the closest affinity to Mulsantina Weise;

however, Cycloneda can be separated by the subspherical antennal club (Fig. 38), and

38 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

the subacute apical angle of the clypeolabrum (Fig. 38). In Mulsantina the antennal

club appears cylindrical or subrectangular block-shaped (Fig. 53), and the apical

angle of the clypeolabrum is rounded (Fig. 33).

Description

Length: 4.5-Smm; width: 3-3.5mm. Body slightly elongate oval and finely

setiferous. |

Head entirely brown with mediolongitudinal line or frontal area pale; surface

smooth to very slightly rugose. Eyes brownish. Antennae (Fig. 38) brown and long,

extending over two-thirds of the distance between the widest lateral margin of the

pronotum and the eye; club subspherical and distinct with four rings of well-

developed papillae; flagellum subquadrate in cross section; scape somewhat flat,

subtriangular and with upper surface slightly rugose. Clypeolabrum subquadrate,

slightly narrowed apically and with the apical margin concave. Apical angles of

labrum subacute (Fig. 38). Mandible bifid at tip. Maxillary palpus brown, large,

trapezoidal or lanceolate. Galeal surface armed with several fine teeth, but less than

five at the anterior inner angle. Labial palpus short and rounded, brown and slightly

rugose at tip.

Pronotum pale yellowish, usually with dark brown spots along anterior and

posterior margins. Lateral margin and especially the anterior margin strongly

marginate (Fig. 39). Prothoracic spiracle elongate oval. Meso- and metanotum pale,

slightly rugose and sometimes maculate. Elytron immaculate, brown to dark brown

along sutural margin from base to about the basal three-fourths, and gradually

lighter toward lateral margin. Apical, subapical, basal and subbasal areas pale

yellowish. Elytral surface finely setiferous and densely micronodulated (as in Fig. 7).

Epipleura wide and slightly concave. Hind wing slightly sclerotized at apex, surface

smooth. Legs light brown except femoral bases and distal ends of tibia which are pale.

Abdominal terga whitish or yellowish, with dorsal dark brown spots. Posterior

margin of terga 3 to 6 finely dentulated. Abdominal spiracles circular or nearly so.

Abdominal pleura subquadrate and slightly rugose. Abdominal sterna pale im-

maculate. In the 2, abdominal sternum 9 is bimammillate and well sclerotized at

the tip.

Key to Species of the Available Pupae of Cycloneda

1 Setae with conspicuous cuticular ring; pronotum with

2 pairs of anterior and 2 pairs of basal

subquadrate dark brown spots; mesonotum with one

pair of large subcircular spots .............. Cycloneda munda (Say)

i Setae with inconspicuous cuticular ring; anterior

and basal spots on pronotum greatly reduced

or wanting; mesonotum usually immaculate

DMO ALE Edie cs OPE se ee ee Cycloneda sanguinea (Linnaeus)

Phuoc and Stehr: Pupae of Coccinellidae oF

Cycloneda munda (Say)

(Figs. 38, 39, 81)

Specimens examined

The study was based on 11 pupae which were reared from adults collected in

Kalamazoo Co., Michigan, 24 May 1971, by Dang T. Phuoc. The specimens are

deposited in the Entomology Museum at Michigan State University and in the U.S.

National Museum.

Diagnosis

Cycloneda munda may be easily separated from C. sanguinea by the conspicuous,

well sclerotized cuticular ring surrounding the base of each seta, and the mesonotum

usually having one pair of dark brown spots. In C. sanguinea the cuticular ring

surrounding the base of the setae is obsolete and the mesonotum is usually im-

maculate.

Description

Length: 4.5-Smm; width; 3-3.5mm. Body slightly elongate oval and finely

setiferous, but the setae with very conspicuous and well sclerotized cuticular rings.

Head light yellow except eyes, antennae and maxillary palpi brownish.

Pronotum pale light yellow with two pairs of apical and two pairs of basal

subquadrate spots. Meso- and metanotum pale yellowish, each with one pair of large

subcircular dark brown spots. Elytron dark brown along sutural area and gradually

lighter toward lateral margin; basal, subbasal, apical and subapical areas light

yellow.

Dorsal surface of abdomen light yellow, abdominal terga 1, 8, and 9 immaculate.

The rest of the terga each have at least one pair of dorsal dark brown spots, except the

second and third which have an additional pair of spiracular spots.

Abdominal pleura light yellow and immaculate, but the third one dark brown

except for the margins.

Cycloneda sanguinea (Linnaeus)

Specimens examined

The study was based on 30 pupae from the U.S. National Museum collected in

Florida on May 1875 and on 31 March 1971.

Description

Similar to Cycloneda munda except that the head is entirely brown except for the

mediolongitudinal pale line, and the pronotum is yellow, with the marginal spots

greatly reduced or wanting. Mesonotum usually pale and immaculate or with very

small spots. Setae with inconspicuous cuticular rings.

60 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Genus MULSANTINA Weise

Mulsantina picta (Randall)

(Figs. 33, 53, 80, 122, 139)

Specimens examined

The study was based on 12 pupae which were reared in the laboratory from adults

collected in Clinton Co., Michigan, 30 May 1971, by Dang T. Phuoc. All specimens

are deposited in the Entomology Museum at Michigan State University and in the

U.S. National Museum.

Diagnosis

As mentioned before, this species is very closely related to Cycloneda (see

Cycloneda diagnosis for separation of these 2 genera). Adalia, Coccinella, Cycloneda,

Mulsantina and Propylaea represent a group of species of Coccinellini without a

mesal process at basal one-third of the urogomphus. Mulsantina differs from

Propylaea by having very fine and normal setae, whereas in Propylaea the setae

appear more coarse and are borne on a small tubercle (chalaza) with the distal end

strongly curved. See Adalia and Coccinella for the diagnostic characters of these two

genera.

Description

Length: 6mm; width: 3.5mm. Body elongate oval, extremely pale yellowish and

finely setiferous.

Head mostly pale and very smooth. Eyes brown, much darker than face. Antennae

long, extending about two-thirds of the distance between the widest lateral margin of

the pronotum and the eyes. Antennae dark brown on upper surface; club distinctly

cylindrical elongate or subrectangular block-shaped with 4 rings of papillae;

flagellum long and subquadrate in cross section; scape somewhat rectangular and

slightly convex (Fig. 53). Clypeolabrum smooth and pale, slightly wider than long,

with lateral margins subparallel. Apical margin concave and apical angles rounded

(Fig. 33). Mandibles bifid at apex and with molar surface well sclerotized. Maxillary

palpi trapezoidal, glabrous, slightly rugose and brownish, usually darker along lateral

sides; surface of the galea armed with more than 5 very fine but conspicuous teeth

(Fig. 80). |

Pronotum deeply marginate along apical margin, surface slightly rugose, mostly

pale and finely setiferous. In dark forms, the pronotum usually with 2 pairs of apical

spots (the inner larger and suboval), and 2 pairs of basal spots (the inner also much

larger than the outer and comma-shaped) (Fig. 122). Prothoracic spiracle elongate

oval. In most cases, mesonotum entirely pale except for very light brown or brown

margins. In dark forms, the mesonotum is light brown, with the anterior angles and

lateral sides of scutellar area brown. Metanotum pale in all cases, usually with one

pair of large boot-shaped spots (Fig. 139). Elytron pale or light brown, surface

micronodulate (as in Fig. 7). Sutural margin dark, and epipleura pale, wide and

slightly concave. Hind wing membranous, inconspicuously micronodulate along mid-

ventrolateral margin but never on apex. Legs dark brown at femorotibial joints and

tarsi; surface of femorotibial joints micronodulate.

Phuoc and Stehr: Pupae of Coccinellidae 61

Abdominal terga pale and maculate, usually with two pairs of subquadrate spots

on each (one pair spiracular and one pair dorsal) except the Ist, 8th and 9th which are

completely pale and immaculate, and the 6th and 7th which lack subspiracular spots.

In light forms, the spiracular spots are greatly reduced in size, and the posterior

margin of terga 3 to 5 is finely dentulate. Abdominal spiracles circular or nearly so,

with the first 5 well sclerotized. Urogomphi with base simple and slightly darker than

abdominal sterna. Abdominal pleura subquadrate, pale and immaculate except for

the 3rd which has a light brown spot. Ventral surface of abdomen completely pale

and immaculate.

Mulsantina hudsonica (Casey)

(Figs. 123, 140)

Specimens examined

The study was based on 4 pupae and an exuvia collected in Chippewa Co.,

Michigan, 19 June 1972, by Dang T. Phuoc. The specimens are deposited in the

Entomology Museum at Michigan State University and in the U.S. National

Museum.

Description and diagnosis

This species is very similar to dark forms of Mulsantina picta, except that the

mesonotum is mostly brownish and the metanotal spots are large and subquadrate

(Fig. 140). Length: 5.5mm; width: 3-Smm. Hind wing apex micronodulated.

Genus OLLA Casey

Olla abdominalis (Say)

(Figs. 34, 138, 225, 226, 245)

Specimens examined

The study was based on a single pupa from the U.S. National Museum collected in

Folsom, California, July 1885, and 8 pupae collected in Gainsville, Florida, on 26 July

1972 by R. E. Waites. Two of the Florida specimens are deposited in the Entomology

Museum at Michigan State University, the remainder have been returned to Dr.

Waites, Department of Entomology and Nematology, University of Florida.

Diagnosis

This is the only species of Coccinellini examined which has the body rounded and

strongly convex dorsally, with the dorsum apparently glabrous and pale yellowish or

silky white with well-defined dark brown rounded spots. Other members of the tribe

usually have a slightly elongate oval body and are finely setiferous. The strongly

convex antennal scape, giving sharp separation between the scape and the flagellum,

and the presence of a mesal process on the urogomphus (Figs. 225, 226) separates this

species from Cycloneda and Mulsantina whose antennal scape appears flat or very

slightly convex anteriorly, and whose urogomphus is without such a mesal process.

62 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Furthermore, Olla can be separated trom Neoharmonia by the mesal process ap-

pearing short and blunt, and the dorsum being smoother. In Neoharmonia the mesal

process of the urogomphi is slender and more pointed, and the dorsum is rugose and

punctate.

Description

Length: 4mm; width: 3.5mm. Body rounded and convex dorsally, yellowish or silky

white, with light brown to dark brown and well defined rounded spots on dorsal

surface, and very finely and sparsely setiferous.

Head yellowish to light brown, surface smooth and somewhat shiny. Eyes brownish.

Antenna yellowish to brown, and extending over two-thirds of the distance between

the widest lateral margin of the pronotum and the eye. Antennal club subspherical

and distinct, with three rings of papillae, the last ring incomplete by lacking some of

the anterior papillae. Flagellum subquadrate in cross section. Scape strongly convex

anteriorly and expanded caudally. Clypeolabrum gradually narrowed and slightly to

deeply concave at apical margin (Fig. 34). Mandibles bifid at apex. Maxillary palpi

large, brownish and trapezoidal; surface slightly rugose, galeal surface smooth and

pale.

Pronotum basically yellowish, strongly marginate along lateral and especially along

anterior margin (Fig. 138). Marginal areas brownish, with one pair of small, rounded,

dark brown anterodiscal spots and 2 pairs of rounded or subquadrate outer and inner

basal spots which are three times as large as the anterodiscal spot (Fig. 138).

Prothoracic spiracle pale and elongate oval. Meso- and metanotum silky white,

rugose with one pair of subcircular well-defined dark brown to black spots on each.

Elytron yellowish at base and apex with three large brown longitudinal sutural,

median and lateral stripes extending from base to the apical one-third (Fig. 245). The

median stripe may be broken at the basal one-third to give a subbasal oval or rounded

spot (Fig. 245). The pupa from the U.S. National Museum has the elytron entirely

pale yellowish (probably faded). Epipleura wide and concave, with lateral angle

slightly expanded anteriorly, and elytral surface densely micronodulate.

Hind wings slightly sclerotized at apex, surface smooth. Legs short and pale except

femorotibial joints and tarsi yellowish or light brown.

Abdominal terga silky white, each with one pair of dorsal and one pair of

spiracular, well-defined, dark brown spots about one-third as big as the metanotal

spots, except that terga 1, 8 and 9 are immaculate, and the dorsal spots on tergum 2

are absent. Urogomphi slightly branched mesally at about basal one-third into a

short and dull spine-like process; urogomphal disk with inner lobe elongate oval and

small, one-fourth as large as the outer one (Fig. 226). Abdominal pleura subquadrate

and yellowish, with posteriolateral angle slightly expanded caudally. Each pleuron

with a rounded brown spot but getting gradually lighter and disappearing in pleura 6

to 9. First two pleura entirely pale and immaculate. Abdominal sterna immaculate

and yellowish.

Phuoc and Stehr: Pupae of Coccinellidae 63

Genus PROPYLAEA Mulsant

Propylaea quatuordecimpunctata (Linnaeus)

(Figs. 44, 56, 172)

Specimens examined

The study was based on 4 pupae of Propylaea quatuordecimpunctata which were

reared in the laboratory from adults given by Dr. R. D. Eikenbary of Oklahoma State

University. The specimens are deposited in the Entomology Museum of Michigan

State University and the U.S. National Museum.

Diagnosis

This is the only known species of Coccinellini with very long antennae with a C-

shaped flagellum and with the tip partly hidden under the front femur. In addition

only abdominal terga S to 7 are “detachable” from their margins, whereas all other

members of the Coccinellini have terga 4 to 7 movable. Furthermore, Propylaea shows

some degree of close affinity to Anisocalvia by the presence of chalaza-type setae on

both forms. However, Propylaea can be separated from Anisocalvia by the lack of

lateral processes on abdominal pleura 3 to 5S, and by the characteristic, long, C-

shaped antennae.

Description

Length: 5mm; width: 4mm. Body oval, slightly elongate, light brown, with poorly

defined dark brown spots on dorsal surface. Body somewhat coarsely and densely

setiferous, and bearing chalaza-type setae whose tips strongly curve to the body

surface.

Head entirely pale and smooth. Eyes light brown. Antennae (Figs. 44, 56) light

brown, very long, and if straight, extending considerably beyond the widest lateral

margin of the pronotum. Antennae actually with six rings of papillae, the first two

rings complete with well developed papillae, but the last four incomplete, with dorsal

and anterior papillae wanting, or greatly reduced in size, especially in the sixth.

Antennal club not so distinct, with the diameter subequal to that of the flagellum and

hidden under the front femur. Flagellum long, ‘‘C’’-shaped and subquadrate in cross

section, with the proximal end somewhat flattened and slightly notched at lower side.

Scape oval and distinct. Clypeolabrum smooth and subquadrate, with the apex

abruptly narrowed apically and with the apical margin slightly concave (Fig. 44).

Mandibles bifid at tip. Maxillary palpi large, trapezoidal and light brown at the apex.

Galea with upper inner surface armed with several very fine teeth. Labial palpi

rounded, smooth and light brown at tip.

Pronotum deeply marginate apically, usually pale at apical angles, dark brown

along the posterior margin, and gradually lighter toward apical margin.

Mediolongitudinal line pale. Prothoracic spiracles elongate oval. Mesonotum light

brown, scutellar area slightly elevated and pale. Metanotum pale with two large,

poorly defined, dark brown spots. Elytron with lateroapical angle obtuse and

rounded, and with a large, irregular, brown, transverse band extending from sutural

64 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

area to the lateral margin at the basal one-fifth to three-fifths, and becoming

gradually lighter along the sutural area toward the base and apex. Epipleura wide

and slightly concave. Legs pale, except light brown at coxae, femorotibial joints and

tarsi. :

Abdominal terga yellowish, immaculate, except terga 3 and 4 each with one pair of

large subquadrate spiracular spots which are considerably lighter or absent on the

fifth or sixth terga. On the whole, the maculation is usually not well-defined.

Urogomphi straight along inner lateral margins and as dark as the previous

tergum. Abdominal spiracles circular or nearly so and very slightly sclerotized.

Abdominal pleura subquadrate, immaculate, light yellow to light brown, with pleura

3, 4, and 5 very coarsely and densely setiferous along the lateral margins (Fig. 172).

Abdominal sterna immaculate and pale. In the 2, abdominal sternum 9 is bipartite

and mammillate, and in the o" sternum 9 is rectangular, flat or very slightly convex.

Genus NEOHARMONIA Casey

Neoharmonia venusta (Melsheimer)

(Figs. 40, 111, 171, 222)

Specimens examined

The study was based on a single pupal exuvia of Neoharmonia venusta from U.S.

National Museum, collected in Brownsville, Texas, 18 April 1944 by Callaghan.

Diagnosis

This species is representative of a group of species of Coccinellini (including Anatis,

Ansocalvia and Synonycha) which possess a mesal process at about the basal one-

third of the urogomphus (Fig. 222). However, the lack of a lateral process on ab-

dominal pleura 3 to 5 separates this species from the others.

Description

Length: 5-7mm; width: 4-Smm. Body slightly elongate and finely setiferous.

Head light brown. Eyes brownish. Antennae long, extending about two-thirds the

distance between the widest lateral margin of the pronotum and the eye. Antennal

club subspherical, with three rings of distinct and well-developed papillae, the dorsal

papillae of the third ring lacking. Flagellum subquadrate in cross section. Scape

somewhat flat and slightly expanded caudad. Clypeolabrum smooth, gradually

narrowed and slightly concave at apex, with rounded apical angles (Fig. 40).

Maxillary palpus large and trapezoidal, with outer lateral side slightly convex.

Pronotum rugose and deeply marginate along lateral and especially along the

anterior margin, with two pairs of large oval brown spots at the base and one pair of

large anterior spots (Fig. 111). Mesonotum rugose and dark brown. Metanotum

yellowish with two large subquadrate dark brown spots. Elytron finely setiferous and

immaculate, brownish, with basal marginal area pale. Epipleura wide, slightly

concave, and with the lateral angle of the elytron slightly expanded anteriorly.

Abdominal terga slightly rugose and punctate, yellowish to brownish, and with

poorly defined dark brown spots. Each tergum usually with one pair of dorsal and one

Phuoc and Stehr: Pupae of Coccinellidae 65

pair of spiracular spots, except that tergum 1 is pale and immaculate. Abdominal

spiracles circular, the first five well-sclerotized. Urogomphus branching mesally into

a long and slender spine-like process at basal one-fourth (Fig. 222). Abdominal pleura

yellowish or brownish, usually paler along marginal areas. Pleura 3 to 6 large,

subquadrate, with the posterior lateral angles slightly expanded caudad and deeply

rugose and punctate (Fig. 171). Sternum 9 bipartite and mammillate in the @.

Genus ANATIS Mulsant

Diagnosis

This genus is representative of a group of three genera of Coccinellini (Anatis,

Anisocalvia and Synonycha) which have a lateral process on abdominal pleura 3 to 5

(Fig. 178). Anatis differs from Anisocalvia by the body being much larger with more

well-defined spots on the dorsum, and by having the apical margin of the labrum

slightly sinuate (Fig. 32). Anatis differs from Synonycha by the pleural process being

much shorter (equal to or shorter than the corresponding pleuron).

Description

Length: 7-10mm, width: 5.5-6mm. Body light yellow with well-defined dark brown

spots on dorsal side and finely setiferous.

Head dark brown, smooth except median line pale. Eyes brownish. Antennae long,

extending to the lateral margin of the pronotum. Club not distinct, with the diameter

of the club subequal to that of the flagellum. Flagellum long and subquadrate in cross

section. Scape somewhat flat and slightly convex along lower margin. Antenna ac-

tually with 6 rings of papillae, the apical 4 rings with well developed papillae and

occupying the “‘club’’ of the antenna, the fifth and the sixth rings located on the

flagellum, with the papillae greatly reduced in size or flattened (Fig. 54).

Clypeolabrum gradually narrowed apically, with the apical margin slightly sinuate,

surface slightly rugose with a few shallow wrinkles (Fig. 32). Mandibles bifid.

Maxillary palpus smooth, large and trapezoidal, with lateral margins parallel (Fig.

32). Surface of galea armed with several fine teeth, especially on the anterior inner

angle (Fig. 79). Labial palpi slightly rugose at tip, and one pair of fine teeth may be

present on the ligula (Fig. 98).

Pronotum pale and strongly marginate along lateral and anterior margins, with

dark brown spots along antero- and basomarginal areas; lateromarginal areas always

pale (Figs. 120-121). Prothoracic spiracle elongate oval. Mesonotum with scutellar

area elevated and usually with one pair of rounded spots anterior to the “‘scutellum.”’

Metanotum slightly elevated along mediolongitudinal line with one large pair of

subquadrate spots. Elytron pale yellowish, maculate and finely setiferous. Epipleura

wide and concave; lateroapical angle obtuse and almost continuous with lateral

margin. Legs light brown except femoral bases and tibiotarsal joints. Tibial papillae

obtuse.

66 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Posterior margins of abdominal terga 4 to 6 smooth. Urogomphi branched

mesally at basal one-third into a long spine-like process. Abdominal spiracles cir-

cular or nearly so and slightly sclerotized. Abdominal ‘pleura 3 to 5 large and with a

spine-like process on each which projects laterocaudad from posteriolateral angle

(Fig. 178). Abdominal sterna pale and immaculate, with the first 2 hidden under the

metacoxae, but visible and distinct medially. The third sternum the largest, with

anterior margin deeply sinuate. In the , abdominal sternum 9 bipartite and

mammillate, and more homogeneously and more strongly sclerotized than the

previous sterna, whereas in the % sternum 9 remains flat and subrectangular.

Key to Species of the Available Pupae of Anatis

1 Elytral apex pale (Fig. 153); lateral margin of

pronotum brown or light brown at posterior end

(Fig. 120); first abdominal tergum maculate

PIGS SEPT cee. bee oe ens Anatis ocellata (Linnaeus)

ss Elytral apex brown (Fig. 154); lateral margin of

pronotum entirely pale (Fig. 121); first abdominal

Terou MItaaciiate . ooo, hn me yn Anatis quindecimpunctata Olivier

Anatis ocellata (Linnaeus)

(Figs. 32, 54, 79, 90, 98, 120, 153, 178, 224)

Specimens examined

The study was based on 14 pupae which were reared in the laboratory from adults

collected in Rose Lake Wildlife Experiment Station, Clinton Co., Michigan, 25 May

1971, by Dang T. Phuoc. The specimens are deposited in the Entomology Museum at

Michigan State University and the U.S. National Museum.

Diagnosis

Two species of Anatis are known as pupae. They can be separated from one

another by the elytral apex being brownish and the first abdominal tergum maculate

in A. gquindecimpunctata, whereas in A. ocellata the elytral apex appears pale and the

first abdominal tergum is immaculate.

Description

Length: 7-10mm, average 9mm; width: 5.5-6mm. Head brown except frontal area

pale; surface smooth except clypeolabral area slightly rugose.

Pronotum (Fig. 120) with two pairs of subquadrate spots along apical margin, one

pair of extremely elongate oval spots at base and two large ‘‘L’’-shaped spots at

posterior angles. Mesonotum brown at anterior angles and along lateral margins, and

with one pair of small circular spots located anterior to the scutellar area. Metanotum

with one pair of large subquadrate spots. Elytron pale yellowish and maculate (Fig.

Phuoc and Stehr: Pupae of Coccinellidae 67

153), with sutural marginal area dark brown except for the subapical portion.

Epipleura pale with a large subquadrate subbasal spot and a lighter distal spot.

Elytral surface with three large circular spots (consisting of a humeral and two

sublateral at basal two-fifths and three-fifths respectively), and with two

mediosutural, elongate oval, large spots at basal one-third and two-thirds respec-

tively. Basal area of elytron light brown.

Each abdominal tergum usually with one pair of large spiracular subquadrate

spots and one pair of large subrectangular dorsal spots except tergum 1 where

spiracular spots are absent. Terga 7, 8 and 9 immaculate. All abdominal pleura

immaculate and yellowish, but pleuron 2 with a small light brown spot and pleuron 3

with a large subquadrate spot. Mesal spine-like process at basal one-third of

urogomphus short and obtuse (Fig. 224).

Anatis quindecimpunctata Olivier

(Figs. 121, 154, 223)

Specimens examined

The study was based on 4 pupae which were reared in the laboratory from adults

collected in Rose Lake Wildlife Experiment Station, Clinton Co., Michigan, on 24

May 1971, by Dang T. Phuoc. The specimens are deposited in the Entomology

Museum at Michigan State University and in the U.S. National Museum.

Description

Length: 7-10mm; width: 5-6mm. Similar to Anatis ocellata, but maculation slightly

different. Lateral margin of the pronotum entirely pale, with the spot at posterior

angle subtriangular (never ‘“‘L’’-shaped or claw-like). Lateral sides and anterior angles

of mesonotum pale. Elytral apex and in some cases epipleura entirely light brown

(Fig. 154); basal area pale. First abdominal tergum entirely pale, spots greatly

reduced in size, and spiracular spots on terga 5 and 6 also reduced in size or wanting.

Mesal spine-like process at base of urogomphus long and slender with apex somewhat

pointed (Fig. 223).

Genus ANISOCALVIA Crotch

Anisocalvia quatuordecimguttata (Linnaeus)

(Figs. 45, 177, 221)

Specimens examined

The study was based on one pupa and one exuvia saved from an emerged adult in

Chippewa Co., near Tahquamenon State Park, Michigan, 18 June 1972 by Dang T.

Phuoc. The specimens are deposited in the Entomology Museum at Michigan State

University.

68 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Diagnosis

This genus is very closely related to Anatis, with a lateral spine-like process at

posteriolateral angle of abdominal pleura 3 to 5, and long antennae. However, the size

is considerably smaller than Anatis, about 5mm in length, the apical margin of

clypeolabrum is deeply concave, and the pleurolateral process is conical and densely

setiferous with chalaza-type setae which are strongly curved distally (Fig. 177).

Description

Very similar to Anatis in many respects except for the following:

Length: 5mm; width: 3.5mm. Head brownish along marginal areas, including eyes,

antennae and maxillary palpi. Clypeolabrum subtrapezoidal, with apical margin

deeply concave and apical angles subpointed (Fig. 45).

Spot pattern on dorsal surface of the body very similar to Anatis ocellata, except

that here the spots are very large, almost filling the segment, and are usually

subquadrate or subrectangular and not as well defined, especially those on the ab-

domen. Elytron light brownish, with 2 large longitudinal brown bands extending from

base to about basal one-half, the sutural band appearing much darker than the

sublateral band. Pleurolateral process on abdominal segments 3 to 5 conical and

densely setiferous, with chalaza-type setae which are strongly curved distally (Fig.

177). Posteriolateral angle of pleuron 6 angulate and densely setiferous.

Genus SYNONYCHA Mulsant

Synonycha grandis (Thunberg)

(Figs. 51, 176)

Specimens examined

The study was based on two broken pupal exuviae of Synonycha grandis from U.S.

National Museum, collected in Buitenzog, Java, by Bryant and Palmer.

Diagnosis

This genus can be quite easily distinguished from Anatis and Anisocalvia by the

short antenna which extends about half way between the widest lateral margin of the

pronotum and the eye, by the presence of only 4 rings of papillae, and by the

pleurolateral spinelike process of abdominal segments 4 to 6 being cylindrical and

slightly curved cephalad, with the length exceeding the greatest width of the

corresponding pleuron. ;

Description

Length: over 10mm (10-12mm); width: 6-8mm. Body larger than Anatis. Head

entirely brown, smooth, except for clypeolabrum and genae which are slightly rugose.

Antennae short, extending about half way between the lateral margin of the

pronotum and the eye. Antennae with four rings of papillae. Flagellum short and

subtriangular in cross section. Scape greatly expanded and somewhat flattened (Fig.

55). Clypeolabrum large with apical margin convex. Mandibles bifid at apex.

Phuoc and Stehr: Pupae of Coccinellidae 69

Maxillary palpi smooth, with outer lateral margin slightly sinuate and very obtuse at

apex. Surface of galea covered with a few very fine teeth. Labial palpi smooth.

Pronotum deeply marginate, especially along apical and lateral margins. At least

one pair of large subquadrate spots at apical margin, one pair of subrectangular spots

at posterior angles, and one pair of subquadrate spots at base. Prothoracic spiracle

elongate oval. Meso- and metanotum pale, each with a pair of large brown spots.

Elytron brown except for pale lateral margin, apex and sutural area. Epipleura light

brown. Hind wing apex glabrous and smooth. Maculation on abdominal terga as in

Anatis ocellata, except that the Ist tergum is immaculate. Urogomphi branched

mesally at about basal one-third into a long spine-like process.

Abdominal spiracles circular or nearly so and slightly sclerotized. Abdominal

pleura immaculate and pale, except the third and the fourth which have a large

subquadrate spot. Pleural spine-like process longer than the greatest width of the

corresponding pleuron, and projecting laterocephalad (Fig. 176). Ventral abdomen

immaculate and pale.

Genus HIPPODAMIA Dejean

Diagnosis

This genus is representative of a group of genera of Coccinellini including

Coleomegilla, Naemia, Paranaemia and Eriopis which possess long and slender legs,

with the front femora considerably extended beyond the widest lateral margins of the

pronotum. This character separates this group from the remaining species of the

Coccinellini. Hippodamia in turn, is distinguished from other members of the group

by the glabrous condition of the exposed intersegmental conjunctivae of abdominal

segments 3 to 7 and by the lateral angle of the elytron which is greatly expanded

anteriorly into a rounded distinct process (Figs. 164-170).

Description

Length: 4mm-6mm; width: 2.5mm-3.5mm. Body elongate oval, finely setiferous,

bright yellow and distinctly maculate. Surface of the body and elytra densely

micronodulated.

Head deeply rugose, dark brown, always with mediolongitudinal pale yellowish line

running the length of the head. Bright yellow frontal spots sometimes present. Eyes

brown. Antennae dark brown and long, extending to the widest lateral margins of the

pronotum, with 4 rings of well developed papillae; club subspherical and distinct,

with the diameter much larger than that of the flagellum which is subquadrate in

cross section; scape strongly convex and expanded caudally (Fig. 47). Clypeolabrum

deeply rugose and dark brown, slightly longer than wide, with apical margin concave

to deeply cleft. Mandibles bifid at tip. Maxillary palpi large, lanceolate, dark brown

and rugose. Labial palpi dark brown and smooth.

Pronotum bright yellow with dark brown spots along marginal areas (Figs. 128-

137). Lateral margin strongly expanded into knife-like edge (Fig. 133). Mesonotum

with rugose anterior angles and elevated scutellar area. Metanotum smooth, with 2

large dark brown spots. Elytron yellow with lateral and sutural areas, lateral angle

70 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

and apex brownish to dark brown. Elytron with a humeral spot and a mediosutural

spot which are subequal, small, well defined and distinct; also with a large medial

spot, three to four times as large as the humeral or mediosutural spot, and often fused

with the sutural dark area. Lateral angle greatly expanded anteriorly into a

prominent thumb-like rounded lobe (Figs. 158-159); lateral margin strongly

marginate and the epipleura wide and slightly concave. Hind wing membranous,

tapered apically and glabrous. Legs long, with front femora extending considerably

beyond lateral margins of the pronotum (Figs. 41, 47), and dark brown except for

femoral bases and distal ends of tibiae which are pale. Tarsi light brown.

Abdominal terga bright yellow with large dark brown spots. Dorsal tubercles well

developed (Fig. 244). Posterior margins of terga 3 to 6 conspicuously spinulate.

Exposed intersegmental conjunctivae of abdominal segments glabrous. Abdominal

spiracles circular or nearly so, except the first which is elongate oval and partly

hidden under the elytra. Abdominal pleura subquadrate. Abdominal sterna im-

maculate pale yellowish or brownish.

s Key to Species of the Available Pupae of Hippodamia

1 Elytron with a subscutellar L-shaped spot

elias 4 aes bane ts ae weias 4) « el ppodamia quinquesignata (Kirby)

ie Elytron witltour a supscutellar L-shaped spot) oS. Be ee re PP Z

ZF) Clypeolabrum subquadrate with apical margin slightly

concave and never notched (Figs. 47, 48);

lateral angle of the elytron longer than wide

ey ee eee ee Hippodamia parenthesis (Say)

2 ~ Clypeolabrum not quite subquadrate, with apical

margin deeply notched (Fig. 41); lateral angle

of the elytron as wide as long or slightly wider

Pi Se) ak a, ulead he Kans sandtaw : dda 3

a2) Scutellar area of mesonotum at least dark brown at

base; lateral margin of the pronotum brownish to

Cae Ge el Oe Aoi cng Sirona ah be cade a Kal outers etintia ins <a eats a

3’ Scutellar area of mesonotum always pale yellowish;

lateral margin of the pronotum always pale yellowish

CF psc) ZOE ise ote vee Hippodamia tredecimpunctata (Linnaeus)

4(3) Elytron with a subbasal transverse band extending

from subsutural area to humeral angle

(rigs: 1664168) foe oh ea Hippodamia quinquesignata (Kirby)

4’ Elytron without such a subbasal transverse band

ee UO ae CE ete oe reine Cree mmr cea tek wer. Bie >

Phuoc and Stehr: Pupae of Coccinellidae 71

5(4’) Dorsal tubercles on abdominal segments very well

developed, with the tip directed caudally as seen

latetally (Pig 244)" a. Sy PEE Hippodamia convergens Guerin

gut Dorsal tubercles on abdominal segments not as well

developed, and slightly elevated above abdominal terga,

the tip never directed caudally as seen

Pater aly Beh SES OR Hippodamia glacialis (Fabricius)

Hippodamia tredecimpunctata (Linnaeus)

(Figs. 128-130, 160, 161)

Specimens examined

The study was based on 20 pupae reared in the laboratory from adults collected in

Fast Lansing, Michigan, 21 August 1968, and eight pupae collected in East Lansing

on 19 July 1972, by Dang T. Phuoc. The specimens are deposited in Entomology

Museum at Michigan State University and the U.S. National Museum.

Diagnosis

This species, along with the other known species of Hippodamia, including H.

convergens, H. glacialis and H. quinquesignata, can be separated from H. parenthesis

by the apical margin of the clypeolabrum being deeply notched, by the apical angles

being subpointed and strongly projected caudally, and by the lateral angle of the

elytron being no longer than the base of the angle. In H. parenthesis, the

clypeolabrum appears subquadrate, with the apical margin slightly concave or

subtruncate, and never with the apical angles projected caudally. In addition, the

lateral angle of the elytron appears longer than the basal width. H. tredecimpunctata

in turn is separated from H. convergens, H. glacialis and H. quinquesignata by the

always pale scutellar area of the mesonotum and the pale lateral margin of the

pronotum. ..these areas of the last three species are brownish or dark brown.

Description

Length: 4.5mm-Smm; width: 2.5mm-3mm. Head rugose, dark brown except for

mediolongitudinal pale yellowish line; frontal area may be pale. Antennae light to

dark brown. Clypeolabrum slightly longer than wide, rugose with lateral margins

subparallel but abruptly narrowed apically, the apical margin deeply notched with

the apical angles projected caudally and pointed (as in Fig. 41).

Pronotum yellowish, rugose along anterior marginal area. In light forms there are

two pairs of subcircular spots at the anterior and posterior angles (Fig. 128) while in

dark forms the anterior and posterior spots are connected anteroposteriorly by a large

longitudinal band into one pair of elongate, hourglass-shaped spots (Fig. 129). One

pair of large subrectangular anterior spots, and one pair of small basal oval spots may

be present. In some cases, all of these spots may be fused together; then the pronotum

becomes black or dark brown with one pair of rounded, oval, lateral, yellow spots,

and one large heart-shaped discodorsal yellow spot (Fig. 130). Mesonotum yellow with

72 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

brown anterior angles; scutellar area always pale yellowish even in very dark forms.

Metanotum yellow with one pair of large oval or kidney-shaped lateral spots which

are greatly enlarged in dark forms so they fill the metanotal surface except for the

pale yellow mediodorsal area. Elytron, in light forms, with a humeral and

mediosutural spot at the basal one-third, and a very large (as large as the other two

spots combined) median spot at the basal one-half (Fig. 160). In dark forms the

elytron is completely dark brown or black.

Each abdominal tergum usually with one pair of spiracular and one pair of dorsal

subquadrate spots, except that tergum 1 is immaculate or with one pair of small

dorsal oval spots, terga 8 and 9 are immaculate and pale, and tergum 4 lacks

spiracular spots. Terga 2 and 3 are in some cases almost entirely dark brown due to

the spiracular and dorsal spots being greatly enlarged and fused except for the

median pale area. Abdominal pleura yellow and immaculate except that the exposed

portions of pleura 2 and 3 are dark brown.

In dark forms, dorsum of abdomen almost entirely dark brown or black except for

a median pale yellow spot which is light-bulb-shaped on segment 2, narrowly elongate

subrectangular on segment 3, and large-mouthed-vase-shaped or subquadrate on

segments 4 and 5. Terga 1, 7 and 8 yellow, each with one pair of subquadrate dorsal

dark brown spots. Tergum 4 with an additional pair of large subquadrate spiracular

yellow spots. Abdominal pleura half pale and half dark except for pleuron 3 which is

entirely black.

Hippodamia parenthesis (Say)

(Figs. 47, 48, 131, 134, 159, 163, 227, 228)

Specimens examined

The study was based on six pupae and one pupal exuvia reared from adults

collected in East Lansing, Ingham Co., Michigan, on 25 July 1971, by Dang T. Phuoc.

The specimens are deposited in the Entomology Museum at Michigan State

University and in the U.S. National Museum.

Diagnosis

This is the only species among the known pupae of Hippodamia with the apical

margin of the clypeolabrum slightly concave and never deeply notched, and with the

lateral angle of the elytron longer than basal width (never shorter than basal width as

in the remaining known pupae of Hippodamia).

Description

Length: 4mm-4.5mm; width: 2.5mm. Head dark brown with one pair of irregular

8-shaped or rounded yellow spots. Clypeolabrum dark brown and subquadrate with

apical margin more or less truncated or slightly concave (Fig. 47).

Pronotum slightly marginate, bright yellow. In light forms, marginal spots are

distinctly present, consisting of two pairs of large subquadrate anterior marginal

spots and one pair of large, poorly defined spots at posterior angles, and one pair of

basal oval spots (Fig. 131). The outer anterior marginal spots and spots at posterior

Phuoc and Stehr: Pupae of Coccinellidae Ve

angles may fuse together along the lateral margins. The inner anterior marginal spots

may expand posteriorly toward the basal spots (Fig. 132, 134). Mesonotum brown at

anterior angle and base. Metanotum with one pair of very large bell-shaped spots.

Elytron dark brown over one-half of the total surface, with an elongate irregular 8-

shaped yellow spot. In light forms, elytral suture and apex brown, two subequal and

small spots respectively at humeral and mediosutural area at the basal one-fourth,

and a large median spot (as large as or larger than the two previous spots combined)

at the basal one-half are distinctly present (Fig. 163).

Abdominal terga bright yellow, each with two pairs of very large subrectangular

dark brown spots except that tergum 1 is immaculate or with one pair of small

rounded spots, and the pygidium is entirely immaculate. The tergal spots on the

abdominal segments are very large, often almost filling the tergal surface except for

the median area. Abdominal pleura brownish with marginal area yellowish; pleuron 3

in dark forms is entirely dark brown. Abdominal sterna yellowish to brownish and

immaculate.

Hippodamia glacialis (Fabricius)

(Figs. 169, 170)

Specimens examined

The study was based on three pupal exuviae from the U.S. National Museum,

collected in association with adults in Massachusetts by Dimmock.

Diagnosis

The maculation pattern of this species and Hippodamia convergens is very similar,

especially on the elytron. However, it can be separated from H. convergens by the

poorly developed dorsal tubercles on the abdomen, where in H. convergens the dorsal

tubercles are greatly developed and directed caudally (Fig. 244). This species also can

be distinguished from H. quinquesignata by the lack of a transverse band at the

subbasal area of the elytron. (See Hippodamia tredecimpunctata diagnosis for the

separation these two species and H. parenthesis).

Description

Length: 6mm; width: 3.5mm. Head rugose and dark brown with two yellowish

rounded frontal spots. Clypeolabrum dark brown and rugose with apical margin

deeply notched and apical angles subpointed and projected caudally. Antennal scape

strongly convex anteriorly.

Pronotum yellowish, slightly marginate along lateral and anterior margins and with

four pairs of spots. . .one pair of large subquadrate spots at medioanterior area which

are often expanded posteriorly, one pair of elongate oval small spots at the anterior

angles, one pair of oval spots at base, and one pair of subquadrate spots at the

posterior angles (Fig. 135).

Meso- and metanotum basically yellowish, mesonotum with one pair of small, and

metanotum with one pair of large, subcircular dark brown spots. Elytron yellowish

(Figs. 169, 170), with a subcircular humeral spot and a mediosutural spot about the

74 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

same size as the previous spot at the basal one-fourth, and a median spot twice as

large as the other two combined at the basal one-half. Lateral margin of the elytron

dark brown; sutural area and lateral angle brown.

All abdominal terga yellowish, each tergum with one pair of large subquadrate

dorsal spots and one pair of spiracular spots except for tergum 1 which is immaculate

or with one pair of small dorsal spots. Pygidium immaculate. Abdominal pleura

immaculate and yellowish except that pleura 3 and the exposed portion of pleuron 2

are dark brown.

Hippodamia convergens Guerin

(Figs. 41, 136, 137, 244)

Specimens examined

The study was based on five pupae and two pupal exuviae reared from adults

collected in East Lansing, Ingham Co., Michigan, on 30 June 1971, by Dang T.

Phuoc. The specimens are deposited in the Entomology Museum at Michigan State

University and the U.S. National Museum.

Diagnosis.

See diagnosis of Hippodamia tredecimpunctata, H. parenthesis, and H. glacialis

for the separation of this species from the group.

Description

Pupae of this species are very similar to Hippodamia glacialis, especially the

maculation pattern on the elytron.

Length: 5mm-5.5mm; width: 3mm-3.30mm. Head entirely dark brown except for

the pale yellowish mediolongitudinal line. The pronotum in dark forms has two pairs

of subequal subquadrate dark brown spots, one pair at the medioanterior marginal

area and the other pair at the base, and two other pairs of spots at the anterior and

posterior angles which are always fused together along the lateral marginal areas

(Figs. 136, 137). Discodorsal area of the pronotum varies from very light brown to

brown. Meso- and metanotum pale yellowish. Mesonotum with one pair of small, and

metanotum with one pair of large subtriangular spots. In dark forms, mesonotum

entirely dark brown except for the pale mediolongitudinal line. Scutellar area of the

mesonotum always brownish to dark brown.

The dorsal tubercles on the abdomen are characteristically well developed in this

species where they are greatly elevated and directed caudally.

Hippodamia quinquesignata (Kirby)

(Figs. 135, 158, 164-168)

Specimens examined

The pupae and adults were collected in Idaho in a cluster, mixed with several

different species, on 31 July 1969 by R. W. Portman. Based on the similarity to the

maculation pattern of the elytra of the adults pupae were carefully selected to have

Phuoc and Stehr: Pupae of Coccinellidae &

the greatest possibility of correct identification. The variation of maculation pattern

within the species is therefore very limited in this study.

The study was based on 16 selected pupae consisting of a single form. Four

specimens are deposited in the Entomology Museum of Michigan State University,

the rest were returned to the University of Idaho.

Diagnosis and description

Hippodamia quinquesignata is very similar to H. glacialis and H. convergens but

differs in the maculation pattern on the elytron. The humeral spot and the

mediosutural spot at the basal one-fourth are transversely fused into a transverse

band extending from the subsutural area to the humeral angle (Figs. 166-168). In

lighter forms, the subbasal transverse band on the elytron becomes obsolete; in this

case, an L-shaped subscutellar spot is characteristic of this species (Figs. 164, 165).

In addition, this species can also be separated from H. glacialis and H. convergens

by the eighth abdominal tergum being glabrous; in H. glacialis and H. convergens the

eighth abdominal tergum is finely setiferous.

Genus COLEOMEGILLA Timberlake

Coleomegilla maculata (DeGeer)

(Figs. 43, 124-126, 155, 156, 174)

Specimens examined

The study was based on 18 pupae and eight pupal exuviae which were reared from

adults collected in East Lansing, Ingham Co., Michigan, 30 June 1972, by Dang T.

Phuoc. The specimens are deposited in the Entomology Museum at Michigan State

University and in the U.S. National Museum.

Diagnosis

This species, along with the closely relaged genera Naemia and Eriopis, form a

group which is separated from Hippodamia by having the exposed intersegmental

conjunctivae finely setiferous, and by having the lateral angle of the elytron obtuse,

rounded, and almost continuous with the lateral margin (Figs. 155-156). Coleomegilla

in turn is separated from Naemia by the first abdominal spiracles being partly hidden

under the elytra (in Naemia the 1st abdominal spiracles are entirely hidden under the

elytra), and from Eriopis by the apical margin of the clypeolabrum being slightly

concave, whereas in Eriopis the apical margin of the clypeolabrum is deeply notched.

Description

Length: 4.5-Smm; width: 2.2-2.8mm. Body very elongate oval, yellowish and finely

setiferous, with well-defined dark brown spots on dorsal surface.

Head yellowish to light brown (supra ocular area sometimes brown) with the frons

always pale; surface smooth or slightly rugose. Antennae long, extending over two-

thirds of the distance between the eye and the widest lateral margin of the pronotum.

76 | Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Scape largely expanded caudally and flat. Flagellum elbowed and subquadrate in

cross section. Club distinctly subspherical and with three rings of papillae, although

the last ring is more or less obsolete. Clypeolabrum subquadrate, as long as or slightly

shorter than wide, with apical margin slightly concave and apical angles rounded

(Fig. 43). Mandible sharply bifid at tip. Maxillary palpi large, trapezoidal, yellowish

to brown and smooth. Galea smooth without teeth.

Pronotum yellowish, large, subrectangular, deeply marginate along anterior and

lateral margins, anterior margin strongly carinate and slightly projected anteriorally

(Fig. 126); a large pair of oval spots usually present at base (Figs. 124-125). Meso- and

metanotum each with one pair of large suboval dark brown spots in dark forms,

whereas in light forms, the spots on the mesonotum are wanting or very light brown.

Elytron smooth, finely setiferous, three times as long as wide. Epipleura wide and

slightly concave; lateral angle of the elytron rounded obtuse and almost continuous

with the lateral margin (Fig. 156). Elytra usually with well-defined dark brown spots,

consisting of a scutellar spot and a subsutural spot at basal two-thirds (these two spots

are usually fused with sutural dark area), a humeral rounded spot, and two median

spots, one at the basal two-fifths, and one at the apical one-fifth. The first medial spot

large and subrectangular, extending from the lateral margin almost to the sutural

dark area; the second medial spot is light brown and much smaller than the humeral

spot; and finally, a distolateral poorly defined light brown spot. In some cases, the

spots may become light or absent. Hind wing membranous, tapered apically and

glabrous. Legs long, with femora slender and somewhat cylindrical, extending

considerably beyond the widest lateral margins of the pronotum, finely setiferous and

brown to dark brown except for the pale femoral bases and tibiotarsal joints. Hind

femorotibial joints visible dorsally.

Abdominal terga yellowish, finely and densely setiferous (including median area).

Dorsal tubercles wanting. The exposed intersegmental conjunctivae finely setiferous.

Most terga usually with one pair of lateral and one pair of dorsal spots which are often

fused together except for the median area between the dorsal spots. Terga 7 to 9 are

immaculate, and tergum | is either immaculate or with 2 small poorly-defined dorsal

spots. Tergum 8, pygidium and urogomphi are glabrous. Abdominal spiracles cir-

cular or nearly so, except for the first which is large, elongate oval and mostly hidden

beneath the elytra. Abdominal pleura subquadrate with lateral margin convex;

surface smooth, yellowish and finely setiferous except for the first 2 pleura which are

pale and hidden beneath the elytra. Abdominal sterna immaculate, pale yellowish,

and finely setiferous except for sterna 8 and 9 which are glabrous.

Genus NAEMIA Mulsant

Naemia seriata (Melsheimer)

(Fig. 42)

Specimens examined

The study was based on two pupae, including one reared exuvia of Naemia seriata

collected in Mayo Beach, Maryland, 30 August 1944 by E. A. Chapin. The specimens

were loaned from the U.S. National Museum.

Phuoc and Stehr: Pupae of Coccinellidae da

Diagnosis

See diagnosis of Coleomegilla maculata for separation of these two species.

Description

The species is similar to Coleomegilla maculata in many respects except the

following:

Length: 4-4.5mm; width: 2.5mm. Clypeolabrum tapering apically. Apical margin

concave and narrow, about one-half as wide as the base (Fig. 42).

Lower margin of anterior edge of the pronotum descending about two-thirds of the

eye length (Fig. 42). Elytral maculation pattern as in Coleomegilla maculata, but

spots with an equal degree of brownness, and a common sutural and a common

scutellar spot present. Legs short, not extended much beyond lateral margin of the

pronotum (Fig. 42). Abdominal tergum 1 with two large but poorly defined dorsal

spots. First abdominal spiracles entirely hidden under the elytra.

Genus ERIOPIS Mulsant

Eriopis connexa (Germar)

(Figs. 7, 49, 127, 157, 175)

Specimens examined

The study was based on 10 pupae of Kriopis connexa reared in Berkeley, California,

byR.L. Tassan. The specimens were loaned from Dr. K. S. Hagen’s collection.

Diagnosis

This genus appears to have some close affinity to Hippodamia because the

pronotum is very slightly marginate and rugose, the antennal scape is strongly convex,

and the clypeolabrum is deeply notched at apical margin. In contrast, the wing form

shows more affinity to Coleomegilla in which the lateral angle is obtuse and almost

continuous with the lateral margin of the elytron. In general, this species is distinctly

separated from Coleomegilla by the sharply notched apical margin of the

clypeolabrum (Fig. 49) and by the slightly angulate lateral margins of abdominal

pleura 3 to 5 (Fig. 175) (in Coleomegilla the apical margin of the clypeolabrum is

slightly concave and the lateral margin of the abdominal pleura 3 to 5 is rounded

convex). It is separated from Hippodamia by the exposed finely setiferous in-

tersegmental conjunctivae on the dorsum of the abdomen, and by the obtuse and

rounded lateral angle of the elytron (in Hippodamia the exposed intersegmental

conjunctivae on the dorsum of the abdomen are glabrous and the lateral angle of the

elytron is greatly expanded anteriorly into a broad rounded lobe).

Description

Length: 5.5mm; width: 3mm. Body elongate, moderately convex dorsally and

apparently glabrous.

Head dark brown along lateral area, including eyes and antennae; discal area

yellow. Antennae long, extending to the widest lateral margins of the pronotum; club

with 4 rings of poorly developed papillae, with the diameter subequal to that of the

flagellum. Scape convex and rugose. Clypeolabrum large, dark brown except

medially, with apical margin angularly and deeply notched; surface slightly rugose.

Maxillary palpi large with lateral margin sinuate (Fig. 49).

78 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Pronotum subquadrate, about one-third wider than long, shiny yellow, slightly and

regularly rugose, very slightly marginate, and with 2 large but not well-defined basal

brownish spots (Fig. 127). Prothoracic spiracle rounded oval. Meso- and metanotum

each with one pair of subtriangular spots. Elytron yellow, elongate (3 times longer

than wide) with lateroapical angle obtuse, and almost continuous with lateral margin.

Sutural and lateral margin dark brown. Hind wings glabrous. Legs long and slender,

with front femora extending considerably beyond lateral margins of the pronotum.

Abdomen yellow and slightly rugose dorsally. Each tergum with one pair of large but

not well-defined subrectangular spots (except terga 1, 7, 8, 9 which are immaculate).

Exposed intersegmental conjunctivae on abdomen finely setiferous. Abdominal

pleura subquadrate, yellow and immaculate, with lateral margin slightly angulate

(Fig, 175),

TRIBE PSYLLOBORINI

Diagnosis

See diagnosis of Coccinellini for separation of these two tribes.

Genus PSYLLOBORA Chevrolat

Psyllobora vigintimaculata (Say)

(Figs. 46, 92, 229)

Specimens examined

The study was based on over 30 pupae of Psyllobora vigintimaculata which were

reared from adults collected in Clinton Co., Michigan, 10 June 1972 by Dang T.

Phuoc. The specimens are deposited in the Entomology Museum at Michigan State

University and in the U.S. National Museum.

Description

Length: 2.5-3mm; width: 1.5-1.8mm. Body slightly elongate oval, finely setiferous,

pale whitish, with a few dark spots on dorsal surface. Head pale except for light

brown eyes. Antennae very long with the tips hidden beneath the front femora, and

more than twice as long as the distance between the antennal bases. Club not distinct

from the flagellum and with 2 entire rings of well developed papillae (Fig. 46).

Clypeolabrum subrectangular, about twice as wide as long, with apical margin

truncated or very slightly concave (Fig. 46). Mandible bifid. Maxillary palpi bell-

shaped, with the apex greatly enlarged (Fig. 46); galea greatly enlarged and rounded

as seen from the top, with the greatest width as wide as the base of maxillary palpus

(Fig. 92). :

Pronotum pale whitish, with anterior margin slightly concave and _ slightly

marginate along anterior and lateral margins. Meso- and metanotum also pale

whitish, each with one pair of subrounded dark brown spots. Elytron subrectangular,

Phuoc and Stehr: Pupae of Coccinellidae 79

immaculate, whitish and finely setiferous. Lateral angle of the elytron obtuse and

lateral margin marginate. Legs short and pale except for terminal “‘segment”’ which is

brownish with pointed ‘“‘claws’’ (Fig. 46).

Abdominal terga pale whitish and immaculate, except that the first 2 terga have

one pair of poorly defined brownish spots on each, and the third tergum with 2 pairs

of more distinct spiracular and dorsal subquadrate spots. Urogomphi well developed

and similar to Coccinellini. Abdominal spiracles circular. Abdominal pleura pale,

subquadrate, and with the lateral margin rounded and convex, except for pleura 2

and 3 which are brownish at the posterior and anterior marginal areas respectively.

Abdominal sterna entirely immaculate and pale.

DISCUSSION AND CONCLUSIONS

Based on adults, the taxonomy of the Coccinellidae has been fairly well worked out.

From this standpoint, several systems of classification within the family have long

been proposed and used. The history of the classification of the family based on adult

characters was presented by Watson (1956) and by Sasaji (1968b, 1971). The system

which has been most accepted by many authors is that proposed by Korschefsky

(1931, 1932) in which the family Coccinellidae contains three subfamilies: the

EPILACHNINAE, the LITHOPHILINAE with the monotypic genus Lithophilus

Frolich, and the COCCINELLINAE which contains the majority of the members of

the family. The system has not been greatly altered except some _ tribal

rearrangements have been attempted. Watson (1956) proposed that the Coccinellini

should be recognized as the three different tribes Coccinellini, Hippodamiini and

Anisostictini after careful study of the morphology of the adults, despite the fact that

Boving (1917), an early author who carefully studied the larval stages of COC-

CINELLIDAE, pointed out that the COCCINELLINI and HIPPODAMIINI can not

be separated by using characters of the larvae.

A number of authors have worked on coccinellid larvae, including Boving (1917),

Gage (1919), Strouhal (1926), Rees (1947, 1948), Van Emden (1949), Kapur (1950) and

recently Savoiskaya (1960, 1962a, 1962b, 1962c, 1963, 1964a, 1964b), Kamiya (1965),

and Sasaji (1968a). Using larval characters Kamiya (1965) proposed new phylogenetic

relationships among coccinellid tribes in which he considered EPILACHNINAE to

have independently evolved from the COCCINELLINAE, the tribes COCCINELLINI

and PSYLLOBORINI to be closely related and highly developed, the tribes

HYPERASPINI, PLATYNASPINI, TELSIMINI, SCYMNINI, STETHORINI,

NOVIINI, and CHILOCORINI to have branched from another stem in which the

members of CHILOCORINI are most advanced, and the SUKUNAHIKONINI,

SERANGIINI and PHARINI to be another stem. He also emphasized that members

of HYPERASPINI and SUKUNAHIKONINI stand as the most primitive forms in

the family.

After careful study of adult and larval characters Sasaji (1968b) (newly adopted

name of Kamiya) came up with a new system of classification in which the family is

divided into six subfamilies, the STICHOLOTINAE, SCYMNINAE,

CHILOCORINAE, COCCIDULINAE, COCCINELLINAE and EPILACHNINAE.

80 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

He proposed a new tribal phylogeny and relationships which do not agree entirely

with his previous work. He thought the EPILACHNINAE were very closely allied with

the COCCINELLINAE, but had evolved divergently with the change of feeding habits

from carnivorous to phytophagous. He also emphasized that these two subfamilies

have evolved from a Sticholotine-like ancestor, with STICHOLOTINAE as the most

primitive group in the family. On the other hand, he believed the CHILOCORINAE

and SCYMNINAE are closely related and have evolved on a separate stem without

any close connections to any other groups. Finally, he stated that although both

groups were primitive, the COCCIDULINAE and STICHOLOTINAE did not have

great affinity. Moreover, within the COCCIDULINAE, the NOVIINI appear to have

a closer affinity to SCYMNINI or ORTALIINI than to the COCCIDULINI.

Therefore, the branching point of COCCIDULINAE was still very much in doubt.

Information on the pupae has never been used in the classification of the family

Coccinellidae. However, from the material examined in this study, it is possible to

arrive at some conclusions concerning the tribal relationships.

By the study of the comparative morphology of the pupae, the relative degree of

advanced or primitiveness of the pupal characters has been determined by the ap-

plication oof Maslin’s concepts (1952) on the use of morphological criteria for phyletic

relationships (see Table 1).

For the following discussion refer to the phylogenetic diagram (Fig. 266).

The COCCINELLINAE contains three tribes COCCINELLINI, PSYLLOBORINI

and DISCOTOMINI (Sasaji, 1971), but pupae for the last tribe were not available.

Based on pupal characters, the COCCINELLINAE is the most highly advanced

subfamily and has evolved along a distinct line in which they, without exception

among known pupae for the family, have distinct maculation on the dorsal body

surfaces, have exposed conjunctivae between the abdominal segments which allow for

expansion of the abdomen by means of a unique folding mechanism, and have fine

and rather inconspicuous setae.

The PSYLLOBORINI are almost identical to the COCCINELLINI except for their

smaller size and having a greatly enlarged galea. As Sasaji (1968b) pointed out, the

PSYLLOBORINI have recently diverged from the predaceous COCCINELLINI in

adopting tthe fungiphagous habit. This opinion is substantiated by the morphology of

the pupae.

The remainder of the family is distinctly separated from the COCCINELLINAE by

their usually pale and immaculate bodies, by the more or less compact abdomen in

which the abdominal terga are firmly attached to one another without the intervening

exposed conjunctivae, and usually by the presence of coarse, long, and very con-

spicuous setae. Therefore, they are probably a monophyletic group which evolved

from a common ancestor which is widely separated from the COCCINELLINAE.

The EPILACHNINAE split off early from this group, and independently acquired

the phytophagous habit and highly evolved mandibles whose tip is bifid and broadly

concave mesally.

Following the EPILACHNINAE, the STICHOLOTINAE-COCCIDULINAE group

diverged from the main stem of branch 1, evolving from a common ancestor which

acquired setae at the apex of the hind wings.

Pupae of Coccinellidae

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The STICHOLOTINAE, according to Sasaji (1971), consists of four tribes:

SHIROZUELLINI, STICHOLOTINI, SERANGIINI and SUKUNAHIKONINI.

Unfortunately, pupae were available for only the STICHOLOTINI and

SERANGIINI, but they appear to be the most primitive group within the family for

they have retained many primitive characters (see Table 1).

According to Sasaji (1971) COCCIDULINAE contains the tribes NOVIINI,

EXOPLECTRINI, LITHOPHILINI, AND COCCIDULINI. Only pupae of the

NOVIINI, COCCIDULINI, (and SCYMNILLINI) were available for this study.

Because of the presence of setae on the pupal hind wing apex, the SCYMNILLINI are

more closely related to the COCCIDULINI than to the SCYMNINI. Therefore, the

tribe SCYMNILLINI should be included in the COCCIDULINAE instead of the

SCYMNINAE. |

The COCCIDULINAE appear to be more advanced than the STICHOLOTINAE;

however, they are far less advanced than the remaining groups of the family. On one

hand, the COCCIDULINI appear to be more closely related to EPILACHNINI than

any other groups of COCCINELLIDAE outside of the COCCIDULINAE by retaining

the first four slightly pedunculated abdominal spiracles, and by having the

urogomphi slender with a simple apex. The NOVIINI, on the other hand, appear to

have a closer relationship to the CHILOCORINI (CHILOCORINAE) and the

HYPERASPINI (SCYMNINAE) than to the SCYMNILLINI and COCCIDULINI by

having short, clubless non-papillated antennae, bipartite urogomphi whose apices are

modified into a complex distal disk, and the subquadrate abdominal pleura in a

subvertical position.

The CHILOCORINAE contains three tribes according to Sasaji (1971), the

CHILOCORINI, PLATYNASPINI, and TELSIMIINI. Only pupae of the

CHILOCORINI were available. The CHILOCORINI appear to have evolved along a

separate branch by acquiring a broadly expanded clypeus. They show some degree of

relationship with the STICHOLOTINAE-COCCIDULINAE group by retaining the

primitive simple mandibles and the clubless non-papillated antennae which may be

found in the STICHOLOTINAE (Microweisea ovalis, Delphastus pusillus) and in the

SCYMNILLINI (Zagloba ornata).

Finally, according to Sasaji (1971) the SCYMNINAE consists of seven tribes, the

ORTALIINI, ASPIDIMERINI, SCYMNINI, HYPERASPINI, CRANOPHORINI,

STETHORINI, and SCYMNILLINI. However, the SCYMNILLINI have been shown

above to belong to the COCCIDULINAE. Only pupae of the SCYMNINI,

HY PERASPINI and STETHORINI were available. The SCYMNINAE have recently

branched from branch 1. The STETHORINI appear to be the most highly evolved

tribe with the frontal area entirely membranous, the well-defined pedunculate first

abdominal spiracles, and the glabrous frons, clypeolabrum and pygidium. By having

the well pedunculated first abdominal spiracles, STETHORINI appear to have close

relationships with the CHILOCORINI. In contrast, the HYPERASPINI are the most

primitive tribe in the SCYMNINAE, even though they have acquired some highly

evolved characters such as the strongly sclerotized and complex distal disk of the

urogomphi.

Phuoc and Stehr: Pupae of Coccinellidae 83

LITERATURE CITED

Binaghi, G. 1941. Larve e pupe di Chilocorini. Note sistematiche e morfogia degle

apparati genitali (Coleoptera, Coccinellidae). Mem. Soc. Ent. Italiana 20: 19-36,

Apls.

Boving, A. G. 1917. A generic synopsis of Coccinellid larvae in the United States

National Museum, with a description of the larva of Hyperaspis binotata Say.

Proc. U.S. Nat. Mus. 51: 621-650.

Coquillet, D. W. 1889. The imported Australian ladybird Vedalia. Ins. Life 2: 71-74.

Gage, J. H. 1919. The larvae of the Coccinellidae. Ill. Biol. Monogr. 6: 232-294, PI. 6.

Gorham, H. S. 1892. On the three new species of Coccinellidae from Scioa. Ann. Mus.

Genov. 2: 910-911.

Kanervo, V. 1941. Zur morphologie der praimaginalen stadien von Calvia 15-gutata

F. (Coleoptera, Coccinellidae). Ann. Ent. Fenn. 7: 52-57.

Kamiya, H. 1965. Comparative morphology of larvae of the Japanese Coccinellidae,

with special reference to the tribal phylogeny of the family (Coleoptera). Mem.

Fac. Lib. Arts, Fukui Univ., Ser. II, Nat. Sci. 14: 83-100.

Kapur, A. P. 1943. On the biology and the structure of the Coccinellid Thea

bisostonotata Mulsant in North India. Indian J. Ent. 5: 165-171.

1950. The biology and external morphology of the larvae of

Epilachninae (Coleoptera, Coccinellidae). Bull. Ent. Res. 41: 161-208.

Kesten, Von U. 1969. Zur morphologie and biologie von Anatis ocellata Linnaeus.

Zeits. Angew. Ent. 63: 412-445.

Korschefsky, R. 1931-1932. Coccinellidae-Coleopterorum Catalogus. Pt. 118 and 120.

W. Junk, Berlin, 659 pp.

Marthur, L. M. L., and B. K. Srivastava. 1966. On the immature stages of Stethorus

gilvifrons Mulsant. Beitr. Entomol. 16: 311-316.

Maslin, T. P. 1952. Morphological criteria of phyletic relationships. Syst. Zool. 1: 49-

70.

Pinto da Fonseca and Autuori, M. 1931. Contribucao para a biologica de Solanophila

clandestina (Mulsant). Rev. Ent., Sao Paulo 1: 219-224.

Palmer, A. M. 1914. Some notes on the life history of ladybeetles. Ann. Ent. Soc.

Amer. 7: 213-238.

Rees, B. E. 1947. Taxonomy of the larvae of some North American Noviini

(Coleoptera: Coccinellidae). Pan. Pacif. Ent. 23: 113-119.

Rees, B. E. 1948. The larva of Catana clausei Chapin, and its comparison with the

larva of Delphastus pusillus (LeConte). Proc. Ent. Soc. Wash. 50: 231-234.

84 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Sasaji, H. 1968a. Descriptions of Coccinellid Larvae of Japan and the cia Mem.

Fac. Educ., Fukui Univ. Ser. II, Nat. Sci. 18: 93-135.

. 1968b. Phylogeny of the family Coccinellidae (Coleoptera). Etizenia 35:

1.37,_13. pls.

. 1971. Fauna Japonica: Coccinellidae (Insecta: Coleoptera). Acad. Press

Jap., 340 pp., 16 pls.

Savoiskaya, G. I. 1960. Morphology and taxonomy of Coccinellid larvae from

Southeast Kazakhstan. Ent. Obozr. 39: 122-133.

. 1962a. Coccinellids of tribe Chilocorini (Coleoptera, Coccinellidae) of

southeastern Kazakhstan. Trudy Akademiia Nauk Kazakh. S. S. R., Alma-Ata,

Inst. Zool. 18: 189-200.

. 1962b. Morphology and Systematic of Coccinella L. (Coleoptera,

Coccinellidae). Trudy Nauch. Issle. Inst., S. S. R. Zashch. Rast., Alma-Ata 7:

299-315.

.1962c. Coccinellid genus Coccinulla Dbzh. in Southeast Kazakhstan.

Trudy Nauch. Issle. Inst., S. S. R. Zashch. Rast., Alma-Ata 7: 323-333.

. 1963. Larvae of ladybugs from the genera Semiadalia, Hippodamia,

Adonia, and Anisosticta. Trudy Akad. Nauk Kazakh, S. S. R., Alma-Ata. Inst.

Zool. 21: 74-95.

. 1964a. Material on morphology and systematic the Coccinellini larvae

(Coleoptera, Coccinellidae). Trudy Nauch. Issle., Inst., S$. S. R. Zashch. Rast.,

Alma-Ata 8: 310-357.

. 1964b. Some larvae of tribes Coelopterini and Hyperaspini with

description of new species, Pharoscymnus balkhashensis Savoiskaya sp. n.

(Coleoptera: Coccinellidae). Trudy Nauch. Issle. Inst. S. S. R. Zashch. Rast.,

Alma-Ata 8: 358-370.

Strouhal, H. 1926. Die Larven der palaearktischen Coccinellini und Psylloborini

(Coleoptera). Arch. Naturg. 92(a) 3: 1-63.

Van Emden, F. I. 1949. Larvae of British beetles. VII. (Coccinellidae). Entomol.

Month. Mag. 85: 265-283.

. 1956. The taxonomic significance of the characters of immature insects.

Ann. Rev. Entomol. 2: 91-106.

Watson, W. Y. 1956. A study of the phylogeny of the genera of the tribe Coccinellini.

Roy. Ontario Mus. Life Sci. Div., Contrib. 42: 1-52.

Pupae of Coccinellidae

Phuoc and Stehr:

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86. Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

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LATEROAPICAL fol

MARGIN APICAL ANGLE 4)

F ANGLE

Figs. 3-4. General areas on the pronotum of coccinellid pupae (dorsal aspect).

Figs. 5-6. General areas on the elytra of coccinellid pupae.

Fig. 7. Eriopis connexa, a close-up portion of the surface of the apical margin of the elytron.

Fig. 8. Coccinella novemnotata, a close-up portion of the surface of apical margin of the elytron.

Phuoc and Stehr: Pupae of Coccinellidae 87

PRONOTUM

CEYREO=

LABRUM

MANDIBLE

1/2mm

MAXILLARY PALPUS

DORSAL

MAXILLARY PALPUS 10 DEPRESSION

PRONOTUM

G4/ & [A

oe 1

ie E

ESS Fe . =

he o Eo CLUB ; me iictae SERPE

& eee 552,

= = 562

& A

CLUB MAXILLARY PALPUS

C /) 12 Lye

MAXILLARY

PALPUS

r\\t Mee PN % ‘ \\ SIN) ]

: ' oe ba see y ae : , Fe meaty nf

5 \ ea: P gE N an: ge eee Sgt

A an a Pine

Bim ~ 29S / ; a SE

LL Pup t VTS |

1/4mm

CLYPEO- i} \

LABRUM

PALPUS 14 15 16

Figs. 9-16. Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae.

Fig. 9. Epilachna borealis

Fig. 10. Epilachna sp.

Fig. 11. Lindorus lophantae

Fig. 12. Zagloba ornata

Fig. 13. Delphastus pusillus

Fig. 14. Microweisea ovalis

Fig. 15. Hyperaspis binotata

Fig. 16. Thalassa montezumae

88 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

PRONOTUM

VERTEX:

FRONTAL AREA —

FRONTAL SUTURE

ya dad 4

FRONTAL

\ — AREA

aA SS |

BASAL TUBERCLES

FRONTAL

AREA

1/10mm

CLYPEOLABRUM

ANTENNA

MAXILLARY PALPUS

21 22

Figs. 17-22: Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae.

Fig. 17. Stethorus punctum

Fig. 18. Stethorus punctum (lateral aspect)

Fig. 19. Stethorus picipes

Fig. 20. Stethorus picipes (lateral aspect)

Fig. 21. Stethorus atomus

Fig. 22. Stethorus atomus (lateral aspect)

PRONOTUM

1/2mm

VERTEX

Figs. 23-31:

Fig.

24.

20%

26.

28.

29;

30.

Ok:

Phuoc and Stehr: Pupae of Coccinellidae

DKS

CLYPEUS

LABRUM CLYPEUS

LABRUM

1/2mm

PRONOTUM

BASAL LOBE SS |

1m eee

ANTENNA <> j \

4 be tl yl

y . LAN

eas

30 MAXILLARY PALPUS

Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae.

Chilocorus bivulnerus

Axion platiatum

Axion plagiatum, a close-up portion of the surface of the pronotum.

Exochomus hoeget

Exochomus cubensis, clypeolabral portion.

Brumoides suturalis

Orcus chalybeus

Scymnus creperus, lateral aspect.

Cryptolaemus montrouzieri

PRONOTUM

1/2mm

90. Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

ANTERIOR MARGIN

1mm

EleY-PEO=

LABRUM

x /

39 TARSUS

MARGIN

CY REO:

LABRUM

5 CLIYPEO=

MAXILLARY = LABRUM

PALPUS

Figs. 32-37: Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae.

Fig. 32 Anatis ocellata

Fig. 33. Mulsantina picta

Fig. 34. Olla abdominalis

Fig. 35. Coccinella novemnotata

Fig. 36. Adalia bipunctata

Fig. 37. Adalia bipunctata, clypeolabrum

Phuoc and Stehr: Pupae of Coccinellidae 91

PRONOTUM ANTERIOR MARGIN

CLUB

CLYPEO-

LABRUM

APICAL

ANGLE MAXILLARY PALPUS

FRONT FEMUR

CLYPEOLABRUM

FRONT FEMUR

1/2mm

Figs. 38-43: Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae.

Fig. 38. Cycloneda munda

Fig. 39. Cycloneda munda (lateral aspect)

Fig. 40. Neoharmonia venusta

Fig. 41. Hippodamia convergens

Fig. 42. Naemia seriata

Fig. 43. Coleomegilla maculata

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Figs. 44-51:

Fig. 44.

Fig. 45.

Fig. 46.

Fig. 47.

Fig. 48.

Fig. 49.

Fig. SO.

Fig. Sl.

1/2mm

U ZB

reset

TF,

le Gs /

‘

aaa tacd)

(Roe) ¢7)

CAS

E AINE Nite op AS

? *

eS -

“ys

ras

oO geteay ~ aa

Pr prt UA Ase z

Frontal aspect of the prothoracic and cephalic portion of coccinellid pupae

Propylaea quatuordecimpunctata

Anisocalvia quatuordecimguttata

Psyllobora vigintimaculata

Hippodamia parenthesis

Hippodamia parenthesis, clypeolabrum

Eriopis connexa

Rodolia cardinalis

1/4mm

Phuoc and Stehr: Pupae of Coccinellidae

CLUB

FLAGELLUM

1/4mm

CLUB

ANTENNAL

54 PAPILLAE

SCAPE

Fig.

Fig,

Fig.

22,

53.

54.

55,

56.

ANTENNAL

PAPILLAE

CLYPEUS

1/4mm

1/10mm

Figs. 52-63: Frontal aspect of left antenna of coccinellid pupae.

Coccinella novemnotata

Mulsantina picta

Anatis ocellata

Synonycha grandis

Propylaea quatuordecimpunctata

Epilachna borealis

Fig.

Fig,

Fig.

38.

59.

60.

61.

62.

63.

Scymnus creperus

Cryptolaemus montrouzieri

Delphastus pusillus

Zagloba ornata

Hyperaspis binotata

Axion plagiatum

Figs. 64-78:

Fig.

Figs. 79-82:

Fig. 81.

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

64.

65.

66.

67.

68.

69.

70.

AQ.

TS:

74,

tes

76.

78.

Left mandible of coccinellid pupae.

Axion plagiatum, dorsal aspect.

Axion plagiatum, mandibular apex as viewed from the top.

Chilocorus bivulnerus, mandibular aspect.

Chilocorus bivulnerus, mandibular apex as viewed from the top.

Hyperaspis binotata, dorsal aspect.

Hyperaspis binotata, mandibular apex as viewed from the top.

Chilocorus bivulnerus, dorsal aspect.

Coccinella novemnotata mandibular apex as viewed from the top.

Epilachna

varivestis, dorsal aspect.

varivestis, mesal aspect.

varivestis, viewed from the top.

borealis, dorsal aspect.

sp., dorsal aspect.

sp., viewed from the top.

borealis, viewed from the top.

Left galea of coccinellid pupae as viewed from the top.

Fig. 79. Anatis ocellata

Fig. 80. Mulsantina picta

Cycloneda

munda

Fig. 82. Adalia bipunctata

Phuoc and Stehr: Pupae of Coccinellidae 95

1/4mm

1/10mm

—_—_—_—

MOLA

POSTERIOR

TYP ANTERIOR

mEP

SES

IMPRESSION 716 17 78

N L

1/10mm

POSTERIOR 80 81 §2

96 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

MAXILLARY

PALPUS

1/4mm

1/10mm

LACINEA

1/10mm

GALEA

LACINEA

MANDIBULAR

IMPRESSION

1/4mm

1/10mm

Figs. 83-92: Dorsal aspect of the maxilla of coccinellid pupae.

Fig. 83. LEpilachna varivestis —

Fig. 84. Zagloba ornata

Fig. 85. Cryptolaemus montrouzieri

Fig. 86. Hyperaspis binotata

Fig. 87. Delphastus pusillus

Chilocorus bivulnerus

Axion plagiatum

Anatis ocellata

Adalia bipunctata

Psyllobora vigintimaculata

Phuoc and Stehr: Pupae of Coccinellidae 97

PALPUS LIGULA

A =

ny HYPOPHARYNX

LABIAL

PALPUS

5 S

HY POPHARYNX

93 94 95 96 97

LABIAL PALPUS

LIGULA

HY POPHARYNX

E 101

tes;

LABIAL PALPUS

102

LABIAL

PALPUS

98 99 100

FEMUR

TARSUS

TIBIA

TARSUS

TERMINAL

SEGMENT

TERMINAL SEGMENT

103 104

‘Figs. 93-102: Labium of coccinellid pupae. Fig. 99. Epilachna varivestis, ventral aspect.

Fig. 93. Chilocorus bivulnerus, ventral aspect. Fig. 100. Epilachna varivestis, viewed from the top.

Fig. 94. Axion plagiatum, ventral aspect. Fig. 101. Delphastus pusillus, ventral aspect.

Fig. 95. Axion plagiatum, viewed from the apex. Fig. 102. Zagloba ornata, ventral aspect.

Fig. 96. Hyperaspis binotata, dorsal aspect. Figs. 103-104: Hind legs.

Fig. 97. Hyperaspis binotata, \ateral aspect. Fig. 103. Axion plagiatum

Fig. 98. Anatis ocellata, ventral aspect. Fig. 104. Brumoides suturalis

1/4mm

BASAL LOBE

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

111

BASAL TUBERCLES

112

Figs. 105-112: Dorsal aspect of the pronotum of coccinellid pupae.

Fig.

105.

106.

107.

108.

109.

110.

111.

12.

Axion plagiatum

Exochomus hoegei

Brumoides suturalis

Epilachna varivestis

Scymnus creperus

Stethorus atomus

Neoharmonia venusta

Coccinella trifasciata

Phuoc and Stehr: Pupae of Coccinellidae 99

117 118 119

120 121

Figs. 113-121: Dorsal aspect of the pronotum of coccinellid pupae.

Fig. 113. Coccinella transversoguttata

Fig. 114. Coccinella transversoguttata

Fig. 115. Coccinella monticola

Fig. 116. Coccinella monticola

Fig. 117. Adalia bipunctata

Fig. 118. Adalia bipunctata

Fig. 119. Adalia bipunctata

Fig. 120. Anatis ocellata

Fig. 121. Anatis quindecimpunctata

100 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

127 128

129 130

Figs. 122-130: Dorsal aspect of the pronotum of coccinellid pupae.

Fig. 122. Mulsantina picta

Fig. 123. Mulsantina hudsonica

Fig. 124. Coleomegilla maculata

Fig. 125. Coleomegilla maculata

Fig. 126. Coleomegilla maculata, \ateral aspect.

Fig. 127. Eriopis connexa

Fig. 128. Hippodamia tredecimpunctata, light form.

Fig. 129. Hippodamia tredecimpunctata, dark form.

Fig. 130. Hippodamia tredecimpunctata, dark form.

102

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Figs. 131-138: Dorsal aspect of the pronotum of coccinellid pupae.

Fig. 131.

Fig. 132.

Fig. 133.

Fig. 134.

Fig. 435:

Fig. 136:

Fag. toy,

Fig: 138.

Hippodamia parenthesis

Hippodamia parenthesis, cross section of lateral margin of the pronotum.

Hippodamia parenthesis

Hippodamia quinquesignata

Hippodamia convergens

Olla abdominalis

Figs. 139-140: Dorsal aspect of the metanotum of coccinellid pupae.

Fig. 139.

Fig. 140.

Mulsantina picta

Mulsantina hudsonica

1/2mm

1mm

Phuoc and Stehr:

131

134

138

Pupae of Coccinellidae

140

103

139

1/2mm

104 Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

SCUTELLAR SPOT

1mm

SCUTELLAR SPOT

141

ey ae CUTELLAR SPOT

MEDIAN SPOT

145 14

MEDIAN SPOT

142 143 144

147 148

149 150 151 152

Figs. 141-152: Dorsal aspect of left elytron of coccinellid pupae.

Fig. 141. Coccinella trifasciata Fig. 147. Coccinella transversoguttata

Fig. 142. Coccinella trifasciata Fig. 148. Coccinella transversoguttata, dark form.

Fig. 143. Coccinella septempunctata Fig. 149. Coccinella novemnotata

Fig. 144. Coccinella septempunctata, dark form. Fig. 150. Coccinella novemnotata

Fig. 145. Coccinella transversoguttata Fig. 151. Coccinella novemnotata, dark form.

Fig. 146. Coccinella transversoguttata Fig. 152. Coccinella monticola

ay,

;

ies en gine

eh Ah in nae ie

PAT) eA - Aa

R a4 7 ihe j aA

: Nor a PADI nor ANNA IY hey Maa ea ecu at ; iG

AN Cte eon ute , so ears Guy 1 wy sbi :

my, hed rset Te. f We Mane MN

106

Contrib. Amer. Ent. Inst., Vol. 10, No. 6, 1974

Figs. 153-163: Dorsal aspect of left elytron of coccinellid pupae.

Fig.

133.

154.

£55,

156.

oo).

158.

io?

160.

161.

162.

163.

Anatis ocellata

Anatis quindecimpunctata

Coleomegilla maculata

Eriopis connexa

Hippodamia quinquesignata, viewed from the apex.

Hippodamia parenthesis, viewed from the apex.

Hippodamia tredecimpunctata, light form.

Hippodamia tredecimpunctata, cross section of the elytron across the lateral margin.

Scymnus creperus, cross section of the elytron across the lateral margin.

Hippodamia parenthesis

Phuoc and Stehr: Pupae of Coccinellidae 107

156

EPIPLEURA

LATERAL

ANGLE

EPIPLEURA

LATERAL ANGLE

159

LATERAL

MARGIN

’

LATERAL

MARGIN

ra 161 162

108 Ganitihe Mee Bow ist. vole Onn 0.1074

Figs. 164-170: Dorsal aspect of the elytron of coccinellid pupae.

Fig. 164. Hippodamia quinquesignata

Fig. 165. Hippodamia quinquesignata

Fig. 166. Hippodamia quinquesignata

Fig. 167. Hippodamia quinquesignata

Fig. 168. Hippodamia quinquesignata

Fig. 169. Hippodamia glacialis

Fig. 170. Hippodamia glacialis

Figs. 171-172: Third left abdominal pleuron of coccinellid pupae.

Fig. 171. Neoharmonia venusta

Fig. 172. Propylaea quatuordecimpunctata

Fig. 173. Epilachna varivestis, dorsal aspect of left elytron.

109

Pupae of Coccinellidae

Phuoc and Stehr

HUMERAL SPOT

SCUTELLAR SPOT

MEDIAN SPOT

yas ett: \

110 Contrib. Amer. Ent. Inst., Vol. 10, No. 6. 1974

ABDOMINAL PLEURON

Carta

+ Aye > ABDOMINAL

SPIRACLE

PLEURAL PROCESS ae =

Q Z

1/2mm

ABDOMINAL

PLEURON

ABDOMINAL

PLEURON

ABDOM INAL 176

SPTRACLE

ABDOMINAL

SPITRACLE

1/4mm

ABDOMINAL

SPTRACLE

ABDOMINAL

PLEURON ABDOMINAL PLEURON

ABDOMINAL :

SP IRACLE &

PLEURAL a

PROCESS

) PLEURAL

3 y) PROCESS

177 178

Figs. 174-178: Third left abdominal pleuron of coccinellid pupae.

Fig. 174. Coleomegilla maculata, third and fourth pleura.

Fig. 175. Eriopis connexa, third and fourth pleura.

Fig. 176. Synonycha grandis

Fig. 177. Anisocalvia quatuordecimguttata

Fig. 178. Anatis ocellata

Phuoc and Stehr: Pupae of Coccinellidae

111

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", Trt e eh

%, F ‘

ti . Me pa hata,

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var ered % So Ri une

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bg ShtG Peet they 4 NATac, 5 ag ee N GIG

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LATERAL Pore te 5

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MARGIN sere

Rs er

teal tated LATERAL

Ret ri MARGIN

Se ry

i SUTURAL

< Pbiae 3

one. '

of 2

syn t Satie AE

Sa ’ Cy ae tt

= =? ° ° wt

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MARGIN pa

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ney 4, ee Ci Pk Bea iy to LATERAL MARGIN

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WR eS oa le A ay 28 ett 8 BS eae

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182

LATERAL MARGIN

SPINULES

184

Figs. 179-183:

Apex of left hind wing.

Fig. 179. Adalia bipunctata, dorsal aspect.

Fig. 180. Adalia bipunctata, ventral aspect.

Fig. 181. Coccinella transversoguttata, ventral aspect.

Fig. 182. Coccinella monticola, ventral aspect.

Fig. 183. Coccinella trifasciata, ventral aspect.

Fig. 184. Aclose-up portion of spinulate area on hind wing apex.