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No.

Contributions.

of the

American Entomological Institute

Volume 15, 1977-1980

Townes, Henry. A revision of the Rhopalosomatidae

(Hymenoptera). 34 pages. August 4, 1977.

Townes, Henry. A revision of the Heloridae (Hymenoptera).

12 pages. August 4, 1977.

Chandler, Donald S. A revision of the Central and South

American Notoxus and a description of a new genus

Plesionotoxus (Coleoptera: Anthicidae). 83 pages. April 24,

1978.

Howden, Henry F. and Antonio Martinez. A review of the

New World genus Athyreus MacLeary (Scarabaeidae,

Geotrupinae, Athyreini). 70 pages. June 5, 1978. |

Helava, Jussi. A revision of the Nearctic species of the

genus Onthophilus Leach (Coleoptera: Histeridae). 43 pages.

June 5, 1978.

Huang, Yiau-Min. Medical entomology studies - XI. The

subgenus Stegomyia of Aedes in the Oriental Region with keys

to the species (Diptera: Culicidae). 79 pages. October 8, 1979.

Faran, Michael E. Mosquito studies (Diptera; Culicidae)

XXXIV. A revision of the Albimanus Section of the subgenus

Nyssorhynchus of Anopheles. 215 pages. March 10, 1980.

Contributions ,

of the

American Entomological Institute

Volume 15, Number 1, 1977

ms, pee

A REVISION OF THE RHOPALOSOMATIDAE

(HY MENOPTERA)

Henry Townes

American Entomological Institute

5950 Warren Road, Ann Arbor, Michigan

SMTASONA

i j i

JUN 13 1980)

TABLE OF CONTENTS

APNG PISS CHOI cae ne Eg ie rea ae Ube ges ied ta 7 Bast ali ake 11000 uate oie“? eva ania te 1

Key to the genera of Rhopalosomatidae. ....... 5... - eee eee 3

$Me WOR Geet RS oa es ee eg eR iA ae hg gl et ca at we ae Be 3

Key to the species Of OLR 4 5 ie a ee ee ek ee ee ee 4

Be ta Se a se ee ee ee We Ge Woh ee a ae ee Ow we cater 5)

2a MOURON TOA IMS ccs Gea alee Be GTR el ane eg BP a er ORC ete eee oe D

as OO Gites ae RR a Se ay Wawra, See es we A 6

@ Ommon TestaCeuat: oo eae Fe Se ce ee ee sR ew 8 7

Oe Mote Bite altene ee a Se ee ee a 7

We OAR BAY PINOS A EIOLINC a ok ae ie, Efe Ss ine ee ee, Se Se 9

Dr "Oricon Centar a ee Fe a Se ret 9

Be OS ORS eee a ee a es alte wl ume ees Ie gs ale el er ae 10

Iney to tne Species 0: Liospiek.. 665. a a ee a 10

1. Liosphex trichopleurum.......... Hane ge age nla ait hg ee Cee 11

Pa ae SEO VAI ee ae ae PN Ge oe eae ee Bere i

a) RES SO SOO oy ae Ske a ww es Re ae a Ewe ie 12

Key to the species 01 Paniscomima: . 2s eee ee ee ee IZ

BS PASCO te se ea eh ee ee a as Be as 13

eo SP ateC Ol ma eS a a en ee ea a ek ee ee 14

Oe oP AesCOM a PhiItICe FIA Ses ee eo ge ee a Se 14

4, <\Paniscomima seyricr Sherer. EMF. seas a 14

ie eee EEE A sg cng we atk we wee Ow OE Seb a eee oe 15

O. VP amiscoimiiiia FilOaneennats. . VS hon ow al ee a ee eS 15

Moe API SCOT Ie CU ee ae el Sn ee ee Sa eS eee 16

4. Genus Rhopalosoma <5 i... i ae OS ie dak SP Sag aI Th ae gor a ee ake 16

Key to the species of Rhopalosoma. .......-+s+eese eee eee 17

CU a ee Se Be Re ee a ee ee Sue ae ee hte ig 5G

i eee roe eoiia Pattie Sete ow ae Oo atta ecw ti et ae oe ee ee ON 19

D5 FRO SOTIA TSODHS oo nak. ei a ea ee em wee ae 19

ee ae a er eee ae 6 4% ee a ay a eke 19

oe wR MODArOGOMIa NIISSIONICUD. 3 sk Gow ce ee ce lee as SS 20

Oy. VaR Ea TAC a ay GS a a tea 5 RET Sa ees 20

Dn RO ALO SONie DOTIVINUEY Ge ee ee we ee 20

B.S OSOIne Se ee ee 2 a ee SS 21

Te CRE ose MEA PCTICUET is ee a we OR ee 21

Be Ns a OTe eis He weed eee a nek ede Means abe ee ees 22

9. Rhopalosoma haitiense. ............. hoes al eet toa ea 20

TO. FRODRIOSOING Oe ies Lr OE EEG 8 ee ee ee a ew ee 23

Th. ANG RaOsOHIn Vani SS NOE ae eee Se em 24

12 PNGB oer Cr ee ye a ee ee ea ele We OR SG 24.

13. “RRO palosoiia SCapostn 665. Be are a DRS 20

14. Rhopalospima suiancweeiiy ye tered cose ew we ee 29

To. ‘hoptioeeina Drecland or. aS ES a ew ee ee ee Oe 26

16. Rhopalosoma alvarengai.......... PrN per aout Rees 26

Panceqlatum: Orowpetes errisg oe ee ey ath BR wa a 27

i1.., Rhepaloseniadanceoistum ci sero. eae ee TE OR. Cae 27

ere eee a ee eS eg Sg Sv ua Ye ok ae a ea CR aI a Cae geet Ug dg, ioe ade 27

Re lt ss ee a ag han eee ee gt Rect mapa rian gt anh a a ACL tS Eo Ae 28

A REVISION OF THE RHOPALOSOMATIDAE

(HY MENOPTERA)

Henry Townes

American Entomological Institute

9950 Warren Road, Ann Arbor, Michigan

The Rhopalosomatidae is a small family of wasps that are parasites of

Gryllidae. Specimens have been scarce in collections because most species

are nocturnal or crepuscular and are not collected on flowers as are most

other wasps. When use of Malaise traps recently became popular these wasps

turned up in the traps in some numbers. Now, there are enough specimens

available for a substantial advance in the systematics of the family. Speci-

mens of the flightless genus Olixon, however, are still in short supply.

Some characteristics of the Rhopalosomatidae, in addition to those com-

mon to other Aculeata, are as follows: Notaulus and prepectal carina absent.

Mesopleurum without a median transverse groove. Mesosternum with a pair

of plates on its hind edge that partially cover the bases of the middle coxae.

Femora without bristles, the tibiae without bristles or with only small weak

bristles. Female tarsi weakly to strongly widened, with dense suberect hairs

beneath. Male tarsal claws with a tooth near apical point. Female tarsal

claws simple or with a postmedian tooth, sometimes also with a subbasal low

rounded tooth. Abdomen flexible between segments 1 and 2. Male clasper

ending in a long upcurved spine. Female subgenital plate roundly folded on

midline, triangular in side view, with short dense hairs. Female pygidium

with a vertical bare area on each side of which the hairs are a little longer

and sparser. Sting upcurved.

There is a wingless or brachypterus genus, Olixon, and three fully winged

genera. In the winged genera the inner margin of the eye is notched, the

flagellar segments (at least the basal segments) have a pair of conspicuous

bristles at the apex of each, and the female tarsi are strongly expanded. In

the flightless genus Olixon the eye margin is not notched, the bristles on the

flagellum are small or absent, and the female tarsi are only weakly expanded.

There has been speculation about the systematic position of the Rhopaloso-

matidae. It is certain that the family belongs in the Aculeata, but I am not

prepared to say to which other aculeate families it is most closely related.

Ashmead in 1895 gave a summary of the opinions expressed up to that date

(Proc. Ent. Soc. Washington 3: 303-309). The interested reader should refer

to this publication. Brothers (1975. Univ. Kansas Sci. Bul. 50: 483-648) re-

lates it to the Psammocharidae.

Some details are known about the biology of one species, the Nearctic

Rhopalosoma nearcticum. It parasitizes crickets of the genera Hapithus and

Orocharis. These crickets are frequently found with a blackish larva adhering

to one side, above the base of a hindfemur. If captured and confined, sucha

parasitized cricket will soon succumb and the full-grown larva will burrow into

the soil, spin a cocoon, and hibernate until it is ready to emerge as an adult

the next summer. Gurney (1953. Proc. U. S. Natl. Mus. 103: 19-34) gives

2 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

biological details about this species. One other species has been reared, the

wingless Olixon australiae, as a parasite of a trigonidiine cricket. It is prob-

able that the entire family has an essentially similar biology.

The family contains four genera: Olixon (wingless or brachypterous) with

species in North America, South America, Africa, and Australia; Liosphex

with one species ranging from Florida to Brazil and another species in Min-

danao; Paniscomima with species in Africa, Madagascar, Sri Lanka, India,

and Nepal; and Rhopalosoma with one species in eastern United States and six-

teen species Neotropical. :

Specimens used in this study were from the following collections. After

the name of each collection is the name of the person who aided this work by

sending the specimens on loan.

Cambridge (Museum of Comparative Zoology, Harvard University).

Prof. E. O. Wilson.

East Lansing (Department of Entomology, Michigan State University).

Prof. Roland Fischer.

Ithaca (Department of Entomology, Cornell University).

Dr. L. L. Pechuman.

La Plata (Facultad de Ciencias Naturales y Museo, Universidad Nacional

de La Plata). Prof. Luis De Santis.

London (British Museum (Natural History)). Mr. Michael Day.

Ottawa (Canadian National Collection). Dr. Lubomir Masner.

Paris (Museum National d'Histoire Naturelle). Mlle. S. Kelner-Pillault.

Philadelphia (Academy of Natural Sciences of Philadelphia).

Dr. David C. Rentz.

Strasbourg (Musée Zoologique de l'Université et de la Ville).

Dr. F. Gouin.

Townes (Collection of Henry and Marjorie Townes).

Washington (National Museum of Natural History).

Dr. Arnold Menke.

All holotypes or lectotypes have been seen except for those of Arnold,

Enderlein, and Berland. Paratypes of all of Berland's species have been

studied. The Arnold and Enderlein species have been identified from their

original descriptions, which in these cases seem to be adequate.

I wish to thank the curators who sent specimens on loan, and to thank

several collectors who sent specimens for the Townes collection: Dr. and

Mrs. Henry Howden, Dr. Sam Breeland, Dr. Lionel Stange, Mr. Luis Pena,

and especially Col. Moacir Alvargena. My wife contributed supportive work

during the study and in the years leading up to it.

Townes: A revision of the Rhopalosomatidae 3

Key to the genera of Rhopalosomatidae

1. Eye margin not notched opposite antennal socket. Wings absent or shorter

than thorax. First tergite 0.33 to 0.6 as long as wide. Male subgenital

plate triangular, its apical 0.5+ margined with short bristles.

1. Olixon (p. 3)

Eye margin notched opposite antennal socket. Wings much longer than

thorax. First tergite 1.3 to 6.8 as long as wide. Male subgenital

plate triangular with apex narrowly truncate, the truncate portion

margined: with.short bristles... \.:d2.hs. susie soe eoma pe ues ces 2

2. Occipital carina lacking. First tergite about 1.5 as long as wide. Anal

lobe reaching 0.35 to 0.6 the distance to nervellus.

2. Liosphex (p. 10)

Occipital carina present on upper 0.6 of head. First tergite 2.4 to 6.8

as long as wide. Anal lobe reaching 0.65 to 1.0 the distance to

Nervellusicis; 662 wale wks wise eee iene ean nal Pe tet 3

3. Postnervulus (= the oblique apical sections of discoideus) arched, the

arching strongest at or below middle and flattened or indented at origin

of subdiscoideus. Female tarsal claws without a tooth. Penis valve with

with its preapical process broadly triangular and overlapped at its apex

by basal part of penis head. Ethiopian and Oriental species.

3. Paniscomima (p. 12)

Postnervulus sinuate. Female tarsal claws with a strong tooth near

middle, often also with a blunt tooth near base. Penis valve with its

preapical process narrow to broad, distant from penis head. American

SWORE Sehr Biavudii and ta Seah Lee P ier Guede oenueies 4. Rhopalosoma (p. 16)

1. Genus Olixon

Olixon Cameron, 1887. Biologia Centrali-Americana. Insecta. Hymenoptera.

1: 412.

Type: Olixon testaceum Cameron. Monobasic.

Harpagocryptus Perkins, 1908. Proc. Hawaiian Ent. Soc. 2: 34.

Type: Harpagocryptus australiae Perkins. Monobasic.

Algoa Brues, 1910. Jour. New York Ent. Soc. 18: 18. Name preoccupied

by Gray, 1840 and by Castelnau, 1861. New synonym.

Type: (Algoa heterodoxa Brues) = dentatum Cameron. Monobasic.

Saphobethyllus Kieffer, 1911. Ann. Soc. Sci. Bruxelles (2) 35: 216.

Type: (Saphobethyllus pallidus Kieffer) = testaceum Cameron. Monobasic.

Algoella Kieffer, 1914. Das Tierreich 41: 473. New name for Algoa. |

Nealgoa Brues, 1922. Psyche 29: 105.

Type: Nealgoa banksii Brues. Original designation.

Psyllosphex Arnold, 1935. Ann. Transvaal Mus. 15: 479.

Type: Psyllosphex saltator Arnold. Original designation.

Body length 2.9 to 7.0 mm. Wings vestigial or absent, not extending

beyond apex of tergite 1. Thorax modified as in other apterous Hymenoptera.

Eye elliptic, its inner margin not notched. Ocelli very small or lacking.

Temple very weakly convex, long or moderately long, the head not or very

little constricted to occipital carina, the occipital carina incomplete below.

4 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Palpi of moderate length. Flagellum short to long, its basal segments some-

times with a short weak bristle at apex on upper side. Hind edge of mesoster-

num with a very large bilobed flange that is neither impressed nor separated

by a groove. Propodeum with a large apicolateral tooth on each side. Front

femur swollen, especially infemale. Female tarsi weakly to moderately

widened and flattened, with dense tenent hairs beneath, the fourth segment

more or less bilobed. Male tarsal claws with a large preapical tooth; female

tarsal claws stout and simple. First tergite 0.33 to 0.6 as long as wide, usu-

ally with a transverse carina near its basal 0.4. Male subgenital plate large,

triangular, its apical 0.5+ margined with small, short bristles.

Eight species are known, one in the United States, one in Central and South

America, four in Africa, and two in Australia. The United States species

(banksii) and one of those from Africa (saltator) are reported as running or

jumping on sand. The United States species has been found active at night as

well as by day. The Neotropic species has been taken in Malaise traps. The

species australiae was reared from a trigonidiine cricket. There is circum-

stantial evidence that O. banksii parasitizes crickets of the genus Nemobius

and that O. testaceum parasitizes crickets of the genus Cycloptilum. This is

the extent of the available biological data.

Key to the species of Olixon

1. Cheek without a groove from the eye to the mandible. ......... 2

Cheek with a groove from the eye to the mandible. ........... 4

2. First flagellar segment 1.8 as long as wide. Upper hind edge of eye

separated from occipital carina by 1.1 the width of the scape. Eastern

Amsifatias ie oe ee es 1. flavibase, new species (p. 5)

First flagellar segment 3.0 to 4.2 as long as wide. Upper hind edge of

eye separated from occipital carina by 0.25 to 1.0 the width of the

3. Body fuscous. Hind femur of female about 2.5 as long as high (male

unknown). Ivory Coast. . . 2. majus, new species (p. 5)

Body fulvous. Hind femur of female about 4.4 as long as high, of male

about 5.5 as long as high. Eastern United States; also Mexico.

3. banksii Brues (p. 6)

4. Upper hind edge of eye separated from occipital carina by about 0.3 the

width of the scape. Head and body mostly or entirely fulvous.

- Neotropic Region. 4. testaceum Cameron (p. 7)

Upper hind edge of eye separated from occipital carina by about 1.1 to 2.6

the width of the scape. Head and body dark brown or blackish, or some-

times the thorax fulvoterruc mods. 47 PAs Mey PO ay a 5)

5. First tergite without a subbasal transverse carina. Propodeum without

a transverse carina between its apicolateral teeth. Eastern Australia.

5. australiae Perkins (p. 7)

First tergite with a subbasal transverse carina. Propodeum with a com-

plete or incomplete transverse carina between its apicolateral teeth.

Townes: A revision of the Rhopalosomatidae 5)

6. Front wing reaching to hind end of thorax. Hairs of head and abdomen very

dense. First flagellar segment about 2.6 as long as wide. South Africa.

6. saltator Arnold (p. 8)

Front wing minute or absent, not longer than tegula. Hairs of head and

abdomen moderately dense (but unusually sparse for the genus Olixon).

First flagellar segment 1.4 to 2.2 as long as wide. ......... 7

7. Thorax fulvoferruginous. Apex of scutellum a 90° rounded angle. Upper

hind edge of eye separated from occipital carina by 1.5 the width of

scape. Front wing present as a small vestige. Rhodesia.

7. myrmosaeforme Arnold (p. 9)

Thorax reddish brown to blackish. Apex of scutellum rounded or sub-

truncate. Upper hind edge of eye separated from occipital carina by

about 2.0 the width of the scape. Front wing absent. Southeast coast

of South Aries.) i iicirarse tL here wet 8. dentatum Cameron (p. 9)

1. Olixon flavibase, new species

Male: Unknown.

Female type: Body 3.8 mm. long. Hind margin of eye separated from

occipital carina by 1.3 the width of scape. Front ocellus lacking, the hind

ocelli present as small vestiges. Vertex with fine, moderately dense hairs

that arise from very small punctures. Cheek without a groove from eye to

mandible. First segment of flagellum 1.6 as long as wide. Propodeum with

a projecting flange on the median 0.4 of the distance between its apicolateral

teeth, rounded on each side between this flange and the apicolateral teeth.

Hind femur 4.4 as long as deep. Fourth tarsal segments short, with long

apicolateral lobes, one lobe about 2.2 as long as the other. Front wing paddle

shaped, reaching to end of first tergite. Hind wing obscured (or absent ?).

Premedian transverse carina of tergite 1 strong, evenly arched. Upper face

of tergite 1 evenly convex.

Fuscous. Scape, pedicel, coxae, trochanters, base of hind femur, and

tibial spurs stramineous or yellowish, the scape brownish medially and the

coxae brownish basally.

Type: %, Cairns [ Australia], Feb. 1902 (London).

2. Olixon majus, new species

Male: Unknown.

Female type: Length 7.7 mm. Eye separated from occipital carina by

0.25 the width of scape. Ocelli lacking. Vertex with fine, short, dense hairs

that arise from dense, very small punctures. Cheek without a groove from

eye to mandible. First flagellar segment 4.2 as long as wide. Propodeum

between its apicolateral teeth with a transverse carina on only the median 0.3

of the distance and this median portion weak at center. Hind femur 2.5 as

long as deep. Front wings paddle-shaped, reaching base of tergite 2. Hind

wing a small, narrow vestige. Tergite 1 with a strong premedian transverse

carina, behind the transverse carina weakly flattened medially and with a

median longitudinal carina.

Fuscous. Mandible, antenna, and legs beyond trochanters brown. Palpi,

coxae, and trochanters pale brown. Front wings weakly infuscate.

6 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Type: 9, ''Lampto" [| biological station], Ivory Coast, Nov. 11, 1965,

Y. Guillon (Paris).

3. Olixon banksii Brues

*Nealgoa banksii pice. 1922. Psyche 29: 106. 2. Type: 2, U. S. A.: Sea

Cliff, Long Island, New York (Cambridge). Type studied in 1973.

Nealgoa banksid Reid, 1939. Proc. Roy. Ent. Soc. London 8: 101. des.

Olixon banksii anksii Krombein, 1950. Jour. Elisha Mitchell Sci. Soc. 65: 264-265.

Oo, N. C.: Kill Devil Hills. "running swiftly over sand barrens”.

Olixon banksii Krombein, 1951. U. S. Dept. Agr. Agr. Monog. 2: 774. N. Y.,

D. C., N. C., Fla., Ohio, Minn., Kans.

Length 2.9to 7.0 mm. Eye separated from occipital carina by about 0.3

to 1.0 the width of the scape. Vertex with fine, dense, short hairs that arise

from small, closely spaced punctures. Ocelli very small, the median ocellus

often lacking. Cheek without a groove from eye to mandible. First flagellar

segment about 3.5 as long as wide. Carina between apicolateral teeth of pro-

podeum present only on median 0.3 of the distance. Hind femur about 5.0 as

long as high in male, about 4.4 as long as high in female. Female tarsi

flattened beneath and widened, the fourth segment short and with apicolateral

angles long, the angle on one side about 1.5 as long as on the other. Front

wing paddle-shaped and reaching about to base of tergite 2, or sometimes

present as a small narrow strap that is about as long as tegula. Hind wing a

tiny linear vestige. Premedian transverse carina of tergite 1 strong, dipped

and irregular at midline. Tergite 1 behind its transverse carina with a weak

median longitudinal impression, often with a short median ridge in the impres-

sion which connects with the transverse carina.

Light fulvous. Flagellum brown apically. Front wing weakly infuscate.

Apical 0.4+ of middle and hind femur often brownish. Subapical 0. 4+ of hind

tibia often darkened in female. Subapical band on tergite 2 and apical tergites

sometimes infuscate, the band on tergite 2 narrowed medially and not reach-

ing sides of the tergite.

Specimens: 160, 12° from the United States in Arkansas (Mississippi Co. );

District of Columbia (Washington); Florida (Arcadia, Gainesville, Hillsboro

Co., and Welaka); Georgia (Mosquito Creek and Spring Creek, both in Decatur

Co. \: Kansas (Pottawatomie Co. and 7 miles west of Laweence). Louisiana

(Chicot State Park in Evangeline Co.); Maryland (Plummers Is. ): New York

(Farmingdale); North Carolina (Kill Devil Hills); and Ohio (Sandusky). One

additional specimen is from ‘'Mexico".

Dates of collection are from May 26 to June 15 for males and from June

28 to Sept. 10 for females; also a female collected March 30 to Apr. 1 at

Arcadia, Fla. One female is labeled as "'collected on sand". Dr. P. A.

es informs me (in litt.) that specimens collected by him in Decatur

Co., Ga. were found while collecting at night (8:30-10:30 P.M.). They were

running around swiftly in leaf litter. Krombein (1950) reported finding males

"running swiftly over sand barrens". In the U. S. National Museum is a nymph

of Nemobius sp. from Plummers Is., Md., that bears a small larval case of

a rhopalosomatid. This case, because of its small size and the small size of

its host, is presumed to be that of the present parasite.

This species occurs in the eastern United States in the Upper and Lower

Austral Zones, and there is a record from ''Mexico".

Townes: A revision of the Rhopalosomatidae 7

4. Olixon testaceum Cameron

*Olixon testaceum Cameron, 1887. Biologia Centrali-Americana. Insecta.

Hymenoptera. 1: 412. 92. des., fig. Type: ¢, Panama: Bugaba

(London). Type studied in 1974.

*Saphobethyllus pallidus Kieffer, 1911. Ann. Soc. Sci. Bruxelles (2) 35: 216.

. Type: &, Mexico: Teapa in Tabasco (London). Type studied in 1974.

Saphobethyllus pallidus Kieffer, 1914. Das Tierreich 41: 288. ©. des.

Olixon testaceum Turner & Waterston, 1917. Ann. Mag. Nat. Hist. (8) 20:

107. syn., des.

Olixon testaceum Reid, 1939. Proc. Roy. Ent. Soc. London 8: 95-97. ©, @.

des., fig.

Olixon pallidum Evans, 1964. Bul. Mus. Comp. Zool. 132: 16. syn.

Body 3.7 to 5.0 mm. long. Hind margin of eye separated from occipital

carina by about 0.3 the width of the scape. Ocelli lacking. Vertex with fine

dense hairs that arise from very dense small punctures. Cheek with a groove

from eye to front condyle of mandible. First segment of flagellum about 4.0

as long as wide. Propodeum with a complete transverse carina between its

teeth. Hind femur about 5.1 as long as deep. Fourth tarsal segment of female

short, with long apicolateral lobes, one lobe about twice as long as the other.

Front wing paddle-shaped, reaching to middle of first tergite. Hind wing a

tiny linear vestige. Premedian transverse carina of tergite 1 strong, evenly

arched. Upper face of tergite 1 evenly convex.

Pale fulvous, the mouth parts, coxae, and trochanters stramineous. First

four incisures between flagellar segments and a line along the vertical groove

in cheek infuscate. Front wing faintly infuscate. Second tergite with a pair

of transverse subapical fuscous areas that make a medially interrupted sub-

apical fuscous band. The type of pallidus, however, has the second tergite

entirely pale fulvous.

Specimens: ©, 2? (including the type of pallidus), Teapa, Tabasco,

Mexico, March, H. H. Smith (London). ©, Botanic Gardens, Georgetown,

British Guiana, Jan. 13, 1926, L. D. Cleare, Jr. (London). ?, Manaus,

Amazonas, Brazil, Jan. 1972, F. M. Oliveira (Ottawa). 39, Represa Rio

Grande, Guanabara, Brazil, Dec. and Feb. 1966 and 1967, M. Alvarenga

(Townes). (type of testaceum), Bugaba, Panama, Champion (London).

The four specimens from Brazil were apparently all collected in Malaise

traps. Inthe U. S. National Museum is an adult male cricket from Honduras,

determined as Cycloptilum trigonipalpum (probably by Caudell), which bears

a small rhopalosomatid larva. Because of the small size of the cricket and its

parasite, the larva is presumed to belong to the present species. ,

This species ranges from southern Mexico to southern Brazil.

5. Olixon australiae Perkins

*Harpagocryptus australiae Perkins, 1908. Proc. Hawaiian Ent. Soc. 2: 34.

@. des. Type: 2, Australia: Queensland (London). "Larva forming

a case on the side of abdomen of small crickets (Trigonidiidae)"". Type

studied in 1973.

Harpagocryptus australiae Kieffer, 1914. Das Tierreich 41: 289. translation

of original des.

8 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Harpagocryptus australiae Reid, 1939. Proc. Roy. Ent. Soc. London 8: 99-100.

notes from original des.

Olixon australiae Perkins, 1976. Handbooks for the identification of British

insects 6 (3a): 1. syn.

Male: Unknown.

Female: Length 3.4mm. Eye separated from occipital carina by 1.5

the width of the scape (this figure estimated as the specimen at hand lacks

antennae). Ocelli very small, the median ocellus vertical. Vertex weakly

mat, with short, fine, rather dense hairs that arise from very fine punctures.

Cheek with a groove from eye to front condyle of mandible. Antennae lacking

in the specimen at hand, described originally by Perkins as being ''much

longer than head and thorax together, the scape is stout and about as long as

the front funicle joint, which is decidedly shorter than the next, while those

following are all subequally elongate, many times as long as thick". Pro-

podeum rounded and not carinate between its apicolateral teeth. Hind femur

4.4 as long as deep. Tarsi moderately widened, flat beneath, their fourth

segments of moderate length and with long apicolateral angles, one angle about

2.2 as long as the other. Front wing present as tiny narrow vestige that is

not longer than tegula. Tergite 1 without a premedian transverse carina,

evenly convex above.

Blackish brown. Clypeus, mouth parts, coxae, and trochanters light

brown. Antennae lacking in specimen at hand. Legs beyond trochanters dark

brown.

Specimen: Described from a female in London that lacks a locality label,

determined by R. C. L. Perkins and believed to be the original type. The type

was described as from Queensland, Australia, and as reared from a small

trigonidiine cricket. ;

6. Olixon saltator Arnold

*Psyllosphex saltator Arnold, 1935. Ann. Transvaal Mus. 15: 480. &. des.,

fig., biol. Type: 2, South Africa: Beach Bush in Durban (Bulawayo).

Identification from description.

Length: 4.6 to 6.2 mm. Eye separated from occipital carina by 1.15 to

1.3 the width of the scape. Ocelli very small. Vertex with slender rather

dense hairs that arise from small dense punctures. Cheek with a groove from

eye to front condyle of mandible. First flagellar segment about 2.6 as long

as wide. Hind femur 4.8 as long as deep in male, 3.8 as long as deep in fe-

male. Female tarsi moderately widened, flattened beneath, the fourth seg-

ments with long apicolateral angles that are equal in length. Front wing

paddle-shaped, reaching to base of tergite 1. Hind wing a small narrow ves-

tige. Tergite 1 with a distinct premedian carina, behind the carina evenly

convex.

Black. Mouth parts, antenna, front legs, and middle and hind legs beyond

trochanters dark brown. Middle andhind coxae and trochanters pale brown.

Front wing moderately infuscate. Free margins of abdominal tergites and

sternites brown.

Specimens: “, ¢, Burman Bush, Durban, South Africa, H. N. Empey,

the o dated Dec. 3, 1960 and ? dated March 18, 1963 (London). ©“, Durban,

South Africa, Apr. 16, 1914, G. B. Cooper (London).

Townes: A revision of the Rhopalosomatidae 9

7. Olixon myrmosaeforme Arnold

*Psyllosphex myrmosaeformis Arnold, 1935. Ann. Transvaal Mus. 15: 482.

. des., fig. Type: %, Rhodesia: Bulawayo (Bulawayo). Identification

from description.

Male: Unknown.

Female: Length 4.0mm. Eye separated from occipital carina by 1.5 the

width of scape. Ocelli very small. Vertex polished, with fine, moderately

dense hairs that arise from very small punctures. Cheek with a groove from

eye to front condyle of mandible. First flagellar segment 2.2 as long as wide.

Apex of scutellum a rounded 90° angle. Propodeum with a strong transverse

carina between its apicolateral teeth. Hind femur 4.0 as long as deep. Tarsi

weakly widened and flattened below, the apicolateral angles of segments 4

moderately long, one angle about 1.2 as long as the other, the angles not tipped

with bristles. Front wing a vestige that is a little longer than tegula. Hind

wing absent. Tergite 1 with a premedian transverse carina, behind the carina

faintly flattened on midline.

Head and abdomen blackish brown, the temple and lower 0. 4+ of the rest

of head reddish brown. Mouth parts and segments 1-3 of antenna light brown,

the rest of antenna dark brown. Thorax and legs fulvoferruginous, the femora

tinged with brown toward apex.

Specimen: ¢, Rhodesia, Oct. 18, 1944, R. H. R. Stevenson (London).

8. Olixon dentatum Cameron

*Apteropompilus dentatus Cameron, 1904. Ztschr. System. Hymen. Dipt.

4:176. [o]. Type: o, South Africa: Dunbrody in Cape Colony

| (London). Type studied in 1973.

Apteropompilus dentatus Cameron, 1905. Ztschr. System. Hymen. Dipt.

9: 1386. &. des. South Africa: Dunbrody in Cape Colony.

*Algoa heterodoxa Brues, 1910. Jour. New York Ent. Soc. 18: 18. 2. des.,

fig. Lectotype: %, (labeled by Townes, 1973 and hereby designated),

South Africa: Algoa Bay, Dec. 25, 1896, H. Brauns (Cambridge).

Lectotype studied in 1973. New synonym.

Algoella heterodoxa Kieffer, 1914. Das Tierreich 41: 273. 2. des., fig.

Algoa heterodoxa Brues, 1922. Psyche 29: 101, 108. ©, ¢. des.

Psyllosphex dentatus Arnold, 1935. Ann. Transvaal Mus. 15: 482. ©. des.

Psyllosphex dentatus Reid, 1939. Proc. Roy. Ent. Soc. London 8: 95-99.

si Fe GCS., Lig. |

Algoa heterodoxa Reid, 1939. Proc. Roy. Ent. Soc. London 8: 97, 99-101.

#,. des. , fie:

Algoella dentata Brues, 1943. Ann. Ent. Soc. America 36: 311. syn., des.

Algoella heterodoxa Brues, 1943. Ann. Ent. Soc. America 36: 311. syn.

des.

Length 3.5to 5.2 mm. Eye separated from occipital carina by about 2.0

the width of scape. Ocelli very small, the median ocellus absent or vestigial.

Vertex polished, with numerous short hairs that are not dense, these arising

from very small punctures. Cheek with a groove from eye to front condyle of

mandible. First flagellar segment about 1.4 to 1.7 as long as wide. Apex of

10 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

scutellum rounded or subtruncate. Propodeum with a complete carina between

its apicolateral teeth, or sometimes the carina interrupted sublaterally. Hind

femur about 4.5 as long as deep in male, 3.5 to 4.2 as long as deep in female.

Tarsi of female only a little widened, their fourth segments moderately long,

with acute but not elongate apicolateral corners, the corners usually tipped

with a long bristle. Wings absent, the tegula very small. First tergite with

a premedian transverse carina, behind the carina without a median impres-

sion or irregularity.

Blackish brown, the antenna, mouth parts, and legs dark brown with the

basal 2.5 or 3.0 segments of antenna fulvous. In a specimen from Durban,

South Africa, the thorax, legs, and first tergite are brownish ferruginous

rather than blackish.

Specimens: 42, Algoa Bay, South Africa, Oct. 11, 1896, Dec. 25, 1896,

and undated, H. Brauns (3, including lectotype of heterodoxa in Cambridge;

12 in Washington). © (type of dentatum), Dunbrody (near Grahamstown),

South Africa, Sept. 20, 1900 (London). 2, Durban, South Africa, Apr. 1914,

G. B. Cooper (London).

This species occurs on the southeast coast of South Africa. It is variable

in structure and color. Possibly it should be subdivided into several species.

2. Liosphex, new genus

Front wing 5.0 to 9.4 mm. long. Body moderately slender. Eye large,

its inner margin with a large notch above antennal socket. Ocelli moderately

small. Temple of moderate width, convex. Occipital carina absent. Palpi

long. Flagellum of moderate length, its basal 7 or 8 segments with an apical

pair of bristles on upper side or upper and inner sides, one of the pair usually

longer than the other. Propodeum smoothly convex. Hind edge of mesoster-

num with a parallel pair of large triangular flanges that are separated from

mesosternum by a groove but are in almost the same plane as the mesoster-

num. Front femur of female weakly to moderately swollen. Segments 2-5

of female tarsi wide, flat beneath, and with dense tenent hairs. Male tarsal

claws with a pair of parallel points at apex and a large angulate lobe a Little

behind apex. Female tarsal claws stout, simple. First tergite about 1.5 as

long as wide. Anal lobe reaching 0.35 to 0.6 the distance to nervellus.

Genotype: Liosphex varius, new species.

The generic name is from Actes (smooth), pluss@n€ (wasp), referring to

the smooth body contures and lack of sculpture.

Two species are known, one ranging from the Gulf shores of southern

United States to southern Brazil and the other known from the Philippines.

Key to the species of Liosphex

1. Nervulus distad of basal vein by about 0.5 its length. Angle between first

recurrent vein and cubitus about 50°. Metapleurum with short fine

hairs all over, the hairs a little sparser medially. Philippines.

1. trichopleurum, new species (p. 11)

Nervulus distad of basal vein by about 1.75 its length. Angle between

first recurrent vein and cubitus about 259. Metapleurum with short

fine hairs on its anteroventral 0.2, the rest bare.

Southern United States to Brazil. ... 2. varius, new species (p. 11)

Townes: A revision of the Rhopalosomatidae 11

1. Liosphex trichopleurum, new species

Male: Unknown.

Type female: Front wing 8.3 mm. long. Side of thorax (including side of

pronotum and of propodeum) covered with short moderately dense hairs, their

distribution even except for a slight thinning on collar of pronotum, near meso-

pleural fovea, and near center of metapleurum. Bristles on hind tibia of mod-

erate length. Longest spur of hind tibia reaching a little past apex of hind basi-

tarsus. Anal lobe reaching about 0.6 the distance to nervellus. Angle between

first recurrent vein and cubitus about 50°. Nervulus distad of basal vein by

about 0.5 its length.

Head stramineous, black on its hind side, vertex, and frons except later-

ally. Antenna blackish above, fulvous below. Mouth parts stramineous ex-

cept that segments 2-5 of maxillary palpus are mostly fuscous. Thorax black,

the tegula and upper half of pronotum stramineous. Front leg stramineous,

its femur with a fuscous mark on base behind and tarsus dark brown above.

Middle leg stramineous, the basal margin of coxa, basal 0.8 of trochanter,

basal 0.15+ of femur, and the upper side of tarsus fuscous. Hind leg black,

the apical 0.4 of coxa, apical 0.2 of trochanter, apical margin of femur, and

under side of tarsus stramineous. Wings faintly brownish. Tergites 1, 4, 5,

and 6 black with brownish apical and lateral stains. Tergites 2 and 3 stramin-

eous, their bases fuscous.

Type: °, Kolambugan, Mindanao, Philippines, C. F. Baker (Washington).

2. Liosphex varius, new species

Front wing 5.0 to 9.4 mm. long. Sides of thorax (including side of pro-

notum and of propodeum) with short moderately dense hairs, the hairs lack-

ing in the following areas: collar except its upper part, mesepisternum ex-

cept for marginal areas, metapleurum except its anteroventral 0.2 and its

upper front area, and central 0.5+ of lateral aspect of propodeum. Bristles

on hind tibia rather long and conspicuous. Longest spur of hind tibia reaching

0.85 the length of hind basitarsus. Anal lobe reaching about 0.5 the distance

to nervellus. Angle between first recurrent vein and cubitus about 25°. Ner-

vulus distad of basal vein by about 1.75 its length.

Body and legs varying from mostly pale fulvous to almost entirely black,

the wings tinged with brown. The lightest and darkest specimens of the series

are colored as follows:

Lightest coloration: Pale fulvous. Head stramineous but vertex black be-

tween the eyes and frons. Face and clypeus pale yellow. Scape and pedicel

each with a fuscous spot above. Median spot on front end and lateral 0.25: of

mesoscutum, spot at base of each wing, and sublateral stripe on propodeum

fuscous. Median part of first tergite light brown. Wings yellowish brown.

Darkest coloration: Black. Area in eye notch, clypeus except for median

apical area, basal 0.65 of mandible, apical part of palpi, hind corner and upper

margin of pronotum, and tibial spurs whitish. Wings weakly infuscate.

The upper margin of the pronotum and the space within the eye notch is

always white, usually also the clypeus, cheek, and mouth parts. The antenna

is usually fulvous below and fuscous above. The tibiae and tarsi are usually

largely or entirely fulvous. Basal 0.3+ of tergites 1-3 and apical 0.2+ of ter-

gite 1 usually fulvous, the abdomen often more extensively fulvous.

12 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Type: 9°, from Malaise trap, Tall Timbers (near Tallahassee), Fla.,

U. S. A., June 29 to July 11, 1971, R. H. Arnett (Townes).

Paratypes: 2, from Malaise trap, Chicot State Park, Evangeline Co., La.,

U. S. A., June 28 to July 5, 1971 (Ottawa). ©, from Malaise trap, Fortin de las

Flores, 3, 400 ft., Mexico, "Aug. 20, 1963, R. "H. and E. M. Painter (Townes).

Oo, Mt. San Salvador, El Salvador, june 24, 1963, M. E. Irwin (Cambridge).

2, Ongota, southeast of Tena, ‘ena, Ecuador, May 1963, L. Petia (Cambridge). ©,

Ecuador or Peru, aes L. Pefia (Townes). 3c, Avispas, 400 m., Peru,

Sept. 10 to 20, 1962, L. Pena (Cambridge). 39, Nova Teutonia, Santa nta Catarina,

Brazil, January 1939, een 10, 1952, and Dec. 4, 1952, Fritz Plaumann

(Cambridge, Ithaca, and Ottawa). °, Repressa Rio Grande, Guanabara, Brazil,

Feb. 1968, M. Alvarenga (Townes). ?, Rio de Janeiro, Brazil, Jan. 1939,

RG Shannon (Washington).

This species ranges from the Gulf shores of the United States to southern

Brazil.

3. Genus Paniscomima

Paniscomima Enderlein, 1904. Zool. Anz. 27: 465.

Type: Paniscomima erlangeriana Enderlein. Monobasic.

Hymenochimaera Brues, 1926. Psyche 33: 19. New synonym.

Type: (Rhopalosoma abnorme ''Cameron") = abnorme Morley.

Rogezia Berland, 1951. Mém. Inst. Sci. Madagascar (A) 15: 300. New

synonym.

Type: Rogezia seyrigi Berland. Monobasic.

Front wing 6.7 to 15 mm. long. Apical 0. 65+ of fifth segment of female

tarsi parallel-sided. Female tarsal claws stout, with a strongly curved apical

tooth, without a preapical tooth. Postnervulus arched and strongly oblique,

most strongly arched at or below middle and flattened or indented at origin of

subdiscoideus. First tergite 2.4 to 3.7 as long as wide. Preapical process of

male penis broadly triangular, its apex narrowly overlapped by base of penis

head.

Species of Paniscomima are Ethiopian and Oriental. They are separable

from Rhopalosoma by the characters given in the key but otherwise are very

Similar to that genus. There are no rearing records.

Key to the species of Paniscomima

1. Nervulus basad or opposite basal vein, or distad by less than 0.25 its

lenothi Oriental epecies 4) ae i PR. SSE TO ee AS 2

Nervulus distad of basal vein by more than 0.4 its length. Ethiopian

Sper ies te Aieg Seal a hes, oe TSI RES Pe Pe GR ok 3

2. Nervulus basad of basal vein by about 0.9 its length. Anal cell of front

wing with a longitudinal spurious vein near its center. India and

Sarria. OU ge Oe ae ee 1. abnormis Morley (p. 13)

Nervulus opposite basal vein or distad by as much as 0.2 its length.

Anal cell of front wing without a spurious vein. Nepal.

2. opposita, new species (p. 14)

Townes: A revision of the Rhopalosomatidae 13

3. Nervulus distad of basal vein by about 0.6 its length, straight and strongly

reclivous. Postnervulus flattened at origin of subdiscoideus. Africa.

3. erlangeriana Enderlein (p. 14)

Nervulus distad of basal vein by 0.8 to 2.0 its length, weakly or moderate-

ly reclivous. Postnervulus indented at origin of subdiscoideus.

Madaeascan Species 5 2iiy Oeste ay eee Oe ee 4.

4. Ocelli small, the lateral ocellus separated from eye by about 2.0 its long

diameter. Wings tinged with yellow, the apex of front wing weakly in-

fuscate. Median vein fulvoferruginous. Madagascar.

4. seyrigi Berland (p. 14)

Ocelli medium sized to large, the lateral ocellus separated from eye by

less than 1.5 its diameter. Wings uniformly subhyaline. Median vein

daik Drow. 6. 0 4 Ue ROPE ee ee oe SR eee 9)

0. Flagellum fuscous. Nervulus distad of basal vein by about 1.6 its length.

Madaeastar. 6 Soe Sse a ae 5. bekilyi Berland (p. 15)

Flagellum brownish fulvous. Nervulus distad of basal vein by 1.0 to 1.45

ibs TERS fps Bg se Re es A ee EOS ch “Gee ea 6

6. Nervulus distad of basal vein by about 1.45 its length. First intercubitus

about 0.7 as long as second section of cubitus. Tergites 4-6 fulvous.

Madacasearty? 642. Gigi ee 5 ae. 6. rufoantennata Berland (p. 15)

Nervulus distad of basal vein by about 1.0 its length. First intercubitus

about 1.0 as long as second section of cubitus. Tergites 4-6 infuscate.

NR OA IBA a OE i a ee ns 7. curta, new species (p. 16)

1. Paniscomima abnormis Morley, new combination

*Rhopalosoma abnorme Morley, 1910. Trans. Ent. Soc. London 1910: 386.

, ¢- des., fig. Type: °, Sri Lanka: Mankulam (London). India:

Calcutta. Type studied in 1974.

Hymenochimaera abnormis Brues, 1926. Psyche 33:19. syn., des.

Male: Not available for study.

Female: Front wing 6.7 mm.long. Lateral ocellus separated from eye

by 1.0 its long diameter. First intercubitus 1.2 as long as second section of

cubitus. Nervulus basad of basal vein by 0.9 its length, vertical. Anal cell

of front wing with a longitudinal spurious vein near its middle. Basal section

of radiella moderately reclivous. First tergite 2.9 as long as wide. |

Light fulvous. Interocellar area infuscate. Flagellum reddish brown.

Wings hyaline, their veins medium brown to dark brown.

Specimen: Redescribed from type, a female labeled Mankulam, N. P.

[ Sri Lanka], Nov. 1908, Wickwar (London). Morley had a second specimen,

a male from Calcutta, India. This I have not seen. Dr. Karl Krombein

informs me that he has recently collected a male and female in Sri Lanka. His

specimens are in Washington. in i

14 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

2. Paniscomima opposita, new species

Male: Front wing 7.8 to 10.3 mm. long. Lateral ocellus separated from

eye by about 0.2 of its long diameter. First intercubitus about 2.2 as long as

second section of cubitus. Nervulus opposite basal vein or distad of it by less

than 0.25 its length, weakly inclivous. Anal cell without a spurious vein.

Basal section of radiella moderately inclivous. First tergite about 2.7 as long

as wide.

Pale fulvous. Interocellar area, flagellum, tergite 7, and sternite 7 dark

brown. Wings hyaline, the wing veins brown.

Female: Unknown. |

Type: %, in Malaise trap, near Simra Adhabar, 600 ft., Nepal, Aug. 23 to

28, 1967, Canadian Nepal Expedition (Ottawa).

Paratypes: 4°, same data as type (Ottawa).

3. Paniscomima erlangeriana Enderlein

*Paniscomima Erlangeriana Enderlein, 1904. Zool. Anz. 27: 466. ¢&. des.,

fig. Type: 2, Somali: Umfudu (Berlin). Identification from description.

Rhopalosoma erlangerianum Schulz, 1906. Spolia hymenopterologica, p. 223-

225. key, syn., des.

Paniscomima erlangeriana Brues, 1926. Psyche 33:18. des. South Africa:

Windenham in Natal.

Rhopalosoma erlangerianum Krombein, 1951. U. S. Dept. Agr. Agr. Monogr.

2: 771. syn.

Front wing 7.3 to 12 mm. long. Lateral ocellus separated from eye by

about 0.15 its long diameter in male, by about 0.18 its long diameter in fe-

male. First intercubitus about 1.0 as long as second section of cubitus. Ner-

vulus distad of basal vein by about 0.6 its length, strongly reclivous. Anal

cell without a spurious vein. Basal section of radiella vertical. First tergite

about 2.3 as long as wide in male, about 3.7 as long as wide in female.

Pale fulvous. Interocellar area black. Flagellum ferruginous brown to

dark brown. Wings subhyaline, the wing veins brown or dark brown.

Specimens: 212, Morogoro, 500 m., Tanzania, Jan. and Feb., 1962,

Gerd Heinrich (Townes). 6%, Uluguru, 700 to 1,800 m., Tanzania, Dec. 1961

and Jan. 1962, Gerd Heinrich (Townes). ©“, Mlanje, Malawi, Dec. 11, 1912,

S. A. Neave (London). 2, Otavifontein, Southwest Africa, Nov. 20, 1933, K.

Jordan (London). 3°, Tzaneen, South Africa, Jan. 11 and 18, 1971, H. and

M. Townes (Townes). ¢, Tongaat, Natal, South Africa, 1909, H. C. Burnap

(London).

This species is widely distributed in tropical Africa.

4. Paniscomima seyrigi Berland, new combination

*Rogezia Seyrigi Berland, 1951. Mém. Inst. Sci. Madagascar (A) 15: 301.

o, &. des., fig. Type: °, Madagascar: Ivondro (Paris). Paratype

studied in 1974. The “ described by Berland is considered to be a

separate species, treated below as P. curta.

Townes: A revision of the Rhopalosomatidae 15

Male: Unknown.

Female: Front wing 10.6 mm. long. Head more swollen than in other

species of the genus. Lateral ocellus separated from eye by 2.0 its long

diameter. First intercubitus 1.25 as long as second section of cubitus. Ner-

vulus distad of basal vein by 1.5 its length, weakly inclivous. Anal cell without

a spurious vein. Basal section of radiella strongly reclivous. First tergite

2.4 as long as wide.

Light fulvous. Interocellar area, tergites 3-6, and sternites 3-6 black.

Flagellum fulvous. Wings tinged with fulvous, the apex of front wing weakly

infuscate. Wing veins fulvous.

Specimen: 2 (paratype), eastern edge of forest, Rogez, Madagascar, 1935,

A. Seyrig (Paris). Besides this paratype Berland had a ¢ from Ivondro, and

another ? from south of the Bay of Antongil (both Madagascar).

5. Paniscomima bekilyi Berland, new combination

*Rhopalosoma bekilyi Berland, 1951. Mém. Inst. Sci. Madagascar (A) 5: 297.

, ¢. des., fig. Type: ¢, Madagascar: Bekily (Paris). Paratypes

studied in 1974.

Front wing 8.3 to 15 mm. long. Lateral ocellus separated from eye by

about 0.12 its long diameter. First intercubitus about 1.25 as long as second

section of cubitus. Nervulus distad of basal vein by about 1.6 its length, weak-

ly inclivous. Anal cell without a spurious vein. Basal section of radiella

weakly reclivous. First tergite about 2.9 as long as wide in male, about 3.8

as long as wide in female.

Fulvous. Ocellar area fuscous. Pedicel and flagellum dark brown. Wings

subhyaline. Wing veins brown or dark brown.

Specimens: 30, 3@ (all paratypes), Bekily, Madagascar, Oct., Nov., and

Dec., 1936, A. Seyrig (Townes, Washington, and London). ©, Isalo, 1, 000

m., Madagascar, R. M. G. and P. D. (London). Berland reported specimens

also from the localities Behara and Antanimora, both in Madagascar.

6. Paniscomima rufoantennata Berland, new combination

*Rhopalosoma bekilyi var. rufoantennata Berland, 1951. Mém. Inst. Sci.

Madagascar (A) 15: 298. Types: Madagascar (Paris). One syntype

studied in 1974.

Male: Unknown.

Female: Front wing 13 mm. long. Nervulus distad of basal vein by 1.45

its length. Pedicel and flagellum brownish fulvous. Otherwise similar to ,

P. bekilyi.

Although Berland described this as a variety of bekilyi, the constant differ-

ence in the color of the antenna and the slight difference in the position of the

nervulus indicate that more probably it is a distinct species.

Specimen: & (syntype), Fort Dauphin, Madagascar, Dec. 1936, A. Seyrig

(Townes). Berland reported 8 additional specimens, from the ''same localities"

as his P. bekilyi.

16 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

7. Paniscomima curta, new species

Rogezia Seyrigi Berland, 1951. Mém. Inst. Sci. Madagascar (A) 15: 301.

only.

Male: Front wing 6.9 to 9.0 mm. long. Lateral ocellus separated from

eye by about 0.45 its long diameter. First intercubitus about 1.0 as long as

second section of cubitus. Nervulus distad of basal vein by about 1.0 its length,

weakly inclivous. Anal cell without a spurious vein. Basal section of radiella

weakly reclivous. First tergite about 2.4 as long as wide.

Fulvous. Interocellar area blackish. Flagellum fulvous. Wings subhya-

line, their veins dark brown. Tergites and sternites 4-7 infuscate.

Female: Unknown.

Berland described this as the male of his P. seyrigi. Several conspicuous

differences in structure and color, however, make it more likely that it is a

separate species. The types designated below were originally paratypes of

seyrigi. There were originally 27 male paratypes. I have not seen the rest

of them.

Type: ©, Ivondro, Madagascar, Dec. 1938, A. Seyrig (Paris).

Paratypes: 20°, same data as type (Townes and Washington). ©“, Périnet,

Madagascar, Feb. 1939, A. Seyrig (Townes).

4. Genus Rhopalosoma

Figures 1-44

Rhopalosoma Cresson, 1865. Proc. Ent. Soc. Philadelphia 4: 58.

Type: Rhopalosoma poeyi Cresson. Monobasic.

Sibyllina Westwood, 1868. Trans. Ent. Soc. London 1868: 329.

Type: (Sibyllina aenigmatica Westwood) = poeyi Cresson. Monobasic.

Front wing 4.8 to 17 mm. long. Body slender, coloration stramineous to

ferruginous. Eye large, its inner margin with a sharp notch just above anten-

nal socket. Ocelli moderately large to very large. Temple short, weakly con-

vex. Occipital carina present on upper 0.6+ of head, absent below. Palpi

long. Flagellum long, its basal 5 to 8 segments with an apical pair of bristles

on upper and inner sides. Propodeum smoothly convex. Hind edge of meso-

sternum with a parallel pair of triangular flanges that are separated from meso-

sternum by a groove and strongly impressed. Front femur of female not swol-

len. Female tarsi very wide, weakly concave beneath and with dense tenent

hairs. Male tarsal claws with a pair of parallel points at apex and a large

angulate lobe a little behind apex. Female tarsal claws long, with a median

or subapical tooth and sometimes with an additional blunt tooth toward base.

First tergite 4.0 to 6.8 as long as wide. Anal lobe reaching approximately

0.65 to 1.0 the distance to nervellus.

This is a moderate sized genus that is restricted to the Neotropic Region,

except for one species in the southeastern United States. The species are

active in late afternoon and at night. They are commonly collected in Malaise

traps and at light. In flight they resemble ichneumonids of the genus Netelia

or the subfamily Ophioninae and like these ichneumonids they rest in low dense

vegetation during the day. The one species whose host is known (nearcticum)

parasitizes crickets of the subfamily Eneopterinae. Its larva lives in a black

Townes: A revision of the Rhopalosomatidae t7

sac on the side of the cricket until it kills its host.

Key to the species of Rhopalosoma

1. Front spur of middle and hind tibiae with basal 0.22+ forming a narrow

basal neck, then suddenly enlarged to an elongate lanceolate distal

portion. Head mostly black. Lanceolatum Group.

17. lanceolatum, new species (p. 27)

Front spur of middle and hind tibiae without a narrow basal neck, the

basal 0.6+ parallel-sided and apical 0.4+ gradually tapered. Head

fulvous or ferruginous, the ocellar area fuscous. .......... 2

2. Nervulus vertical, straight or weakly curved. Apical 0.65 of fifth seg-

ment of female tarsi parallel-sided. Isopus Group. ......... 3

Nervulus reclivous, moderately to strongly curved. Apical 0.65 of fifth

segment of female tarsi tapered, about 0.8 as wide at apex as at basal

O-S0° Peeyr Grouse. Wa ee a eg is ee

3. Basal section of radiella reclivous at about 40°. Front wing 6 to 8 mm.

long. Body ochraceous. Northwestern Argentina.

1. minus, new species (p. 19)

Basal section of radiella almost vertical. Front wing 11 to 14 mm. long.

Body ferrupinous. Brazil... . 0.3 2. isopus, new species (p. 19)

4. Lateral ocellus separated from eye by 1.0 to 1.25 its long diameter. . 5

Lateral ocellus separated from eye by 0.25 to 0.9 its long diameter. . 6

0. Nervulus distad of basal vein by about 2.0 its length, about 2.0 as far from

basal vein as from first recurrent vein.

3. missionicum Ogloblin (p. 20)

Nervulus distad of basal vein by about 2.7 its length, about 3.1 as far

from basal vein as from first recurrent vein. |

4. impar, new species (p. 20)

6. Basal section of radiella with a bend of 85° to 110°. Cubitella meeting

bees section of radiclla at 259 10 30". Se 7

Basal section of radiella with a curve rather than a bend, its two ends

30° to 80° from each other. Cubitella meeting basal section of radiella

CU CO ee ee, 10

7. Notch in eye margin 0.8 as deep as width of median ocellus. Bend in

basal section of radiella rather broad. Bolivia. :

5. bolivianum Brues (p. 20)

Notch in eye margin about 0.55 as deep as width of median ocellus. Bend

in basal section of radiella sharper. . 6°.:. ee ee, 8

8. Flagellar segments 1-5 fuscous. Mexico to Panama.

6. simile Brues (p. 21)

Flagellar segments 1-5 fulvous to brown. ............e.e0e6-% 2)

9. Bend in basal section of radiella about 90°. Lateral ocellus separated from

eye by about 0.75 its long diameter. Body ochraceous. Flagellum

18 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

fulvous. Southeastern United States. .. 7. nearcticum Brues (p. 21)

Bend in basal section of radiella about 100°. Lateral ocellus separated

from eye by about 0.55 its long diameter. Body ferruginous. Flagellum

dark fulvous to brown. Brazil. ... 8. angulare, new species (p. 22)

10. Flagellar segments 1-5 fulvous to ferruginous. ............. 11

Plapeliay Geumients 1-9 Drown tO.01ACk. {sos .s ale Goes ale ole 13

11. Lateral ocellus separated from eye by 0.25 its long diameter. Haiti.

9. haitiense, new species (p. 23)

Lateral ocellus separated from eye by 0.3 to 0.6 its long diameter. . 12

12. Second section of cubitus about 0.65 as long as third section. Basal sec-

tion of radiella strongly arched, its two ends divergent at about 70°.

Postnervulus moderately sinuate. Caribbean islands.

| 10. poeyi Cresson (p. 23)

Second section of cubitus about 0.5 as long as third section. Basal section

of radiella moderately arched, its two ends divergent at about 45°.

Postnervulus strongly sinuate. Brazil.

11. bahianum Schulz (p. 24)

13. Basal section of radiella inclivous at about 18°. Second cubital cell 2.1

as wide as high. Mexico..... 12. obliquum, new species (p. 24)

Basal section of radiella approximately vertical. Second cubital cell 2.6

Be es a Rs eee ta de sk Ga Se pie ee 8 leo 14

14. Scape entirely fuscous. Nervulus weakly curved. Bolivia.

13. scaposum, new species (p. 25)

Scape fulvous or ferruginous, usually infuscate apically. Nervulus weakly

Ry eee CIE, il a a na ame a eae a ae db uel onsale wagers. . « 15

15. Eye notch small, about 0.45 as deep as width of median ocellus. Panama

BO OUI Vie ONG SUT TAG oe aoe at co. seco 14. guianense Schulz (p. 25)

Eye notch larger, about 0.8 as deep as width of median ocellus. .... 16

16. Nervulus distad of basal vein by about 1.55 its length, the front end of

nervulus a little closer to basal vein than to first recurrent vein.

Thorax ochraceous. Panama... 15. breelandi, new species (p. 26)

Nervulus distad of basal vein by about 2.1 its length, the front end of

nervulus a little farther from basal vein than from first recurrent vein,

or equidistant. Thorax fulvous to feruginous. Brazil.

16. alvarengai, new species (p. 26)

Isopus Group

Front spur of middle and hind tibiae without a narrow basal neck, the

basal 0.6+ parallel-sided and apical 0.4+ gradually tapered. Apical 0.65 of

fifth segment of female tarsi parallel-sided. Nervulus vertical, straight or

weakly curved.

The parallel-sided segment 5 of the female tarsi is a character shared with

the Old World genus Paniscomima, which on other characters is definitely

more primitive than Rhopalosoma. This is evidence that the Isopus Group

Townes: A revision of the Rhopalosomatidae 19

is the primitive species group of Rhopalosoma.

There are two species of this group, occurring in Brazil and Argentina.

1. Rhopalosoma minus, new species

Figures 1 (head); 17 (venation); and 32 (“ genitalia)

Front wing 5.6 to 8.7 mm. long. Notch in eye about 0.66 as deep as width

of median ocellus. Lateral ocellus separated from eye by about 0.3 its long

diameter. Female tarsal claws moderately long, with a postmedian acute

tooth, the apical tooth stout. First intercubitus about 1.0 as long as second

section of cubitus. Nervulus vertical, almost straight, distad of basal vein

by about 2.0 its length. Basal section of radiella almost straight, reclivous

at about 40° from the vertical. Male genitalia as in figure 32.

Ochraceous, including the flagellum. Ocellar area dark brown. Wings

tinged with ochraceous, the veins light brown.

Type: ?, 11 kilometers west of Las Cejas, Tucuman, Argentina, March 9

to 26, 1967, ‘Lionel Stange (Townes).

Paratypes: 30°, same locality and collector as type, dated Jan. 15 to Feb.

14, 1967, March 9 to Apr. 11, 1968, and April 26 to 29, 1967 (Townes). §,

Las Cejas, Tucuman, Argentina, Jan. 21 to Feb. D1, 1968, C. C. Porter

(Cambridge). ©, San Pedro Colalao, Tucuman, Argentina, Feb. 1966, Lionel

Stange eee

This species occurs in northwestern Argentina.

2. Rhopalosoma isopus, new species

Figures 2 (head); 18 (venation); and 33 (% genitalia)

Front wing 11 to 13 mm. long. Notch in eye about 0.75 as deep as width

of median ocellus. Lateral ocellus separated from eye by about 0.45 its long

diameter. Female tarsal claws short, with a large postmedian tooth and with

avery stout, short, strongly curved apical point. First intercubitus about

0.82 as long as second section of cubitus. Nervulus vertical, weakly curved,

distad of basal vein by about 1.7 its length. Basal section of radiella weakly

curved, subvertical or slanted about 10° from the vertical. Male genitalia as

in figure 33.

Ferruginous, including the flagellum. Ocellar area blackish. Wings

tinged with brown, their veins medium brown.

Type: 9°, Jatai, Goias, Brazil, Nov. 1972, F. M. Oliveira (Townes).

Paratypes: 2%, 72, same data as type (Townes). 49, Sinop, 12° 31'S

55° 37' W, Brazil, Oct. 1975, M. Alvarenga (Townes).

Poeyi Group

Front spur of middle and hind tibiae without a narrow basal neck, the

basal 0.6+ parallel-sided and apical 0.4+ gradually tapered. Apical 0.65 of

fifth segment of female tarsi tapered, about 0.8 as wide at apex as at basal

0.35. Nervulus reclivous, strongly curved.

This group contains the majority of the genus.

20 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

3. Rhopalosoma missionicum Ogloblin

*Rhopalosoma missionicum Ogloblin, 1951. Rev. Soc. Ent. Argentina 15: 11.

. des., fig. Type: “, Argentina: Loreto in Misiones (La Plata).

Type studied in 1976.

Male type: Front wing 6.9 mm. long. Lateral ocellus separated from eye

by 1.0 its long diameter. First intercubitus 1.0 as long as second section of

cubitus. Nervulus distad of basal vein by 2.0 its length, 2.0 as far from basal

vein as from first recurrent vein. Otherwise similar to R. impar. The male

genitalia of this species have not been examined.

Specimen: & (type), Loreto, Misiones, Argentina, Feb. 15, 1934 (La Plata).

4. Rhopalosoma impar, new species

Figures 3 (head); 19 (venation); and 34 (% genitalia)

Front wing 6 to 11 mm. long. Eye notch about 0.65 as deep as width of

median ocellus. Lateral ocellus separated from eye by about 1.25 its long

diameter. Female tarsal claws long, with a postmedian tooth, the apical

tooth of moderate length, rather weakly curved. First intercubitus about 1.25

as long as second section of cubitus. Nervulus sinuate, strongly reclivous,

separated from basal vein by about 2.7 its length, relatively close to first

recurrent vein, about 3.1 as far from basal vein as from first recurrent vein.

Basal section of radiella bowed, subvertical or weakly inclivous, joining cubi-

tella at an angle of about 40°, its two ends divergent at about 7 } Male geni-

talia as in figure 34.

Fulvoferruginous, the flagellum brownish ferruginous and interocellar

area blackish. Wings tinged with brown, their veins brown.

Type: ¢, Pedra Azul, 800 m., Minas Gerais, Brazil, Nov. 1972, Alva-

renga and Seabra (Townes).

Paratypes: 29, same data as type (Townes). °%, 2°, Piracicaba, Sao

Paulo, Brazil, Jan. 12, 20, and 22, 1965, C. A. and W. E. Triplehorn

(Ithaca and Townes). ©, Nanawa, Chaco, Paraguay, Oct. to Nov. 1926, A.

Pride (London).

This species occurs in Paraguay and southern Brazil.

5. Rhopalosoma bolivianum Brues

Figures 4 (head) and 20 (venation)

*Rhopalosoma bolivianum (as boliviense in Fig. 3) Brues, 1943. Ann. Ent.

Soc. America 36: 316. 2. key, des., fig. Type: ¥, Bolivia: Buena

Vista near Santa Cruz (Ithaca). Type studied in 1974.

Male: Unknown.

Female: Front wing 12 mm. long. Eye notch 0.8 as deep as width of

median ocellus. Lateral ocellus separated from eye by 0.67 its long diameter.

Tarsal claws moderately long, with a postmedian tooth, the apical tooth of

moderate length and moderately curved. First intercubitus 0.78 as long as

second section of cubitus. Nervulus sinuate, reclivous, distad of basal vein

Townes: A revision of the Rhopalosomatidae 21

by 1.6 its length. Basal section of radiella weakly inclivous, with a median

90° bend, joining cubitella at an angle of 30°.

Fulvous, the flagellum medium brown. Interocellar area brown. Wings

tinged with fulvous brown, the veins brown.

Specimen: 2? (type), Buena Vista, near Santa Cruz, Bolivia, 1928, J.

Steinbach (Ithaca).

6. Rhopalosoma simile Brues

Figures 5 (head) and 35 (° genitalia)

*Rhopalosoma simile Brues, 1943. Ann. Ent. Soc. America 36: 317. 9%, ¢.

key, des., fig. Type: ¢, Barro Colorado Island, Panama (Washington).

Type studied in 1974.

Front wing 8.2 to 16.5 mm. long. Scape dark brown. In all other respects

similar to R. nearcticum. Male genitalia as in figure 30.

This will probably prove to be only subspecifically distinct from R. nearcti-

cum.

~ Specimens: 150°, 532 from Mexico (Atoyak in Vera Cruz, Catemaco at

1,100 ft. in Vera Cruz, 20 miles north of Huixtla in Chiapas, Mazatlan in

Sinaloa, Minatitlan in Vera Cruz, and Merida in Yucatan); Guatemala (Cayuga

and Quirieud): British Honduras (Camp Sibim in Coyo District and Middlesex

at 125 m.); El Salvador (Lake Ilopango near Apulo and Alta Marina at 2, 500 ft.

in San Salvador); Costa Rica (Turrialba at 600 m.); and Panama (Barro Col-

orado Island and Trinidad ee) Collection dates are distributed from March

25 to October 6. Two of the specimens were collected at lights, others in

Malaise traps. Specimens are in the collections of East Lansing, London,

Ottawa, Townes, and Washington.

This species ranges from Mexico to Panama.

7. Rhopalosoma nearcticum Brues

Figures 6 (head) and 21 (venation)

Rhopalosoma poeyi Ashmead, 1896. Proc. Ent. Soc. Washington 3: 304.

Mo.: St. Louis. N. C. Fla.: St. Augustine. Ky.: Louisville. Misde-

termination.

Rhopalosoma poeyi Hood, 1913. Proc. Ent. Soc. Washington15: 145-147. fig. ,

biol. Md.: Plummers Island. Host: Orocharis saltator. Misdetermination.

Rhopalosoma poeyi Rohwer, 1913. Proc. Ent. Soc. Washington 15: 147- sii

discussion, distribution. Misdetermination.

Rhopalosoma poeyi Cresson, 1916. Mem. Amer. Ent. Soc. 1: 71. N.C.

Misdetermination.

Rhopalosoma poeyi Brimley, 1938. The insects of North Carolina .

p. 436. N. C.: Raleigh. Misdetermination.

*Rhopalosoma nearcticum Brues, 1943. Ann. Ent. Soc. America 36: 316.

oa, 2. key, des., fig. Type: ¢, Florida: Kissimmee (Cambridge).

Fla.: Seminole Co., Hillsboro Co., Ft. Meade, Gulfport, St. Augustine.

Md.: Plummers Island. Type studied in 1973.

Rhopalosoma nearcticum Gurney, 1953. Proc. U. S. Natl. Mus. 103: 19-34.

biol... figs.:oGa,, Fla: ; Ind., Middl, Migs... Mo Ne Cy 6. Cl Va:

Hosts: Hapithus agitator, H. brevipennis.

e “3

22 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Front wing 7.0 to 13.2 mm. long. Eye notch about 0.44 as deep as width

of median ocellus. Lateral ocellus separated from eye by about 0.75 its long

diameter. Female tarsal claws long, with a postmedian pointed tooth and sub-

basally a smaller rounded tooth, its apical tooth short, stout, and with a mod-

erately strong curvature. First cubital vein about 0.90 as long as second sec-

tion of cubitus. Nervulus sinuate, strongly reclivous, distad of basal vein by

about 1.9 its length. Basal section of radiella strongly inclivous, with a bend

of about 90° that is centered above the middle, the radiella meeting cubitella

at an angle of about 259. Male genitalia similar to those of R. simile (figure

$5). 7

Ochraceous. Ocellar area black. Flagellum fulvous. Wings tinged with

ochraceous, the veins light brown.

Specimens: 460°, 662 from Florida (Fort Myers, Gulfport, Homestead,

Jacksonville, Kissimmee, Lake Placid, Martin Co., Orange Co., Palm Beach

Co., St. Augustine, Seminole Co., and Tall Timbers near Tallahassee);

Georgia (Atlanta and Stone Mt.); Kentucky (Golden Pond); Louisiana (Bayou

Chicot in Evangeline Co.); Maryland (Berlin, Mayo, and Plummers Island);

Missouri (Columbia and Williamsville); North Carolina (Long Beach and Oxford);

South Carolina (Greenville); Tennessee (Knoxville ?); and Texas (Sinton). Col-

lection dates in Florida range from April 23 to August 28, with a peak of col-

lections in May and June. Farther north, the collection dates are from July 9

to October 29, and from San Patricio Co., Texas, there is a specimen dated

June 17. About half of the total number of specimens were collected in Malaise

traps. Two of the specimens are labeled as having been collected at lights.

In my own experience this species is locally common in the south Atlantic

states, in dense shrubby vegetation where the humidity is high. Adults fly

shortly before sunset and at least until dark, just above the bushes. In flight

they resemble Ophion and Netelia except for their somewhat stouter appear-

ance. Crickets of the genera Hapithus and Orocharis are often found with a

bulging blackish larva of this parasite stuck on one side near the base of a

hind leg, forcing the leg outward in an unnatural position. From such para-

sitized crickets there have been rearings by Hood (from Orocharis saltator)

and by Gurney (from Hapithus agitator). Both of these workers published their

observations (see synonymy above) and Gurney gave a detailed account of the

biology with figures of the larva.

This species occurs in the Upper and Lower Austral zones of the eastern

United States. It parasitizes crickets of the genera Orocharis and Hapithus.

8. Rhopalosoma angulare, new species

Figures 7 (head); 22 (venation); and 36 (“ genitalia)

Front wing 11 to 17 mm. long. Eye notch about 0.63 as deep as width of

median ocellus. Lateral ocellus separated from eye by about 0.55 its long

diameter. Female tarsal claws long, with a postmedian pointed tooth and sub-

basally a smaller rounded tooth, its apical tooth stout, weakly curved. First

intercubitus about 1.15 as long as second section of cubitus. Nervulus sinuate,

reclivous, separated from basal vein by about 2.2 its length. Basal section of

radiella inclivous, with a sharp bend of about 100° that is centered just above

the middle, the radiella meeting cubitella at an angle of about 25°. Male geni-

talia as in figure 36.

Fulvoferruginous. Ocellar area blackish, the blackish mark extending a

Townes: A revision of the Rhopalosomatidae 23

little beyond the ocellar interspace. Flagellum dark fulvous to brown. Wings

with a fulvoferruginous tinge, the veins ferruginous brown. Abdomen often

somewhat infuscate beyond segment 3.

Type: °, Encruzilhada, 960 m., Bahia, Brazil, Nov. 1972, M. Alvarenga

(Townes).

Paratypes: 50°, 222, same data as type (Townes). %, 89, Encruzilhada,

980 m., Bahia, Brazil, Nov. 1974, M. Alvarenga (Townes). ¢, Caruaru,

900 m., Pernambuco, Brazil, Apr. 1972, M. Alvarenga (Townes). 5%, Lin-

hares, Espirito Santo, Brazil, Nov. 1972, M. Alvarenga (Townes). ?, Pedra

Azul, Minas Gerais, Brazil, Nov. 1972, Alvarenga and Seabra (Townes).

This species occurs in eastern and southeastern Brazil.

9. Rhopalosoma haitiense, new species

Figures 8 (head) and 23 (venation)

*Rhopalosoma aenigmaticum Brues, 1943. Ann. Ent. Soc. America 36: 315.

. Key, des., fig. Haiti. Misdetermination of aenigmaticum Westwood.

Male: Unknown.

Female type: Front wing 10.2 mm. long. Eye notch 0.75 as deep as width

of median ocellus. Lateral ocellus separated from eye by 0.25 its long dia-

meter. Tarsal claws moderately long with a postmedian tooth and near the

base with a very low rounded tooth, its apical tooth of moderate stoutness and

curvature. First intercubitus 0.88 as long as second section of cubitus. Ner-

vulus sinuate, reclivous, distad of basal vein by 1.7 its length. Basal section

of radiella inclivous, with a strong even curvature, its two ends divergent at

75°, the radiella meeting cubitella at 30°.

Ochraceous, the flagellum fulvous. Interocellar area dark brown. Wings

subhyaline, the veins light brown.

Type: ¢, Manneville, Haiti (Cambridge). This specimen is one of those

reported by Brues in 1943 incorrectly as R. aenigmaticum. Brues had an

additional 20°, 12 from the same locality. These I have not seen.

10. Rhopalosoma poeyi Cresson

Figures 9 (head); 24 (venation); and 37 (“ genitalia)

*Rhopalosoma Poeyi Cresson, 1865. Proc. Ent. Soc. Philadelphia 4: 58.

, ¢. des., fig. Lectotype: 2 (designated by Cresson, 1916), Cuba

(Philadelphia). Lectotype studied in 1974.

*Sibyllina aenigmatica Westwood, 1868. Trans. Ent. Soc. London 1868: 329.

@. des. Lectotype: 2 (labeled by Townes, 1973, hereby designated),

Haiti (London). Lectotype studied in 1973. New synonym.

Sibyllina aenigmatica Smith, 1869. Trans. Ent. Soc. London 68 (proc.): LI.

des.

Rhopalosoma aenigmatica Westwood, 1874. Thesaurus entomologicus oxonien-

sis p. 1380-132. des., fig. Cuba. Haiti.

Rhopalosoma poeyi Schulz, 1906. Spolia hymenopterologica, p. 226. key.

Rhopalosoma poeyi Cresson, 1916. Mem. American Ent. Soc. 1: 71. Lecto-

type designation.

Rhopalosoma poeyi Brues, 1943. Ann. Ent. Soc. America 36: 315. 9. key,

des. Cuba.

24 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

Rhopalosoma poeyi Krombein, 1953. Amer. Mus. Novitates 1633: 7. des.

Bahamas: South Bimini.

Front wing 4.8 to 8.2 mm. long. Notch in eye about 0.70 as deep as width

of median ocellus. Lateral ocellus separated from eye by about 0.5 its long

diameter in male, by about 0.4 its long diameter infemale. Female tarsal

claws long, with a postmedian tooth, the apical tooth strongly curved and rather

short. First intercubitus about 0.83 as long as second section of cubitus. Ner-

vulus moderately sinuate or curved, weakly reclivous, distad of basal vein by

about 2.0 its length. Basal section of radiella strongly curved, meeting cubi-

tella at an angle of about 30°, its two ends divergent at about 70°. Male geni-

talia as in figure 37.

Fulvoferruginous, including the flagellum. Interocellar area partly or en-

tirely dark brown. Wings tinged with fulvous brown, their veins brown. |

Specimens: 30°, Man-o-War Cay, Bahamas, Aug. 23 and 30, 1971, H. and

A. Howden (Townes). &%, South Bimini Island, Bahamas, June 8, 1950, Cazier

and Rindge (Washington). 2, 62, in light trap, interior of Cayman Island,

behind Stakes Bay, Apr. 17 to Aug. 26, 1938, C. B. Lewis and C. H. Thompson

(London and Townes). 2 (type of poeyi), Cuba (Philadelphia). ¢, Jamaica (Lon-

don). 2 (lectotype and paratype of aenigmatica), Santo Domingo, [ Haiti] (Lon-

don).

This species occurs in the Caribbean Islands, from the Bahamas to His-

paniola.

11. Rhopalosoma bahianum Schulz

Figures 10 (head); 25 (venation); and 38 (% genitalia)

*Rhopalosoma bahianum Schulz, 1906. Spolia hymenopterologica, p. 222, 226.

2. key, des. Type: 2, Brazil: Orobo in Bahia (Strasbourg). Type

studied in 1974.

Front wing 8.8 to 11.3 mm. long. Eye notch about 0.67 as deep as width

of median ocellus. Lateral ocellus separated from eye by about 0.36 its long

diameter. Female tarsal claws of moderate length, with a postmedian tooth

but no subbasal tooth, the apical tooth moderately stout and weakly curved.

First intercubital vein about 1.1 as long as second section of cubitus. Nervulus

strongly curved and strongly reclivous, distad of basal vein by about 1.8 its

length. Basal section of radiella long, weakly inclivous, evenly curved, its

two ends divergent at about 45°, the radiella meeting cubitella at about 60°.

Male genitalia as in figure 38.

Fulvous, the flagellum fulvous brown. Interocellar area black. Wings

with a weak fulvous brown tinge, the veins brown.

Specimens: 20, 4°, Encruzilhada, 960 m., Bahia, Brazil, Nov. 1972, M.

Alvarenga (Townes). 2, Barialba, 400 m., Ceara, Brazil, May 1969, M.

Alvarenga (Townes).

12. Rhopalosoma obliquum, new species

Figures 11 (head); 26 (venation); and 39 (° genitalia)

Male type: Front wing 10.2 mm. long. Eye notch 0.70 as deep as width

Townes: A revision of the Rhopalosomatidae 20

of median ocellus. Lateral ocellus separated from eye by 0.47 its long dia-

meter. First intercubitus 1.25 as long as second section of cubitus. Nervulus

sinuate, reclivous, distad of basal vein by 1.44 its length. Basal section of

radiella inclivous at 35° from the vertical, moderately curved, meeting cubi-

tella at 70°. Genitalia as in figure 39.

Pale fulvous. Flagellum black. Pedicelfuscous. Scape stramineous, in-

fuscate above. Ocellar area fuscous and the fuscous area reaching laterad to

eye. Wings with a fulvous tinge, the veins light brown. Abdomen infuscate

beyond segment 3.

Female: Unknown.

Type: ©, 12 miles south of Valle Nacional, 3,000 ft., Mexico, May 17,

1971, Henry Howden (Townes).

13. Rhopalosoma scaposum, new species

Figures 12 (head); 27 (venation); and 40 (° genitalia)

Male type: Front wing 8.7 mm. long. Eye notch 0.56 as deep as width of

median ocellus. Lateral ocellus separated from eye by 0.80 its long diameter.

First intercubitus 0.85 as long as second section of cubitus. Nervulus weakly

curved, strongly oblique, basad of basal vein by 1.12 its length. Basal sec-

tion of radiella almost vertical, strongly bowed at middle with the two ends

divergent at 75°, radiella meeting cubitella at 459. Genitalia as in figure 40.

Pale fulvous. Ocellar area and frons fuscous, the frons fulvous along its

lateral and lower edges. Antenna entirely black. Wings tinged with pale ful-

vous, the veins light brown.

Female: Unknown.

Type: ©“, Crystal Mayu, 300 m., Cochabamba, Bolivia, Dec. 5, 1949, L.

Pefia (Townes).

14. Rhopalosoma guianense Schulz

Figures 13 (head); 28 (venation); and 41 (% genitalia)

*Rhopalosoma guianense Schulz, 1906. Spolia hymenopterologica, p. 221, 226.

@. key, des. Type: 9, British Guiana: Oyapok [ in Para] (Strasbourg).

Type studied in 1974.

*Rhopalosoma martinezi Ogloblin, 1951. Rev. Soc. Ent. Argentina 15: 14. 2.

des., fig. Type: 2, Bolivia: Santa Cruz (La Plata). Type studied in

1976. New synonym.

Front wing 9.8 to 16.0 mm. long. Eye notch small and shallow, about >

0.50 as deep as width of median ocellus. Lateral ocellus separated from eye

by about 0.90 its long diameter. Female tarsal claws long, with a postmedian

tooth and a low rounded tooth near base, the apical point moderately stout and

curved. First intercubitus about 0.80 as long as second section of cubitus.

Nervulus strongly curved, reclivous, distad of basal vein by about 2.6 its

length. Basal section of radiella weakly inclivous, weakly curved, its two

ends divergent by about 60°, radiella meeting cubitella at about 70°. Note:

The radiella figured is less curved than normal, and less inclivous than nor-

mal. Male genitalia as in figure 41.

Fulvoferruginous. Head black between ocelli and the black area extending

26 Contrib. Amer. Ent. Inst., vol. 15, no. 1, 1977

ventrolaterally to reach or almost reach the eye margins. Flagellum blackish.

Scape and pedicel infuscate above. Wings tinged with brown, the veins brown.

Abdomen sometimes infuscate beyond segment 2 or beyond segment 3.

Specimens: 12°, 372 from Costa Rica (Sixola River); Panama (Almirante,

Barro Colorado Island, Darien, Margarita, Pipeline Road near Gamboa, San

Jose in the Pearl Islands, and Trinidad River); Colombia (Coqueto on the Rio

Orteguaza near Rio Peneya); Ecuador (Coca); Peru (Avispas at 400 m. and

Chanchamayo at 1,300 m.); Bolivia (Alto Palmar at 1,100 m. in Cochabamba);

Trinidad (Morne Bleu at 2, 700 ft.); British Guiana (Blairmont, Demarara in

Georgetown, King Frederick William IV Falls on the upper Courantyne River,

and Moraballi Creek of the Essequibo River); and Surinam (45 kilometers south

of Paramaribo). One of the specimens is labeled as collected at light. The

specimens from Trinidad River, Panama and from Sixola River, Costa Rica

are paratypes of R. simile Brues (misdetermined). Specimens are in the col-

lections of Cambridge, London, Ottawa, Townes, and Washington.

This species ranges from Central America to Bolivia and Surinam.

15. Rhopalosoma breelandi, new species

Figures 14 (head); 29 (venation); and 42 (“ genitalia)

Male: Front wing 9.3 to 10.3 mm. long. Eye notch about 0.93 as deep as

width of median ocellus. Lateral ocellus separated from eye by about 0.60 its

long diameter. First intercubitus about 0.82 as long as second section of cubi-

tus. Nervulus sinuate, reclivous, distad of basal vein by about 1.55 its length.

Basal section of radiella subvertical, bowed at center, its two ends divergent

at about 70°, radiella joining cubitella at about 60°. Genitalia as in figure 42.

Ochraceous. Pedicel brown. Flagellum dark brown. Interocellar area

fuscous and the fuscous area extending laterally to reach the eye margin.

Wings with an ochraceous tinge, the veins light brown.

Female: Unknown.

Type: “, Margarita, Canal Zone, Panama, Jan. 1960, Sam Breeland

(Townes).

Paratype: ©, same locality and collector as type, dated June, 1960

(Townes).

16. Rhopalosoma alvarengai, new species

Figures 15 (head); 30 (venation); and 43 (“ genitalia)

Front wing 7.7 to 14 mm. long. Eye notch about 0.70 as deep as width of

median ocellus. Lateral ocellus separated from eye by about 0. 43 its long

diameter. Female tarsal claws long, with a postmedian tooth and subbasally

with a low rounded prominence, the apical point stout and moderately curved.

First intercubitus about 0.74 as long as second section of cubitus. Nervulus

reclivous, strongly curved, distad of basal vein by about 2.1 its length. Basal

section of radiella weakly inclivous, strongly curved, its two ends divergent at

about 60°, radiella meeting cubitella at about 60°. Male genitalia as in figure

43. |

Ferruginous. Ocellar area dark brown, the brown mark extending laterally

to eye margin, often this dark area covering also the upper part of frons. Pedi-

cel and flagellum blackish. Scape infuscate apically. Wings tinged with brown,

Townes: A revision of the Rhopalosomatidae 27

the veins brown. Abdomen of male often infuscate beyond second segment.

Type: °, Represa Rio Grande, Guanabara, Brazil, Jan. 1968, M. Alvar-

enga (Townes).

Paratypes: 33°, same locality and collector as type, dated Jan., Feb.,

Mar., Sept., Oct., Nov., and Dec., 1967, 1968, 1969, 1970, and 1972

(Townes). 22, Alto da Serra, Moretes, Brazil, Feb. 12 and 18, 1966, H. and

M. Townes (Townes). 9°, Campina Grande (near Curitiba), Brazil, Feb. 16,

1966, H. and M. Townes (Townes). 3%, Caruaru, Brazil, Apr. 1972, M.

Alvarenga (Townes). 4°, 13°, Encruzilhada, 960 m., Bahia, Brazil, Nov.

1972, M. Alvarenga (Townes). 2°, 6%, Encruzilhada, 980 m., Bahia, Brazil,

Nov. 1974, M. Alvarenga (Townes). 60, 112, Linhares, Espirito Santo,

Brazil, Sept. 1972, M. Alvarenga (Townes). 22, Margaritiba, Muriqui, Rio

de Janeiro, Brazil, July 1969, M. Alvarenga (Townes). 3°, Rio de Janeiro,

Brazil, Jan. 1939, R. C. Shannon (Washington). &% (paratype of R. simile

Brues, misdetermined), Tuis, 2,400 ft., Costa Rica, C. H. Lankester

(Washington).

This species is known from eastern and southeastern Brazil and from

Costa Rica. .

Lanceolatum Group

Front spur of middle and hind tibiae with a narrow basal neck that is about

as long as diameter of their basitarsi, distad of the neck swollen and continued

in a narrowly lanceolate shape, bent about 15° at the distal end of the neck.

Nervulus straight, vertical.

One species is known, and this only in the male sex.

17. Rhopalosoma lanceolatum, new species

Figures 16 (head); 31 (venation); and 44 (“ genitalia)

Male type: Front wing 10.9 mm. long. Eye notch 0.65 as deep as width

of median ocellus. Lateral ocellus separated from eye by 0.5 its long diameter.

First intercubitus 1.2 as long as second section of cubitus. Nervulus weakly

inclivous, almost straight, distad of basal vein by 1.55 its length. Basal sec-

tion of radiella vertical, weakly curved. Genitalia as in figure 44.

Fulvous. Antenna entirely black. Head black, the cheek, clypeus, and

occiput fulvous. Maxillary palpus weakly infuscate except basally. Wings with

a fulvous tinge. Abdomen blackish beyond the first segment except that seg-

ment 2 is stained with fulvous, more strongly fulvous toward base.

Female: Unknown. |

Type: “, Nova Teutonia, 279 11' S, 52° 23' W, 300-500 m., Brazil, Nov.

1968, Fritz Plaumann (Ottawa).

List of figures

1-16. Rhopalosoma, front view of head (figures 11, 12, 13, 15, and 17 of

males, the rest of females).

17-31. Rhopalosoma, venational features (2nd cubital cell, nervulus and post-

nervulus, and radiella).

32-44. Rhopalosoma, male genitalia (penis valve, and volsella and digitus).

5. Rhopalosoma

simile

1. Rhopalosoma

minus

6. Rhopalosoma

nearcticum

2. Rhopalosoma

isopus

7. Rhopalosoma

angulare

8. Rhopalosoma

haitiense

4, Rhopalosoma

bolivianum

9. Rhopalosoma

poeyi

13. Rhopalosoma

guianense

14. Rhopalosoma

breelandi

10. Rhopalosoma

bahianum

15. Rhopalosoma

11. Rhopalosoma alvarengai

obliquum

12. Rhopalosoma

scaposum

Rhopalosoma

lanceolatum

‘ a i@

, ae

2nd. cubital cell a 2nd. cubital cell ,

se pies ene eae tle NS ETE SAW

eS oer ae

—a ee

17. Rhopalosoma minus

20. Rhopalosoma bolivianum

2nd. cubital cell

ez Nervulus —

Nervulus ns L

18. Rhopalosoma isopus Zh

. Rhopalosoma nearcticum

2nd. cubital cell

Ce Oe

SO ee

ee ee, aa

oe :

19, Rhopalosoma impar 22. Rhopalosoma angulare

2nd, cubital cell

2nd. cubital cell

————————— ——_

Nervulus cet

ee ued vA

ee ee

immer en

yee

23. Rhopalosoma haitiense

26. Rhopalosoma obliquum

2nd. cubital cell oe

m8 ee : A RSs

Bs : :

oe eae :

ea Nervulus < ———

Nervulus

Oa be ee :

— aT Pa es acme gencee

24. Rhopalosoma poeyi

2nd. cubital cell

ao. : |

Me a Sey Nervulus

Se ee,

PR 9p Wena oe ene een

Radiella |

Ean eee

i ea

25. Rhopalosoma bahianum 28. Rhopalosoma guianense

Radiella

2nd. cubital cell

ee es a

Nervulus

4 :

2nd. cubital cell

Nervulus

a en ee

Radiella Testun hee Deer

eee es Radiella

29. Rhopalosoma breelandi i NE RIT Ei

31. Rhopalosoma lanceolatum

Nervulus

Radiella

30. Rhopalosoma alvarengai wr

Rhopalosoma minus

35. Rhopalosoma simile

Rhopalosoma isopus 34. Rhopalosoma impar

44. Rhopalosoma lanceolatum

42. Rhopalosoma breelandi 43. Rhopalosoma alvarengai

34 Contrib: Amer. Ent. Inst. , vol. 15, no. 1, 1977

INDEX TO RHOPALOSOMATIDAE

*An asterisk indicates a synonym, misspelling, or misidentification.

abnormis (Paniscomima, Rhopalosoma, Hymenochimaera) 12 (key), 13

*aenigmaticum (Rhopalosoma, Sibyllina) 23

*Algoa 3

*Algoella 3

alvarengai (Rhopalosoma) 18 (key), 26, 29, 32, and 33 (figs. )

angulare (Rhopalosoma) 18 (key), 22, 28 30, and 33 (figs. )

australiae (Olixon, Harpagocryptus) 2, 3, 4 (key), 7

bahianum (Rhopalosoma) 18 (key), 24, 29, 31, and 33 (figs. )

banksii (Olixon, Nealgoa) 3, 4 (key), 6

bekilyi (Paniscomima, Rhopalosoma) 13 (key), 15

bolivianum (Rhopalosoma) 17 (key), 20, 28 and 30 (figs).

*boliviense (Rhopalosoma) 20

breelandi (Rhopalosoma) 18 (key), 26, 29, 32, and 33 (figs. )

curta (Paniscomima, Rogezia seyrigi) 13 (key), 16

dentatum (Olixon, Apteropompilus, Psyllosphex, Algoella) 3, 5 (key), 9

erlangeriana (Paniscomima, Rhopalosoma) 12, 13 (key), 14 .

flavibase (Olixon) 4 (key), 5 ao

guianense (Rhopalosoma) 18 (key), 25, 29, 31, and 33 (figs. )

haitiense (Rhopalosoma) 18 (key), 23, 28 and 31 (figs. )

*Harpagocryptus 3 ae

*heterodoxa (Algoa, Algoella) 3, 9

*Hymenochimaera 3

impar (Rhopalosoma) 17 (key), 20, 28, 30, and 32 (figs. )

isopus (Rhopalosoma) 17 (key), 19, 28, 30, and 32 (figs. )

Isopus Group 17 (key), 18 ae

lanceolatum (Rhopalosoma) 17 (key), 27, 29, 32, and 33 (figs. )

Lanceolatum Group 17 (key), 27

Liosphex 3 (key), 10 a

majus (Olixon) 4 (key), 5

*martinezi (Rhopalosoma) 25

minus (Rhopalosoma) 17 (key), 19, 28, 30, and 32 (figs. )

missionicum (Rhopalosoma) 17 (key), 20

myrmosaeforme (Olixon, Psyllosphex) 5 (key), 9

*Nealgoa 3 =

nearcticum (Rhopalosoma) 1, 18 (key), 21, 28 and 30 (figs. )

obliquum (Rhopalosoma) 18 (key), 24, 29, 31, and 33 (figs. )

Olixon 3 (key) oi

opposita (Paniscomima) 12 (key), 14

*pallidus (Saphobethyllus, Olixon) 3, 7

Paniscomima 3 (key), 12

poeyi (Rhopalosoma) 16, 18 (key), 23, 29, 31, and 33 (figs. )

*poeyi (Rhopalosoma) 21

Poeyi Group 17 (key), 19

*Psyllosphex 3 a

Rhopalosoma 3 (key), 16

*Rogezia 3 os

rufoantennata (Paniscomima, Rhopalosoma bekilyi) 13 (key), 15

saltator (Olixon, Psyllosphex) 3, 5 (key), 8

*Saphobethyllus 3 os

scaposum (Rhopalosoma) 18 (key), 25, 29, 31, and 33 (figs. )

seyrigi (Paniscomima, Rogezia) 12, 13 (key), 14, 16

*Sibyllina 3

simile (Rhopalosoma) 17 (key), 21, 28 and 32 (figs. )

testaceum (Olixon) 3, 4 (key), 7

trichopleurum (Liosphex) 10 (key), 11

varius (Liosphex) 10 (key), 11

Ae a)

Ye of

lM, a0

La ee

i ae

i: a

‘Lemna Weriiiene Sa

ee eer Te en ai

: 7

on 7

7. |

tna f

Contributions

of the

American Entomological Institute

Volume 15, Number 2, 1977

ny te

A REVISION OF THE HELORIDAE

(HYMENOPTERA)

Henry Townes

American Entomological Institute

0950 Warren Road, Ann Arbor, Michigan

TABLE OF CONTENTS

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A REVISION OF THE HELORIDAE

(HYMENOPTERA)

By Henry Townes

American Entomological Institute,

5950 Warren Rd., Ann Arbor, Michigan, U. S. A.

The only living genus of the Heloridae is Helorus. The genera Ropronia,

Vanhornia, Tetraconus, Monomachus, and Austronia were placed in the Helori-

dae by Riek (1955. Australian Jour. Zool. 3: 258-259). These have venations

somewhat similar to that of Helorus but have conspicuous differences in body

structure. The resemblances in venation are due to their having unspecialized

venation of the serphoid type rather than to a close phyletic relationship with

Helorus or between themselves. I exclude them from the Heloridae. The

genera Ropronia, Monomachus, and Austronia may be considered the only

living representatives of the families Roproniidae, Monomachidae, and Aus-

troniidae, at least till there is clearer evidence as to their individual relation-

ships. Vanhornia is believed to be an aberrant and primitive member of the

Serphidae. Judging from the original description, Tetraconus is a synonym of

Monomachus.

All species of Helorus are parasites of Chrysopidae. Oviposition is into a

chrysopid larva. The host larva is killed after it has spun a cocoon. Helorus

pupates within the chrysopid cocoon and the adult cuts a circular hole in the

cocoon for emergence. Clancy (1946. Univ. California Pub. Ent. 7: 407-418)

gives details of the life history of Helorus anomalipes (under the name H. para-

doxus).

This revision was substantially assisted by a revision of the European

Helorus by Pschorn-Walcher (1955. Mitt. Schweizerischen Ent. Gesell. 38:

233-250). Pschorn-Walcher's paper contains discussions of the types, vari-

ability, distribution, hosts, and some of the previous confusions in the deter-

minations of European species. These discussions will not be repeated here.

I have differed from Pschorn-Walcher's revision in treating corruscus as a

synonym of ruficornis, rugosus as a synonym of nigripes, and meridionalis as

a synonym of striolatus.

The following collections were studied:

British Museum (Natural History), London, England.

Curator: Mr. N. Fergussen.

Canadian National Collection, Ottawa, Ontario, Canada.

Curator: Dr. Lubomir Masner.

Instituto Miguel Lillo, Tucumdan, Argentina. Curator: Dr. Abram Willink.

Michigan State University, East Lansing, Michigan.

Curator: Prof. Roland Fischer.

National Museum of Natural History, Washington, D. C.

Curator: Mr. C. F. W. Muesebeck

Townes Collection, Ann Arbor, Michigan.

Curators: Henry and Marjorie Townes

2 Contrib. Amer. Ent. Inst., vol. 15, no. 2, 1977

I wish to thank the curators who loaned specimens, and especially to thank

my brother, George Townes, who collected a substantial amount of material.

My wife has given assistance in a number of ways. Dr. D. W. Clancy gener-

ously gave permission to reproduce his figure of Helorus anomalipes.

Genus Helorus

Helorus Latreille, 1802. Histoire naturelle . . . des Crustaces et des

insectes 3: 309. Type: (Helorus ater Latreille) = anomalipes Panzer.

Monobasic.

Copelus Provancher, 1881. Nat. Canad. 12: 206. Type: (Copelus paradoxus

Provancher) = anomalipes Panzer. Monobasic.

The genus is best described by giving a figure, as below. The venation,

body shape, and pectinate claws make it easy to recognize. The size varies

from a front wing length of 2.5 mm. to a front wing length of 4.3 mm. The

head and body are black in all species except in the Neotropic H. brethesi, in

which the thorax is partly red.

Figure 1. Helorus anomalipes 9. Republished from Clancy, 1946.

Townes: A revision of the Heloridae 3

Key to the species of Helorus

1. First abdominal segment 2.5 to 3.8 as long as wide. Face and mesocutum

essentially smooth, their punctures so fine and shallow as to be incon-

SOICUOUS c's gos paces elle Ga See te ek ee pea ck eta eee age 2

First abdominal segment about 1.8 as long as wide (about 2.3 as long as

wide in H. australiensis). Face and mesoscutum with punctures that

are small but deep, or very coarsely rugoso-punctate. ....... 5)

2. Concave part of side of pronotum smooth or with a few weak wrinkles.

Nervulus basad of basal vein by about 0.3 its length. Mesoscutum

partly or entirely ferruginous. Neotropic Region.

1. brethesiOgloblin (p. 3)

Concave part of side of pronotum with strong horizontal wrinkles. Nervu-

lus opposite basal vein, or distad, or basad of basal vein. Mesoscutum

3. Trochanters and middle femur blackish. Stigma about 3.1 as long as deep.

Mesopleurum with a wide foveolate band behind prepectal carina. Base

of tergite 1 with a relatively large oblique truncation. Palearctic

ROGION. ea a ee 4, striolatus Cameron (p. 6)

Trochanters and middle femur pale fulvous. Stigma about 2.2 as long as

deep. Mesopleurum with a narrow foveolate band behind prepectal

carina. Base of tergite 1 with a small obliquely truncate area... 4

4. Nervulus basad of basal vein by 0.5 its length. First tergite about 3.8 as

fone 26 wide, Kenya. . 2 oa ae ee 2. elgoni Risbec (p. 4)

Nervulus opposite basal vein or basad of it by up to 0.23 its length. First

tergite about 3.0 as long as wide. Holarctic and Ethiopian regions.

3. ruficornis Foerster (p. 4)

0. Face, vertex, and median lobe of mesoscutum very coarsely rugoso-

pierce,” PULODe. 403 6 ees ee 7. nigripes Foerster (p. 9)

Face, vertex, and median lobe of mesoscutum with moderately small

POUT Oe ee iy ae a eh ge re Gira ign IGE gS HG ae Pe aera as ator, AS ss 6

6. Trochanters reddish brown. Front and middle femora reddish brown.

Bastern Australia . co. ere Se Ue es 5. australiensis New (p. 7)

Trochanters black or blackish. Front and middle femora fuscous near

base, apically fulvous. Holarctic Region.

6. anomalipes Panzer (p. 7)

1. Helorus brethesi Ogloblin

Figure 2 (side view)

Helorus Brethesi Ogloblin, 1928. Rev. Soc. Ent. Argentina 1928: 77. des.,

figs. Type: sex?, Loreto, Misiones, Argentina (La Plata). Type not

seen. Description adequate for identification.

Front wing 2.6 to 3.0 mm. long. First flagellar segment about 5.7 as

long as wide in male, about 6.0 as long as wide infemale. Second flagellar

segment about 3.5 as long as wide in male, about 4.7 as long as wide in fe-

4 Contrib. Amer. Ent. Inst., vol. 15, no. 2, 1977

male. Face and mesoscutum polished, with very fine faint punctures. Lateral

part of pronotal collar rugose and rounded, surmounted by an irregular and

incomplete longitudinal carina. (In all other species of Helorus the lateral

part of pronotal collar is surmounted by a strong longitudinal carina.) Con-.

cave part of side of thorax smooth or with a few irregular wrinkles (in all

other species of Helorus with strong, horizontal, parallel wrinkles). Meso-

pleurum with a single row of moderately small foveae behind prepectal carina.

Scutellum smooth, polished, with sparse hairs that arise from almost invisible

punctures. Stigma about 3.1 as long as deep. Nervulus basad of basal vein by

about 0.3 its length. First tergite about 3.6 as long as wide, tapered to apex

from widest point at basal 0.3, its baso-dorsal slope an almost vertical trunca-

tion and surrounded by a rim. Basal 0.3+ and subapical 0.2+ of surface of

first tergite with longitudinal wrinkling and sparse shallow punctures, the rest

of its surface almost smooth.

Black. Mouth parts yellowish white, the apical part of mandible dark.

Antenna fulvous or stramineous at base, gradually darkening to dark brown at

apex. Hind margin and hind corner of pronotum, usually upper part of prono-

tum, much or all of mesoscutum, and axillary areas ferruginous, sometimes

the thorax more extensively ferruginous with only the under side and most of

propodeum blackish. Tegula whitish. Legs stramineous or pale fulvous, the

hind coxa and hind femur except its base and apex medium brown. Basal 0.25

of hind tibia whitish. Wings hyaline.

Specimens: ©, Horco Molle (near Tucumdn), Argentina, May 6 to 14, 1966,

Lionel Stange (Townes). -2°, Horco Molle (near Tucuman), Argentina, Decem-

ber 10 to 23, 1967, C. C. Porter (Cambridge). “, °, Horco Molle (near

Tucumdn), Argentina, May 15-21 and June 12-25, 1966, Lionel Stange (Tucu-

man). “, 9, Tucumdan, Argentina, December 1968 and December 1972, J.

Foerster and F. M. Oliveira (Ottawa). ©“, Jatai, Goias, Brazil, November

1972, F. M. Oliveira (Ottawa). ¢, Motozinha, 3,000 ft., Chiapas, Mexico,

June 6, 1969, W. R. M. Mason (Ottawa). ¢, Yerba Buena, 6,500 ft., 20

miles north of Bochil, Chiapas, Mexico, June 9 to 24, 1969 (Ottawa).

This species ranges from southern Mexico to northern Argentina.

2. Helorus elgoni Risbec

Helorus elgoni Risbec, 1950. Trav. Lab. d'Ent. Sect. Soudanais Rech.

Agron. 2: 514. ¢. des., fig. Type: ¢, Kenya: Elgon Sawmill on

Mt. Elgon (Paris). Type studied in 1975.

Similar to H. ruficornis, from which it differs in having nervulus basad of

basal vein by 0.5 its length; first tergite about 25% longer than in ruficornis;

flagellum a little longer than in ruficornis; and front and middle tarsus entirely

fulvous.

Specimen: Known only from the type, a from Mt. Elgon in Kenya.

3. Helorus ruficornis Foerster

Figure 3 (side view)

Helorus ruficornis Foerster, 1856. Hymenopterologische Studien, p. 143. :.

key, des. Type: °, Germany: Aachen (Vienna). Type studied in 1975.

Townes: A revision of the Heloridae 5)

Helorus coruscus Haliday, 1857. Nat. Hist. Review 4 (proc. ): 168. ©. des.

Types: ©, England and Germany: Westphalia (Dublin). Study of types

reported by Pschorn-Walcher, 1955. New synonym.

Helorus flavipes Kieffer, 1907. In André? Species des hymenopteres d'Europe

et d'Algérie... 10: 267. &. Lectotype: % (labeled by Townes, 1975

and hereby designated), France: Chanville (Paris). Lectotype studied in

1975.

Front wing 2.8 to 3.7 mm. long. First flagellar segment about 2.8 as long

as wide in male, about 4.2 as long as wide in female. Second flagellar seg-

ment about 2.7 as long as wide male, about 3.8 as long as wide in female.

Face and mesoscutum polished, with very small punctures that are separated

by about 2.5 their diameter on face, by about 3.5 their diameter on mesocutum.

Mesopleurum polished, smooth, its punctures very small, behind the prepectal

carina with a narrow band of foveae, the band one fovea wide. Scutellum pol-

ished, smooth, its punctures very small, and moderately sparse. Stigma about

2.2 as long as deep. Nervulus opposite basal vein or basad by as much as 0.23

its length. First tergite about 3.0 as long as wide, its baso-dorsal truncation

small, about 50° from the vertical, not bordered by a ridge, the surface of

tergite coarsely punctato-rugose with a longitudinal bias, or with strong irregu-

larly longitudinal wrinkling. Punctures on female pygidium small, deep, sep-

arated from each other by about 0.7 their diameter.

Black. Mouth parts whitish, the mandibular teeth dark brown. Antenna

light brown to dark brown, rarely fuscous. Tegula pale fulvous. Front and

middle coxae brown, paler apically. Front and middle legs beyond coxae ful-

vous. Hind coxa blackish. Hind trochanters fulvous. Hind femur brown,

paler basally. Fifth segment of front and middle tarsi brown. Hind tibia and

tarsus light brown. Wings hyaline.

Palearctic specimens: 6°, 3, Skane, Sweden, June and July, 1969, Bo

Svenson (Townes). ©, Dorking, Surrey, England, July 11, 1964, H. and M.

Townes (Townes). 20", New Forest, England, July 12, 1964, H. and M. Townes

(Townes). 6, 48, Leicester, England, June, July, and August, 1972 and

1973, Jennifer Owen (Townes). §, reared from Chrysopa prasina, Roserntal,

Bale- Liestal, Switzerland, June 13, 1935, W. Elgin (London). 9, Seegraeben,

Switzerland, June 7, 1951, V. Delucchi (Ottawa). ee Pizzighettone, Italy,

June 11, and May 31 to June 7, 1973, Franco Frilli (Townes). ?, near Linz,

Austria, July 6, 1947, H. Priesner (Washington). ¢, Igls, 900 m., Tirol,

Austria, Aug. 31, 1953, J. R. Vockeroth (Ottawa). 92, Rawalpindi, Pakistan,

June 16, 1958, Murree (Washington). ?, reared from chrysopid on Abies

pindrow, Pakistan, June 6, 1959, Murree, (London). “, Pulchauki, 6, 800 ft.,

Katmandu, Nepal, July 21, 1967 (Ottawa).

Ethiopian specimens: oC, Grahamstown, South Africa, Dec. 1971, Fred

Gess (Townes). 2, Kloof, i, 000 ft., Natal, South Africa, Sept. 1926, R. E.

Twiner (London).

Nearctic specimens: 132, Dryden, Me., Aug. 4, 6, 7, and 8, 1971, H.

and M. Townes (Townes). 2, Pinkham Notch, N. H. . Aug. a3. 1951, e. M:.,

and D. Townes (Townes). &, Mt. Washington, 5, 100 “ft, N. H. . Aug. 25,

1951, H. and D. Townes (Townes): oe Aylmer West, Ont. tuly 20 to 94

and Aug 8 to 14, 1972 (Ottawa). &, Point Pelee, Ont., Oct. : 29, 1968 (Ottawa).

tac... 32, Cleveland, S. C., May 23, June 1, 7, 13, and 29, and July 4, 5, 7,

15, and 19, all in 1971, G. “Townes tamile (Uemee).

Pschorn- Walcher records specimens from Finland, Sweden, Ireland,

Czechoslovakia, Austria, Switzerland, and Tirol. He records Chrysopa

6 Contrib: Amer. Ent. Inst., vol. 15, no: 2, 1977

ventralis and Chrysopa prasina as hosts.

This species is widespread in Eurasia, eastern part of South Africa, and

eastern North America. In North America it is a recent introduction, as the

earliest collection there was in 1951. Probably its occurrence in Africa is

also due to introduction.

4. Helorus striolatus Cameron :

Figure 4 (side view)

Helorus striolatus Cameron, 1906. Jour. Bombay Nat. Hist. Soc. 7: 98. ¢,

des. Type: © (lacking head), Pakistan: [ Quetta] in Baluchistan

(London). Type studied in 1975.

Helorus meridionalis Pschorn-Walcher, 1955. Mitt.Schweizerischen Ent.

Gesell. 28: 247. ©, 2. key, des., figs. Type: 2 (reared from Chryso-

pa flavifrons), Italy: near Bologna (Geneva ?; not in Vienna in 1975).

Type not seen but description adequate. New synonym.

Front wing 2.6 to 3.4 mm. long. First flagellar segment about 2.9 as

long as wide in male, about 3.4 as long as wide in female. Second flagellar

segment about 2.8 as long as wide in male, about 3.2 as long as wide in fe-

male. Face and mesoscutum polished, with very small punctures that are

separated by about 2.5 their diameter on face, by about 3.5 their diameter on

mesoscutum. Mesopleurum polished, with moderately small punctures on its

lower 0.3, the rest impunctate and variously smooth or weakly wrinkled, be-

hind the prepectal carina with a wide, mostly single row of strong foveae.

Scutellum polished, smooth except for moderately sparse very small punctures.

Stigma about 3.1 as long as deep. Nervulus opposite basal vein or basad by

as much as the width of nervulus. First tergite about 3.0 as long as wide, its

baso-dorsal truncation of moderate size, about 40° from the vertical, and

bordered by a rim, the surface of tergite with strong wrinkles that have a

longitudinal bias, its setiferous punctures very small and hardly visible.

Punctures on female pygidium moderately small, deep, separated by about

0.7 their diameter.

Black. Mouth parts whitish to medium brown. Tegula, apical 0.65+ of

front femur, apical 0.1+ of middle and hind femora, and front and middle tibi-

ae and tarsi light brownish fulvous. Hind tibia and tarsus brown. Wings sub-

hyaline.

Specimens: ©, Helsinki, Finland, Aug. 7, 1962, O. Ranin (Townes).

o, 9, Helsinki, Finland, Aug. 14, 1968, H. and M. Townes (Townes). 4°,

Eberswalde, East Germany, July 5 and 6, 1964, H. K. Townes (Townes). 2,

"Kopl. Aschachtal", Austria, Aug. 29, 1962, H. Priesner (Washington). 2,

Marchtrenk, Austria, July 15, 1938, H. Priesner (Washington). ¢, Naturno,

Trentino, Italy, July 18, 1958, David Townes (Ottawa). 30°, 22, reared from

Chrysopa, in forest, ''Sh. Neumark", Israel (Washington).

Pschorn-Walcher records this from Italy, Hungary, and Finland, and as

having been reared from Chrysopa flavifrons and Chrysopa septempunctata.

Kozlov has recorded it from Mongolia (1972. Insects of Mongolia 1: 646).

This species is widely distributed in Eurasia.

Townes: A revision of the Heloridae 7

5. Helorus australiensis New

Helorus australiensis New, 1975. Jour. Australian Ent. Soc. 14:15. o%, &.

des., figs. Type: 2, reared from Chrysopa cocoon, Australia: Bun-

doora in Victoria (Canberra). Australia: Murphy's Creek near Helidon in

southwestern Queensland. Hosts: Chrysopa ramburi, Chrysopa near

dispar, and Chrysopa near signata. Types not seen.

Front wing 2.5 to 2.7mm. long. First flagellar segment of female about

3.2 as long as wide. Second flagellar segment of female about 3.4 as long as

wide. Face with dense white pubescence. Mesoscutum with fine punctures

and hairs that are less dense than on face. Scutellum mostly smooth. Stigma

about 3.2 as long as deep. Nervulus opposite basal vein. First tergite slightly

furrowed longitudinally, about 2.3 as long as wide.

Black. Scape and pedicel reddish. Flagellar segments 1-3 usually reddish,

sometimes darker. Mandible reddish toward apex. Tegula reddish. Coxae

mostly black, the rest of legs reddish brown but hind femur sometimes dark-

ened.

This species has not been seen. It is placed in the key and redescribed

from data in the original description.

Specimens: Known only from the types (7%, 11°) from eastern Australia.

6. Helorus anomalipes Panzer

Figures 5 and 6 (side view)

Sphex anomalipes Panzer, 1798. Faunae insectorum germaniae . . . heft 52,

plate 23. Type: 2?, Germany (lost). Description and figure sufficient

for identification.

Helorus ater Latreille, 1802. Histoire naturelle . . . des crustacés et des

insectes. 3: 309. sex? Type: France? (lost). Description by mention

of name under the new genus Helorus, but without distinguishing specific

characters. First revisor was Latreille, 1809, who synonymized his ater

with anomalipes.

Copelus paradoxus Provancher, 1881. Nat. Canad. 12: 207. 2. des., fig.

Lectotype: & (designated by Gahan and Rohwer, 1916), Quebec: Cap

Rouge (Sainte Foy). Lectotype and paratype studied in 1975. New

synonym.

?Helorus anomalipes var. bifoveolata Gregor, 1938. Casopis Cesk. Spol. Ent.

34:15. o. des. Type: %, Czechoslovakia: Novy Ji@in (Prague?). Type

not seen.

?Helorus coruscus nigrotibia Hellen, 1941. Notulae Ent. 21: 30. Types: Fin-

land: localities and sexes not specified (location of types unknown).

Types not seen.

Front wing 3.1 to 3.8 mm. long. First flagellar segment about 2.9 as long

as wide in male, about 3.9 as long as wide infemale. Second flagellar seg-

ment about 2.4 as long as wide in male, about 2.8 as long as wide in female.

Face and mesoscutum polished, with medium sized or small deep punctures

that are separated by about 0.7 their diameter on face, by about 1.5 to 2.5

their diameter on mesoscutum. Mesopleurum with moderate sized punctures

that usually are crowded, with some rugosity, smoother in specular area.

8 Contrib. Amer. Ent. Inst., vol. 15, no. 2, 1977

Behind the prepectal carina a band of coarse foveae, the band about 2 foveae

wide. Scutellum foveolate or rugose near apical margin, or with more exten-

sive rugosity that covers its hind half or more, sometimes the rugosity rudi-

mentary and the scutellum almost smooth. Non-rugose part of scutellum with

moderately sparse small weak punctures. Stigma about 3.6 as long as deep.

Nervulus opposite basal vein or basad or distad by as much as 1.5 the width of

nervulus. First tergite about 1.8 as long as wide, its baso-dorsal truncation

large, about 45° from the vertical, bordered by a strong rim. First tergite

in dorsal view tapered to apex from a little behind its basal truncation, its

surface with a strong rugose reticulation and with strong longitudinal wrinkles

or carinae and not distinctly punctate, or in some European specimens its

surface smoother and distinctly punctate. Punctures on female pygidium deep,

moderately small to medium sized.

There is much variation in the sculpture of the first abdominal segment,

sculpture of the scutellum, size of the punctures on lower part of mesopleurum,

and size of punctures on the pygidium. Figure 5 shows a specimen with nor-

mal scupture; figure 6 shows one with the first tergite unusually smooth.

Specimens similar to figure 6 are frequently found in Europe but never in

North America.

Black. Mandible, tegula, and tibiae fulvous brown. Front and middle fe-

mora fuscous or black basally, fulvous brown apically, the front femur less

extensively fuscous than middle femur, sometimes the front femur entirely

fulvous brown. Tarsi brown.

Specimens from Europe: 25, 302 from Austria (Winden and near Linz);

Czechoslovakia (Cernovice); England (Aldeburgh in Suffolk, Cambridge,

Dartmoor, Faversham in Kent, Horsell Common in Surrey, Mill Hill, Notting-

ham, Tunbridge Wells, and Windsor Forest); France (Landes on Cap Breton):

East Germany (Beekow); Germany (Eisen and Mainz); ie ry; Spain (Grenada);

Sweden (Degaberga in Skane and Loderup); and Turkey nkara).

~ Specimens from North America: 850, 1002 from Alberta (Jumping Pound

20 miles west of Calgary, Lancaster Park, Lethbridge, and Rockyford);

British Columbia (Agassiz, Atlin, Garibaldi Park at 3, 400 to 4, 800 ft., Hudson

Hope, Robson, Salmon Arm, and Terrace); California (Topaz Lake); Colorado

(Boulder, Estes Park, and Morley); District of Columbia (Washington): Georgia

(Ringgold); Iowa (Sioux City); Kansas (Lawrence); Kentucky (Golden Pond);

Louisiana (Tallulah); Maryland (Laurel, Takoma Park, and Plummers Is. ).

Massachusetts (Arlington, North Saugus, and Holliston): Michigan (Ann Arbor,

Antrim Co., Bay City, Benzie Co., Crystal Falls, East Lansing, Galien, Glen

Haven, Grand Traverse, Gull Lake Biological Station in Kalamazoo Co., Hart,

Hickory Corners, Iron River, Ontonagan Co., Roscommon Co., St. Charles,

and Wexford Co. ); Manitoba (Brandon and Riding Mt. National Park near Nor-

gate); Minnesota (Cloquet and Eaglenest); Missouri (Webster Groves); Nebraska

(Halsey); Nevada (Wine Cup Ranch in Elko Co.); New Brunswick (Fredericton

and Nerepis); New York (Bemus Point, Farmingdale, Northwest, and River-

head); North Carolina (Hickory); North Dakota (McHenry Co.); Ontario (Ban-

croft, Belleville, Chatterton, North Bay, Ottawa, St. Davids, Simcoe, and

Toronto): Pennsylvania (North East and Inglenook); Prince Edward Island

(Atherton); Quebec (Aylmer, Cross Point on the Gaspé Peninsula, Forestville,

Hemmingford, and Hull); Rhode Island (Westerly); Saskatchewan (Dunblane,

Minton, and Saskatoon); South Carolina (Greenville); Virginia (Charlottesville

and Winchester); Washington (Yakima); Wisconsin (Lake ake Madeline); Wyoming

(6 miles northwest of Newcastle); and Yukon Territory (mile 2 on the Dawson

Road and Whitehorse). Collection dates are from early summer to mid fall,

Townes: A revision of the Heloridae 9

with the majority in July and August. There are two reared specimens, one

from Chrysopa sp. and one from ''Chrysopa sp. (probably rufilabris)".

There is a good account of the biology of this species by Clancy (1946.

Univ. California Pub. Ent. 7: 407-418), who reared it from Chrysopa majus-

cula. Koslov has recorded the species from Mongolia (1972. Insects of Mon-

golia 1: 645).

This species is widespread in Europe and in North America. It occurs over

the entire extent of the United States and southern Canada but is rare or absent

in arid habitats. It is uncertain whether it is native to both Europe and North

America or was introduced from one to the other.

7. Helorus nigripes Foerster

Figure 7 (side view)

Helorus nigripes Foerster, 1856. Hymenopterologische Studien, p. 143. ©.

key, des. Type: “, Germany: Aachen (lost, not found in Vienna in

1975). Foerster's statement ''Mesonotum ganz grob runzlig'’ makes

the specific identity clear. _

Helorus rugosus Thomson, 1858. Ofvers. Svenska Vet.-Akad. Forh. 15: 380.

des. Type: 2, Sweden: Skane (lost, not found in Stockholm in 1975).

Description sufficient for identification.

Front wing 3.7 to 4.3 mm. long. Face and mesoscutum with strong, very

coarse punctato-rugosity, but the sublateral 0.2+ of mesoscutum nearly smooth.

First flagellar segment about 3.3 as long as wide in male, about 3.7 as long

as wide infemale. Second flagellar segment about 2.5 as long as wide in

male, about 3.2 as long as wide in female. Mesopleurum very coarsely and

strongly rugose except for a large specular area. Scutellum coarsely,

strongly punctato-rugose. Stigma about 3.3 as long as deep. Nervulus oppo-

site basal vein or basad of it by as much as 2.0 the width of nervulus, or

sometimes a little distad of basal vein. First tergite about 1.85 as long as

wide, its baso-dorsal truncation large, about 15° from the vertical and bor-

dered by a projecting rim. General surface of tergite 2 strongly and coarsely

reticulate-rugose, with a median dorsal carina or wrinkle and traces of other

longitudinal wrinkles. Punctures on female pygidium small and deep, separ-

ated by about 0.6 their diameter.

Black. Mandible dark brown to blackish. Palpi brown. Flagellum black,

brownish below. Tegula black. Legs black or blackish brown, the front leg

with tibia, tarsus, and apical 0.3+ of femur brown. Wings subhyaline to faintly

infuscate.

Specimens: 60, 8? from Austria (Winden); England (Leicester); East

Germany (Eberswalde); and France (Hyeres).

Seven specimens from Hyéres, France, were reared from Chrysopa

cocoons.

Pschorn-Walcher (1955. Mitt. Schweizerischen Ent. Gesell. 28: 247)

records 16 specimens of this species from Sweden, Austria, Czechoslovakia,

and Hungary. One of these specimens was reared from Chrysopa nigricortala

and one from Chrysopa sp.

9. Helorus bréthesi

Helorus ruficornis

4, Helorus striolatus Co

Helorus anomalipes

Helorus nigripes

12 Contrib. Amer. Ent. Inst., vol. 15, no. 2, 1977

INDEX TO HELORIDAE

* An asterisk indicates a synonym, misspelling, or misidentification.

anomalipes (Helorus, Sphex) 2 (fig.), 3 (key), 7, 11 (figs. )

*ater (Helorus) 2, 7

australiensis (Helorus) 3 (key), 7

*bifoveolata (Helorus anomalipes) 7

bréthesi (Helorus) 3 (key), 10 (fig. )

*Copelus 2

*coruscus (Helorus) 1, 5

elgoni (Helorus) 3 (key), 4

*flavipes (Helorus) 5

*meridionalis (Helorus) 1, 6

nigripes (Helorus) 1, 3 (key), 9, 11 (fig.)

*nigrotibia (Helorus coruscus) 7

*paradoxus (Copelus) 2, 7

ruficornis (Helorus) 1, 3 (key), 5, 10 (fig. )

*rugosus (Helorus) 1, 9 a

striolatus (Helorus) 1, 3 (key), 6, 10 (fig. )

Contributions

of the

American Entomological Institute

Volume 15, Number 3, 1977

A REVISION OF THE CENTRAL AND SOUTH

AMERICAN NOTOXUS AND DESCRIPTION OF A

NEW GENUS, PLESIONOTOXUS

(Coleoptera: Anthicidae)

Donald S. Chandler

TABLE OF CONTENTS

1d PRODUC TION | gc se ee a ee

Acknowledements 2 26603 Wow a ee

TAXONOMIC: HISTORY 2. co oo as in ee a ee ea ee

TOTO Y ia 5 ck a ea ae ae ee a ee

UOtletChs Pic rs a a a De et A

HOY COT URVOIICN) as i ee ee

Wouneliet UE) ie ee eee

Mens (PC ee a a

GENUS NOTOXUS PABRICIUS 106k si Viele a ee es a ee ee

“a axonomic: GHATaCtere. 6055 a a ae ee ae ae

Key to the Central and South American Notoxus ............

FUCO PONS IS PT OUG i he Gs eek a ee ee eee

Bebionsis CRAIG i

LPORCHS IS BOBO CAA a ee hee a ee er ee

SCS Ts BD Ab ls Ae a a ee

Wa ROY oo ce eo a ee eee

COLCA TUS FOP aig ou i hw i ee I ee

EAI POND isin ich in uk esa ee eck oe gee nee ey ee

Pie ON OU a a ee ee aw

EOE 6A i i sa es eA a ee ea ee

PU ONMOEN SIS Be BO act ke a me hie eee ae Ee

PU AER i Dh a i ak GG a a Be oe ae

OH Bi ls a a a

BEL OO ag eee ed he Wale ha lates) igi he ele Le ace Om

PIG CO ae are a a ae ol ew a we ae a i eee

eR ag ek Wk i ea AO Oe ee ss

Be ee ay big ie ee ee ee I te ee

Ver aHe A MOAI os i ei ee a we a a ees

CEUMTIATH® TRING Le Whe ea we a WS

ele Ee lig, A a Rn Leta. areGuMta Ny iat sem acpi i a. Serer nse iar anan ay

cumanensis LaFerte

fon fg UR TG age ME Sean em eA i i a CaN AE Meme erireyn Caria Pay a) aac Tang

CEES ELE ADAG) eh ae eee Ten RNA fe ak ng Mae ORAM m ee P ELA Mri as Nien Me Un ic?

DEP UCS CAOV OA i ogi acute dda woe Biante ial ae

SETI AEG ROM ins itt aie GN elim ae ibs aN BN A aC a SOR A al ea

murinepennis KLBCONe), ci wa We a ee

VUTEC OILS SGT UOD 55h Es i inl ML ae Ae aan a ed aad

DOVUUTINMS FOV ie a eR GE MES lO I ese iia alee

PUD PEG 2TOU i ON ee Oa Ung ieee eR a Oe ee teat

WUD ETUS TORI i OE Re RE OE A EI ee eg ae

BEUSTFUSR BOBO SOO Ga EE AE Se A a

POSTICLUS . Bi BB Ee aay eam eL IL ah

OVLCRTOIES | Ri Bi A ME Oe CR at tata dal Orel g

lil

OMDMUMDUODWDAWAAADRWNHH FH

AuICa ie ev0W0. . 2. a. a eS.

Cpr es eC Oue ee e s

pie eu s Leone a ee

Wee eas CHAINDION. 6. a ee

Pe ONO a a ee

oe, ke

Pues, 6. ee a a ee Py A a

Pome. 1 Bb. ea

WIOHOICO TOR + SOND 6 ee a ee

WHCTO? BOWS a a

Be eS OPO ee a ee

SCP SUS DECOM a a a

DO AB a a ee a a a i

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INTRODUCTION

Notoxus is one of the last large anthicid genera of the New World which is

badly in need of revision. The North American (the United States and Canada)

Species have been adequately collected and described, with only a few species

remaining still unnamed. The species from Central America (Mexico through

Panama) are less well known. Recent collecting in Mexico has revealed that

the Central Plateau is the center of species diversity for the New World.

This revision of the Central and South American species includes some

twenty-seven previously named species and twenty which are described as new.

The relationships within the genus are primarily derived from a numerical

analysis of all the New World species (a total of seventy-two species). Most of

the South American species which had been placed in Notoxus have been re-

moved to Mecynotarsus (Chandler 1977) or are placed in a new genus, Plesio-

notoxus (p. 8).

Acknowledgments

Many individuals deserve my deepest thanks for their help in this study.

The following list indicates those curators of collections and private collectors

who cooperated with the loan of specimens. Most of the abbreviations of the

collections are from Arnett & Samuelson (1969). The abbreviations are used

to indicate the present location of types or of the specimens studied for the re-

vision.

Public collections:

AMNH: American Museum of Natural History; Lee H. Herman, Jr.

BMNH: British Museum (Natural History); C. M. F. von Hayek.

CASC: California Academy of Sciences; H. B. Leech, P. H. Arnaud,

Jr., D. H. Kavanaugh.

CDAE: California Department of Agriculture; F. G. Andrews.

CISC: University of California, Berkeley; J. T. Doyen.

CNCI: Canadian National Collection; E. C. Becker.

CUIC: Cornell University; L. L. Pechuman.

FMNH: Field Museum of Natural History; H. S. Dybas.

LACM: Los Angeles County Museum of Natural History; J. P. Donahue.

MCZC: Museum of Comparative Zoology; J. F. Lawrence,

oe. C. Sent,

MNHN: Museum National d'Histoire Naturelle, Paris.

MSUC: Michigan State University, R. L. Fischer.

OSUC: Ohio State University; C. A. Triplehorn.

current address: Rt. 1, Box 275-19, Safford, Arizona 85546

27 his paper is a portion of the dissertation submitted to the Ohio State Univer-

sity in partial fulfillment of the requirements of the Doctor of Philosophy

Degree at that institution.

2 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

TAMU: Texas A & M University; H. R. Burke, S. J. Merritt.

TTCC: Texas Tech University; D. E. Foster.

UCDC: University of California, Davis; R. O. Schuster.

UMMC: University of Michigan; R. D. Alexander.

USNM: United States National Museum; T. J. Spilman.

Private collections:

DSC: Donald 8. Chandler, Safford, Arizona.

FGW: Floyd G. Werner, Tucson, Arizona.

HAHC: Henry Howden, Ottawa, Ontario, Canada.

KSC: Karl Stephan, Tucson, Arizona (now placed in the Florida

Department of Agriculture, Gainesville, Florida).

MGF: Gerhard Frey, Tutzing bei Munchen, West Germany.

C. M. F. von Hayek of the British Museum (Natural History) helped great-

ly by lending several of the Champion types. J. C. van Hille of Rhodes Uni-

versity, Republic of South Africa, graciously sent specimens of Pseudonotoxus

Pic for comparison with Plesionotoxus. A portion of a trip to Mexico was

funded by a Sigma Xi Grant-in-Aid in 1974. Dr. Alejandro Ortega of C.I.M.

M.Y.T., Texcoco, Mexico, graciously arranged an insect collecting permit

for the summer of 1975 in Mexico. Dr. D. E. Johnston deserves my thanks

for his guidance of the numerical analysis, Drs. D. J. Hornand D. J. Borror

for reading this paper and offering numerous suggestions, and Dr. C. A.

Triplehorn for his help and encouragement in the preparation of this paper.

Dr. F. G. Werner, University of Arizona, offered suggestions on the general

organization and the key to species. Lastly, I would like to thank my wife,

Christine A. Janus-Chandler, for her help and encouragement in the prepara-

tion of this paper.

TAXONOMIC HISTORY

Fabricius (1801) described Notoxus monodon from "Virginia" for the first

record of the genus from the New World. Say (1817), LeConte (1824, 1847)

and Hentz (1827) added four more species from the eastern United States.

LaFerte-Senectere (1848) treated the family for the world and described four

new species from the United States, two from Mexico and two from South

America.

In the period following LaFerte's work, many of the United States species

were described (LeConte 1851, 1852, 1876; Horn 1884, 1892; Casey 1895). No

new species have been added since the works of Fall (1902, 1916, 1932) and

Blaisdell (1929, 1936). Recently Chandler and Hagen (1977) listed a number of

new synonymies for North America.

In contrast the Central American fauna was largely ignored until near the

end of the nineteenth century. Two species were described from the West

Indies (Chevrolat 1877; Quedenfeldt 1886), and fourteen new species were

added to the Central American fauna by Champion (1890) in his revision of the

Anthicidae for the Biologia Centrali-Americana. The remainder of the neo-

tropical species were added by Pic (1901a, 1901b, 1904, 1912, 1913, 1914a,

1914b, 1916, 1918) and Heberdey (1936).

Champion's work (1890) is still useful as he described the male characters

of the species and had excellent illustrations. Pic's descriptions are almost

completely useless since he seldom mentions any characters of use in separ-

ating species. Many of Pic's species have been removed to Mecynotarsus

(Chandler 1977) or to Plesionotoxus, new genus. Of the four Pic names re-

Chandler: Notoxus of Central and South America 3

tained in Nofoxus, one is a senior synonym, two are junior synonyms, and the

last is placed as Incertae sedis,

The last paper dealing with the Central American Notoxus was Pallister's

(1955) identification of the Anthicidae collected on the Central Plateau of

Mexico. This work is of some importance in that several species from the

United States were recorded from Mexico for the first time.

BIOLOGY

Most of the available biological information on Notoxus is restricted to

the types of habitat in which the adults have been collected. No life histories

have been described for any of the species. The only larvae figured have been

of N. monoceros (Boving & Craighead 1930), an unknown species of Notoxus

(Peterson 1951) and of N. monodon (Cuthbert 1967). The larva described by

Peterson was collected in soil from a peach orchard, while the monodon larvae

(misidentified as N. calcaratus in Cuthbert) apparently will bore into the roots

of sweetpotatoes. The larvae have a well-developed mola on the mandible and

probably feed on plant material. The pupa of monodon is also figured in Cuth-

bert.

The adults are known to be associated with vegetation and flowers (Blatchley

1910). Ihave collected the majority of my specimens by beating vegetation or

examining the ground beneath low vegetation. Other collection methods are

based on the adults' attraction to white or ultra-violet light and to cantharidin

or meloid beetles. Almost all of the New World species have fully developed

wings, with both sexes of obesulus and females of other species in the sparsus-

group being exceptions. The majority of the winged species have been attracted

to white, arc or ultra-violet lights. One of the more recent methods of collec-

ting Notoxus is based on their attraction to cantharidin, which is found in

meloid beetles. Abdullah (1964) lists the Old World Anthicidae which have

been attracted to cantharidin and dead meloids, while van Hille (1954) used

cantharidin as a collecting bait. Chandler (1976) used both cantharidin and

meloid beetles as baits and found that both were very effective in attracting

various species of Notoxus. Only two species, N. eximius and N. opacus,

were known to be near these baits and not attracted. Blaisdell (1936) mentions

that he used ''chemical" traps to collect several species of California Notoxus,

but the chemical used was not indicated.

The adults are known to eat a wide variety of foods. Abdullah (1969) ob-

served N. monoceros feeding on bread, sugar water, minced beef, freshly cut

grasshoppers and a banana skin. Hinton (1945) notes that N. monodon was

found on some apples sent from North America to Germany. Records in this

revision indicate that N. apicalis is attracted to freshly harvested crops.

Essig (1926) mentions that the ''fruit Notoxus", N. constrictus (his figure of

this species is actually N. calcaratus), was common in orchards feeding on

damaged or drying fruit. Also mentioned is that N. calcaratus acts similarly

and that N. monodon (= desertus) was collected boring into twigs of apple trees

in California. This latter record is almost certainly incorrect as the beetles

were probably just resting in these holes. In Colorado Gill (1913) noted that

N. monodon (= desertus) would feed on the pupae of the fruittree leafroller,

Archips argyrospila Walker. Notoxus often have been observed feeding on the

dead insects which accumulate at lights. Loan (1972) mentions that N. anchora

was common in and feeding on alfalfa in Ontario, although it apparently is not

considered a pest.

4 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

There have been two instances where the biological control value of N.

calcavatus has been examined. Butler (1966) in Arizona noted that this species

would feed on the eggs of the cotton bollworm in the field. Orphanides ef al

(1971) found that calcavatus in California would feed on the eggs and first instar

larvae of the pink bollworm in the laboratory. Both studies noted that although

the adults did not eat many eggs or larvae, this species could be very common

in the field and thus may have a noticable effect on a pest population.

The use of the pronotal horn apparently has not been previously recorded.

Specimens of several species collected in New Mexico and Arizona were placed

in a vial containing sand, water and food. As midday approached the beetles

would become very active even though the vials had been placed in some shade.

Many individuals were observed to move about until encountering a depression

or crack inthe sand. The head and pronotum were then worked alternately up

and down as the beetles burrowed down into the depression forming a tunnel

which could be more than an inch long. Species of the genus are often found

in or near sandy areas. Another Notoxine genus, Mecynotarsus, is known to

be associated with dunes, and individuals will quickly burrow into loose sand

when disturbed using a similar motion of the head and pronotum.

There is one known case of parasitism of Notoxus. Loan (1972) noted that

a braconid wasp, Syrrhizus agilis (Cresson), was an internal parasite of adults

of N. anchora. The only mating behavior known was recorded by Abdullah

(1969) for N. monoceros.

ZOOGEOGRA PHY

Notoxus is recorded from all the continents except Antarctica, with very

few species being known from South America and Australia. The majority of

the species are recorded from three general areas: the southeastern quarter

of Africa, the eastern Mediterranean and central Mexico. These regions are

generally semiarid and have an important grassland component with various

interrupting mountain ranges. Species diversity is apparently highest at the

erassland-forest interface.

Only one group, the monoceros-group in Eurasia, has received a recent

thorough taxonomic treatment (Heberdey 1936). Recent faunal works have

been by Bonadona (1974) for France, Kaszab (1969) for central Europe and

van Hille for South Africa (1961). The relationships between the faunas of the

different continents have been unexplored and there have been few attempts to

define the genera.

The area under consideration in this study, Central and South America,

includes all of the Neotropical faunal province and the portion of the Nearctic

faunal province present in northern Mexico. The genus in the New World is

predominantly Nearctic with all of the species-groups (see p. 7 and Figure 69

for discussion of species-groups) having most of their members in Mexico or

the United States.

Two species-groups, the monoceros- and sparsus-groups, have the majori-

ty of their species in the western United States. Previously the monoceros-

group was known only from the Palearctic region. Five Nearctic species are

here placed in this group with four of the five species occurring in an area

extending from northern California to Washington. Most of the species in this

eroup have asymmetrical genitalia, which is a characteristic of the Nearctic

species, with a few of the Palearctic species possessing symmetrical genitalia.

The sparsus-group is almost completely restricted to the western coastal

Chandler: Notoxus of Central and South America 9)

states. This group possesses symmetrical genitalia. Both groups have in

common robust genitalia and lateral apical tubercles on the elytra of the male.

The form of the symmetrical genitalia in both groups is similar. The distri-

bution pattern of both groups, principally in the western and northern United

States, is a pattern common to many groups introduced to the New World from

Eurasia via the Bering Land Bridge. Taking into account the fact that the two

groups are more difficult to separate in the Old World, it appears likely that

both groups originated in Eurasia and were introduced across the Bering Land

Bridge which was periodically present during the late Pliocene (Gressitt 1974).

A re-examination of the Old World species in the monoceros-group is needed

to firmly delineate the group characteristics.

The remaining species-groups appear to have originated in the region of

central and southern Mexico. The greatest species diversity is found on the

southern half of the central plateau where there is an intermixing of the

mesquite-grassland and pine-oak forest zones (Leopold 1950). At least one

member of all the species-groups (except the montanus-group and excluding

the sparsus- and monoceros-groups) and most of the species which were not

placed in groups are found in this area. Many of the species-groups exhibit a

simple pattern in which one species is found on the eastern half of the Central

Plateau, another on the western half, and often a third species in the south-

western United States.

Rosen (1975) recently analyzed the patterns of animal distribution in the

Caribbean region. Notoxus species-groups appear to fit the North American-

Caribbean Track (distribution pattern) fairly well, even though Rosen had

applied it only to lowland groups. Groups placed in this pattern have Mexico

containing the greatest number of species and with a few species ranges ex-

tending into the southern United States, the most northern portion of South

America, and the Antilles. The monodon-group, which is primarily found in

the lowlands, exhibits this pattern perfectly with two species in the Greater

Antilles, two reaching northern South America, three in the United States and

four in Mexico.

Continental land bridges have been of variable importance in influencing

the distribution of Notoxus. The Bering Land Bridge was probably the means

of introduction of members of the monoceros- and sparsus-groups, while the

land bridge between Central and South America apparently has been of little

importance in Notoxus movement. Rosen (1975) believes that the Central

America region was mainly occupied by various islands from the Mesozoic

until the Pliocene when the first complete land bridge was formed. Two mem-

bers of the monodon-group are isolated in the Greater Antilles, but, using

Rosen's model of Caribbean continental drift, they probably were isolated on

these islands when they drifted east from their alignment near the North Ameri-

can Plate. Since only three species are present in South America, with two of

these being widespread through Central America, it appears that Notoxus could

not traverse this island chain and was probably introduced in or after the Plio-

cene when the Panama Land Bridge was formed.

Climate has probably been the greatest influence in determining the dis-

tribution of Notoxus. Perhaps the most important change occurred in the

middle and late Cenozoic when extensive mountain building began taking place

along the axis of the present-day Sierra Madre Occidental. Asa result of this

mountain formation rain shadows were formed in central Mexico and the Madro-

Tertiary Geoflora (Axelrod 1958) evolved rapidly. By the late Miocene this

flora extended from the current southern apex of the Central Plateau to the

northern edge of the Great Basin. This flora is typical of the semiarid condi-

6 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

tions which Notoxus favors.

Other climatic changes of lesser influence were the pluvials and inter-

pluvials of the Pleistocene, and the following Xerothermic Interval of the

Recent. All of these have undoubtedly influenced Notoxus distributions by

providing suitable climates for dispersal between mountain ranges, and then

following with drier periods which would then isolate populations in the different

major mountain ranges. The major separations apparently are between the

Sierra Madre Oriental, the area around the Mexican state of Puebla, the

Sierra Madre Occidental, and the southern apex of the Rocky Mountains in

New Mexico-Arizona.

A reconstruction of the past history of the New World species can be

derived from these events and current distributions. Notoxus probably evolved

from seashore species which moved inland and eventually became isolated from

the other groups of the Notoxinae. The genus is characterized by having few

seashore species. A portion of the genus was then isolated in the Mexico area

following the breakup of the supercontinents in the late Mesozoic. With the rise

of the Madro-Tertiary Geoflora, extensive areas of Mexico gradually became

drier and the conditions which Notoxus species appear to favor became wide-

spread. Notoxus populations expanded northward with the geoflora into the

United States.’ These populations were then confined to the margins of the

mountain ranges when the grassland-forest areas were reduced by the contin-

ued drying on the plateau. This led to some degree of isolation of populations

between the major mountain ranges, and eventually to speciation. Possibly the

ancestors from which the species-groups are derived were present at this time.

Many of the present species which are not placed in a group are found at

the southern apex of the Central Plateau, as are members of every group except

the montanus group. This area apparently is the most heterogeneous favorable

habitat for Notoxus.

With the onset of the Pleistocene pluvials, species could disperse between

mountain ranges and were then isolated in the following dry periods. With

this fragmentation the present species-groups appeared as populations which

were isolated during one of the dry periods following dispersal. This would

explain the distribution of many of the species-groups which have a different

Species on each half of the plateau and one or more species isolated in the

southwestern United States. With following moister periods several species

in a few of the groups have expanded their ranges and are now sympatric in

the Chihuahua-Durango area and the Mexico City region.

METHODS AND MATERIALS

Over 3,000 specimens of Notoxus from Central and South America were

studied. The specimens were separated into species by comparison of their

external appearance and genitalia. Names were assigned to these species fol-

lowing the examination of all the types of the species, except for the LaFerte,

Chevrolat, and Pic types in the Museum National d'Histoire Naturelle, Paris.

Fortunately, specimens compared to most of the LaFerte types were available

from the F. G. Werner collection, and the description or locality of most of the

Chevrolat and Pic types was sufficient to associate names with species.

Fach previously described species is represented by a general statement

on the variation and characters of the species and the measurements of a re-

described male. Types were not used to represent the species in the descrip-

tion since they were available for only short periods of time and were seldom

Chandler: Notoxus of Central and South America fi

in the best of condition. The redescribed males were considered to be typical

of the species and were generally collected near the middle of the species

range. Males were used since they have more unique characters than females.

Comparisons were made using a Leitz Wetzlar stereo microscope at 64X.

Male genitalia were prepared and examined following the procedure of Werner

(1964). All genitalia were placed in genitalia capsules containing glycerine

following study.

Measurements were taken using an ocular reticule and are entered in

millimeters in the species descriptions. Drawings of the elytra and male fifth

sternite were made using the ocular reticule at 64X. Drawings of the genitalia

were made by sketching the general outlines as projected by a Ken-a- Vision

slide projector and then filling in the fine details while observing the genitalia

with the Leitz.

The distribution of each species is mapped only for Central America, un-

less it is a new species. In this case the distributions are continued into the

United States as far as the map permits.

The classification used is based on a phenogram produced by a numerical

analysis of the New World species (Fig. 65, Chandler 1976a). The limits of

the species-groups were delineated through the patterns of groupings in the

phenogram. Several species were not placed in a species-group as they either

were not very similar to other species or the clusters produced contained a

heterogeneous assemblage of species.

The similarity measure used was correlation and the phenogram was con-

structed using UPGMA. The numerical analysis was performed on the IBM

350 at the Ohio State University Computer and Research Center using the NT-

SYS program package developed by F. J. Rohlf, J. Kishpaugh and D. Kirk.

NEW WORLD NOTOXINAE

The Notoxine anthicids are easily recognized by the protruding horn ex-

tending anteriorly over the head, and also by the lateral expansion of the mandi-

bles. Three of the current Notoxinae genera (Pseudonotoxus Pic, Hypaspistes

Waterhouse, and Leptoprion Krekich) are found only in the Old World. Both of

the Notoxine genera known from the New World (Mecynotarsus LaFerte, and

Notoxus Fabricius) are also found in the Old World. A new genus, Plesio-

notoxus, is proposed for a number of South American species which were

formerly placed in Notoxus.

The following characterization of the genus Notoxus is based on all of the

species available, which includes some twenty species from the Palearctic,

Oriental and Ethiopian regions. The genus Notoxus is here defined as those

Notoxinae with the following characters: erect tactile setae on the head out-

lining the horn when head is raised, mandibular explanation often with posteri-

or margin, eyes with short aciculate setae between the facets, setae over body

aciculate, not noticeably thickened or flattened; pronotum with distinct pit above

the cervical notches, these pits densely setate, antebasal transverse impres-

sion distinct; lateral mesosternal margins straight or slightly curved outwardly;

hind wings with 1A and 2A parallel, with cross vein at middle (Fig. 1); male

genitalia with distinct phallobase, parameres large, either free or fused near

base, penis or median lobe well developed (Fig. 2); the genitalia laterally

flattened in general appearance.

Plesionotoxus is here defined as those Notoxinae with the following charac-

ters: erect tactile setae on the head outlining the horn when head is raised,

8 Contrib. Amer. Ent. Inst., vol..15, no. 3, 1977

mandibular explanation with faint posterior margin, eyes without any setae

between facets; pronotum with nude pit above the cervical notches, anteriorly

directed setae near ventral margin of pronotum obscuring pits; horn with crest

indistinct, lateral carinae from horn base to the cervix anterior to the pronotal

pits, pronotum with antebasal transverse impression distinct; setae over body

aciculate; lateral mesosternal margins straight to slightly curved inwardly;

wing venation similar to Notoxus, with large sclerotized patch lateral to apex

of Cuy; male genitalia with phallobase and penis closely joined (Fig. 3, 4),

partially fused, parameres distinct and joined to genitalia in apical third;

genitalia flattened dorso-ventrally. This genus is easily separated from Nof-

oxus by the anteriorly directed setae on the lateral margins of the pronotum

and the carinae extending laterally from the horn base to the cervix.

The genus Mecynotarsus is very difficult to characterize on a world basis,

or even for only the New World. There are several well-defined groups placed

in the genus which are sufficiently distinct to be placed in new genera (Chandler

1977). This genus can be separated from the others in the New World by the

lack of the pronotal pit above the cervical notches and the metatarsi being

longer than the metatibiae.

Notoxus is represented by seventy-two species in the New World with most

of these occurring in Mexico. Plesionotoxus consists of seven species with all

of these being restricted to South America. Mecynotarsus contains thirteen

species with the majority occurring in Mexico.

GENUS PLESIONOTOXUS, NEW GENUS

The type-species of Plesionotoxus is designated as Notoxus lebasi LaFerte.

The genus is restricted to South America and is easily distinguished from

Notoxus by the lateral carinae extending from the horn base to the cervix and

the anteriorly directed setae over the pronotal pits. The species included in

the genus are very similar externally and a revision of the group will require

that the genitalia of the types be examined.

Plesionotoxus is perhaps most similar to Pseudonotoxus Pic by the form

of the genitalia and the lateral carinae at the base of the pronotal horn. It is

easily separated by the lack of setae between the eye facets, the absence of

any modified setae on the body, and the distinct antebasal impression of the

pronotum.

Plesionotoxus lebasi (LaFerté)

(Fig. 3, 4)

Notoxus Lebasii (sic) LaFerte 1848: 51-2, Table 21, fig. 16. Type locality:

Colombia. Generotype of Plesionotoxus. Werner 1965: 22. Also known

from Bolivia, Brazil, Panama and Peru.

Plesionotoxus argentinus (Pic)

Notoxus argentinus Pic 1912: 456. Type locality: Mendoza, Argentina.

Bruch 1928: 193; 1938: 165.

Chandler: Notoxus of Central and South America 9

Plesionotoxus boliviensis (Pic)

Notoxus boliviensis Pic 1914a: 250. Type locality: Bolivia.

v. Germaini (sic) Pic 1914b: 61. Type locality: Bolivia.

v. imnotatipennis Pic 1914b: 61. Type locality: Peru.

Plesionotoxus informicornis (Krekich)

Notoxus informicornis Krekich 1913: 135-6. Type locality: Mendoza,

Argentina. Bruch 1928: 193; 1938: 167.

Plesionotoxus gounellei (Pic)

Notoxus Gounellei (sic) Pic 1901b: 227. Type locality: Brazil.

Plesionotoxus venustus (Pic)

Notoxus venustus Pic 1901b: 227. Type locality: Argentina Pic 1911: 13.

Bruch 1928: 193; 1938: 166.

GENUS NOTOXUS FABRICIUS

Notoxus Geoffroy 1762: 356 (not binomial, see Hemming 1954). Fabricius

1792: 210; 1801: 289. Say 1817: plate IV; 1824: 21; 1827: 38. LeConte

1824: 170 (Anthicus); 1851: 152; 1852: 92; 1876: 518. Hentz 1827: 375.

LaFerte 1848: 21. Chevrolat 1877: ix. Horn 1884: 165; 1892: 47.

Casey 1884: 189; 1895: 756. Quedenfeldt 1886: 121. Champion 1890: 203

1893: 461. Pic 1894: 46; 1897: 215; 1901la: 227; 1901b: 89; 1904: 228;

1911: 5; 1912: 456; 1913: 8; 1914a: 61; 1914b: 250; 1916: 11; 1918: 117.

Fall 1902: 257; 1916: 33; 1932: 56. Blatchley 1910: 1334. Krekich 1913:

135. Leng 1920: 162. Blaisdell 1929: 57; 1936: 144. Bruch 1928: 193;

1938: 165. Barber 1941: 25. Pallister 1955: 5. Werner 1964: 200;

1965a: 124; 1965b: 123. Generotype Meloe monoceros Linnaeus (cited as

generotype in Bonadona 1971).

Monocerus Faldermann 1837: 106. LeConte 1847: 89. Generotype Monocerus

brachycerus Faldermann, fixed by monotypy.

Geoffroy's (1762) work was rejected by the International Commission on

Zoological Nomenclature (Hemming 1954) on the basis that it was not binomial.

The first use of the genus Notoxus with a specific name was by Fabricius

(1792), according to the literature dates in Pic (1911).

Three of the species named by LaFerte were collected by Piccolomini in

"California.'' Ags has been noted by Werner (1964: 200), this locality is erro-

neous and the material was probably collected in the area around Mexico City.

The three species of Notoxus collected by Piccolomini are commonly collected

around Mexico City and this is probably the type locality.

10 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Taxonomic Characters

Color and patterns: Color is expressed in simple English rather than

Latin. The elytra exhibit the most useful color patterns. The basic pattern

consists of light elytra with a median or postmedian band and darkened omo-

plates (a hump centered between the humeri, elytral suture and postbasal

band, Fig. 7A). There is often an apical or postbasal band. For a species

the general position of the bands does not vary, but the smoothness of the mar-

gins and the degree that they may extend anteriorly may vary considerably.

One species has completely dark elytra with the elytral pattern being formed

by dark and light setae. Several species have all dark or light elytra without

any setal patterns.

The pronotal horn may be abruptly darker than the remainder of the pro-

notum. The head is usually slightly darker or the same color as the pronotum.

The light areas of the elytra are lighter than the pronotum and head, and vary

from tan to brown.

Setation: The setae of the head, pronotum and elytra are often raised and

their appearance is one of the best characters for recognition of species. The

lengths of the raised setae of the head which outline the pronotal horn are

usually highly correlated with the lengths of the raised pronotal and elytral

setae, but in a few species these setae are very reduced in the male and nor-

mal in the female.

The pronotum of most species will have some setae which are raised,

while for a few species all the setae are appressed. The average lengths of

the raised elytral setae and their average angles from the elytral surface are

characteristic for each species. The setae vary from appressed (about 8°) to

decumbent (about 14°) to distinctly raised (20° to 85°).

Head: The mandibles have the peculiar lateral explanation which is a

characteristic of the Notoxinae (Fig. 6). The posterior margin of the explana-

tion may be very distinct or obsolete. The distance between the two margins

in a ventral view is useful in the separation of species-groups.

The last segment of the maxillary palpus varies slightly in shape, but is

best characterized as being subangulate. The length/width ratios of the anten-

nomeres were recorded and the results from the numerical analysis indicated

that the ratios of I, I], Il, V-IX, X and XI were significantly different from

each other for many of the species.

Pronotum: The pronotal horn provides a number of useful characters. On

the pronotal horn is a shorter, median elevated area, the crest, which extends

posteriorly to the pronotal disc (Fig. 5). The horn proper either has several

ventro-lateral pits or lacks them completely. The horn margin may be smooth

or possess up to twelve denticles or teeth. The apex is usually smooth, with the

teeth being on the lateral margins or near the base. The crest usually has a

smooth margin, but may have a series of sharp denticles representing the mar-

gin which will often indistinctly merge anteriorly with the horn. The crest

apex varies from rounded to pointed with the apically pointed crests generally

depressed anteriorly. Some species may have a smooth, median longitudinal

ridge on the crest (Fig. 5).

There is a deep pubescent pit dorsal to each pronotal cervical notch. The

size of these pits is stable for a species and is useful in recognizing species-

eroups. One species has a lateral carina on each side of the pronotum extend-

ing from the horn base to the cervix just anterior to the pronotal pits. In dor-

sal view the general shape of the pronotal disc is globose, but may be shortened

Chandler: Notoxus of Central and South America 11

to appear transverse.

Elytra: Elytra of many species are distinctly widest at the middle, but a

few species are only slightly wider medially and have the lateral margins of

the elytra appearing almost parallel. The shape of the elytral apices is diffi-

cult to characterize except for the more extreme variations. A few species

have the elytra distinctly narrowed in the apical half with the apices acuminate

(Fig. 21A). In some species the females have the elytra protruding at the ely-

tral suture (Fig. 42F) while others have the elytra somewhat shortened and

truncate. Males of some species have an acute protuberance on the lateral

apical angles (Fig. 50A). Males of two species have a median preapical im-

pression which is densely setate.

Mesosternum: The lateral margins of the mesosternum vary from straight

to slightly curved outwardly. The anterior margin is pointed and reaches the

prothorax. The median process between the mesocoxae is narrow and truncate

at the apex.

Abdomen: In the females of all but one species, the last tergite, the mor-

phological seventh, is in the form of a broadly rounded triangle. The excep-

tion has a large apical tubercle arising from this tergite which extends beyond

the elytral apices (Fig. 43F). The last sternite in the males is rounded. The

morphological eighth tergite is usually barely visible between the apparent

last tergite and the last sternite. In females this eighth tergite is not visible.

The sternites in the females are generally unmodified with the apex varying

slightly from rounded-triangular to triangular. Modifications in the male are

almost always confined to the last (morphological seventh) sternite, but in one

species the first four sternites may be flattened. The apex of the last sternite

varies from broadly rounded to truncate to deeply emarginate. There may be

an oval or transverse impression before the apex or no impression. The im-

pression may be shallow or deep. In two cases there is a transverse carina at

the center of the sternite. The apex may be straight or sharply downcurved in

lateral view.

Males are best separated from females by the form of the last sternite.

This sternite in the females will always be pointed to some degree and lacking

in other modifications. In all males this sternite will never be simply pointed,

but will be broadly rounded or have various modifications.

Legs: The first and second protarsomeres may have a distinct spur at the

lateral apical margin, or these spurs may be reduced or lacking in either or

both sexes. A few males have a median tubercle on the protibiae. One species

has a prominent tubercle on both the pro- and mesofemora. Another species

has the first tarsomere of the last leg enlarged and flattened.

Hind wings: The hind wings are usually fully developed. The wing venation

did not vary in all of the species examined (Fig. 1).

Genitalia: The genitalia of the females was found to be membraneous and

no characters could be found which are taxonomically useful. The male geni-

talia are heavily sclerotized and exhibit many valuable specific characters.

The parameres may have lobes at various points and vary in general shape.

The median lobe or penis exhibits a variety of modifications, particularly near

the apex. Before the apex there may be a dorsal notch (Fig. 52E), segments,

or an oval opening. The apex may be pointed, rounded, truncate, forked,

split dorso-ventrally (Fig. 13E), split laterally, with an apical hook (Fig. 50D)

or with a combination of these characters. The internal sac is membraneous

and lacking in obvious signs of sclerotization.

rE.

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 19077

Key to the Central and South American Notoxus

In the following key I have attempted to use characters which will enable

the user to identify both sexes. However, several species can be positively

identified using only male genitalia. Several United States species which occur

near the Mexican border are included in the key as they probably will be found

in the northern mountains and deserts of Mexico.

(3)

. (4)

(6)

oh)

Lateral carina extending from horn base to pronotal pits at neck;

elytra unicolorous; southern Brazil. ..... Notoxus sp. (p. 63)

Lateral margins of pronotum adjacent to horn smoothly rounded. . 2

Venter of horn with lateral pile... .. 2. 2.6 6 we ec ws 3

Venter of horn smoothly rugulose, without pits. .......... 14

Elytral apices acuminate (Fig. 22A); crest margin crenulate; male

fifth sternite emarginate at apex, transverse ridge before apex

(Fig. 22B); Chiapas to Guatemala. . acuminatus Champion (p. 30)

Elytral apices rounded (Fig. 7A); crest margin smooth; male fifth

Steriite VavinAvle, «0 6 ee ae 4.

Elytra with distinct dark median band (Fig. 7A); crest often with

MIOGIOn PICCC a ee 9)

Elytra with dark postmedian band or spots, or unicolorous; crest

WithOUL Wiedian ridge. 5 2. 2 re es 6

Midband with very irregular margins (Fig. 10A); male fifth sternite

slightly emarginate; males with setaceous impression near elytral

apices and lacking a medial tubercle on the profemora; throughout

MesicO. 6 6 hirsutus Champion (p. 21)

Midband with smoother margins (Fig. 11A); male fifth sternite with

apical triangular impression; medial tubercle on male protibiae

and lacking any setaceous impressions on elytra; northwestern

Meteo... 6 ee a calcaratus Horn (p. 22)

Horn with three or less teeth on margins; mandibular explanation

narrow in ventral view, hind margin meeting mandible at lower

level than front; male fifth sternite emarginate.

pallidus Fall (p. 44)

Horn with three or more teeth; both margins of mandibular explana-

tion meeting mandible at same level; male fifth sternite broadly

VOUNUGG. 5 a er ee ce ee 7

MAyira UnicGlorous, Without MiarRUlGS 2. co ie eee es 8

Elytra with dark band or spots on elytra. . 6 6 6 we ee 10

Raised elytral setae bristling, average angle 69°; Peru and Bolivia.

peruvianus Pic (p. 38)

Raised elytral setae at average angle of 359; Central America. . . 9

Horn abruptly darker than rest of pronotum; Guatemala to Panama

ruficollis Champion (p. 38)

10.

il.

12.

13.

14.

15.

16.

Tf,

16.

19.

20.

(10)

(12)

(2)

(14)

(15)

(14)

(17)

(19)

Chandler: Notoxus of Central and South America 13

Horn and rest of pronotum same color; northeastern Mexico to

Guatemala, 2. 9 ee murinipennis (LeConte) (p. 37)

Present in the West Indies: . 2 2. 286 ee a ee, 11

Present on the maininnd. 2 3 Se ee 12

Penis apex truncate, with small indentation; Jamaica.

jamaicus Pic (p. 36)

Penis apex deeply split; Puerto Rico. |

bipunctatus Chevrolat (p. 35)

Penis simple, not split at apex (Fig. 23D); through much of Mexico

no Clinteiala ee ee desertus Casey (p. 33)

Penis split dorso-ventrally at apex (Fig. 24D). ......... 13

Penis with strong dorsal tooth at apex (Fig. 24D); central Mexico

north 6 Chitudiaa. 44... ee. celatus, n. sp. (p. 32)

Penis without dorsal tooth (Fig. 24D); northeastern Mexico to

Venewiela 2 cumanensis LaFerte (p. 31)

Pronotum with antebasal pubescent band unbroken; males with later-

al apical tubercles On Clyita:. 2 oe ee ss 15

Pronotum with antebasal pubescent band widely broken at center;

male elytra with lateral tubercles in two rare species. .... 17

Elytral setae appressed; crest elevated slightly above horn margin;

penis with deep dorsal notch at apex; aja California.

vonformis LeConte (p. 54)

Some elytral setae erect, average angle greater than 459; crest and

DENIS BVCk VATIAIO. 406 eh Ae a we 16

Crest width at base one-half greatest width; penis with narrow dor-

sal notch at apex; Baja California Sur. . . bajae, n. sp. (p. 53)

Lateral margins of crest about parallel; penis with broad dorsal

noteh at apex: Baja California. . .. . sparsus LeConte (p. 52)

Elytral setae in postbasal impression directed laterally (Fig. 19A);

elyiva narrowed inapical pall... 6 ve oe ee a we a ss 18

Setae on elytra directed posteriorly; elytral apices variable. . . 19

Crest margin crenulate, apex indistinctly merging into horn; central

Memeo to Colombia. « 2S ieee lateralis, mn. sp. (p. 28)

Crest margin smooth, apex distinct and elevated above horn mar-

ein) Baja Calivornia. <6. 8 a ee solus, n. sp. (p. 29)

Crest margin roughly crenulate, apex indistinctly merging into

NOP 6 a I OB ae. 20

Crest margin smooth, apex distinct. . ..% 2.34097 24. 22

Raised elytral setae bristling, about 62°; mandibular explanation

narrow in ventral view, hind margin obsolescent; northern

MeONiOO. 665 Se ea anchora Hentz (p. 50)

21.

22.

23.

24.

20.

26.

27.

(20)

(19)

(22)

(23)

(24)

(23)

(26)

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Raised elytral setae appressed; mandibular explanation wide in ven-

tral view, with both margins distinct.......... ae 21

Elytra with postmedian light markings evenly curved anteriorly to

near elytral suture (Fig. 18A); setae dense on elytra; elytra

slightly narrowed in apical half; central Mexico to Panama.

eximius Champion (p. 27)

Elytra with postmedian light band meeting broadly at elytral suture

(Fig. 21A); setae sparse on elytra; elytra narrowed in apical

half: Chiapas to El] Salvador... . . impressus Champion (p. 29)

Elytra with all setae appressed to decumbent, or with a few setae

slightly raised (ignore very short tactile setae at 80° and less

than one-third length of other setae)... 2... 6... ee 23

Elytra with some setae raised, distinct from other setae, average

angle of raised setae greater than 20°, usually much longer than

One? Sette, 6 Ee a Re Rae 34

Pronotum with all setae appressed; males with fifth sternite emar-

ginate, circular impression before apex (Fig. 42B); females with

elyiral apices spinose (Pie. 42F)o oe eke 24

Pronotum with some setae raised, distinct from other setae, may

be sparse-if apparently absent then males with large circular

impression of fifth sternite (Fig. 32B) and females with elytral

NGS Pe ae ee ee 26

Pronotum unicolorous, only slightly darker than the light areas on

elytra; females with pygidium simple; northwestern Mexico.

photus, n. sp. (p. 49)

Pronotum distinctly darker than light areas on elytra, horn lighter

than Pest Of OYONOUIN 2.6 ek a ee a ae as 20

Females with pygidium apex slightly produced; males with shallow

impression before apex of fifth sternite, never densely setate;

Conta Memeo, 2 ee talpa LaFerte (p. 47)

Females with apical protuberance from pygidium (Fig. 43F); males

with deep circular impression on fifth sternite, margin of impres-

sion often densely setate; south-central Mexico.

pygidialis, n. sp. (p. 48)

Body brownish-black, rarely with faint lateral light markings;

brownish setae forming median band, outlined by white setae

(Fig. 45A); males with fifth sternite emarginate, slight to deep

transverse impression before apex (Fig. 45B); central Mexico.

fraternus Champion (p. 56)

Body with distinct elytral ground color pattern, if completely dark

then setae are all whitish and males with circular or oval impres-

Sion, On Viith Slernibe, 0G Oe eee ee 27

Elytra with slightly postmedian band, apices dark (Fig. 33A); males

with fifth sternite distinctly emarginate, circular impression at

apex; mandibular explanation usually narrow in ventral view

(hie. SoG a ae ea ee eG 28

28.

ao.

30.

31.

32.

33.

34.

35.

(27)

(28)

(29)

(27)

(31)

(32)

(22)

(34)

Chandler: Notoxus of Central and South America 15

Elytra trifasciate with light apices, to completely dark; mandibular

explanation moderately wide in ventral view (Fig. 16C); males

with fifth sternite truncate to slightly emarginate, no impressions

DETOTO ANOR o. e s a as e 31

Elytra with postbasal band rarely broken at elytral suture; male

fifth sternite with shallow circular impression; northeastern

Mexteo. 4. Ghd Be orientalis, n. sp. (p. 42)

Elytra with only omoplates dark near base; male fifth sternite with

CCD TMDTeSsSION 70 ee ee a, 29

Often with some elytral setae slightly raised giving slightly ruffled

appearance; horn margins rarely extending posteriorly past

crest apex; northeastern Mexico. . . . postictus, n. sp. (p. 41)

All setae appressed; horn margins often extending well past crest

mex: hortiwestern Mexiro. 2 6k a a 30

Crest narrowly rounded at apex; horn margins extending past first

quarter of crest; horn slightly depressed before apex, not scoop-

like; males with at most fourth sternite slightly flattened

ILI oe Oe a nuperus Horn (p. 39)

Crest usually broadly rounded at apex; length of horn margins vari-

able; horn often centrally depressed before apex to form scoop-

like structure; males with sternites I-IV often flattened medially.

haustrus, n. sp. (p. 40)

Elytra dark or with faint lateral light areas (Fig. 14A), not trifasci-

ate; head and pronotum about same color; male fifth sternite dis-

tinctly emarginate; Durango. : . . durangoensis, n. sp. (p. 24)

Elytra trifasciate (Fig. 16A); male fifth sternite truncate or very

Sligntly GMIATCINGle. 6. 6 ee ae elas a 32

Elytra obscurely trifasciate; pronotum much lighter than head; male

fifth sternite slightly emarginate; ventral portion of penis same

width through length; Durango. ..... nubilus, n. sp. (p. 25)

Elytra markedly trifasciate; pronotum about same color as head;

wiale 71h SIAPNITe PINCAtey bk a aw Ge a ie es 33

Ventral portion of penis same width (Fig. 15E); Arizona.

balteatus Casey (p. 23)

Ventral portion of penis enlarged at apex (Fig. 16E); central Mexico

AO a ae ee Nakata a ya wigs toltecorum, n. sp. (p. 26)

Elytra completely brown, setae unicolorous; raised setae long,

about two-thirds as long as eye, average angle 329; male fifth

sternite emarginate, not impressed; northeastern Mexico.

campus, n. sp. (p. 60)

Elytra brown with tan areas; setal length variable; usually not occur-

ring in northeastern MeRiCO. 6 ner eee ey as es 30

Elytra with raised setae bristling, average angle greater than 50°;

often with median ridge on crest; apex of crest distinct and well

elevated above NOTH MGPTIN bhi aie Ww in 4 ke

36.

37.

38.

39.

40.

41.

42.

43.

(35)

(36)

(37)

(35)

(39)

(40)

(41)

(42)

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Raised elytral setae at average angle of 45° or less, if greater than

mandibular explanation narrow in ventral view (Fig. 47C); median

ridge lacking in all except one rare species (caudatus). .... 39

Crest about as long as width at base; males with fifth sternite broadly

rounded, small triangular impression at apex; northwestern

MeGXICO) 5.6 6 es gelidus, n. sp. (p. 20)

Crest nearly or greater than twice as long as width at base. .. 37

Crest apex broadly pointed; raised elytral setae almost as long as

eye; northwestern Mexico. ...... occidentalis, n. sp. (p. 55)

Crest apex narrowly rounded to broadly rounded; raised elytral

setae not more than two-thirds eye length. ........... 38

Parameres bulging laterally near base, not swollen near apex in

lateral view (Fig. 7E); central Mexico.

pueblensis Champion (p. 18)

Parameres parallel at base, swollen near apex in lateral view (Fig.

BE): Nueva U6on. . 2 leonensis, n. sp. (p. 19)

Slight longitudinal ridge on crest; males with shallow densely setate

impression near apex of elytra, elytra truncate near apex; New

NMORIGG 2 a ee -. caudatus Fall (p. 18)

Without trace on longitudinal ridge on crest; males lacking setate

Oe ON CIV. 8 ee ek 40

Elytra with preapical band or spots (Fig. 51A); mandibular explana-

tion narrow in ventral view, hind margin obsolescent; raised

setae on elytra bristling; male elytra with lateral apical tubercles;

Chihtahad 0 2. ee serratus (LeConte) (p. 51)

Elytra with median, postmedian or apical bands, or unicolorous;

mandibular explanation with hind margin distinct; male elytra

lacking lateral apical tubercles. 2... 2 Fie a ee a 41

Male protibiae with median tubercle; male fifth sternite slightly

emarginate; raised elytral setae more than two-thirds eye length,

raised at 40-459; elytra with median band; Arizona to New Mexico.

montanus Casey (p. 23)

Males lacking median tubercle on protibiae; if setae long, then

elytra with postmedian band; male fifth sternite variable. .. 42

Mandibular explanation narrow in ventral view (Fig. 47C), hind mar-

gin recessed; raised elytral setae at average angle of 30-65°,

often bristling; elytra with postmedian band; males with fifth

sternite truncate to slightly emarginate, usually with shallow

transverse impression before apex. .........4.4-+4s86-6 43

Mandibular explanation wide to moderately wide in ventral view

(Fig. 41C), hind margin prominent; average angle of raised ely-

tral setae 20-459; elytra usually with median band; males with

filth sternite usually emarcinare. . 6... wk ee 45

Raised elytral setae relatively short, less than half of eye length

(0.15-0.16), average angle 30-40°; northern Mexico.

marginatus LeConte (p. 45)

44.

49.

46.

47.

48.

49.

00.

ol.

o2.

(43)

(42)

(45)

(46)

(45)

(49)

(50)

(51)

Chandler: Notoxus of Central and South America 17

Raised elytral setae long, bristling, about two-thirds eye length

(0. 18-0. 22), average anole 50-059... 8 be ws 44

Horn margin with 4-8 teeth; central Mexico to Sonora.

mexicanus Champion (p. 46)

Horn margin with 1-4 teeth; northern Mexico.

apicalis LeConte (p. 44)

With 3-8 teeth on horn margin; crest margin smooth to smoothly

CVONUIOLC.:6 50 a oo wea eo ee ree Se ee 46

Postmedian band reflexed anteriorly along elytral suture (Fig. 40A);

males with fifth sternite deeply emarginate, transverse impres-

sion before apex; first metatarsomere enlarged and flattened;

COntval WMeRIOO oo eg ea opacus Champion (p. 62)

Median band straight across elytra (Fig. 41A); males with fifth

sternite slightly emarginate, oval impression before apex; meta-

TATSOMIOTOS CYNE ICA cio ie kk ee a a 47

Penis apex truncate; Chiapas and Guatemala.

crucifer Champion (p. 61)

Penis apex pointed; central Mexico to Sinaloa.

zapotecorum, n. sp. (p. 60)

Elytra with preapical band (Fig. 13A); crest elevated well above

horn; male fifth sternite slightly emarginate; northwestern

Wie a ee ee arizonensis Fall (p. 24)

Elytra with apical band; crest and male fifth sternite variable. . 49

Males with apex of fifth sternite rounded, no impression before apex;

mandibular explanation wide; western Texas to Arizona.

brevicornis Fall (p. 55)

Males with apex of fifth sternite emarginate; mandibular explanation

PID OR CRT We tO WA ge a a he ek co ee a 20

Males with fifth sternite emarginate into large circular impression

(Fig. 39B); phallobase longer than wide; Jalisco to Sonora.

stephani, n. sp. (p. 57)

Males with transverse impression or flat before apex; phallobase

wicer than lone: CONtrAl WIGRIC(.. coin so) eo ck kk ew oO

Penis widely forked at apex (Fig. 38D); male fifth sternite flattened;

MO re ey aa ee nr a desperatus, n. sp. (p. 58)

Penis truncate at apex; male fifth sternite variable. ....... O2

Male fifth sternite with transverse impression before apex.

truncatipennis Champion (p. 59)

Male fifth sternite flattened before apex.

aztecorum, n. sp. (p. 57)

18 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Pueblensis-group

The pueblensis-group is defined as those Notoxus with the following char-

acters: mandibular explanation wide, prominent hind margin meeting mandible

at same level as front margin; raised pronotal setae distinct, pubescent ante-

basal band widely separated; crest margin smooth, apex rounded and distinct,

median crest usually present; elytra with median band, raised elytral setae

longer than half eye length; males without lateral apical projections on elytra;

fifth sternite without impression before apex; parameres with lobes or penis

divided dorso-ventrally.

Central American species included are calcavatus Horn, hirsutus Champi-

on, pueblensis Champion, gelidus n. sp. and leonensis n. sp. There is only

one United States species in this group, caudatus Fall. It is included in the

key to the species because it has been collected near the Mexico-New Mexico

border and will probably be found in the northern montane regions of Mexico.

Notoxus pueblensis Champion

(Pigs... 755)

Notoxus pueblensis Champion 1890: 214, Table IX, fig. 24. Type locality:

Puebla, Puebla, MEXICO, type series in BMNH.. Pic 1894: 47; 1911: 11.

Length 2.78-3.48. Head and pronotum tan to red-brown, elytra tan with

brown to dark brown markings: oval mark on omoplates, irregular preapical

band, flanks of elytra dark posteriorly to midband. Setae brownish when aris-

ing from dark areas, whitish over rest of body. Raised setae long and erect

on pronotum and elytra, average angle on elytra 649. Mandibular explanation

moderately wide, front and hind margins meeting at same level at mandible.

Pronotum with antebasal pubescent band widely broken. Horn without pits

beneath, 0-2 teeth on margins. Crest distinct, elevated well above horn, with

short median ridge, rounded at apex. First and second protarsomeres with

spur on posterior apical margin.

Males with fifth sternite emarginate at apex. Penis divided dorso-ventrally

at apex. Spurs on protarsomeres small.

Females with fifth sternite rounded-triangular. Spurs on protarsomeres

large and easily distinguished.

Redescribed °: 7 mi NE Atlixco, Puebla, MEXICO (DSC). Length 2.82.

Head 0.58 long, eye 0.26 long, raised setae above eyes average 0.14 long;

antennomere ratios: 11.94, I1 1.42, Il] 2.33, IV 1.85, V 1.86, VIII 1.60,

X 1.28, XI 1.94; fourth maxillary palpus 0.20 long, 0.11 wide; mandibular

explanation 0.09 long, 0.05 wide. Pronotum 0.90 long, length from base to

point of greatest width 0.32, break in antebasal band 0.24 wide; horn 0.37 long,

0.27 wide at base, 1 tooth on margin; crest 0.21 long, 0.10 wide at base, with

short median ridge; crest apex 0.10 above horn margin, horn apex 0.20 from

crest apex; pronotal pit 0.10 long. Raised elytral setae 0.15 long, average

angle 65°. Emargination of fifth sternite 0.14 wide, 0.01 deep.

Specimens examined, 31: MEXICO: Puebla: May, A. Fenyes coll. (CASC);

San Martin, 26 May 1922, E. G. Smyth (USNM); Puebla, 28 May 1922, E. G.

Smyth (USNM): Santa Maria, May, A. Fenyes coll. (CASC): Tehuacan, A.

Fenyes coll. (CASC); 7 mi NE Atlixco, 26 July 1974, R. L. Mangan & D. S.

Chandler, under low vegetation in corn field (DSC); same locality, 26 June

Chandler: Notoxus of Central and South America 19

1975, D. S. Chandler, under low plants in corn field (DSC). Mexico: Chapingo,

22 July 1964, T. R. E. Southwood (BMNH). 1 specimen with 'Mex.", F. C.

Bowditch coll. (MCZC) and two specimens with ''X" and ''IX"' (USNM).

Distribution: The semiarid region in and around the state of Puebla. Col-

lected underneath and on vegetation in corn fields. Present from May through

July.

Discussion: Very similar to leonensis in general appearance and form of

the genitalia. It is separated by the apically narrowing parameres and its

distribution in central Mexico.

Notoxus leonensis, n. sp.

(Fies. 8, 65)

Length 2.70-3.62. Head and pronotum orange, pronotum brown laterally,

elytra orange with brown markings: band across omoplates interrupted at

elytral suture, usually meeting median band. Pubescence brownish when aris-

ing from markings, yellowish over rest of elytra. Raised setae long and erect

on pronotum and elytra, average angle on elytra 559. Mandibular explanation

moderately wide, front and hind margins meeting mandible at same level.

Pronotum with antebasal pubescent band widely broken. Horn without pits

beneath, 0-2 teeth on margin. Crest margin smooth, apex rounded and dis-

tinct. First and second protarsomeres with strong spur on posterior apical

margin.

Males with fifth sternite slightly emarginate. Penis divided dorso-ventrally

at apex, apices bilobed.

Females with fifth sternite rounded-triangular.

% holotype: 18 mi SW Linares, Nuevo Leon, MEXICO. Length 3.25.

Head 0.60 long, eye 0.26 long, raised setae above eyes average 0.13 long;

antennomere ratios: 12.51, 11.91, Il 2.26, IV 1.67, V 2.04, VII 1.69,

X 1.51, XI 2.42; fourth maxillary palpus 0.22 long, 0.10 wide; mandibular

explanation 0.10 long, 0.04 wide. Pronotum 0.97 long, length from base to

point of greatest width 0.34, break in antebasal pubescent band 0.24 wide;

horn 0.38 long, 0.28 wide at base, 1 tooth on margin; crest 0.23 long, 0.08

wide at base; crest apex 0.12 over horn margin, horn apex 0.21 from crest

apex; pronotal pit 0.08 long. Raised elytral setae 0.18 long, average angle

49°. Emargination of fifth sternite 0.17 wide, 0.01 deep.

Specimens examined, 5: HOLOTYPE “, 18 miSW Linares, Nuevo Leon,

MEXICO, 2 July 1974, Clark, Murray, Ashe, Schaffner, at light (USNM).

PARATYPES: MEXICO: Nuevo Leon: 1%, 19, eutopotypical (DSC, TAMU);

222, 16 mi W Linares, on Mexico 60, 2200', 26-29 May 1971, A. Newton, |

canyon bottom, UV trap (FGW).

Distribution: Known from the Sierra Madre Oriental in Nuevo Leon.

Adults collected at light and ultra-violet light. Present in June and July.

Discussion: Very similar to pueblensis. Separated by the apically en-

larged parameres and its distribution at the northern end of the Sierra Madre

Oriental.

20 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus gelidus, n. sp.

(Figs. 9, 55)

Length 2.81-3.69. Head and pronotum tan to red-brown, elytra tan with

brown to dark brown markings: circular mark on omoplates, irregular medi-

an band, irregular preapical band which may be broken at middle. Setae

brownish when arising in markings, whitish over rest of body. Raised setae

long and erect on pronotum and elytra, average angle on elytra 539. Mandi-

bular explanation moderately wide, front and hind margins meeting at same

level at mandible. Pronotum with antebasal pubescent band widely broken.

Horn without pits beneath, 0-3 teeth on margins. Crest distinct, elevated well

above horn, with short median ridge, widely rounded at apex. First and second

protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite broadly rounded, small triangular impression at

apex. Parameres with long medial flange near middle.

Females with fifth sternite rounded-triangular.

S holotype: Sunnyside, Arizona. Length 3.30. Head 0.68 long, eye 0.34

long, raised setae over eyes average 0.12 long, antennomere ratios: I 2.35,

Ii 2.00; Hi2.67, 1V 2.23, V 2.21, VUl2.20, X 1.76, X12. 88; fourth maxil-

lary palpus 0.21 long, 0.10 wide; mandibular explanation 0.08 long, 0.05 wide.

Pronotum 0.95 long, length from base to point of greatest width 0.36, break

in antebasal pubescent band 0.29 wide; horn 0.35 long, 0.24 wide at base, one

tooth on margin; crest 0.15 long, 0.13 wide at base, with short median ridge;

crest apex 0.15 over horn margin, horn apex 0.23 from crest apex; pronotal

pit 0.10 long. Elytra with raised setae 0.16 long, average angle 56°.

Specimens examined, 33: HOLOTYPE ©, Sunnyside, Cochise County,

Arizona, 27 August 1973, K. Stephan (CNCI). PARATYPES: Arizona: Co-

chise Co.: 50°, 499, eutopotypical (DSC, KSC); 1%, Sunnyside, 20 June 1970,

K. Stephan (DSC). Chiricahua Mountains - 1°, 6000', 29 June 1968, K.

Stephan (DSC); 1“, 292, 26 July 1952, D. J. & J. N. Knull (OSUC); 1°, Turkey

Creek Camp, 6400', 14 June 1964, J. Burger, UV light (FGW); 1%, Southwest

Research Station, 5 mi W Portal, 1 September 1959, J. R. Powers, at light

(CISC); same locality, 1°, 4 August 1961, J. M. Linsley (CISC); same locality,

1, 5 August 1967, F. G. Andrews and 12, 28 August 1967, F. G. Andrews

(CDAE); 1“, Stewart Camp, 1 miS Portal, 18-20 July 1971 (CISC). Huachuca

Mountains - 1°, Miller Canyon, 2.7 mi W parking area, 5800', 6 June 1964,

J. Burger, UV light (FGW); same locality, 12, 5700', 10 August 1974, D. S.

Chandler (DSC); 12, Ramsey Canyon, 13 July 1955, F. G. Werner & G. D.

Butler (FGW). Santa Cruz Co.: 12, Madera Canyon, 9 August 1961, Werner

& Nutting, UV light (FGW); 12, same locality, 4880', 25 September 1963, V.

L. Vesterbey (UCDC); 1°, same locality, 2 September 1960, F. G. Andrews

(CDAE); 1% same locality, 8 September 1970, E. A. Kane, UV light (CDAE).

MEXICO: Sonora: 12, Sierra del Ajos, Canyon de Evans, 15 July 1970, V.

Roth (DSC). Durango: 2°%o, 26 mi W Durango, 13 July 1974, R. L. Mangan

& D. S. Chandler, beating oak (DSC).

Distribution: Montane areas in southern Arizona south through the Sierra

Madre Occidental. Apparently associated with the oak zone. Karl Stephan

noted that a portion of the type series was collected on a cold night at light,

while little else was attracted. Present from June through September.

Discussion: Most similar to pueblensis and leonensis in appearance.

Distinct by its high, almost semicircular crest and the medial flanges on the

parameres.

Chandler: Notoxus of Central and South America 21

Notoxus hirsutus Champion

(Figs. 10, 55)

Notoxus hirsutus Champion 1890: 211, Table IX, fig. 20. Type locality: near

Mazatlan, Sinaloa, MEXICO, BMNH Lectotype 2, designated by Chandler,

1975. Pic 1894: 47; 1911: 8. Not Pallister 1955: 10.

Notoxus cristatus Champion 1890: 213-4, Table IX, fig. 23. Type localities:

Guanajuato, Guanajuato, and Cuernavaca, Morelos, MEXICO, type

series in BMNH and F. G. Werner collection. Pic 1894: 46; 1911: 7.

Pallister 1955: 10, in part. NEW SYNONYMY.

Length 2.95-3.70. Head and pronotum tan to dark reddish-brown, elytra

tan with brown markings: oval mark on omoplates, irregular median band,

irregular preapical band, marks on flanks behind humeri in few specimens.

Pubescence brownish when originating in darkened areas, whitish over rest

of body. Raised setae long and erect on pronotum and elytra, average angle

on elytra 709. Mandibular explanation wide, with front and.hind margins

meeting at same level at mandible. Pronotum with pubescent antebasal band

widely broken. Horn with pits beneath, 1-8 teeth on margin. Crest elevated

and distinct, apex rounded, usually with distinct median ridge. First and

second protarsomeres with strong spur on posterior apical margin.

Male with circular impressed area near apex of elytron, densely setate.

Fifth sternite slightly emarginate at apex. Penis split dorso-ventally, with

dorsal part bilobed at apex.

Female without setate impressed area on elytron, fifth sternite rounded-

triangular.

Redescribed “: 2 mi N Jojutla, Morelos, MEXICO (DSC). Length 3.50.

Head 0.75 long, eye 0.34 long, raised setae over eyes average 0.20 long;

antennomere ratios: 12.30, 111.69, Il 2.20, IV 1.94, V 1.79, VIIL 1.58, X

1.35, XI 2.39; fourth maxillary palpus 0.24 long, 0.12 wide; mandibular

explanation 0.08 long, 0.06 wide. Pronotum 1.16 long, length from base to

point of greatest width 0.38, break in antebasal pubescent band 0.25 wide;

horn 0.46 long, 0.38 wide at base, 2 teeth on margin; crest length 0.25, width

at base 0.12, with median ridge; crest apex 0.15 above horn margin, crest

apex 0.24 from horn apex; pronotal pit 0.14 long. Raised elytral setae 0.22

long, average angle 71°. Emargination of fifth sternite 0.18 wide, 0.01 deep.

Specimens examined, 79: MEXICO: Chihuahua: Ciudad Jimenez;

Catarinas, 5800'. Durango: Rio Florido, nr. Las Nieves, 5200'. Sinaloa:

Presidio (Mazatlan), Forrer, Lectotype female of hirsutus (BMNH); 5.5 mi

NW Choix. Nayarit: Arroyo Santiago, nr. Jesus Maria. Zacatecas: 28 mi

S Jalpa. San Luis Potosi: Tamazunchale, Rt. 1, km. 365; 1 mi SW Tamazun-

chale. Queretaro: 1 mi NW Ayutla. Guanajuato: Guanajuato, type of crista-

tus; Silao. Guerrero: Xalitla; 17 mi E Acapulco. Morelos: 2 mi N Jojutla;

Cuernavaca, type of crisiatus; 3 mi N Alpuyeca, 3400'. Puebla: Tepexco, Rt.

115, 4000'; 12 mi SE Izucar de Matamoros. Oaxaca: Oaxaca; Yagul ruins;

10 mi N Miltepec. Chiapas: 31 mi SE Comitan; 29 mi SW Cintalapa. Speci-

mens present in collections of: AMNH, BMNH, CDAE, CISC, CUIC, DSC,

FGW, TAMU, UCDC.

Distribution: From New Mexico through most of Mexico. Specimens have

been collected by beating vegetation and are attracted to ultra-violet light and

cantharidin. Present from February through August.

22 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Discussion: Hirsutus is most similar to caudatus Fall, which also has the

setate impressions on the male elytra. N. hirsutus can be separated from

caudatus and the others in the group by the very irregular midband. N. cau-

datus males have a deeply emarginate fifth sternite and an apically pointed

fifth tergite, which will also separate this species from the others in the group.

The size of the elytral impression appears to diminish in a northerly direction

as the specimens of hirsutus from New Mexico have the impressions small

and easily overlooked in a quick examination.

Notoxus calcaratus Horn

(Figs. 11, 55)

Notoxus calcaratus Horn 1884: 170-1. Type locality: Arizona, MCZC type

#3041, “. Champion 1890: 211, Table IX, fig. 21. Pic 1894: 46; 1911:

6. Casey 1895: 759. Leng 1920: 163. Pallister 1955: 9.

Length 2.65-4.15. Head and pronotum tan, elytra tan with brown mark-

ings: circular mark on omoplates, irregular midband and band at apex.

Pubescence brownish when arising in markings, whitish over rest of body.

Raised setae long and erect on pronotum and elytra, average angle on elytra

71°. Mandibular explanation wide, with front and hind margins meeting at

same level at mandible. Pronotum with antebasal pubescent band widely

broken. Horn with pits beneath, 3-8 teeth on margin. Crest elevated and dis-

tinct, usually with distinct median ridge, rounded at apex. First and second

protarsomeres with strong spur on posterior apical margin.

Male protibiae with medial tooth on proximal margin, fifth sternite trun-

cate at apex, triangularly impressed anteriorly from apex. Parameres with

outer lobe near apex and strong outer angulations at apex.

Female without modifications of the protibiae, fifth sternite rounded-

triangular.

Redescribed “: 3 miN La Puerta, Baja California, MEXICO (DSC).

Length 4.00. Head 0.74 long, eye 0.28 long, raised setae over eyes average

0.29 long; antennomere ratios: 12.50, 12.00, IIT 2.20, 1V 2.13, V 2.38,

VII. 1.84, X 1.65, XI 2.64; fourth maxillary palpus 0.24 long, 0.13 wide;

mandibular explanation 0.14 long, 0.08 wide. Pronotum 1.28 long, length from

base to point of greatest width 0.42, break in antebasal pubescent band 0.31

wide; horn 0.54 long, 0.46 wide at base, 4 teeth on margin; crest 0.32 long,

0.08 wide at base, with median ridge; crest apex 0.22 above horn margin, horn

apex 0.38 from crest apex; pronotal pit 0.15 long. Raised elytral setae 0.30

long, average angle 70°. Truncation of fifth sternite 0.22 wide.

Specimens examined, 217: MEXICO: Baja California: 3 mi N La Puerta;

Rancho El Topo, 3 mi NE Sierra Juarez; 26 miS San Felipe. Baja California

Sur: 15 mi N San Ignacio; 3 mi W San Miguel de Commondu; 10 mi SW San

Jose del Cabo; 6 mi N San Jose del Cabo; 6 mi SW Santiago; 4 mi N Todos

Santos; La Paz; 12 mi NW La Paz; 3.5 mi NE San Pedro. Sonora: Alamos;

10 mi N Santa Ana; Saric; Rancho La Floresta, 8 mi E Tastiota; Imuris; 10 mi

NE Cananea; Navojoa; 20 mi NNE Obregon; 4 mi NE Altar; 2 mi E Huatabampo.

Sinaloa: Los Mochis; 5.5 mi NW Choix; Wollamo. Chihuahua: 12 mi N

Escalon; Ciudad Jimenez; 5 mi N Matamoros; Delicias; 6 mi NE Meoqui; Buena

Chandler: Notoxus of Central and South America 23

Vista; Samalayuca; Catarinas; Valle de Olivos; Primavera; Santa Clara Canyon,

5 mi W Parrita; 11 mi W Gran Morelos; Rio Conchos, 2 km W Oginaga.

Durango: 26 mi W Durango; Tepehuanes; 25 mi NE Bermejjillo; 2 mi N Cerro

Gordo; 2 mi NW Nombre de Dios; Sombrerte; Gomez Palacio; 11 mi W

Cuencame. Zacatecas: 16 mi NW Fresnillo; 15 mi NE Fresnillo. Coahuila:

Boquillas del Carmen. Nuevo Leon: Monterrey; 5 miS Monterrey. Specimens

present in collections of: AMNH, CISC, DSC, FGW, LACM, MSUC, TAMU,

TTCC, USNM.

Distribution: Throughout the southern half of the United States and south

into the northern semiarid portions of Mexico. Specimens have been collected

on Opuntia invicta flowers, Prosopis, Hoffmannseggia, cotton, under low plants

in corn fields and are attracted to white light, ultra-violet light and drying

meloid beetles. This species is fairly common on cotton and is of some econ-

omic importance in that it will eat the eggs of lepidopterous pests. Present

in Mexico from April through September.

Discussion: This species and hirsutus are the only species in the group

with lateral pits on the ventral surface of the horn. Only calcavatus has the

medial tooth on the protibiae of the males. The smooth median band on the

elytra will separate this species from similar appearing species of the

monodon-group, which also have the pits beneath the horn.

Montanus-group

The montanus-group is defined as those Notoxus with the following char-

acters: mandibular explanation wide; raised pronotal setae distinct, pubescent

antebasal band widely broken; horn without median ridge, lacking pits beneath

horn; raised elytral setae distinct and elevated, elytra with slightly postmedi-

an band. Males with simple elytra; protibiae with a median tubercle; fifth

sternite emarginate, without impressions; penis either dorsoventrally or

laterally split at apex. ,

Three United States species are included: montanus Casey, intermedius

Fall and schwarzi Horn. Of these species montanus occurs in the montane

regions of eastern Arizona and probably occurs in northern Mexico. The other

two species are found from the upper Sonoran desert of California to Alberta,

Canada. N. montanus is separated in the key to species.

Arizonensis-group

The avizonensis-group is defined as those Notoxus with the following char-

acters: mandibular explanation moderately wide, hind margin meeting mandible

at lower level than front margin; raised pronotal setae distinct, pubescent

antebasal band widely broken; horn without lateral pits, crest margin smooth,

apex rounded, no median ridge; raised elytral setae slightly longer than other

setae; males with simple elytra and legs, fifth sternite truncate to slightly

emarginate; parameres simple, penis divided dorso-ventrally through apical

half, dorsal thorn at apex.

Central American species included are arizonensis Fall, durangoensis n.

sp., nubilus n. sp. and toltecorum n. sp. One United States species, N.

balteatus Casey, is included. It has been most commonly collected in Arizona

near the Mexico border and probably occurs in northern Mexico. For this

reason it is separated in the key to species.

24 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus arizonensis Fall

(Figs. 13, 39)

Notoxus arizonensis Fall 1916: 36-7. Type locality: Chiricahua Mountains,

Cochise Co., Arizona. MCZC type #24326, %. Leng 1920: 163.

Notoxus talpa, Pallister 1955: 7, misidentification.

Length 3.30-4.00. Head and pronotum orange to red-brown, elytra tan to

orange with brown markings: oval mark covering omoplates, median band

rarely connected to mark on omoplates, preapical band often advancing anteri-

orly along elytral suture, rarely meeting median band. Pubescence brownish

when arising from markings, whitish over rest of body. Raised setae moder-

ately long and distinct on pronotum and elytra, average angle on elytra 31°.

Mandibular explanation moderately wide, hind margin meeting mandible at

lower level than front margin. Pronotum with pubescent antebasal band widely

broken. Horn without pits beneath, 0-2 teeth on margin. Crest margin smooth.

Apex rounded and distinct. First and second protarsomeres with strong spur

on posterior apical margin.

Males with fifth sternite slightly emarginate or often appearing only trun-

cate. Penis split dorso-ventrally near apex, with dorsal thorn at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: Nogales, Santa Cruz Co., Arizona (DSC). Length 3.31.

Head 0.68 long, eye 0.24 long, raised setae over eyes average 0.14 long;

antennomere ratios: 12.75, 111.67, I12.25, IV 1.69, V 1.93, VIII 1.44,

X 1.20, XI 2.11; fourth maxillary palpus 0.18 long, 0.09 wide; mandibular

explanation 0.11 long, 0.05 wide. Pronotum 1.08 long, length from base to

point of greatest width 0.34, break in antebasal pubescent band 0.30 wide;

horn 0.46 long, 0.32 wide at base, 1 tooth on margin; crest 0.32 long, 0.13

wide at base; crest apex 0.11 above horn margin, horn apex 0.21 from crest

apex; pronotal pit 0.08 long. Raised elytral setae 0.11 long, average angle

339. Truncation of fifth sternite 0.15 wide.

Specimens examined, 17: MEXICO: Chihuahua: Chihuahua, Wickham

(MCZC); Primavera, 550-6000', 30 June 1947, Cazier (AMNH); Santa Clara

Canyon, 5 mi W Parrita, 6 July 1954, J. W. MacSwain, Baccharis (CISC);

same locality and collector, 1 September 1956, Salix (CISC). Nayarit: La

Mesa de Navarit, 19 July 1955, B. Malkin (CISC).

Distribution: Southeastern Arizona south through the Sierra Madre

Occidental. Collected on Baccharis and Salix. Adults present from June

through August.

Discussion: Most similar to duvangoensis and nubilus by the form of the

genitalia. Separated from them by the raised elytral setae and the largely light

elytra. The lack of a postbasal band separates it from toltecorum and balteatus.

Notoxus durangoensis, n. sp.

(Figs. 14, 59)

Length 2. 78-4.35. Head and pronotum brown, elytra brown, rarely with

faint postbasal and preapical lateral markings. Pubescence whitish over body.

Raised setae distinct on pronotum, on elytra setae appressed, average angle

8°. Mandibular explanation moderately wide, hind margin meeting mandible

at lower level than front margin. Pronotum with pubescent antebasal band

widely broken. Horn without pits beneath, 0-1 teeth on margin. Crest margin

smooth, apex rounded and distinct. First and second protarsomeres with

Chandler: Notoxus of Central and South America 25

strong spur on posterior apical margin.

Males with fifth sternite slightly emarginate. Penis with apex divided

dorso-ventrally, with dorsal thorn at apex.

Females with fifth sternite rounded-triangular.

oS holotype: 25 mi W Durango, Durango, MEXICO. Length 3.63. Head

0.75 long, eye 0.26 long, raised setae above eyes average 0.10 long; antenno-

mere ratios: 12.50, 111.72, 112.14, IV 1.80, V 1.88, VUl 1.60, X 1.53, Al

2.37; fourth maxillary palpus 0.20 long, 0.12 wide; mandibular explanation

0.14 long, 0.05 wide. Pronotum 1.16 long, length from base to point of great-

est width 0.34, break in antebasal pubescent band 0.26 wide; horn 0.52 long,

0.32 wide at base, no teeth on margin; crest 0.26 long, 0.14 wide at base;

crest apex 0.12 above horn margin, horn apex 0.29 from crest apex; pronotal

pit 0.10 long. Elytral setae 0.08 long, average angle 10°. Emargination of

fifth sternite 0.20 wide, 0.01 deep.

Specimens examined, 49: HOLOTYPE %, 25 mi W Durango, 29 June 1964,

H. F. Howden (CNCI). PARATYPES: MEXICO: Durango: 20°, 10%2, eutopo-

typical (CNCI); 50°c’, 62°, same data except 23 June 1964 (CNCI); 1%, 6%, 10

mi W El Salto, 29 June 1964, H. F. Howden (CNCI); 1°, same data except 23

July 1964 (CNCI).

Distribution: Only known from the Sierra Madre Occidental in Durango.

Adults present in June and July.

Discussion: Most similar to nubilus by the appressed elytral setae and the

general form of the genitalia. It can be separated by the larger dorsal thorn

on the penis. Distinct in the group by the unicolorous elytra which rarely have

faint light lateral markings.

Notoxus nubilus, n. sp.

(Figs. 15, 59)

Length 2.88-3.48. Head brown, pronotum orange, elytra red-brown with

brown markings: basal, medial and preapical bands, basal band interrupted

along elytral suture. Pubescence brownish when arising from bands, whitish

over rest of body. Raised setae distinct on pronotum, setae on elytra appres-

sed, average angle 6°. Mandibular explanation moderately wide, hind margin

meeting mandible at lower level than front margin. Pronotum with pubescent

antebasal band widely broken. Horn without pits beneath, 1 tooth on margin.

Crest margin smooth, apex distinct, usually rounded. First and second pro-

tarsomeres with strong spur on posterior apical margin.

Males with fifth sternite truncate. Penis with apex divided dorso-ventrally,

with dorsal thorn at apex.

Females with fifth sternite rounded-triangular.

So holotype: 24 mi W La Ciudad, Durango, MEXICO. Length 3.23. Head

0.67 long, eye 0.22 long, raised setae over eyes average 0.08 long; antenno-

mere ratios: 12.33,.01.64, IN 2.27, IV 1.77, V 1.60, VOL 1.90, & 1.23,

XI 2.06; fourth maxillary palpus 0.15 long, 0.10 wide; mandibular explanation

0.10long, 0.04wide. Pronotum 0. 96long, lengthfrom base to point of greatest

width 0. 28, break in antebasal pubescent band 0. 24 wide; horn 0. 40 long, 0.25 wide

at base; crest 0.20long, 0.10 wideat base, apex pointed; crest apex 0. 08 above horn

margin, hornapex 0.24from crest apex; pronotal pit 0.08 long. Elytral setae 0.07

long, average angle 5°. Truncation of fifth sternite 0.20 wide.

Specimens examined, 3: HOLOTYPE ©, 24 mi W La Ciudad, Durango,

MEXICO, 4 June 1964, H. F. Howden (CNCI). PARATYPES: MEXICO: Dur-

ango: 10°, 1%, 24 mi W La Ciudad, 30 July 1964, H. F. Howden (CNCI, DSC).

26 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Distribution: Only known from the Sierra Madre Occiental in Durango.

Adults present from June through July.

Discussion: Similar to toltecorum and balteatus by the banding on the

elytra, but most similar to duvangoensis by the form of the genitalia. Separ-

ated from durangoensis by the faint bands on the elytra, the light pronotum and

the narrow thorn at the penis apex. Separated from the other species by the

more slender genitalia.

N. nubilus may be a population of durangoensis with larger light elytral

markings, but the genitalic differences, although slight, have been consistent

in the specimens examined.

Notoxus toltecorum, n. sp.

(Figs. 16, 59)

Length 2.82-3.50. Head and pronotum orange to brown, elytra orange.to

tan with brown markings: basal, median and preapical bands, basal band usu-

ally interrupted at elytral suture. Pubescence brownish when arising from

marks, whitish over rest of elytra. Raised setae distinct on pronotum, on

elytra appressed, average angle 109. Mandibular explanation moderately

wide, hind margin meeting mandible at lower level than front margin. Pro-

notum with pubescent antebasal band widely broken. Horn without pits beneath,

0-1 teeth on margin. Crest margin smooth, apex distinct, usually rounded.

First and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite truncate. Penis with apex divided dorso-ventrally,

with dorsal thorn at apex.

Females with fifth sternite rounded-truncate.

S holotype: 6 mi NE Zacatepec, Puebla, MEXICO. Length 3.30. Head

0.70 long, eye 0.24 long, raised setae above eyes average 0.10 long; antenno-

mere ratios: 12.50, I1 1.82, I11 2.09, I1V 1.54, V 1.65, VIII 1.28, X 1.32,

XI 2.00; fourth maxillary palpus 0.18 long, 0.10 wide; mandibular explanation

0.12 long, 0.05 wide. Pronotum 1.07 long, length from base to point of great-

est width 0.32, break in antebasal pubescent band 0.30 wide; horn 0. 46 long,

0.28 wide at base, no teeth on margin; crest 0.28 long, 0.11 wide at base,

apex pointed; crest apex 0.08 above horn margin, horn apex 0.24 from crest

apex; pronotal pit 0.08 long. Elytral setae 0.08 long, average angle 10°.

Truncation of fifth sternite 0.18 wide.

Specimens examined, 13: HOLOTYPE @, 6 mi NE Zacatepec, Puebla,

MEXICO, 27 June 1975, D. S. Chandler, meloid trap (USNM). PARATYPES:

MEXICO: Durango: 1°, Palos Colorados, 8000', 5 August 1947, Schramel

(AMNH). Distrito Federal: 12, Valley of Mexico, July 1929, Y. Mexia

(CASC). Puebla: 300°, 222, eutopotypical (DSC); 32°, same data except

cantharidin trap (DSC); 222, same data except sweeping vegetation (DSC).

Distribution: Montane areas of central and west Mexico. Adults collected

sweeping vegetation, and are attracted to cantharidin and freshly killed meloid

beetles. Present from June through August.

Discussion: Most similar to balteatus Casey by the trifasciate elytra and

the general form of the genitalia. It is separated by the abruptly swollen ven-

tral portion of the penis apex and the apically thickened parameres.

Eximius- group

The eximius-group is defined as those Notoxus with the following charac-

ters: mandibular explanation wide, the hind margin prominent; setae on pro-

Chandler: Notoxus of Central and South America ay

notum usually appressed, pubescent antebasal band widely broken, crest with-

out median ridge, margin variable; elytra with median band, elytra narrowed

in apical half, apices often acuminate. Males without modified elytra; fifth

sternite truncate to slightly emarginate; penis simple.

The species included are eximius Champion, lateralis n. sp., impressus

Champion, solus n. sp. and acuminatus Champion.

There is also an undescribed species known from the southwestern United States.

Notoxus eximius Champion

(Figs. 18, 56)

Notoxus eximius Champion 1890: 206-7, Table XI, fig. 15. Type localities:

GUATEMALA: near Guatemala City; Duenas; Zapote. NICARAGUA:

Chinandega. PANAMA: Tole. Types in BMNH, FGW, MCZC.

Pallister 1955: 6.

Notoxus solavii (sic) Pic 1916: 11. Type locality: NICARAGUA.

NEW SYNONYMY.

Length 2.07-2.95. Head and pronotum reddish-brown to dark brown,

elytra brown to dark brown with tan to light brown markings: thin, light band

across base of elytra, posthumeral band from lateral margin inward and

posterior to near elytral suture, postmedian light band curving anteriorly

from lateral margins to near suture, in lighter specimens these bands join to

form outward facing semicircles. Pubescence white when arising from mark-

ings, rest of pubescence on elytra brownish. Setae on pronotum and elytra

appressed, on elytra average angle 59. Mandibular explanation wide, with

front and hind margins on same level at mandible. Pronotum with antebasal

pubescent band widely broken. Horn without pits beneath, 6-10 teeth on mar-

gin. Crest crenulate, anterior margin pointed and indistinct. First and

second protarsomeres lacking strong spur on posterior apical margin. Elytra

slightly narrowed in apical half.

Males with fifth sternite emarginate at apex, with very short raised setae

above eyes, average length 0.02. Parameres with lateral protuberances near apex.

Females with fifth sternite narrowed to apex, with raised setae above eyes

averaging 0.12.

Redescribed “: 6 km San Jose, San Jose Province, COSTA RICA (DSC).

Length 2.54. Head 0.55 long, eye 0.18 long, raised setae over eyes average

0.02 long; antennomere ratios: I 2.00, I1 1.67, III 2.56, IV 1.90, V 2.00,

VII 1.33, K 1.19, XI 2.14; fourth maxillary palpus 0.18 long, 0.09 wide;

mandibular explanation 0.08 long, 0.05 wide. Pronotum 0.88 long, length

from base to point of greatest width 0.32, break in antebasal pubescent band

0.20 wide; horn 0.34 long, 0.22 wide at base, 6 teeth on margin; crest 0.24

long, 0.10 wide at base; crest apex 0.02 above horn margin, horn apex 0.12

from crest apex; pronotal pit 0.10 long. Elytral setae 0.06 long, average

angle of setae 69. Emargination of fifth sternite 0.18 wide, 0.04 deep.

Specimens examined, 362: MEXICO: Ixtlan del Rio. Morelos: 2 miN

Jojutla; Xochitepec. Tamaulipas: Matamoros. Veracruz: Alvarado; Vera-

cruz; Tampico; Laguna Verde. Oaxaca: La Ventosa, 72 mi E Oaxaca.

GUATEMALA: Zapote, types of eximius; Yepocapa; Gualan; Los Amates; El

Rancho. HONDURAS: La Ceiba; 12 km W Olanchito. EL SALVADOR: La

Libertad; Cerro Verde; San Salvador. NICARAGUA: San Marcos. COSTA

RICA: San Jose, 6 km NW San Jose, Rio Virilla; San Pedro de Montes de Oca.

PANAMA: Tole, part of type series (not seen). Present in all collections.

Distribution: From the neotropical portions of Mexico through central

28 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

America. Found throughout the year on a variety of plants (Cordia alba,

Sechium edrile, Inga edulis, Musa paradisiaca, Citrus sp.). Collected by

beating vegetation, sweeping shrubs on beach, and is attracted to light.

Discussion. It is the most similar to lateralis by the crenulate crest

merging into the horn and the elytral pattern. Separated from lateralis by all

the elytral setae being directed posteriorly.

Notoxus lateralis, n. sp.

(Figs. 19, 56)

Length 2.30-2.91. Head, pronotum and markings on elytra reddish-brown

to dark brown, remainder of elytra tan; thin mark arising near humeri passing

over omoplates and joining irregular midband near elytral suture, preapical

band often incomplete. Pubescence brownish when arising from dark portions

of elytra, yellowish in light areas. Setae on pronotum and elytra appressed,

on elytra average angle 79. Mandibular explanation wide, with front and hind

margins on same levelat mandible. Pronotum with antebasal pubescent band widely

broken. Horn without pits beneath, 5-9teethonhornmargin. Crestserrate, an-

terior margin usually pointed and indistinct. First and second protarsomeres lack-

ing strong spur on posterior apical margin. Elytra narrowed in apical halt.

Males with fifth sternite emarginate at apex, very short raised setae above

eyes, average length 0.02.

Females with fifth sternite triangular, raised setae over eyes longer,

average length 0.03. |

& holotype: 5 mi N Cuernavaca, Morelos, MEXICO. Length 2.56. Head

0.54 long, eye 0.17 long, raised setae over eyes average 0.02 long; antenno-

mere ratios: 12.00, 111.89, III2.10, I1V2.40, V2.00, VIII 1.40, X 1.33,

XI 2.13; fourth maxillary palpus 0.18 long, 0.10 wide; mandibular explanation

0.10 long, 0.06 wide. Pronotum 0.90 long, length from base to point of great-

est width 0.32, break in antebasal pubescent band 0.27 wide; horn 0.34 long,

0.22 wide at base, 6 teeth on margin; crest 0.20 long, 0.08 wide at base;

crest apex at same level as horn margin, horn apex 0.14 from crest apex;

pronotal pit 0.10 long. Elytral setae 0.06 long, average angle of setae 5°.

Emargination of fifth sternite 0.19, 0.02 deep.

Specimens examined, 46: HOLOTYPE ~, 5 mi N Cuernavaca, Morelos,

MEXICO, 28 August 1958, H. F. Howden (CNCI). PARATYPES: MEXICO:

Nayarit: 40°o', 322, San Blas, 6 October 1964, W. L. Nutting, at light 200

yards from beach (FGW); 422, same locality, 5 July 1972, K. Stephan (KSC).

Michoacan: 50°, 929, Uruapan, 10 August 1949, L. J. Bottimer (CNCI); 12,

Tancitaro, 6000', 27 June 1941, H. Hoogstraal (FMNH). Mexico: 192, Real

de Arriba, Temescaltepec, 6-7000', 1932 (BMNH). Morelos: 1%, Cuernavaca,

12/19 July 1961, R. & K. Driesbach (MSUC); 12, eutopotypical, beating oak

(CNCI); 722, 7 miS Tres Cumbres, 7 July 1975, D. S. Chandler, cantharidin

trap (DSC). San Luis Potosi: 1°, Xilitla, 23 July 1954, R. R. Driesbach

(MSUC). Veracruz: 1°, El Palmar, 16 km W Tetzonapa, 600', 9/15 June

1948, F. Werner & W. Nutting, jungle rubber area, beaten from vegetation

(FGW). Chiapas: 1°, 222, Junction Highways 190-195, 6 June 1969, H. F.

Howden (CNCI). No state, 1°, 12, Texpan, 7500', 12 August 1954, R. R.

Driesbach (MSUC). GUATEMALA: 198, Escuintla Tiquisate, 7 km from coast

near Junction Zanjons del Mico de Noria, 8 May 1956, T. H. Hubbell (UMMZ).

COLOMBIA: 19, Rio Frio, Magdalena, Darlington (MCZC).

Distribution: Through neotropical Central America to Colombia. This

species has been collected by beating oak and vegetation, and is attracted to

light and cantharidin. Present from May through October.

Chandler: Notoxus of Central and South America 29

Discussion: It is most similar to eximius, but it can be separated by the

presence of the laterally directed setae in the elytral postbasal impression.

The crenulate crest separates it from solus, the other species with laterally

directed setae.

Notoxus solus, n. sp.

(Figs. 20, 56)

S holotype: 10 miS Catavina, Baja California, MEXICO. Head and pro-

notum tan, elytra tan with brown markings: thin band at base of elytra, smooth

median band, wide apical band. Pubescence brownish when arising from dark

areas, tan over rest of body. Setae on pronotum and elytra appressed, on

elytra average angle 10°. Mandibular explanation moderately wide, with front

and hind margins on same level at mandible. Pronotum with pubescent ante-

basal band widely broken. Horn without pits beneath, 2 teeth on horn margin.

Crest margin smooth, anterior margin pointed and distinct. First and second

protarsomeres with strong spur on posterior apical margin. Elytra narrowed

in apical halt.

Male with fifth sternite emarginate.

Length 2.64. Head 0.62 long, eye 0.18 long, raised setae over eyes aver-

age 0.14 long; antennomere ratios: 12.33, 111.90, II 2.00, IV 2.00, V 2.10,

VIII 1.738, X 1.05, XI 1.95; fourth maxillary palpus 0.20 long, 0.12 wide;

mandibular explanation 0.12 long, 0.06 wide. Pronotum 1.00 long, length

from base to point of greatest width 0.30, break in antebasal pubescent band

0.26 wide; horn 0.40 long, width at base 0.24; crest 0.30 long, 0.15 wide at

base; crest apex 0.04 above horn margin, horn apex 0.15 from crest apex;

pronotal pit 0.06 long. Elytral setae 0.08 long. Emargination of fifth sternite

0.14 wide, 0.01 deep.

Specimens examined, 1: HOLOTYPE“, 10 miS Catavina, Baja California,

MEXICO, 29 July 1938, Michelbacher & Ross (CASC).

Distribution: The single specimen is known from the southern portion of

Baja California.

Discussion: Most similar to an undescribed species from Arizona by the

laterally directed setae in the elytral postbasal impression and the smoothly

margined crest. It can be separated by the apex of the fifth sternite being

emarginate rather than bisinuate.

Notoxus impressus Champion

(Figs. 21, 56)

Notoxus impressus Champion 1890: 206, Table IX, fig. 14. Type series

localities: GUATEMALA: Aceytuno; Guatemala City; Capetillo. Type

series in BMNH and FGW. Pic 1874: 47; 1911: 8. |

Length 1.62-3.18. Head and pronotum reddish-brown to dark brown,

elytra tan with brown markings: omoplates dark, in some specimens with

irregular midband proceeding anteriorly along elytral suture to join mark on

omoplates, mark on flanks of humeri joining midband, wide apical band.

Pubescence brownish when arising from marks on elytra, yellowish in lighter

areas. Setae on elytra and pronotum appressed, on elytra average angle 10°.

Mandibular explanation wide, with front and hind margins on same level at

mandible. Pronotum with antebasal pubescent band widely broken. Horn

30 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

without pits beneath, 6-9 teeth on horn margin. Crest crenulate, anterior

margin pointed and indistinct. First and second protarsomeres lacking strong

spur on apical posterior margin. Elytra narrowed in apical half.

Males with fifth sternite emarginate, slightly bisinuate, transverse depres-

sion immediately anterior to apical margin. Penis acutely pointed at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: Cerro Verde, EL SALVADOR (DSC). Length 3.24. Head

0.68 long, eye 0.25 long, raised setae over eyes average 0.12 long; antenno-

mere ratios: [2.28,.0.1.67, 112.50, 1V 2.50, V 2.23, VUL2.33, X 1. 90,

XI 1.94; fourth maxillary palpus 0.21 long, 0.12 wide; mandibular explanation

0.12 long, 0.07 wide. Pronotum 1.10 long, length from base to point of great-

est width 0.38, break in antebasal pubescent band 0.25 wide; horn 0.48 long,

0.28 wide at base, 6 teeth on margin; crest 0.32 long, 0.12 wide at base; crest

apex 0.01 above horn margin, horn apex 0.07 from crest apex; pronotal pit

0.14 long. Elytral setae 0.07 long, average angle 14°. Emargination of fifth

sternite 0.18 wide.

Specimens examined, 14: GUATEMALA: Capetillo, G. C. Champion,

paratype (FGW); Finca San Rafael, Sacatepeque, 6900', 28 June 1948, R. D.

Mitchell (FMNH). EL SALVADOR: Cerro Verde, 6800', 29 June 1963, D. Q.

Cavagnaro & M. E. Irwin (CASC); same locality, 18 June 1958, L. J. Bottimer

(USNM); same locality, 2000 meters, 1 May 1971, H. F. Howden (CNCI).

Distribution: The central plateau in Guatemala and El Salvador. Present

from May to June.

Discussion: It is actually quite distinct in appearance, but keys to near

eximius by the crenulate crest merging into the horn. It can be separated by

the sparse setae on the elytra, the more acuminate elytral apices and the com-

plete postmedian light band.

Notoxus acuminatus Champion

(Figs. 22, 56)

Notoxus acuminatus Champion 1890: 210-11, Table IX, fig. 19. Type locality:

Quezaltenango, 7800', GUATEMALA. BMNH. Lectotype “, designated

by Chandler 1975. Pic 1894: 46; 1911: 5.

Length 3.05-3.80. Head and pronotum tan to brown, elytra tan with brown

markings: oval marks on omoplates, often connected to irregular midband

along elytral suture, preapical band present with apices tan, mark beneath

humeri on flanks. Pubescence brownish when arising from dark areas, yellow-

ish over rest of body. Raised setae long and erect on pronotum and elytra,

average angle on elytra 42°. Mandibular explanation wide, with front and

hind margins at same level at mandible. Pronotum with pubescent antebasal

band widely separated. Horn with pits beneath, 5-9 teeth on margins. Crest

crenulate, pointed at apex, anterior margin indistinct. Elytra narrowed in

apical half.

Males with fifth sternite widely emarginate at apex, with transverse ridge

across center. Penis short, with apex slightly split into lateral parts. Lack-

ing strong spur on posterior apical margin of first and second protarsomeres.

Protibia with medial spine near middle.

Females with fifth sternite triangular, with median longitudinal ridge; first

and second protarsomeres with strong spur on posterior apical margin.

Redescribed male: Junction of Highways 190 & 195, Chiapas, MEXICO

Chandler: Notoxus of Central and South America 31

(DSC). Length 3.79. Head 0.79 long, eye 0.28 long, raised setae over eyes

average 0.19 long; antennomere ratios: 12.65, I1 1.50, HI 2.33, IV 2.26,

V 2.12, VUI1.89, X 1.47, XI 2.03; fourth maxillary palpus 0.24 long, 0.12

wide; mandibular explanation 0.12 long, 0.07 wide. Pronotum 1.28 long,

length from base to point of greatest width 0.50, break in antebasal pubescent

band 0.29 wide; horn 0.52 long, 0.39 wide at base, 6 teeth on margin; crest

0.32 long, 0.13 wide at base; crest apex 0.20 above horn margin, horn apex

0.18 from crest apex; pronotal pit 0.12 long. Raised elytral setae 0.18 long,

average angle 32°. Emargination of fifth sternite 0.22 wide, 0.02 deep.

Specimens examined, 32: MEXICO: Chiapas: Junction of Highways 190-

195, 8 May 1969, H. F. Howden (CNCI); same locality 6 June 1969, H. F.

Howden (CNCI); same locality, 24 June 1973, G. Ekis (DSC, USNM); same

locality, 32 mi W San Cristobal, 12 May 1969, H. J. Teskey (CNCI); 19 mi

SE Tuxtla Gutierrex, 23 July 1963, J. Doyen (CISC). GUATEMALA: |

Quezaltenango, 7800', Champion, Lectotype and Syntype of @cuminatus

(BMNH); Antigua, 2000 meters, 20 June 1973, G. Ekis (DSC).

Distribution: Chiapas and Guatemala. Present from May through July.

Discussion: It is distinct in the group by the presence of lateral pits on

the horn, the raised setae on the pronotum and the medial tubercle of the male

protibiae.

Monodon- group

The monodon-group is defined as those Notoxus with the following char-

acters: mandibular explanation wide, with hind margin prominent; pronotum

with distinct raised setae, pubescent antebasal band widely broken; horn with

lateral pits; crest without median ridge, margin smooth or crenulate; elytra

unicolorous or with postmedian band, some setae raised. Males lacking modi-

fications of elytra and legs; fifth sternite broadly rounded or truncate, without

impressions, parameres simple, penis often split laterally at apex.

The Central American species are desertus Casey, cumanensis LaFerte,

celatus n. sp., jamaicus Pic, bipunctatus Chevrolat, murinipennis (LeConte),

ruficollis Champion and peruvianus Pic. The United States species included

are monodon Fabricius, planicornis LaFerte and filicornis Casey.

This group is the most difficult to identify using color patterns. Examina-

tion of the genitalia and the collecting locality are the two tools used here for

species identification, particularly for the maculate species near N. monodon,

The genitalic differences are subtle, but consistent in the populations examined.

Also most of these species are allopatric and restricted to islands or to separ-

ate areas on the mainland. It is possible that these species are in reality

subspecies or ‘incipient species:'' The numerical analysis recognized these

very similar species as being distinct, which can be justified here on the

basis of the genitalic differences. There are names available for all of these

species. Further collecting may reveal areas of sympatry, which would lend

credence to the assumption that they are distinct species.

Notoxus cumanensis LaFerte

(Figs. 25, 57)

Notoxus cumanensis LaFerte 1848: 38-9. Type locality: Cumana,

VENEZUELA. Type series in MNHN.

32 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus monodon var. cumanensis, Champion 1890: 212. Pic 1911: 10.

Werner 1965b: 22.

Notoxus monodon, auctorum.

Length 2.83-3.83. Head and pronotum tan, elytra tan with brown markings:

circular mark on omoplates, postmedian irregular band protruding anteriorly

along elytral suture, anterior flanks of elytra may be darkened. Pubescence

brownish when arising from markings, whitish over rest of body. Raised

setae long and erect on pronotum and elytra, average angle on elytra 70°.

Mandibular explanation wide, with front and hind margins meeting at same

level at mandible. Pronotum with antebasal pubescent band widely broken.

Horn with pits beneath, 5-9 teeth on margin. Crest elevated and distinct,

rounded at apex. First and second protarsomeres with strong spur on posteri-

or apical margin.

Males with fifth sternite broadly rounded. Penis split laterally at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: Rio Hato, PANAMA (DSC). Length 3.14. Head 0.64 long,

eye 0.28 long, raised setae over eyes average 0.19 long; antennomere ratios:

12,47, 114.73, th 2-06, 1V 1.92, Vilet, Vill 1.13, % 1. 29, Al 2.22; fourth

maxillary palpus 0.19 long, 0.09 wide; mandibular explanation 0.10 long,

0.06 wide. Pronotum 1.07 long, length from base to point of greatest width

0.36, break in antebasal pubescent band 0.22 wide; horn 0.42 long, 0.28 wide

at base, 6 teeth on margin; crest 0.24 long, 0.11 wide at base; crest apex

0.08 above horn margin, horn apex 0.24 from crest apex; pronotal pit 0.16

long. Raised elytral setae 0.20 long, average angle 71°.

Specimens examined, 39: MEXICO: Tamaulipas: Victoria; 3 mi N Cd.

Victoria, 800'; 47 km S Cd. Victoria, 900'; 8.5 miS Soto la Marina; Santa

Engracia; Gomez Farias. San Luis Potosi: Cuesta de los Cedros, 36 km E

Cd. del Maiz, 2250'; El Salto de Agua, 100'; 8.7 miS Santa Mario del Rio.

Veracruz: El Palmar, 16 km W Tetzonapa, 600'; Jalapa; Orizaba; Cotaxtla

Experiment Station, Cotaxtla. Tabasco: 59.4 mi SE Villahermosa, 100'.

Oaxaca: Salina Cruz. BELIZE: Rio Hondo, Bluncaneau. HONDURAS: La

Ceiba. PANAMA: Old Panama; Rio Hato: Canal Zone, Ancon. Collected in

quarantine on orchids from COLOMBIA. Specimens in collections of: AMNH,

CASC, CISC, CNCI, DSC, FGW, FMNH, MCZC, TAMU, TTCC, USNM.

Distribution: The moister areas of the east slope of the Sierra Madre

Oriental in Mexico, south through Central America to Venezuela. Apparently

present throughout the year. Attracted to light, ultra-violet light, arc light

and cantharidin.

Discussion: Very similar in appearance to the other banded species in the

eroup. The genitalia are similar to monodon and jamaicus by the laterally

split apex of the penis. N. jamaicus is an island species with the parameres

as long as the penis. N. cumanensis has the penis distinctly longer than the

parameres and only a small dorsal hump near the penis apex. N. monodon

has the parameres protruding beyond the penis and has a small dorsal tubercle

near the penis apex.

Notoxus celatus, n. sp.

(Figs. 24, 57)

Length 2. 70-3.68. Head and pronotum tan, elytra tan with brown mark-

ings: circular mark on omoplates, irregular postmedian band protruding

Chandler: Notoxus of Central and South America 33

anteriorly along elytral suture, mark on flanks of elytra behind humeri.

Pubescence brownish when originating in markings, yellowish over rest of

body. Raised setae long and erect on pronotum and elytra, average angle on

elytra 649. Mandibular explanation wide, with front and hind margins meeting

on same level at mandible. Pronotum with antebasal pubescent band widely

broken. Horn with pits beneath, 3-8 teeth on margin. Crest elevated and

distinct, rounded at apex. First and second protarsomeres with strong spur

on posterior apical margin.

Males with fifth sternite broadly rounded. Penis split dorso-ventrally at

apex, with strong dorsal tooth near apex.

Females with fifth sternite rounded-triangular.

So holotype: 2 mi N Jojutla, Morelos, MEXICO. Length 2.99. Head 0.64

long, eye 0.25 long, raised setae over eyes average 0.18 long; antennomere

ratios: 12,47, 11 1,64, Tb 2.27, 1¥ 2.09, V¥ 1.92, VUlL1. 67, 21.01, Al2.1G

fourth maxillary palpus 0.20 long, 0.10 wide; mandibular explanation 0. 09

long, 0.06 wide. Pronotum 0.96 long, length from base to point of greatest

width 0.30, break in antebasal pubescent band 0.20 wide; horn 0.38 long, 0.26

wide at base, 6 teeth on margin; crest 0.18 long, 0.10 wide at base; crest apex

0.08 above horn margin, horn apex 0.22 from crest apex; pronotal pit 0.14

long. Raised elytral setae 0.17 long, average angle 62°.

Specimens examined, 241: HOLOTYPE GC, 2 mi N Jojutla, Morelos,

MEXICO, 20 July 1974, R. L. Mangan & D. S. Chandler, cantharidin trap

(USNM). PARATYPES: Chihuahua: Catarinas, 5800'. Jalisco: 3 mi SE

Plan de Barrancas. Michoacan: 12 miS Tzitzio on Huetamo Road, 1050

meters; Tuxpan, 6720'. Guerrero: 8 mi SE Iguala; Iguala. Mexico: 2 mi

NE Ixtapan de la Sal; Real de Arriba, Temescaltepec, 6-7000'; Tejupilco, ca.

400'; Temescaltepec, ca. 5000'. Morelos: Xochitepec; Cuernavaca; 3 mi N

Alpuyeca, 3400'; 5 mi N Axochiapan; 2 mi N Jojutla; 4.4 mi E Cuernavaca;

10 mi E Cuernavaca; 7 mi E Cuernavaca. Puebla: Atlixco; Puebla; 4 mi NW

Tehuitzingo, 3500'; 13.3 mi NE Tehuitzingo; 1.1 mi W Acatlan; 11 mi SE

Acatlan; 7 mi SW Izucar de Matamoros; 12 mi SW Izucar de Matamoros; 11.8

mi W Izucar de Matamoros; 5 mi SW Chipilo; Tehuacan, ca. 5200'; 6 mi SW

Tehuacan; 4 miS Petalcingo, 5150'. Oaxaca: Huajuapan de Leon; 10 mi NE

Huajuapan de Leon. One specimen, state unknown, Rio Blanco. Paratypes

in collections of: AMNH, BMNH, CASC, CISC, CNCI, CUIC, DSC, FGW,

FMNH, MCZC, MSUC, TAMU, TTCC, UCDC, UMMZ, USNM.

Distribution: The higher elevations of central Mexico, extending north to

Chihuahua. It has been collected from April to August by beating vegetation,

mesquite, acacia and looking under recently uprooted weeds. Attracted to

light, ultra-violet light, cantharidin and drying meloid beetles.

Discussion: It is similar in external appearance to the other banded

species in the group. It is separated from these species by having the penis

split laterally and dorsoventrally at the apex. There is a dorsal, anteriorly

directed process near the apex of the penis which is also not found in the other

species.

Notoxus desertus Casey

(Pics, 33.37)

Notoxus desertus Casey 1895: 767-8. Type locality: Tucson, Arizona. USNM

Type #36536. Leng 1920: 163.

34 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus constrictus Casey 1895: 768. Type locality: coast regions of Cali-

fornia. USNM Type #36527. Leng 1920: 163. Fall 1902: 33, 182.

Length 2.83-4.18. Head and pronotum tan, elytra tan with brown mark-

ings: circular mark on omoplates, irregular postmedian band protruding an-

teriorly along elytral suture, form of band variable, may reach anteriorly

almost to omoplates or may have straight or irregular margins, mark may be

present along anterior flanks. Pubescence brownish when arising from mark-

ings, yellowish over rest of body. Raised setae long and erect on pronotum

and elytra, average angle on elytra 72°. Mandibular explanation wide, with

front and hind margins on same level at mandible. Pronotum with antebasal

pubescent band widely broken. Horn with pits beneath, 4-9 teeth on margin.

Crest elevated and distinct, usually faintly crenulate, rounded at apex. First

and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite broadly rounded.

Females with fifth sternite rounded-triangular.

Redescribed “: Rio Vista, California (DSC). Length 3.22. Head 0.66

long, eye 0.23 long, raised setae over eyes average 0.21 long, antennomere

ratios: -[f2.05,-1) 1.91, Tl 2.27, IV 2.29, V 2.00, VUlL1. 67, Xx 1.24, Xl 2.35;

fourth maxillary palpus 0.20 long, 0.10 wide; mandibular explanation 0.10

long, 0.06 wide. Pronotum 1.02 long, length from base to point of greatest

width 0.30, break in antebasal pubescent band 0.24 wide; horn 0. 41 long,

0.30 wide at base, 8 teeth on margin; crest 0.26 long, 0.12 wide at base;

crest apex 0.11 above horn margin, horn apex 0.19 from crest apex; pronotal

pit 0.18 long. Raised setae 0.21 long, average angle 66%.

Specimens examined, 344: MEXICO: Baja California: Mision; Hamilton

Ranch; Ensenada. Baja California Sur: Mulege; 15 mi N San Ignacio; La Paz;

12 mi NW La Paz; 3.5 mi NE San Pedro; 2 mi SE Todos Santos. Sonora:

Alamos: 13 mi SE Alamos; Navajoa; 10 mi E Navajoa; 16 mi NE Cd. Obregon;

Saric; Sinaloa: San Javier; Mazatlan; 5 mi N Mazatlan; Los Mochis; 13 mi N

Los Mochis; Choix; 5.5 mi NW Choix; 35 mi S Escuinapa; 21 mi E Villa Union;

Venedio; 26 mi N Pericos; Los Mayos; 20 mi E Guasave; 5 mi E Concordia;

1 mi W San Blas; Culiacan. Nayarit: San Blas; Rio de las Canyas, 8 mi NW

Acaponeta; Tepic; Arroyo Santiago, nr. Jesus Maria; Arroyo Canaveral, nr.

Jesus Maria; La Mesa de Nayarit. Chihuahua: Delicias; 10 miS Delicias.

Durango: 2 mi NW Nombre de Dios. Zacatecas: 4 mi E Zacatecas; 15 mi

NE Fresnillo; 28 miS Jalpa. Coahuila: 10 miS Allende; Boquillas del

Carmen; 12 mi N Hermanas. Nuevo Leon: Monterrey; 5 miS Monterrey;

Apodaca; 6 km § Galeana; Linares, Rio Camacho; Mamulique Pass.

Tamaulipas: Tampico; 39.5 miS Jaumave. Hidalgo: Zimapan; 1 mi SE

Yolotepec. Queretaro: 12 mi N Vizarron; Tequisquiapan. Guanajuato: 3 mi

N Irapuato. Aguascalientes: Aguascalientes. Jalisco: Lagos de Moreno;

Oblatos Canyon, Guadalajara; Chapala; Tecalitlan; Catalitlan. Oaxaca:

Oaxaca; 15 mi NW Oaxaca; Mitla; 11.3 mi SE Totolapan; Yagul ruins. Vera-

cruz: Veracruz; Sontecomapan; Jicacal, 7 mi N Sontecomapan; 2 mi NE

Catemaco; 2 mi SE Nautla. Tabasco: Frontera. GUATEMALA: Atitlan-see.

Specimens in collections of: AMNH, CASC, CDAE, CISC, CNCI, DSC, FGW,

FMNH, LACM, MCZC, MSUC, OSUC, TAMU, UCDC, USNM.

Distribution: Found through all of the United States except Southeast,

extending south through most of Mexico to Guatemala. Present year-round.

This species does not appear to occur in the moister areas of the Sierra Madre

Oriental and the region centered by Mexico City. It has been collected sweep-

ing, under low vegetation in corn fields, beating, on Hoffmannseggia, on

Chandler: Notoxus of Central and South America 35

Helenium nudifloris, on Medicago, on roses and on shelled corn. It is

attracted to light, ultra-violet light, drying meloid beetles and cantharidin.

Discussion: It is similar to all of the banded species in the group due to

the variation in the elytral color pattern. The penis is not split laterally at

the apex and is most similar to bipunctatus. It is separated from that species

by being present on the mainland and lacking the more extensive dark areas on

the elytra.

Notoxus bipunctatus Chevrolat

(Figs. 26, 57)

Notoxus bipunctatus Chevrolat 1877: ix. Type locality: Puerto Rico. Type in

MNHN. Pic 1894: 46; 1911: 6.

Notoxus Krugi (sic) Quedenfeldt 1886: 121-2. Type locality: Puerto Rico.

Pic 1894: 47; 1911: 47.

Length 2.80-3.38. Head and pronotum yellow-brown, elytra yellow-brown

with brown markings: usually with oval mark on omoplates, very irregular

postmedian band, usually anterior flanks of elytra darkened. Pubescence

brownish when arising from marks, yellowish over rest of body. Raised setae

long and erect on pronotum and elytra, average angle on elytra 44°. Mandibu-

lar explanation moderately wide, with front and hind margins meeting on same

level at mandible. Pronotum with antebasal pubescent band widely broken.

Horn with pits beneath, 4-6 teeth on margin. Crest distinct and elevated,

pointed or rounded at apex. First and second protarsomeres with strong spur

on posterior apical margin.

Males with fifth sternite broadly rounded. Penis slightly split laterally

at apex.

Females with fifth sternite rounded-triangular.

Redescribed ©: Ponce, PUERTO RICO (DSC). Length 3.22. Head 0.62

long, eye 0.28 long, raised setae over eyes average 0.16 long; antennomere

ratios: 12.00, 121,86, Ul2.00, 1V 4,.99,.V 4.08, Vil 1.71, %-1,-98,, 41.2. 60;

fourth maxillary palpus 0.18 long, 0.10 wide; mandibular explanation 0.08

long, 0.06 wide. Pronotum 0.92 long, length from base to point of greatest

width 0.29, break in antebasal pubescent band 0.30 wide; horn 0.37 long, width

at base 0.28, 4 teeth on margin; crest 0.22 long, 0.12 wide at base; crest

apex 0.07 above horn margin, horn apex 0.16 from crest apex; pronotal pit

0.12 long. Raised elytral setae 0.13 long, average angle 40°.

Specimens examined, 18: PUERTO RICO: Ponce, 1/15 June 1945, J. M.

Capriles, light trap (USNM); same locality, 21 September 1933, R. G. Oakley,

Scirpus validus (DSC, USNM); same data except, San Juan, 31 August 1933

(USNM); Coamo Springs, 2 April 1931, S. T. Danforth (MCZC); same locality,

11 February (AMNH); Rio Piedras, July 1931, Alsina (MCZC); Anasco district,

3 July 1917, H. Morrison (USNM).

Distribution: The island of Puerto Rico. Collected on Scirpus validus and

attracted to light. Present year-round.

Discussion: It is most similar to desertus in having the penis without a

deep lateral split, although it does have a notch at the apex. Separated from

the other species by its occurrence on the island of Puerto Rico.

36 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus jamaicus Pic

(Wies, 27, 57)

Notoxus jamaicus Pic 1913: 8-9. Type locality: Jamaica. Type in MNHN.

Notoxus quinquemaculatus Pic 1918: 117. Type locality: Jamaica. Type in

MNHN. NEW SYNONYMY.

Length 2.70-3.50. Head and pronotum yellow-brown to dark red-brown,

pronotum dark brown in lower half, elytra yellow-brown with dark red-brown

markings: oval mark on omoplates, rectangular mark along sides of elytral

suture at middle, lateral oval marks at end of this mark, these marks often

connected together and with mark on omoplates, mark on flanks of elytra

curving inward to elytral suture near apex. Pubescence brownish when arising

from dark portions of body, yellowish over rest of body. Raised setae long

and erect on pronotum and elytra, average angle on elytra 539. Mandibular

explanation wide, with front and hind margins on same level at mandible.

Pronotum with antebasal pubescent band widely broken. Horn with pits be-

neath, 3-7 teeth on margin. Crest distinct and elevated, usually rounded at

apex. First and second protarsomeres with strong spur on posterior apical

margin.

Male fifth sternite broadly rounded. Penis split laterally at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: Alligator Pond, JAMAICA (DSC). Length 2.95. Head

0.66 long, eye 0.28 long, raised setae over eyes average 0.15 long; antenno-

mere ratios: 12.18, I1 1.73, Ill 2.00, IV2.00, V 2.00, VII1 1.89, X 1.44,

XI 2.24; fourth maxillary palpus 0.18 long, 0.12 wide; mandibular explanation

0.09 long, 0.06 wide. Pronotum 0.93 long, length from base to point of great-

est width 0.28, break in antebasal pubescent band 0.28 wide; horn 0.38 long,

0.32 wide at base, 4 teeth on margin; crest length 0.22, width at base 0.12;

crest apex 0.08 above horn margin, horn apex 0.17 from crest apex; pronotal

pit 0.16 long. Raised elytral setae 0.15 long, average angle 49°.

Specimens examined, 23: JAMAICA: Alligator Pond, 20 February,

Chaplin & Blackwelder (DSC, USNM); Alligator Pond Bay, 25 February 1937,

Station 410, Chaplin & Blackwelder (USNM); Santa Cruz, 24 February 1937,

Station 421, Chaplin & Blackwelder (USNM); Morant Bay, 6 February 1937,

Station 391, Chaplin & Blackwelder (DSC, USNM); Spanish Town, 2 February

1936, Station 377, Chaplin & Blackwelder, flying at dusk (USNM); Milk River,

25 February 1937, Station 415, Chaplin & Blackwelder (USNM); Trelawney,

Duncans, 4 August 1966, Howden & Becker (CNCI); same locality, 7 August

1966, A. T. Howden, at light (HAHC); St. Andrew Parish, Bull Run, 19 April

1959, Farr & Sanderson, on Agave americana (FGW).

Distribution: The island of Jamaica. Collected on Agave and attracted to

light. Probably present year-round.

Discussion: It is most similar to cumanensis and monodon by the penis

being laterally split at the apex. The parameres are as long as the penis

which places it as an intermediate between the other two species. There is no

trace of a dorsal hump near the penis apex.

Chandler: Notoxus of Central and South America 3t

Notoxus murinipennis (LeConte)

(Figs. 28, 57)

Anthicus bicolor Say 1817: plate IV (not bicolor Oliver). Type locality: New

Jersey and Pennsylvania. Say 1824: 21; 1827: 38.

Anthicus murinipennis LeConte 1824: 170. Type locality: Georgia. MCZ

Type #4889.

Monocerus bicolor, LeConte 1847: 90.

Notoxus bicolor LaFerte 1848: 53. LeConte 1852: 94. Horn 1884: 167.

Champion 1890: 205. Casey 1895: 756. Pic 1911: 6. Leng 1920: 162.

Notoxus murinipennis, Barber 1941: 26.

Length 2.91-3.90. Head and elytra varying from tan to blue-brown,

pronotum varying from tan to orange. Pubescence whitish over all of body.

Raised setae distinct on pronotum, of moderate length on elytra, average

angle 359. Mandibular explanation wide, with front and hind margins meeting

on same level at mandible. Pronotum with antebasal pubescent band widely

broken. Horn with pits beneath, 6-10 teeth on margin. Crest margin crenu-

late, front rarely indistinct and merging with horn, front pointed. First and

second protarsomeres with strong spur on posterior apical margin.

Male with fifth sternite broadly rounded at apex. Penis slightly split

laterally at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: 2 mi E Roper, North Carolina (DSC). Length 3.34.

Head 0.67 long, eye 0.28 long, raised setae over eyes average 0.14 long;

antennomere ratios: 12.92, 112.00, Il 2.45, 1V1.61, V 1.78, VIII 1.86,

X 1.37, XI 2.33; fourth maxillary palpus 0.18 long, 0.10 wide; mandibular

explanation 0.08 long, 0.05 wide. Pronotum 1.10 long, length from base to

point of greatest width 0.38, break in antebasal pubescent band 0.26 wide;

horn 0.46 long, 0.32 wide at base, 10 teeth on margin; crest 0.32 long, 0.15

wide at base; crest apex 0.05 above horn margin, horn apex 0.15 from crest

apes pronotal pit 0.18 long. Raised elytral setae 0.11 long, average angle

aot.

Specimens examined, 19: MEXICO: San Luis Potosi: Tamazunchale, Rt.

1, km 365, 31 May 1948, F. Werner & W. Nutting, river in tropical area,

light trap (FGW); same locality, 1 May 1955, Buliofer & Guelo (USNM).

Veracruz: Cordoba, 19 April 1908, A. Fenyes (USNM); same data except,

April 30, 1908 (USNM); same locality (CASC); Fortin de las Flores, 24 De-

cember 1963, L. B. & C. W. O'Brien (CASC); same locality, 28 June 1975,

D. S. Chandler, meloid trap, ultraviolet light (DSC). Oaxaca: 11 miN

Matias Romero, 6 June 1971, Clark, Murray, Hart, Schaffner (TAMU).

GUATEMALA: Gualan, 13 February 1906, J. S. Hine (OSUC). Collected in

peecee os at Brownsville, Texas, with avocados and on bag of green pepper

USNM).

Distribution: From the southeastern portion of the United States through

the moist eastern slope of the Sierra Madre Oriental to Guatemala. Appar-

ently present throughout the year. Collected by beating vegetation and

attracted to harvested crops, ultra-violet light and drying meloid beetles.

Discussion: It is most similar to ruficollis by the short raised elytral

setae and the unicolorous elytra. It can be separated by the unicolorous pro-

notum and the parameres paralleling the penis.

38 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus ruficollis Champion

(Figs. 29, 57)

Notoxus ruficollis Champion 1890: 205, Table XI, fig. 13. Type locality:

San Lorenzo, Chiriqui, PANAMA. Type series in BMNH. Pic 1894:

47; 1911: 11.

Notoxus atripennis Champion 1890: 205. Type locality: Chacoj, Polochic

Valley, GUATEMALA. Type in BMNH, ¢. Pic 1894: 46; 1911: 6.

NEW SYNONYMY.

Length 2.41-2.46. Head, horn on pronotum and elytra blue-orange, rest

of pronotum dusky-orange. Pubescence whitish over all of body. Raised setae

distinct on pronotum, of moderate length on elytra, average angle 35°. Mandi-

bular explanation wide, with front and hind margins meeting at same level at

mandible. Pronotum with antebasal pubescent band widely broken. Horn with

pits beneath, 9-11 teeth on margin. Crest margin crenulate, front indistinctly

merging with horn. First and second protarsomeres with strong spur on pos-

terior apical margin.

Male with fifth sternite broadly rounded. Apical half of parameres bent

laterally.

Female with fifth sternite rounded-triangular.

Redescribed “: Trece Aguas, Alta Vera Paz, GUATEMALA (USNM).

Length 2.41. Head 0.72 long, eye 0.28 long, raised setae over eyes average

0.14 long; antennomere ratios: 12.15, 111.78, I] 1.78, 1V 1.44, V 1.409,

VII 1.67, X 1.28, XI 2.05; fourth maxillary palpus 0.20 long, 0.11 wide;

mandibular explanation 0.12 long, 0.07 wide. Pronotum 1.12 long, length

from base to point of greatest width 0.46, break in antebasal pubescent band

0.22 wide; horn 0.38 long, 0.28 wide at base, 11 teeth on margin; crest 0.295

long, 0.13 wide at base; crest apex merging with horn; horn apex 0.10 from

crest apex; pronotal pit 0.24 long. Raised elytral setae 0.07 long, average

angle 35°.

Specimens examined, 2: GUATEMALA: Cacao, Trece Aguas, Alta Vera

. Paz, April 1923, Schwarz & Barber (USNM); Chacoj, Vera Paz, Champion,

Holotype of Notoxus atripennis Champion (BMNH).

Distribution: From Guatemala to Panama.

Discussion: It is most similar to murinipennis. It can be separated by

the horn being abruptly darker than the pronotum and the laterally spread

parameres.

Notoxus peruvianus Pic

(Figs. 30, 58)

Notoxus peruvianus Pic 1904: 228. Type locality: Cuzco, PERU. Type in

MNHN. Pic 1914: 183.

Length 2.70-3.74. Head and pronotum tan, elytra tan to blue-orange.

Pubescence yellowish. Raised setae long and erect on pronotum and elytra,

average angle on elytra 699. Mandibular explanation wide, with front and

hind margins meeting on same level at mandible. Pronotum with antebasal

pubescent band widely broken. Horn with pits beneath, 6-11 teeth on margin.

Chandler: Notoxus of Central and South America 39

Crest crenulate, anterior margin usually distinct, apex usually pointed. First

and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite broadly rounded at apex.

Females with fifth sternite rounded-triangular.

Redescribed “: 16 km N Barranca, PERU (DSC). Length 3.60. Head

0.75 long, eye 0.32 long, raised setae over eyes average 0.20 long; antenno-

mere ratios: 12.30, 111,40, I1.53, 1V-1.39,7¥' 2-07, Vill 1. 70... 145.

XI 2.15; fourth maxillary palpus 0.20 long, 0.11 wide; mandibular explanation

0.12 long, 0.06 wide. Pronotum 1.14 long, length from base to point of great-

est width 0.38, break in antebasal pubescent band 0.26 wide; horn 0.48 long,

0.28 wide at base, 6 teeth on margin; crest 0.30 long, 0.11 wide at base;

crest apex 0.02 above horn margin, horn apex 0.18 from crest apex; pronotal

pit 0.18 long. Raised elytral setae 0.19 long, average angle 72°.

Specimens examined, 39: PERU: 16kmN Barranca, 15 March 1951,

Ross & Michelbacher (CASC); Casma, 24 March 1951, Michelbacher & Ross

(CASC); 10 km S Chiclayo, 19 March 1951, Ross & Michelbacher (CASC);

Amotape, 13 November 1938, D. & H. Frizzell (FGW); Chancay, 25 March

1951, Ross & Michelbacher, river valley (CASC); Lima, J. Wille (USNM);

Lima, 13 March 1931, F. Wille (USNM); Yantan, September 1929, Wille

(USNM); Canete, May and June 1941, C. P. Clause (USNM); Montevideo,

South America Paras Laboratory, 30 December 1941, Berry (USNM).

BOLIVIA: Chaco-Chalumani, 25 May, G. L. Harrington (CASC).

Distribution: Peru and Bolivia. Apparently present throughout the year.

Discussion: It is most similar to murinipennis and ruficollis by the uni-

colorous elytra. It can be separated by the longer and more erect setae of the

elytra.

Nuperus-group

The nuperus-group is defined as those Notoxus with the following char-

acters: mandibular explanation narrow to wide; pronotal tactile setae short

and distinct, usually uncommon, pubescent antebasal band widely broken;

horn without lateral pits; crest margin smooth, apex pointed or rounded; ely-

tral setae appressed; males with simple elytra and legs; fifth sternite emar-

ginate, circular impression before apex.

The species included in the group are nuperus Horn, haustrus n. sp.,

posticus n. sp. and orientalis n. sp.

Notoxus nuperus Horn

(Figs. 32, 60)

Notoxus nuperus Horn 1884: 168. Type locality: Arizona. MCZC Type

#3043, So. Pic 1894: 47; 1911: 10. Casey 1895: 757. Leng 1920: 162.

Pallister 1955: 11, in part.

Length 3.30-4.48. Head and pronotum tan to red-brown, elytra tan with

brown markings: postmedian band extending anteriorly along elytral suture to

omoplates, apical band extending along elytral suture to postmedian band,

flanks often dark. Pubescence brownish when arising from marks, whitish

over rest of body. Raised setae uncommon but distinct on pronotum, setae on

elytra appressed, average angle 99. Mandibular explanation moderately wide,

40 Contrib, Amer. Ent. Inst., vol. 15, no. 3, 1977

often front and hind margins meet mandible at same level. Pronotum with

antebasal band widely broken. Horn without pits beneath, 0-2 teeth on margin.

Crest margin smooth, apex distinct, pointed or rounded. First and second

protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite deeply emarginate into deep circular impression.

Fourth sternite often slightly flattened medially.

Females with fifth sternite rounded-triangular.

Redescribed “: Texas Canyon, Cochise County, Arizona (DSC). Length

3.596. Head 0.67 long, eye 0.26 long, raised setae over eyes average 0.10

long; antennomere ratios: 12.67, 111.83, Il 2.00, IV 1.78, V 2.00, VIII

1.40, X 1.13, XI 2.24; fourth maxillary palpus 0.20 long, 0.10 wide; mandibu-

lar explanation 0.12 long, 0.05 wide. Pronotum 1.09 long, length from base

to point of greatest width 0.32, break in antebasal pubescent band 0.28 wide;

horn 0.48 long, 0.33 wide at base, no teeth on margin; crest 0.35 long, 0.14

wide at base, apex pointed; crest apex 0.08 above horn margin, horn apex

0.16 from crest apex; pronotal pit 0.12 long. Elytral setae 0.08 long, average

angle 8°. Emargination of fifth sternite 0.18 wide, 0.06 deep.

Specimens examined, 78: MEXICO: Chihuahua: 5 mi NE Hidalgo del

Parral, 6000'; Catarinas, 5800'; Valle de Olivos, 5500'; Primavera, 5500-

6000'; 6 miS Villa Matamoros; 15 mi E Cuauhtemoc, 6600'; Santa Clara Can-

yon, 0 mi W Parrita. Durango: Tepehuanes; Durango; 5 mi W Durango; 23

miS Durango. Jalisco: 5 km W Lagos, 6000'. Specimens in collections of:

AMNH, CASC, CISC, CNCI, DSC, FGW, MCZC, TTCC, UCDC, USNM.

Distribution: From southeastern Arizona through the western half of the

central plateau of Mexico. Attracted to white and ultra-violet light. Present

from June through August.

Discussion: Very similar to haustrus by the deep impression of the male

fifth sternite and the general external appearance. It is separated by having

at the most the fourth sternite slightly flattened, the presence of some erect

setae on the pronotum, and the mandibular explanation being moderately wide

to wide. Similar to orientalis in the appressed setae on the elytra, but easily

separated by the deep impression on the male fifth sternite.

Notoxus haustrus, n. sp.

(Figs. 33, 60)

Notoxus nuperus, Pallister 1955: 11, in part.

Length 2.90-4.38. Head and pronotum tan to orange-brown, elytra tan

with brown markings: irregular postmedian band extending anteriorly along

elytral suture, flanks often dark, apical band extending anteriorly along ely-

tral suture. Pubescence brownish when arising from marks, whitish over

rest of body. Setae on pronotum and elytra appressed, on elytra average

angle 109. Mandibular explanation narrow, with hind margin indistinct and

meeting mandible at lower level than front margin. Pronotum with antebasal

pubescent band widely broken. Horn without pits beneath, 0-2 teeth on mar-

gin. Crest margin smooth, apex rounded and distinct. First and second pro-

tarsomeres with strong spur on posterior apical margin.

Males with fifth sternite deeply emarginate into large circular impression.

All sternites usually medially flattened.

Females with fifth sternite rounded-triangular.

S holotype: Colossal Cave County Park, Pima County, Arizona. Length

Chandler: Notoxus of Central and South America 41

3.59. Head 0.70 long, eye 0.30 long, raised setae above eyes average 0.10

long; antennomere ratios: 12.35, 111.54, 111.92, 1V 1.33, V 1.47, VIII

1.14, X 0.83, XI 1.70; fourth maxillary palpus 0.20 long, 0.10 wide; mandibu-

lar explanation 0.10 long, 0.05 wide. Pronotum 1.05 long, length from base

to point of greatest width 0.35, break in antebasal pubescent band 0.38 wide;

horn 0.50 long, 0.33 wide at base, no teeth on margin; crest 0.26 long, 0.16

wide at base; crest apex 0.12 above horn margin, horn apex 0.19 from crest

apex; pronotal pit 0.10 long. Elytral setae 0.08 long, average angle 8°.

Emargination of fifth sternite 0.24 wide, 0.08 deep.

Specimens examined, 225: HOLOTYPE %, Colossal Cave County Park,

Pima Co., Arizona, 8 July 1973, K. Stephan, UV light (USNM). PARATYPES:

Montana: Broadwater Co.: Toston. Utah: San Juan Co.: Monticello; Poverty

Flats. New Mexico: McKinley Co.: Tohatchi; Gallup. Bernalillo Co.: Albu-

querque; 7 mi NE Albuquerque. Dofia Ana Co.: Las Cruces. Luna Co.: 40

miS Deming. Arizona: Apache Co.: Ganado. Gila Co.: Cutter; Globe;

Claypool. Graham Co.: Bonita. Yavapai Co.: Dewey; Kirkland; Skull Valley.

Pinal Co.: 5 miN Mammoth. Pima Co.: Tucson; Corona de Tucson; I. B.

P. site, Santa Rita Range Reserve; Continental; 7 mi SE Continental; 5 mi E>

Sahuarita; 4 mi NNW Redington; Arizona-Sonora Desert Museum; mouth of

Bear Canyon, Santa Catalina Mtns.; Chutum Vaya Canyon, Baboquivari Mtns. ;

Brown's Canyon, Baboquivari Mtns.; Bog Springs, Madera Canyon. Santa

Cruz Co.: Tumacacori; 8 mi N Nogales; Sycamore Canyon, nr. Ruby; Pefia

Blanca, 10 mi W Nogales; 1 miS Patagonia; 10 miS Patagonia; Canelo.

Cochise Co.: Pearce; Willcox; Cochise Stronghold, Dragoon Mtns. ; Bowie;

2 mi W entrance Chiricahua National Monument. MEXICO: Sonora: Hermo-

sillo; 40 mi N Hermosillo; Saric; Navajoa; 20 mi NE Obregon. Sinaloa: Los

Mochis. Chihuahua: Santa Clara Canyon, 5 mi W Parrita; 3 miS Villa

Ahumada; Chihuahua; 5 mi N Cerro Campana; Primavera; Delicias; 25 mi SW

Camargo; 5 miS Galeana. Zacatecas: Laguna Balderama, 25 mi W Fresnillo.

Paratypes in collections of: AMNH, CASC, CDAE, CISC, DSC, FGW, MCZC,

OSUC, TTCC, USNM.

Distribution: The Great Plains to northwestern Mexico. Collected on

cotton, alfalfa, Salix, Baccharis glutinosa, Prosopis glandulosa, Chilopsis

lineavis and Sapindus saponaria. Attracted to light, ultra-violet light and

drying meloid beetles (Werner 1964 as N. nuperus). Present from June to

September.

Discussion: It is most similar to nuperus. Separated by the narrow man-

dibular margin, the appressed setae on the pronotum, and by usually having

the first four sternites medially flattened in the male, along with the differences

in horn structure used in the key to species. Nuperus and haustrus are the

only species of the nuperus-group found on the western half of the Mexican

central plateau.

Notoxus postictus, n. sp.

(Figs. 34, 60)

Length 2.80-3.39. Head and pronotum orange to brown, elytra tan with

brown markings: circular mark on omoplates, postmedian band slightly

curved anteriorly to elytral suture, occasionally reaching marks on omo-

plates, apical band often meeting postmedian band along elytral suture, flanks

usually dark. Pubescence brownish when arising from marks, whitish over

rest of body. Raised setae distinct on pronotum, setae on elytra decumbent,

42 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

average angle of setae at greatest elevation 209. Mandibular explanation nar-

row, hind margin meeting mandible on lower level than front margin. Prono-

tum with antebasal pubescent band widely broken. Horn without pits beneath,

0-4 teeth on margin. Crest margin smooth, apex distinct, usually rounded.

First and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite deeply emarginate into deep, slightly transverse

impression.

Females with fifth sternite rounded-triangular.

% holotype: Santa Engracia, Tamaulipas, MEXICO. Length 3.48. Head

0.72 long, eye 0.30 long, raised setae above eyes average 0.12 long; antenno-

mere ratios: 12.17, 111.67, IIl 2.31, 1V 1.47, V 1.47, VUI1.20, X 0.87,

XI 1.75; fourth maxillary palpus 0.20 long, 0.11 wide; mandibular explanation

0.10 long, 0.04 wide. Pronotum 1.10 long, length from base to point of great-

est width 0.37, break in antebasal pubescent band 0.27 wide; horn 0. 44 long,

0.31 wide at base, 1 tooth on margin; crest 0.26 long, 0.12 wide at base,

apex pointed; crest apex 0.09 above horn margin, horn apex 0.22 from crest

apex; pronotal pit 0.13 long. Raised setae 0.10 long, average angle 19°.

Emargination of fifth sternite 0.16 wide, 0.04 deep.

Specimens examined, 38: HOLOTYPE ©, Santa Engracia, Tamaulipas,

MEXICO, 23 June 1969, J. M. Mathieu & M. W. Sanderson, black light trap

(USNM). PARATYPES: Texas: Brewster Co.: 2°°', 12, 9 mi W Alpine, 17

August 1969, J. C. Schaffner, at light (TAMU). Crosby Co.: 1°, 19, Post

& Crosbyton, 27 June, T. C. Clark & L. J. Muchmore (LACM). Culberson

Co.: 1°, Pine Springs, 18 August 1970, C. W. O'Brien (TTCC). Randall

Co.: 1°, Palo Duro State Park, nr. Amarillo, 27 June 1968, J. E. Slansky

(UCDC). Uvalde Co.: 2c‘, Garner State Park, 22/24 June 1961, R. L.

Westcott (LACM). MEXICO: Coahuila: 1°, 10 miS Allende, 11 August 1958,

H. F. Howden (CNCI). Nuevo Leon: 12, Mamulique Pass, 18 June 1975, D.

S. Chandler, UV light (DSC); 12, Linares, Rio Camacho, 21/22 June 1965,

O. S. Flint (USNM); 1°, 15 mi W Linares, 27 August 1969, J. Doyen & J.

Haddock (CISC); 12, 20 mi W Linares, 8 November 1946, E. S. Ross (CASC);

2co°, 229, Huasteca Canyon, nr. Monterrey, 11 July 1963, H. F. Howden

(CNCI); 1°, 6 km S Galeana, 25/26 June 1969, M. W. Sanderson & J. M.

Mathieu, black light trap (FGW). Tamaulipas: 400", 12, eutopotypical; 4 mi

SW Cd. Victoria, 1100'; 3°, 10 July 1963 and 1%, 5 August 1963, Duckworth

& Davis (USNM); 1c‘, 29 km S Hidalgo, 11 July 1952, F. & F. Werner (FGW).

San Luis Potosi: 1°, 12, Tamazunchale, Rt. 1, km 365, 31 May 1948, F.

Werner & W. Nutting, river in tropical area, light trap (FGW); 1°, same

locality, 1 May 1955, Buliofer & Guelo (USNM); 529, 1 mi SW Tamazunchale,

7 July 1966, P. M. Wagner, at light (TAMU).

Distribution: Western Texas into northern Mexico east of the Sierra

Madre Oriental. Attracted to light and ultra-violet light. Present from May

to November.

Discussion: It is similar to haustrus and nuperus in the deep impression

of the male fifth sternite. Easily separated from the other species in the

group by the slightly raised elytral setae. The only species of this group

which are on the east side of the Central Plateau are postictus and orientalis.

Notoxus orientalis, n. sp.

(Figs. 35, 60)

Length 2.82-3.55. Head brown to dark brown, pronotum orange to red-

Chandler: Notoxus of Central and South America 43

brown, elytra tan with brown markings: postbasal band across omoplates often

interrupted at elytral suture, irregular postmedian band advanced along elytral

suture, rarely meeting postbasal band, apical band extending anteriorly along

elytral suture. Pubescence brownish when arising from dark areas, whitish

over rest of body. Raised setae distinct on pronotum, setae on elytra ap-

pressed, average angle 89. Mandibular explanation narrow, hind margin

meeting mandible at lower level than front margin. Pronotum with antebasal

pubescent band widely broken. Crest margin smooth, apex rounded and dis-

tinct. First and second protarsomeres with strong spur on posterior apical

margin.

Males with fifth sternite emarginate, shallow circular impression from

apex.

Females with fifth sternite rounded-triangular.

Oo holotype: 6 km S Galeana, Nuevo Leon, MEXICO. Length 3.58. Head

0.65 long, eye 0.24 long, raised setae above eyes 0.08 long; antennomere

ratios: 12.25, 111.50, I112.00, IV 1.64, V 1.59, VIII 0.96, X 0.86, XI 1.54;

fourth maxillary palpus 0.20 long, 0.10 wide; mandibular explanation 0.11

long, 0.04 wide. Pronotum 1.14 long, length from base to point of greatest

width 0.38, break in antebasal pubescent band 0.26 wide; horn 0.44 long, 0.32

wide at base, no teeth on margin; crest 0.28 long, 0.14 wide at base; crest

apex 0.12 above horn margin, horn apex 0.22 from crest apex; pronotal pit

0.10 long. Elytral setae 0.08 long, average angle 79. Emargination of fifth

sternite 0.16 wide, 0.01 deep.

Specimens examined, 69: HOLOTYPE ©, 6 km S Galeana, Tamaulipas,

MEXICO, 25/26 June 1969, M. W. Sanderson & J. M. Mathieu, black light

trap (USNM). PARATYPES: MEXICO: Coahuila: 19, 21 mi SE Saltillo,

6000’, 18 July 1963, H. F. Howden (CNCI). Nuevo Leon: 5c, 19, 3 miS

Galeana, 5400', 29 May 1971, A. Newton, blacklight-thorn scrub (FGW); 80,

eutopotypical; 1°, 16 mi W Linares on Mex. 60, 2200', 26/29 May 1971, A.

Newton, canyon bottom, UV trap (FGW); 20's", 40 mi NW Junction Highway

of & 60, 22 June 1971, C. W. O'Brien & Marshall (TTCC); 1°, Cerro Potosi,

10,200' Schaffner (TAMU); 999, 18 N La Escondida, 4 July 1974, Clark,

Murray, Ashe, Schaffner (TAMU); 12, 3 miS Pacheco, 3/4 July 1974, Clark,

Murray, Ashe, Schaffner (TAMU). San Luis Potosi: 30c, 399, km 710, High-

way 97 N of San Luis Potosi, 11 July 1969, Ward, Tenorio, Bennett, host:

mesquite (TTCC); 1%, 222, 19 mi N San Luis Potosi, 1 September 1958, H. F.

Howden (CNCI); 32°, 11 mi N Matehuala, 2 September 1958, H. F. Howden

(CNCI). Hidalgo: 1300, 12, Zimapan, 11/14 June 1951, P. D. Hurd, H. E.

Evans, at light (CASC); 10°, 2 miSE Zimapan, 24 June 1975, D. S. Chandler,

beating vegetation (DSC). :

Distribution: The drier areas around the Sierra Madre Oriental. Collected

on mesquite and by beating vegetation. Attracted to light and ultra-violet light.

Present from May through September. |

Discussion: Similar to nuperus and haustrus in the appressed elytral setae.

It is very distinct in the shallow impression of the male fifth sternite and the

very irregular postmedian band.

Apicalis- group

The apicalis-group is defined as those Notoxus with the following charac-

ters: mandibular explanation narrow, hind margin meeting mandible ata

lower level than the front margin; pronotal tactile setae distinct, pubescent

44 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

antebasal band widely broken; crest margin smooth, apex rounded and distinct;

elytra with postmedian band, raised setae present; males without modifications

of the elytra and legs; fifth sternite emarginate; penis and parameres simple.

The Central American species included are apicalis LeConte, marginatus

LeConte and mexicanus Champion. One United States species, pallidus Fall,

is also included. This species is found along the lower Colorado River and

probably also occurs in Mexico along the river. It is included in the key to

species.

Notoxus apicalis LeConte

(Figs. 47, 63)

Notoxus apicalis LeConte 1852: 93. Type locality: Detroit, Michigan (probably

incorrect). MCZC type #4892, °. Horn 1884: 170. Casey 1895: 758.

Pic 1911: 5. Leng 1920: 163. Not Pallister 1955: 9, misidentification.

Notoxus nuperoides Fall 1916: 33-4. Type locality: Silver City, New Mexico.

MCZC type #24331, “. Leng 1920: 162.

Length 3.37-4.40. Head and pronotum tan to brown, pronotum often with

lateral areas darker, elytra tan with brown markings: circular mark on omo-

plates, postmedian band often advancing anteriorly along elytral suture to omo-

plates, flanks rarely dark, apical band often narrowly extended along elytral

suture to postmedian band. Pubescence brownish when arising from dark

areas, yellowish over rest of body. Raised setae distinct on pronotum, long

and erect on elytra, average angle 50°. Mandibular explanation narrow, an-

terior margin meeting mandible at higher level than posterior margin. Pro-

notum with antebasal pubescent band widely broken. Horn without pits beneath,

1-4 teeth on margin. Crest margin smooth, rounded anteriorly. First and

second protarsomeres with strong spur on posterior apical margin.

Male with fifth sternite slightly emarginate, transverse depression anterior

to apical margin.

Females with fifth sternite rounded-triangular.

Redescribed “: 18 mi E El Puerto, Sonora, MEXICO (DSC). Length 4. 02.

Head 0.79 long, eye 0.32 long, raised setae over eyes average 0.20 long;

antennomere ratios: 12.39, 11.85, Il 2.14, IV 2.00, V 1.76, VIII 1.50, X

1.20, XI 2.24; fourth maxillary palpus 0.22 long, 0.11 wide; mandibular ex-

planation 0.14 long, 0.05 wide. Pronotum 1.34 long, length from base to point

of greatest width 0.40, break in antebasal pubescent band 0.36 wide; horn 0.60

long, 0.40 wide at base, 3 teeth on margin; crest 0.41 long, 0.12 wide at base;

crest apex 0.08 above horn margin, horn apex 0. 24 from crest apex; pronotal

pit 0.12 long. Raised elytral setae 0.20 long, average angle 55°.

Specimens examined, 83: MEXICO Sonora: 18 mi E El Puerto, 7 August

1960, P. H. Arnaud, Jr., E. S. Ross, D. C. Rentz (CASC). Chihuahua: 20

mi SW Camargo, 13 July 1947, Gertsch (AMNH); 25 mi SW Camargo, 14 July

1947, Cazier, Gertsch (AMNH); Santa Barbara Dist., Santa Barbara, 7500",

18 July 1947, Gertsch (AMNH). Durango: 6 miS Durango, 4 July 1964, H. F.

Howden (CNCI); Rio Florido, nr. Las Nieves, 5200', 3 August 1966, G. E.

Ball & D. R. Whitehead, blacklight (FGW).

Distribution: The Great Plains and Great Basin south into Sonora and the

Mexican central plateau. It has been collected on alfalfa and cotton and is.

yeni to white light and ultra-violet light. Present from July through

ugust.

Chandler: Notoxus of Central and South America 45

Discussion: This species is most similar to mexicanus and pallidus. It

is separated from mexicanus by the horn having less than four teeth on the

margin, and can be separated from pallidus by the absence of ventro-lateral

pits on the horn.

Notoxus marginatus LeConte

(Figs. 48, 63)

Notoxus marginatus LeConte 1852: 93. Type locality: Detroit, Michigan

(probably incorrect). MCZC type 34893.

Notoxus apicalis var. marginatus, Pic 1911: 6. Leng 1920: 163.

Notoxus apicalis, auctorum.

Length 3.20-4.79. Head and pronotum tan, elytra tan with brown mark-

ings: circular mark on omoplates, postmedian band advanced somewhat along

elytral suture, mark on posterior flanks of elytra extending to apex and anteri-

orly along elytral suture. Pubescence brownish when arising from marks,

yellowish over rest of body. Raised setae distinct on pronotum, long and

erect on elytra, average angle on elytra 359. Mandibular explanation narrow,

hind margin joining mandible at lower lower level than front. Pronotum with

antebasal pubescent band widely broken. Horn without pits beneath, 1-3 teeth

on margin. Crest elevated, apex rounded and distinct. First and second pro-

tarsomeres with strong spur on posterior apical margin.

Males with fifth sternite slightly emarginate at apex, with shallow trans-

verse impression before apex.

Females with fifth sternite rounded-triangular.

Redescribed “: 3 mi N Las Cruces, Dofia Ana Co., New Mexico (DSC).

Length 4.50. Head 0.80 long, eye 0.32 long, raised setae over eyes average

0.13 long; antennomere ratios: 12.37, 111.53, III 1.93, IV 1.69, V 1.84,

VIII 1.60, X 1.32, XI 2.48; fourth maxillary palpus 0.22 long, 0.12 wide;

mandibular explanation 0.12 long, 0.04 wide. Pronotum 1.42 long, length from

base to point of greatest width 0.44, break in antebasal pubescent band 0. 32

wide; horn 0.64 long, 0.36 wide at base, 1 tooth on margin; crest 0.38 long,

0.11 wide at base; crest apex 0.12 over horn margin, horn apex 0.36 from

crest apex; pronotal pit 0.12 long. Raised setae 0.14 long, average angle 30°.

Emargination of fifth sternite 0.17 wide, 0.01 deep.

Specimens examined, 126: MEXICO: Chihuahua: Samalayuca, 24 June

1947, Gertsch, Cazier, Michener (AMNH); same locality, 6 August 1950, R.

F. Smith (AMNH); 25 mi SW Camargo, 14 July 1947, Gertsch, Cazier,

Michener (AMNH); Delicias, 4150', 11 July 1947, Schramel, Michener (AMNH).

Coahuila: Saltillo, 9 July 1952, F. & F. Werner (FGW); Laguna Coahuila, 26

June 1905, A. L. Herrera (USNM); San Pedro de Colonias, 3700', 20 August

1947, Michener (AMNH); Torreon, 18 July 1918, MacKinney & Loftin (USNM);

Boquillas del Carmen, 1850', 23 May 1959, Howden & Becker, at light (CNCI).

Durango: Lerdo, 31 June 1918, MacKinney & Loftin (USNM); same data ex-

cept, 16 July 1918 (USNM); Gomez Palacio, 14 July 1975, D. S. Chandler,

meloid trap (DSC). Collected in quarantine at Laredo-on grocery bag, on

dry shelled beans, on corn, on chick peas, in shelled corn, in dry corn, on

string beans, on dry shelled beans, on Tamarinds, on zinnias, on dry garlic,

on apple stem, on garlic; in quarantine at El Paso-on green beans, with Avo-

cado, on vegetable bin, on husk tomatoes, on green corn ears, in seed, on

Verbena plants, with carrots, with sugar cane leaves, on mango seeds; in

46 Contrib. Amer. Ent: Inst., vol; 15, no: 3, 1977

quarantine at Eagle Pass-on roots.

Distribution: From the Great Basin south to the northern portion of the

Central Plateau of Mexico. Collected on a variety of harvested crops, also

on Tamarix and sorghum in the United States. Attracted to white light, ultra-

violet light and drying meloid beetles. Present from May through August.

Discussion: It is distinct from the other species in the group by having

the raised elytral setae shorter than half the eye length, and the setae ata

lower angle of about 359. Similar to apicalis in the transverse impression on

the male fifth sternite and the elytral color pattern.

Notoxus mexicanus Champion

(Figs. 49, 63)

Notoxus mexicanus Champion 1890: 213, Table IX, fig. 22. Type series

localities: Mazatlan, Sinaloa; Acapulco and Iguala, Guerrero;

Guanajuato, Guanajuato, MEXICO. Type series in BMNH and FGW.

Pic 1894: 47; 1911: 9.

Length 3.45-6.18. Head and pronotum tan to red-brown, elytra tan with

brown markings: oval mark on omoplates usually joined together and to post-

median band along elytral suture, flanks of elytra dark, apical band advancing

anteriorly along elytral suture, often reaching postmedian band. Pubescence

brownish when arising from marks, whitish over rest of elytra. Raised setae

long and erect on pronotum and elytra, average angle on elytra 609. Mandibu-

lar explanation narrow, posterior margin meeting mandible at lower level than

anterior margin. Pronotum with antebasal band widely broken. Horn without

pits beneath, 4-8 teeth on margin. Crest margin smooth, anterior margin

distinct, usually rounded. First and second protarsomeres with strong spur

on posterior apical margin.

Males with fifth sternite truncate, shallow transverse impression before

apex.

Females with fifth sternite rounded-triangular.

Redescribed %: Arroyo Cuyuchaba, 11.5 mi E Alamos, Sonora, MEXICO

(DSC). Length 3.91. Head 0.89 long, eye 0.32 long, raised setae over eyes

average 0.21 long; antennomere ratios: 12.74, I1 1.67, III 3.18, IV 2.07,

V1.79, VII1.62, X 1.37, XI 2.46; fourth maxillary palpus 0.25 long, 0.14

wide; mandibular explanation 0.15 long, 0.05 wide. Pronotum 1.40 long,

length from base to point of greatest width 0.39, break in antebasal pubescent

band 0.38; horn 0.60 long, 0.34 wide at base, 6 teeth on margin; crest 0.36

long, 0.24 wide at base, rounded at apex; crest apex 0.06 above horn margin,

horn apex 0.17 from crest apex; pronotal pit 0.16 long. Raised elytral setae

0.18 long, average angle 73°.

Specimens examined, 144: MEXICO: Sonora: 5 mi E Alamos; 13 mi SE

Alamos; Arroyo Cyyuchaba, 11.5 mi E Alamos. Sinaloa: Choix; 5.5 mi NW

Choix; 8 miS Elota; 21 mi E Villa Union; Los Mayos; Rosario. Nayarit:

San Blas; 3 mi SE San Blas; Jesus Maria; Arroyo Santiago, nr. Jesus Maria;

Tepic. Zacatecas: 28 miS Jalpa. Jalisco: 8 mi N Guadalajara. Guerrero:

2 miS Mexcala; Iguala, Hoge, type of N. mexicanus (FGW). Morelos: 5 mi

N Axochiapan; Tejalpa. Puebla: 7 mi NE Atlixco; 34 miS Atlixco; 45 miN

Acatlan; Tepexco, Rte. 115, 4000'; Puente Estudo, nr. Tepexco, Rte. 115,

4000'; 5 mi SW Chipilo; 6 mi NE Zacatepec. Oaxaca: Huajuapan de Leon; 10

mi NE Huajuapan de Leon; 25 mi NE Oaxaca; Yagul ruins. All collections con-

Chandler: Notoxus of Central and South America 47

taining specimens.

Distribution: Throughout Mexico except the northern desert and the south-

ern tropical regions. Present from June to November. Collected under re-

cently uprooted weeds, under low vegetation in a corn field and sweeping low

vegetation. Attracted to light, ultra-violet light and drying meloid beetles.

Discussion: It is most similar to apicalis in general appearance. Separ-

ated by having four or more teeth on the horn margin.

Talpa-group

The talpa-group is defined as those Notoxus with the following characters:

mandibular explanation narrow to moderately wide; pronotum with tactile setae

appressed and not apparent, pubescent antebasal band widely broken; horn

without lateral pits; crest margin smooth, apex distinct, no median ridge;

elytra with median band, setae appressed. Males with elytra and legs unmodi-

fied; fifth sternite emarginate, circular impression before apex; penis and

parameres simple. Females with elytra prolonged at the apices.

The species included are talpa LaFerte, photus n. sp. and pygidialis n.

sp. :

Notoxus talpa LaFerte

(Figs. 42. 63)

Notoxus talpa LaFerté 1848: 50. Type locality: "California" (probably the

area near Mexico City). Type series in MHNH. Pic 1911: 12.

Notoxus elegantulus LaFerte 1848: 52, Table 21, fig. 17. Type locality:

"California" (probably the area near Mexico City). Type in MNHN.

NEW SYNONYMY.

Notoxus ventralis Champion 1890: 207-8, Table IX, fig. 17. Type series

localities: Guanajuato, Guanajuato; and Jalapa, Veracruz, MEXICO.

Types in BMNH. Champion 1893: 461. Pic 1894: 47; 1911: 13. NEW

SYNONYMY.

Length 2.75-3.78. Head and pronotum orange to brown, elytra tan with

brown markings: irregular median band usually reaching circular mark

covering omoplates and base of elytra, apical band and median band usually

forming tan postmedian W-shaped band, specimens from Michoacan with

elytra all brown. Pubescence brownish when arising from dark areas, whit-

ish in light areas. Setae appressed on pronotum, decumbent on elytra, aver-

age angle 13°. Mandibular explanation moderately wide, hind margin meeting

mandible at lower level than front margin. Pronotum with pubescent antebasal

band widely broken. Horn without pits beneath, 0-4 teeth on margin. Crest

margin smooth, apex distinct, usually pointed. First and second protarso-

meres with strong spur on posterior apical margin.

Males with fifth sternite emarginate, small circular impression at apex in

larger, poorly-defined circular impression.

Females with fifth sternite rounded-triangular. Elytra with apices abruptly

pointed.

Redescribed “: 4 mi E Zacatecas, Zacatecas, MEXICO (DSC). Length

3.82. Head 0.80 long, eye 0.29 long, raised setae above eyes average 0.08

long; antennomere ratios: 12.47, I1 1.62, III 2.21, IV 1.73, V 1.81, VII

48 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

1.22, X 1.22, XI 2.26; fourth maxillary palpus 0.26 long, 0.12 wide; mandibu-

lar explanation 0.10 long, 0.06 wide. Pronotum 1.31 long, length from base

to point of greatest width 0.39, break in antebasal pubescent band 0.37 wide;

horn 0.61 long, 0.41 wide at base, apex pointed, no teeth on margin; crest

0.34 long, 0.14 wide at base; crest apex 0.08 above horn margin, horn apex

0.24 from crest apex; pronotal pit 0.14 long. Elytral setae 0.11 long, average

angle 159. Emargination of fifth sternite 0.16 wide, 0.02 deep.

Specimens examined, 106: MEXICO: Durango: 26 mi W Durango.

Zacatecas: Sombrerete, 6000'; 4 mi E Zacatecas. Aguascalientes: 8 mi N

Aguascalientes, 6700'. Guanajuato: Guanajuato, type of N. ventralis.

Queretaro: 10 mi E San Juan del Rio; 1 mi S Palmillas. Mexico: Texcoco;

Atlacomulco; Tepotzotlan; 15 mi NW Toluca. Distrito Federal: Chapultepec;

Coapa, Mexico, Tlalpan. Michoacan: 11 mi SE Carapan; Uruapan; 10 mi E

Zamora on Highway 15; Tancitaro; 3 miS Carapan. Veracruz: Jalapa, type

of N. ventralis. Oaxaca: Yagul ruins; 10 mi N Miltepec. Specimens in col-

lections of: CASC, CNCI, DSC, FGW, LACM, MSUC, TAMU, TTCC, USNM.

Distribution: The central plateau to the higher areas of Oaxaca. Collected

on beans, under low vegetation in corn field, and in squash flowers. Attracted

to light, cantharidin and freshly killed meloid beetles. Present from June

through August.

Discussion: This species is most similar to pygidialis. The males can

be separated by the shallow impression of the fifth sternite which is never

obscured by setae. The females have the pygidium simple, not projecting.

Notoxus pygidialis, n. sp.

(Figs. 43, 63)

Length 2.65-3.48. Head and pronotum red-brown to brown, elytra tan

with brown markings: median band advancing anteriorly along elytral suture to

cover omoplates, apical band advancing anteriorly along elytral suture, usually

meeting postmedian band, anterior flanks dark. Pubescence brownish when

arising from markings, yellowish in light areas. Setae appressed on prono-

tum, decumbent on elytra, average angle 16°. Mandibular explanation moder-

ately wide, hind margin meeting mandible at lower level than front margin.

Pronotum with pubescent antebasal band widely broken. Horn without pits

beneath, 0-2 teeth on margin. Crest margin smooth, apex distinct, usually

pointed. First and second protarsomeres with strong spur on posterior apical

margin.

Males with fifth sternite emarginate, circular impression before apex,

may be shallowly or deeply impressed with dense setae around margin.

Females with fifth sternite rounded-triangular. Elytra slightly truncated,

last tergite prolonged apically into distinct protuberance.

S holotype: Zacatelco, Tlaxcala, MEXICO. Length 2.81. Head 0.68 long,

eye 0.22 long, raised setae above eyes average 0.06 long; antennomere ratios:

12.53, I 1.55, U0 3.18, 1V 1.67, ¥ 1,67, Volk 1.30, % 1,05, Al 2.25;

fourth maxillary palpus 0.19 long, 0.10 wide; mandibular explanation 0.10

long, 0.06 wide. Pronotum 1.00 long, length from base to point of greatest

width 0.28, break in antebasal pubescent band 0.23 wide; horn 0. 46 long,

0.28 wide at base, no teeth on margin; crest 0.30 long, 0.11 wide at base;

crest apex 0.06 above horn margin, horn apex 0.19 from crest apex; pronotal

pit 0.09 long. Elytral setae 0.07 long, average angle 12°. Emargination of

fifth sternite 0.10 wide, 0.04 deep.

Chandler: Notoxus of Central and South America 49

Specimens examined, 124: HOLOTYPE ©, Zacatelco, Tlaxcala, MEXICO,

26 July 1974, R. L. Mangan & D. S. Chandler, under low vegetation in corn

field (USNM). PARATYPES: MEXICO: Tlaxcala: 21°, 2422, eutopotypical

(DSC); 20°o', 522, Tepeyanco, 25 June 1975, D. S. Chandler, under low plants

in corn field (DSC); 3°, Xicohtzinco, 25 June 1975, D. S. Chandler, under

low plants in corn field (DSC). Puebla: 19, Puebla, 3 July 1952, C. D. Mac-

Neill (CISC); 60°", 892, 3 mi E Acatepec, 26 June 1975, D. S. Chandler, under

low plants in corn field (DSC); 10°, 49°, same data except, cantharidin trap

(DSC); 1°, 5 mi SW Chipilo, 26 June 1975, D. S. Chandler, sweeping low vege-

tation (DSC); 70°", 32°, same data except, cantharidin trap (DSC); 10°°, same

data except, meloid trap (DSC); 100s", 29°, 1 mi E Cholula, 26 June 1975,

D. S. Chandler, cantharidin trap (DSC). Morelos: 1°, Cuernavaca (MCZC);

1c‘, 9km W Tres Cumbres, 7600', Rt. 3, km 60, 24 June 1948, F. Werner

& W. Nutting, pine clearing (FGW). Oaxaca: 2°, 10 mi NE Huajuapan de

Leon, 25 June 1965, Burke, Meyer, Schaffner (TAMU).

Distribution: South-central Mexico. Collected sweeping low vegetation

and under low vegetation in corn field. Attracted to cantharidin and freshly

killed meloid beetles. Present in June and July.

Discussion: It is most similar to falpa and cannot be separated except by

the sexual characters. The males can usually be separated by the deeper

and densely setate margin of the impression of the fifth sternite. The females

are very distinctive in having the apically produced pygidium.

Notoxus photus, n. sp.

(Figs. 44, 63)

Length 3.00-3.60. Head and pronotum tan to red-brown, elytra tan to red-

brown with brown markings: median band often reaching circular mark cover-

ing omoplates, apical band extending anteriorly along elytral suture. Pubes-

cence brownish in markings, whitish over rest of body. Setae appressed on

pronotum and elytra, average angle on elytra 10°. Mandibular explanation

narrow, front and hind margins meeting at mandible on same level. Pronotum

with pubescent antebasal band widely broken. Horn without pits beneath, 0-4

teeth on margin. Crest margin smooth, apex distinct, usually rounded. First

and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite slightly emarginate, circular impression before

apex.

Females with fifth sternite rounded-triangular. Elytra truncated, apices

spinose.

S holotype: Superior, Boyce-Thompson Arboretum, 2500', Gila County,

Arizona. Length 3.16. Head 0.71 long, eye 0.26 long, raised setae above

eyes average 0.09 long; antennomere ratios: 12.54, I1 1.67, II 2.23, IV 1.85,

V 1.74, VIII 1.50, X 1.27, XI 2.11; fourth maxillary palpus 0.20 long, 0.10

wide; mandibular explanation 0.10 long, 0.06 wide. Pronotum 1.02 long,

length from base to point of greatest width 0.34, break in antebasal pubescent

band 0.30 wide; horn 0.44 long, 0.31 wide at base, 1 tooth on margin; crest

0.32 long, 0.10 wide at base, apex pointed; crest apex 0.05 above horn mar-

gin, horn apex 0.17 from crest apex; pronotal pit 0.10 long. Elytral setae

0.08 long, average angle 10°. Emargination of fifth sternite shallow, 0.16

wide.

Specimens examined, 199: HOLOTYPE °C, Superior, Boyce-Thompson

Arboretum, 2500', Gila Co., Arizona, 31 July 1948, H. K. Gloyd, white light

20 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

trap (USNM). PARATYPES: Arizona: Pima Co., Tucson; Santa Catalina

foothills; I. B. P. site, Santa Rita Range Reserve; Colossal Cave County

Park; Santa Catalina Mtns., Molino Canyon and Sabino Canyon; Brown Canyon,

E slope Babaquivari Mtns. Santa Cruz Co.: Patagonia; 2 mi SW Patagonia;

Pajarito Mtns., Pefla Blanca Canyon and Sycamore Canyon; Madera Canyon,

Santa Rita Mtns. Cochise Co.: Wood Canyon, Bisbee; Cochise Stronghold and

West Stronghold, Dragoon Mtns. ; Southwest Research Station, Chiricahua

Mtns. MEXICO: Chihuahua: 12 mi N Chihuahua. Paratypes in collections of:

CASC, CDAE, DSC, FGW.

Distribution: Associated with the oak-juniper zone through southeastern

Arizona and south into Chihuahua. The only collection data is that most speci-

mens were collected at light or ultra-violet light. Females are poorly

attracted to light as only two are included in the type series. Present from

June through September.

Discussion: It is most similar to falpa. Easily distinguished by the lighter

appearance of the species. The shallow circular impression of the male fifth

sternite is similar to that of falpa. The female elytral apices are very acute,

while those of falpa are usually more rounded.

Monoceros- group

The monocervos-group is defined as those Notoxus with the following char-

acters: mandibular explanation narrow, hind margin obsolete; raised pronotal

setae distinct, pubescent antebasal band widely broken; horn without lateral

pits; crest margin and apex variable, no median ridge; elytra with preapical

band, raised setae present. Males with lateral apical tubercles on the elytra;

fifth sternite truncate to shallowly emarginate; penis greatly modified, asym-

metrical with dorsal surface as a plate or as a left-directed hook; parameres

simple.

Species represented in Mexico are anchora Hentz and serratus LeConte.

Only a single specimen of each has been collected. North American species

included are nevadensis Casey, pictus Casey and robustus Casey. There are

thirteen Palearctic species which were covered by Heberdey in 1936.

Notoxus anchora Hentz

(Figs. 50, 64)

Notoxus anchora Hentz 1827: 375, Table 13, fig. 4. Type locality: Massachu-

setts. LaFerté 1848: 33. LeConte 1852: 92. Horn 1884: 173. Casey

1895: 757. Pic 1911: 5. Leng 1920: 162.

Monocerus anchora, LeConte 1847: 89.

Length 2.99-4.38. Head and pronotum tan to brown, elytra tan with brown

markings: preapical band extending anteriorly along elytral suture over omo-

plates to base of elytra, median lateral marks often attached to preapical band.

Pubescence brownish when arising from markings, yellowish over rest of ely-

tra. Raised setae long and erect on pronotum and elytra, average angle on

elytra 62°. Mandibular explanation narrow, hind margin indistinct, meeting

mandible at lower level than front margin. Pronotum with antebasal pubescent

band widely broken. Horn without pits beneath, 3-10 teeth on margin. Crest

margin crenulate, anterior margin pointed and indistinctly merging with horn.

Chandler: Notoxus of Central and South America 51

First and second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite truncate at apex, elytra with lateral apical tuber-

cles. Penis with ventral surface modified into a left-directed hook, dorsal

surface forming two broad lobes which have two smaller lobes at each apex.

Females with fifth sternite rounded-triangular.

Redescribed “: 1 mi W Nichols, New York (DSC). Length 3.15. Head

0.67 long, eye 0.26 long, raised setae above eyes average 0.12 long: antenno-

mere ratios: 12.50, 184:67, 12.00, IV 1.77, V 1.80, VIE 1.02,;:X%-1, 82;

XI 2.37; fourth maxillary palpus 0.20 long, 0.10 wide; mandibular explanation

0.10 long, 0.03 wide. Pronotum 1.02 long, length from base to point of great-

est width 0.31, break in antebasal pubescent band 0.24 wide; horn 0.43 long,

0.24 wide at base, 6 teeth on margin; crest 0.34 long, 0.18 wide at base;

crest apex merging with horn, horn apex 0.16 from crest apex; pronotal pit

0.15 long. Raised elytral setae 0.15 long, average angle 729. Truncation of

fifth sternite 0.10 wide.

Specimens examined, 1: 'Mex.'' (MCZC).

Distribution: Northeastern United States south through the Rocky Mountains.

This species is probably found in the northern mountains near the United States

border. Collected in the United States by sweeping alfalfa and is attracted to

cantharidin. Present in the United States from June through August.

Discussion. It is most similar to nevadensis by the crenulate crest mar-

gin and the indistinct crest apex. It can be separated by the narrow horn and

the presence of a left-directed hook as the ventral portion of the penis. JN,

nevadensis has two narrow lobes at the apex of the ventral plate. N. robustus

has the crest margin rounded-crenulate, the apex never breaks down anteriorly

and merges with the horn.

Notoxus serratus (LeConte)

(Figs. 51, 64)

Monocerus serratus LeConte 1847: 90. Type locality: Rocky Mountains.

MCZC Type #4894, ©.

Notoxus serratus, LeConte 1852: 93. Horn 1884: 172. Pic 1911: 11. Casey

1895: 758. Leng 1920: 163.

Notoxus digitatus LeConte 1876: 518. Type locality: southern Colorado.

MCZC Type #4895, ©.

Notoxus vandykei Blaisdell 1929: 57-9. Type locality: Little Cottonwood

Creek, Inyo County, California. CASC Type #2616, “~. Leng 1933: 26.

Length 3.19-4.60. Head and pronotum red-brown, elytra tan with brown

markings: circular mark on omoplates, preapical band often broken along

elytral suture, extending anteriorly along elytral suture, rarely reaching omo-

plates, often with lateral marks in median portion of elytra, all marks may be

joined, separated or absent. Pubescence brownish in dark areas, yellowish

over rest of body. Raised setae long and erect on pronotum and elytra, aver-

age angle on elytra 58°. Mandibular explanation narrow, with hind margin

meeting mandible at lower level than front margin. Pronotum with antebasal

pubescent band widely broken. Horn without pits beneath, 6-10 teeth on mar-

gin; crest margin smooth, apex distinct, usually pointed. First and second

protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite emarginate at apex. Penis with ventral surface

modified into a left-directed hook, dorsal surface forming two broad lobes.

D2 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Each elytron with lateral apical tubercle.

Females with fifth sternite rounded-triangular.

Redescribed “: Hallelujah Junction, Lassen County, California (DSC).

Length 4.40. Head 0.89 long, eye 0.32 long, raised setae above eyes average

0.20 long; antennomere ratios: 12.50, 112.00, Il 2.50, IV 1.57, V 2.00,

VIII 1.90, X 1.67, XI 2.85; fourth maxillary palpus 0.25 long, 0.14 wide;

mandibular explanation 0.16 long, 0.06 wide. Pronotum 1.33 long, length

from base to point of greatest width 0.45, break in antebasal pubescent band

0.30 wide; horn 0.58 long, 0.32 wide at base, apex pointed, 7 teeth on mar-

gin; crest 0.35 long, 0.12 wide at base, apex pointed; crest apex 0. 12 above horn mar-

gin, hornapex 0.30from crest apex; pronotal pit0.11long. Raised elytral setae

0.25long, averageangle 689. Emargination of fifth sternite 0.23 wide, 0.01 deep.

Specimens examined, 1 MEXICO: Chihuahua: 48 km S Juarez, 28 April

1974, D. Giuliani, sand dunes, (CDAE).

Distribution: Montane regions of the western United States and northern

Mexico. Collected on grass and Salix, Attracted to light, ultra-violet light

and cantharidin. Present in the United States from April to September.

Discussion: It is most similar to pictus by the high smooth crest margin

and is separated by having more than four teeth on the horn margin. It is also

similar to vobustus, but can be separated by the non-crenulate crest margin.

Some specimens of vobustus have crests approaching the smoothness of ser-

vatus, which can then only be separated by the males of vobustus possessing a

tubercle on the pro- and mesofemora.

Sparsus-group

The sparsus-group is defined as those Notoxus with the following char-

acters: mandibular explanation wide, hind margin prominent; pronotum with

raised setae distinct, pubescent antebasal band complete to widely broken;

horn without lateral pits; crest margin and apex variable. Males with lateral

apical tubercles on the elytra. The fifth sternite rounded or truncate at apex,

with oval impression before apex; parameres simple; penis often thickened

dorsoventrally and with preapical notch.

The group is basically Californian with two species, sparsus LeConte and

conformis LeConte, extending south into Baja California. N. bajae n. sp. is

restricted to Baja California Sur. The United States species included in the

group are cavicornis LeConte, denudatus Horn, lustrellus Casey, spatulifer

Casey, breviusculus Fall and obesulus Blaisdell.

Notoxus sparsus LeConte

(Figs. 52, 64)

Notoxus sparsus LeConte 1859: 284-5. Type locality: Point Reyes, Califor-

nia. MCZC Type #4898, *. Horn 1884: 176 (synonym of conformis

LeConte). Casey 1895: 757. Pic 1911: 7 (synonym of conformis).

Leng 1920: 162 (synonym of conformis).

Length 2.42-3.08. Head and pronotum red-brown, elytra yellow-brown

with brown markings: posthumeral marks on flanks usually joining oval mark

on omoplates to form postbasal band, postmedian band usually advanced nar-

rowly along elytral suture, elytral apices obscurely darkened. Pubescence

Chandler: Notoxus of Central and South America 53

brownish when arising from marks, whitish over rest of body. Raised setae

long and erect on pronotum and elytra, average angle on elytra 48°. Mandibu-

lar explanation wide, with front and hind margins on same level at mandible.

Pronotum with antebasal pubescent band complete. Horn without pits beneath,

0-3 teeth on margin. Crest margin smooth, rounded and indistinct anteriorly,

entire crest only slightly elevated above distinct margin. First and second

protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite truncate, slight circular impression anterior to

apical margin. Penis swollen dorsoventrally, with shallow dorsal notch near

apex.

Females with fifth sternite triangular, elytra truncated at apex.

Redescribed ©: 12 mi SW Red Bluff, Tehama County, California (DSC).

Length 3.08. Head 0.72 long, eye 0.24 long, raised setae above eyes average

0.21 lone; antennomere ratios: 12.53, 111.84, 012.59, TV 2.08, V 2.15,

VIII 1.71, X 1.34, XI 2.36; fourth maxillary palpus 0.19 long, 0.10 wide;

mandibular explanation 0.10 long, 0.05 wide. Pronotum 1.10 long, length

from base to point of greatest width 0.32; horn 0.44 long, 0.22 wide at base,

1 tooth on margin; crest 0.29 long, 0.10 wide at base; crest apex 0.04 above

horn margin, horn apex 0.20 from crest apex; pronotal pit 0.09 long. Raised

elytral setae 0.20 long, average angle of setae 459. Truncation of fifth ster-

nite 0.12 wide.

Specimens examined, 1: MEXICO: Baja California: Sierra Juarez, 1 mi

N El Compadre, 23 January 1965, E. L. Sleeper (CDAE).

Distribution: Through central and coastal California to the northern por-

tion of Baja California. Collected in oak litter in California. Attracted to

white light and is present throughout the year in southern California.

Discussion: It is most similar to an undescribed species from California

by the long, raised setae on the elytra and the globular (rather than transverse)

pronotum. It is best separated by the genitalia with the penis having a distinct

shallow preapical notch. |

Despite being synonymized under conformis by several authors, spayrsus is

easily separated by the elevated tactile setae on the elytra.

Notoxus bajae, n. sp.

(Figs. 54, 64)

CS holotype: 25 mi W La Paz, Baja California Sur, MEXICO (USNM).

Head and pronotum tan, elytra tan with brown markings: mark on omoplates,

irregular postmedian band extending anteriorly along elytral suture to near

omoplates, posthumeral mark on flanks. Pubescence brownish when arising

from markings, yellowish over rest of body. Raised setae long and erect on

pronotum and elytra, average angle on elytra 67°. Mandibular explanation

wide, with front and hind margins on same level at mandible. Pronotum with

antebasal pubescent band complete. Horn without pits beneath, no teeth on

margin. Crest margin smooth, rounded and distinct anteriorly. First and

second protarsomeres with strong spur on posterior apical margin.

Male with fifth sternite truncate at apex, circular depressed area anterior

to margin. Penis with dorsal notch near apex.

Length 4.04. Head 0.86 long, eye 0.28 long, raised setae over eyes aver-

age 0.28 long; antennomere ratios: I 2.04, I1 2.46, III 2.00, IV 2.33, V 2.06,

Vill 1.55, KX 1.60, XI 2.37; fourth maxillary palpus 0.21 long, 0.08 wide;

mandibular explanation 0.14 long, 0.09 wide. Pronotum 1.32 long, length

o4 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

from base to point of greatest width 0.40, horn 0.58 long, 0.29 wide at base,

no teeth on margin; crest 0.40 long, 0.10 wide at base; crest apex 0.15 above

horn margin, horn apex 0.28 from crest apex; pronotal pit 0.13 long. Raised

elytral setae 0.26 long. Truncation of fifth sternite 0.18 wide.

Specimens examined, 1: HOLOTYPE ©, 25 mi W La Paz, Baja California

Sur, MEXICO, 4 September 1959, K. W. Radford & F. G. Werner, light

trap (USNM).

Distribution: The single specimen is from southern Baja California Sur

and was collected at light.

Discussion: It is distinct in the group by having the crest half as wide at

base as at the greatest width and having the crest well-elevated above the

horn margin. The penis is similar to that of spatulifer by the dorsal preapical

slit, but the apex is simple, not flattened and dilated.

Notoxus conformis LeConte

(Figs. 53, 64)

Notoxus conformis LeConte 1851: 152. Type locality: Gila River, Arizona.

MCZC Type #4896, °. LeConte 1852: 92-3. Horn 1884: 174. Casey

1605" (91. Fall $907: 33, 162. Pic 1911: 7. Leng 1920: 162.

Length 2.60-3.46. Head and pronotum reddish-brown, brown markings

on reddish-brown elytra: mark on omoplates offset toward elytral suture,

frequently postmedial band extends along elytral suture to meet omoplate

marks, posthumeral marks on flanks on elytra, often reaching to postmedian

band, obscure darkened band near elytral apices. Pubescence brownish when

arising from dark areas, whitish over rest of body. Short scattered raised

setae on pronotum, on elytra setae appressed, average angle 10°. Mandibular

explanation moderately wide, with front and hind margins on same level at

mandible. Pronotum with antebasal pubescent band complete. Horn without

pits beneath, 0-2 teeth on margin. Crest smooth, anterior margin rounded,

only slightly elevated above horn margin. First and second protarsomeres

lacking spur on apical posterior margin.

Males with fifth sternite truncate, circular impression before apex which

may be faint, elytra with lateral apical tubercles, anterior margin of crest

usually distinct. Penis swollen dorso-ventrally, with deep dorsal notch near

apex.

Females with fifth sternite rounded-triangular; anterior margin of crest

indistinct, breaking down and merging with horn; elytra appearing slightly

truncated; body appearing stouter than males.

So: Hemet, Riverside County, California (DSC). Length 3.46. Head 0.76

long, eye 0.26 long, raised setae over eyes average 0.16 long; antennomere

ratios: 12,55, 1. t, 64.. 11 2.50, 1¥ 2.01, V 2.20,. VI L.63, X 1.387, AT 2.10;

fourth maxillary palpus 0.21 long, 0.12 wide; mandibular explanation 0.12

long, 0.06 wide. Pronotum 1.24 long, length from base to point of greatest

width 0.34; horn 0.55 long, 0.32 wide at base, 1 tooth on margin; crest 0.31

long, 0.15 wide at base; crest apex 0.02 above horn margin, horn apex 0.23

from crest apex; pronotal pit 0.08 long. Elytral setae 0.10 long, average angle

13°. Truncation of fifth sternite 0.12 wide.

Specimens examined, 19: MEXICO: Baja California: Sierra San Pedro

Martir, La Sanja, 6500', 27 May 1958, J. Powell (CISC); 2.4 mi SE El Consu-

elo, 24 April 1963, H. B. Leech & P. H. Arnaud, Jr. (CASC).

Chandler: Notoxus of Central and South America 55

Distribution: Found in hilly and montane areas of California, Arizona and

Baja California. Collected by beating vegetation, sifting oak duff, and has been

found on dead yucca and in a swallow nest. Present from February to Novem-

ber in California.

Discussion: It is most similar to lustrellus by the decumbent elytral setae.

It can be easily separated by the smooth crest margin and few teeth on the horn

margin. N. lustrellus has the crest crenulate and about five teeth on the horn

margin.

Species Not Placed In Groups

The Central American species not placed in any group are crucifer

Champion, ofacus Champion, fvaternus Champion, truncatipennis Champion,

aztecorum n. sp., campus n. sp. stephanin. sp., despevatus n. sp., zapote-

corum n. sp. and occidentalis n. sp. United States species not placed in any

eroup are brevicornis Fall and an undescribed species from Texas. An uni-

dentified Notoxus female of probable Old World origin is included in this sec-

tion.

Notoxus occidentalis, n. sp.

(Figs: 12; 55)

Notoxus bifasciatus, Pallister 1955: 6, misidentification.

Length 3.00-3.30. Head and pronotum red-brown to brown, elytra red-

brown with brown markings: median band extending anteriorly along elytral

suture to cover omoplates, apical band may extend anteriorly along elytral

suture to join median band. Pubescence brownish when arising from dark

areas, whitish in lighter areas. Raised setae long and erect on pronotum and

elytra, average angle on elytra 559. Mandibular explanation moderately wide,

front and hind margins meeting at mandible on same level. Pronotum with

pubescent antebasal band widely broken. Horn without pits beneath, no teeth

on margin. Crest margin smooth, apex pointed and distinct. First and second

protarsomeres with strong spur on posterior apical margin.

Males unknown.

Females with fifth sternite rounded-triangular.

? holotype: San Jose Babicora, Chihuahua, MEXICO. (AMNH). Length

3.00. Head 0.70 long, eye 0.23 long, raised setae over eyes average 0.20

long; antennomere ratios: I 2.47, IL 1.73, HI 2.08, 1V 1.79, V 1.67, VIII

1.50, X 1.33, XI 1.84; fourth maxillary palpus 0.17 long, 0.09 wide; mandibu-

lar explanation 0.10 long, 0.06 wide. Pronotum 1.04 long, length from base

to point of greatest width 0.35, break in antebasal pubescent band 0.20 wide;

horn 0.42 long, 0.28 wide at base; crest 0.28 long, 0.10 wide at base; crest

apex 0.10 above horn margin, horn apex 0.25 from crest apex; pronotal pit

0.08 long. Raised elytral setae 0.20 long, average angle 49°.

Specimens examined, 2: HOLOTYPE 2, San Jose Babicora, Chihuahua,

MEXICO, 5 June 1947, Michener (AMNH). PARATYPE: 19, Palos Colorados,

8000', Durango, MEXICO, 5 August 1947, Cazier (DSC).

Distribution: The Sierra Madre Occidental. Adults present from June to

August.

Discussion: Collection of the males will be necessary before this species

56 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

can be placed positively ina group. The species is either a member of the

montanus- or pueblensis-groups. Pallister (1955) identified the type series

as Notoxus bifasciatus (LeConte), a species which occurs only in northern

and eastern United States.

Notoxus fraternus Champion

(Figs. 45, 61)

Notoxus fraternus Champion 1890: 208-9. Type series localities: Guanajuato,

Guanajuato (BMNH) and Jalapa, Veracruz (FGW), MEXICO. Pic 1894:

47; 1911: 8. Not Pallister 1955: 8.

Length 2.58-3.45. Head and pronotum brown to dark brown, elytra brown

to dark brown, rarely with faint lighter marks laterally behind omoplates and

laterally in apical half. Brownish setae forming vague median band, extending

anteriorly near elytral suture to cover omoplates, narrow band of white setae

behind median band, followed by brownish setae to apex. Upraised setae

present on pronotum, setae on elytra decumbent, average angle 12~. Mandibu-

lar explanation narrow, with hind margin often meeting mandible at lower level

than front margin. Pronotum with pubescent antebasal band widely broken.

Horn without pits beneath, 0-1 teeth on margin. Crest margin smooth, apex

distinct, usually rounded. First and second protarsomeres with strong spur

on posterior apical margin.

Males with fifth sternite emarginate at apex, transverse impression before

apex.

Females with fifth sternite rounded-triangular.

Redescribed ©: 4 mi SW Perote, Veracruz, MEXICO (DSC). Length 3. 04.

Head 0.68 long, eye 0.20 long, raised setae above eyes average 0.12 long;

antennomere ratios: 12.65, I1 1.59, Il 2.34, 1V 1.84, V 1.93, VIII 1.35,

X 1.21, XI 2.15; fourth maxillary palpus 0.16 long, 0.10 wide; mandibular

explanation 0.11 long, 0.05 wide. Pronotum 1.00 long, length from base to

point of greatest width 0.34, break in antebasal pubescent band 0.28 wide;

horn 0.38 long, 0.28 wide at base, no teeth on margin; crest 0.28 long, 0.11

wide at base, apex rounded; crest apex 0.07 above horn margin, horn apex

0.14 arom crest apex; pronotal pit 0.10 long. Elytral setae 0.09 long, average

angle 14°. Emargination of fifth sternite 0.20 wide, 0.01 deep.

Specimens examined, 266: MEXICO: Guanajuato: Guanajuato, type of

fraternus. Michoacan: 2 mi SE Los Reyes. Mexico: 7 miS Amecameca,

8000'. Distrito Federal: 2 km S Mexico, 8600'; 16 mi S Mexico City, 7400';

37 km S Mexico City; Guadalupe; Tlalpan. Tlaxcala: Xicohtzinco. Puebla:

7 mi NE Atlixco; 13.8 mi SW Alchichica. Veracruz: 4 mi SW Perote; 10 mi

SW Parote; Jalapa. Specimens in collections of: CASC, CISC, CNCI, CUIC,

DSC, FGW, MCZC, TAMU, USNM.

Distribution: The higher areas of south-central Mexico. Collected on

corn, on the ground in a moist agriculatural area and under low plants in

corn field. Attracted to freshly killed meloid beetles. Present from June

through August.

Discussion: It is generally similar to truncatipennis by the smooth crest

margin and the emarginate fifth sternite. It is separated by the decumbent

elytral setae, the more elevated crest and the transverse impression before

the down-curved apex of the male fifth sternite.

Chandler: Notoxus of Central and South America 57

Notoxus stephani, n. sp.

(Figs. 39, 61)

Length 2.99-4.37. Head and pronotum orange to brown, elytra tan to red-

brown with brown markings: oval mark covering omoplates often joined to

slightly postmedian band along elytral suture, apical band joined to postmedian

band along elytral suture. Pubescence brownish when arising from dark areas,

whitish in light areas. Raised setae distinct on pronotum, and elytra, average

angle on elytra 32°. Mandibular explanation moderately wide, front and hind

margins often meeting on same level at mandible. Pronotum with antebasal

pubescent band widely broken. Horn without pits beneath, 0-1 teeth on mar-

gin. Crest margin smooth apex usually pointed, apex indistinct and merging

with horn when pointed. First and second protarsomeres with strong spur on

posterior apical margin.

Males with fifth sternite emarginate into large circular impression.

Females with fifth sternite rounded-triangular.

% holotype: 5 mi E Alamos, Sonora, MEXICO. Length 4.09. Head 0.84

long, eye 0.30 long, raised setae above eyes average 0.10 long; antennomere

patios: 12.90. 1) 1.50, Bio. 13, 7V 1.81, Ve.00 Vil 1.05, 2 4.10, Al 4, 11;

fourth maxillary palpus 0.18 long, 0.12 wide; mandibular explanation 0.12 long,

0.04 wide. Pronotum 1.32 long, length from base to point of greatest width

0.42, break in antebasal pubescent band 0.30 wide; horn 0.58 long, 0.36 wide

at base, 1 tooth on margin; crest 0.42 long, 0.14 wide at base, apex pointed;

crest apex 0.02 over horn margin, horn apex 0.24 from crest apex; pronotal

pit 0.12 long. Raised elytral setae 0.11 long, average angle 41°. Emargina-

tion of fifth sternite 0.15 wide, 0.01 deep.

Specimens examined, 19: HOLOTYPE ©, 5 mi E Alamos, Sonora,

MEXICO, 11 August 1973, K. Stephan & D. S. Chandler, blacklight (USNM).

PARATYPES: MEXICO: Sonora: 90°, 1%, eutopotypical (DSC). Jalisco:

12, Guadalajara, 12 August 1903, J. F. McClendon (MCZC); 1°, same locality,

1 August 1903 (MCZC); 12, same locality, 8 August 1903 (MCZC); 1°, 9 mi W

Guadalajara, 1550 meters, 16 July 1947, T. H. Hubbell, on Tequila pd.

(CASC); 10°, 13 mi W Guadalajara, 21 August 1964, H. R. Burke & J. Apperson

(TAMU); 1“, 4 mi SW Jalostotitlan, 4 August 1954, E. G. Linsley, J. W.

MacSwain, R. F. Smith (CISC); 1°, 2 miS Tlaquepaque, 11 July 1953, C. &

P. Vaurie (AMNH); 1°, 6 mi SE Sayula, 23 July 1966, P. M. & P. K. Wagner

(TAMU). Zacatecas: 1°, 10 mi WSW Huejucar, 7200', 22 July 1954, R. H.

Brewer (CASC).

Distribution: The semi-tropical regions on the west coast of Mexico. Col-

lected by sweeping and found on a Tequila pod. Attracted to ultra-violet light.

Present from July to August.

Discussion: It is most similar to truncatipennis by the low-lying, smooth-

ly margined crest and the emarginate male fifth sternite. Easily separated in

the males by the deep circular impression at the apex of the fifth sternite.

Notoxus aztecorum, n. sp.

(Figs. 36, 61)

Length 2.91-3.34. Head and pronotum red-brown to brown, elytra tan

with brown markings: postmedian band extending anteriorly along elytral

08 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

suture to base, apical band extending anteriorly along elytral suture to post-

median band. Pubescence brownish when arising in dark areas, yellowish in

tan areas. Raised setae distinct on pronotum and elytra, average angle on

elytra 18°. Mandibular explanation moderately wide, front and hind margins

meeting mandible at same level. Pronotum with antebasal pubescent band

widely broken. Horn without pits beneath, no teeth on margin. Crest margin

smooth, apex rounded and distinct. First and second protarsomeres with

strong spur on posterior apical margin.

Males with fifth sternite emarginate at apex.

Females with fifth sternite rounded-triangular.

% holotype: Mixcoac, Distrito Federal, MEXICO. Length 3.05. Head

0.65 long, eye 0.24 long, raised setae above eyes average 0.12 long; antenno-

mere ratios: 12.19, 111.67, 112.08, IV 1.70, V 1.74, Vill 1.05, X 0.92,

XI 1.71; fourth maxillary palpus 0.18 long, 0.10 wide; mandibular explanation

0.10 long, 0.06 wide. Pronotum 0.98 long, length from base to point of great-

est width 0.38, break in antebasal pubescent band 0.29 wide; horn 0.36 long,

0.30 wide at base; crest 0.28 long, 0.12 wide at base; crest apex 0.04 over

horn margin, crest apex 0.14 from horn apex; pronotal pit 0.10 long. Raised

setae 0.11 long, average angle 17°. Emargination of fifth sternite 0.23 wide,

0.05 deep.

Specimens examined, 3: HOLOTYPE, Mixcoac, Distrito Federal,

MEXICO, Wickham (USNM). PARATYPES: MEXICO: Distrito Federal: 1¢,

Barrett (CASC); 19, Teotihuacan, 3 June 1948, F. Werner, ground (FGW).

Distribution: The region around Mexico City.

Discussion: It is most similar to trvuncatipennis and desperatus. Separated

from the former by the lack of an impression on the male fifth sternite. It is

separated from the latter by the lack of a carina on the male fifth sternite and

the apically rounded penis.

Notoxus desperatus, n. sp.

(Figs. 38, 61)

Length 2.98-3.20. Head and pronotum orange to red-brown, elytra tan

with brown markings: postmedian band extending anteriorly along elytral

suture, covering omoplates and reaching base of elytra, apical band meeting

postmedian band along elytral suture. Pubescence brownish when arising

from markings, yellowish over rest of body. Raised setae present on pronotum

and elytra, average angle on elytra 20°. Mandibular explanation moderately

wide, front and hind margin meeting mandible at same level. Pronotum with

antebasal pubescent band widely broken. Horn without pits beneath, no teeth

on margin. Crest margin smooth, apex distinct, often pointed. First and

second protarsomeres with strong spur on posterior apical margin.

Males with fifth sternite emarginate, flattened and with transverse carina

at center. Apex of penis widely split.

Females with fifth sternite rounded-triangular.

Oo holotype: Durango, Durango, MEXICO. Length 3.20. Head 0.68 long,

eye 0.22 long, raised setae above eyes average 0.12 long; antennomeres miss-

ing; fourth maxillary palpus missing; mandibular explanation 0.12 long, 0.06

wide. Pronotum 1.02 long, length from base to point of greatest width 0. 34,

break in antebasal pubescent band 0.28 wide; horn 0.42 long, 0.32 wide at

base; crest 0.30 long, 0.13 wide at base, apex pointed; crest apex 0.04 above

horn margin, horn apex 0.12 from crest apex; pronotal pit 0.11 long. Raised

Chandler: Notoxus of Central and South America 59

elytral setae 0.13 long, average angle 24°. Emargination of fifth sternite

0.26 wide, 0.04 deep.

Specimens examined, 3: HOLOTYPE“, Durango, Durango, MEXICO

(USNM). PARATYPES: MEXICO: Durango: 1, eutopotypical (DSC); 19,

Durango, Wickham (MCZC).

Distribution: Only known from near the city of Durango.

Discussion: It is most similar to truncatipennis and aztecorum. Separ-

ated from both by the medial transverse carina of the male fifth sternite and

the apex of the penis being forked.

Notoxus truncatipennis Champion

(Pigs, 37, G1)

Notoxus truncatipennis Champion 1890: 209. Type locality: Guanajuato,

Guanajuato, MEXICO. BMNH Lectotype 2, designated by D. S. Chandler,

1975. Pic 1894: 47; 1911: 13.

Length 2.78-2.94. Head and pronotum brown, elytra brown with red-

brown markings: median band extending anteriorly along elytral suture over

omoplates to base, apical band extending along elytral suture to median band.

Pubescence brownish in dark areas, whitish in light areas. Raised setae

present on pronotum and elytra, average angle on elytra 31°. Mandibular

explanation wide, front and hind margins meeting mandible at same level.

Pronotum with pubescent antebasal band widely broken. Horn without pits

beneath, no teeth on margin. Crest margin smooth, apex pointed, usually

distinct. First and second protarsomeres with strong spur on posterior api-

cal margin.

Males with fifth sternite emarginate, shallow transverse impression before

apex.

Females with fifth sternite rounded-triangular.

Redescribed 7: Ridge NW of Jocoque Dam, Aguascalientes, MEXICO (DSC).

Length2.78. Head 0.62 long, eye 0.21 long, raised setae above eyes average 0. 09

long; antennomere ratios: 12.15, 111.55, 1112.28, IV 1.59, V 1.58, VII

1.11, X 1.10, XI 2.00; fourth maxillary palpus 0.16 long, 0.09 wide; mandibu-

lar explanation 0.10 long, 0.07 wide. Pronotum 0.96 long, length from base

to point of greatest width 0.32, break in antebasal pubescent band 0.25 wide;

horn 0.42 long, 0.30 wide at base; crest 0.32 long, 0.11 wide at base, apex

indistinctly merging with horn; crest apex 0.02 above horn margin, horn apex

0.16 from crest apex; pronotal pit 0.08 long. Raised elytral setae 0.09 long,

average angle 32°. Emargination of fifth sternite 0.24 wide, 0.02 deep.

Specimens examined, 3: MEXICO: Aguascalientes: Ridge NW of Jocoque

Dam, 19 August 1960, P. H. Arnaud, Jr., E. S. Ross, D. C. Rentz (CASC,

DSC). Guanajuato: Guanajuato, Lectotype of truncatipennis (BMNH).

Distribution: On the southern margin of the Mexican Central Plateau.

Discussion: Similar to aztecorum, desperatus, fraternus and stephani by

the low-lying, smoothly margined crest and the emarginate fifth sternite.

Separated by the shallow medial impression of the fifth sternite of the male.

60 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Notoxus zapotecorum, n. sp.

(Figs. 46, 62)

Length 2.75-3.38. Head and pronotum orange to red-brown, elytra tan

with brown markings: circular marks covering omoplates, often meeting

postmedian band extending along elytral suture, apical band extending along

elytral suture to postmedian band. Pubescence brownish when arising from

markings, yellowish in light areas. Raised setae distinct on pronotum and

elytra, average angle on elytra 32°. Mandibular explanation narrow, often

with margins not meeting mandible at same level. Pronotum with antebasal

pubescent band widely broken. Horn without pits beneath, 2-7 teeth on mar-

gin. Crest margin smooth, apex usually rounded. First and second protarso-

meres with strong spur on posterior apical margin.

Males with fifth sternite slightly emarginate, transverse impression be-

fore apex.

Females with fifth sternite rounded-triangular.

S holotype: Jesus Maria, Nayarit, MEXICO. Length 3.28. Head 0.68

long, eye 0.27 long, raised setae over eyes average 0.10 long; antennomere

ratios; 12.50, 11.6%, 111 2,25, 1V 2.00, V 1.74, VOL1.82, X.1.21, X1 2.35;

fourth maxillary palpus 0.20 long, 0.11 wide; mandibular explanation 0.10

long, 0.05 wide. Pronotum 1.09 long, length from base to point of greatest

width 0.35, break in antebasal pubescent band 0.26 wide; horn 0.46 long, 0.32

wide at base, 2 teeth on margin; crest 0.29 long, 0.12 wide at base, apex

rounded; crest apex 0.04 above horn margin, horn apex 0.16 from crest apex;

pronotal pit 0.12 long. Raised elytral setae 0.11 long, average angle 33°.

Emargination of fifth sternite 0.10 long, 0.01 deep.

Specimens examined, 23: HOLOTYPE ©, Jesus Maria, Nayarit, MEXICO,

6 July 1955, B. Malkin (CASC). PARATYPES: MEXICO: Sinaloa: 2c,

Choix, 5 August 1968, T. A. Sears, R. C. Gardner, C. S. Glaser (UCDC);

aco’, 5.5 mi NW Choix, 14 July 1968, T. A. Sears, R. C. Gardner, C. S.

Glaser (UCDC); 3°, 8 miS Elota, 26 August 1963, L. A. Stange (UCDC); 1,

Culiacan, 17 August 1962, A. E. Michelbacher (CISC). Nayarit: 2o', 329,

Jesus Maria, 26 June 1955, B. Malkin (CISC). Puebla: 222, Tepexco, Route

115, 4000’, 9 August 1965, G. E. Ball & D. R. Whitehead, blacklight (FGW);

1°, 2 mi NW Tehuacan, 25 April 1953, R. C. Bechtel & E. I. Schlinger,

sweeping alfalfa (CISC); 1°, 6 mi SW Tehuacan, 7 July 1973, Mastro & Schaf-

fner, at light (TAMU). Oaxaca: 300’, 10 mi NW Oaxaca, 22 July 1974, R. L.

Mangan & D. S. Chandler, attracted to box of drying Meloidae (DSC); 1°,

Huajuapan de Leon, 28 June 1971, C. W. O'Brien & G. B. Marshall (TTCC).

Distribution: Western and south-central Mexico. Collected sweeping

alfalfa. Attracted to light, ultra-violet light and freshly killed meloid beetles.

Present from April through August.

Discussion: It is most similar to marginatus of the apicalis-group. It

can be separated by the shorter raised elytral setae, the moderately wide

mandibular explanation and the smaller size.

Notoxus campus, n. sp.

(Figs. 31, 62)

Length 2.48-3.63. Head and pronotum orange to red-brown, elytra red-

Chandler: Notoxus of Central and South America 61

brown, elytra red-brown to brown. Pubescence whitish over body. Raised

setae long and erect on pronotum and elytra, average angle on elytra 32°.

Mandibular explanation narrow, hind margin meeting mandible at lower level

than front margin. Pronotum with pubescent antebasal band widely broken.

Horn without pits beneath, 0-2 teeth on margin. Crest margin smooth, apex

rounded and distinct. First and second protarsomeres with strong spur on

posterior apical margin.

Males with fifth sternite emarginate.

Females with fifth sternite rounded-triangular.

So holotype: km 210, Highway 57 N of San Luis Potosi, San Luis Potosi,

MEXICO. Length 2.89. Head 0.62 long, eye 0.23 long, raised setae over

eyes average 0.16 long; antennomere ratios: 12.25, I11.54, II 1.77, IV

1.75, V 1.50, VU1L1.56, X 1.25, XI 2.26; fourth maxillary palpus 0,17 long,

0.10 wide; mandibular explanation 0.12 long, 0.04 wide. Pronotum 1.00 long,

length from base to point of greatest width 0.32, break in antebasal pubescent

band 0.22 wide; horn 0.42 long, 0.28 wide at base, 1 tooth on margin; crest

0.25 long, 0.10 wide at base; crest apex 0.12 above horn margin, horn apex

0.27 from crest apex; pronotal pit 0.11 long. Raised elytral setae 0.17 long,

average angle 35°. Emargination of fifth sternite 0.18 wide, 0.03 deep.

Specimens examined, 24: HOLOTYPE ©“, km 710, Highway 52 N of San

Luis Potosi, San Luis Potosi, MEXICO, 11 July 1969, Ward, Tenorio, Ben-

nett (USNM). PARATYPES: Coahuila: 1, 19, 21 mi SE Saltillo, 6000', 18

July 1963, H. F. Howden (CNCI); 20°o, 399, 33 mi SE Saltillo, nr. Jame,

7500', 18 July 1963, A. T. Howden (CNCI). Nuevo Leon: 2, 1°, 10 miS

Junction 57 & 60, 23 June 1971, G. Brothers, acacia (TTCC). San Luis

Potosi: 50°, 829, eutopotypical (TTCC).

Distribution: East side of the Mexican central plateau.

Discussion: It is distinctive in the long erect setae on the unicolorous

elytra. The mandibular explanation is narrow and the fifth sternite is emar-

ginate without any impression.

Notoxus crucifer Champion

(Figs. 41, 62)

Notoxus crucifer Champion 1890: 210, Table IX, fig. 18. Type locality: San

Geronimo, Guatemala. BMNH Type &. Pic 1894: 46; 1911: 7.

Length 2.92-3.75. Head and pronotum red-brown to brown, elytra tan

with brown markings: circular mark covering omoplates, median band often

reaching omoplates along elytral suture, apical band often reaching median

band along elytral suture. Pubescence brownish when arising from dark areas,

whitish in light areas. Raised setae distinct on pronotum and elytra, average

angle on elytra 429. Mandibular explanation moderately wide, rarely with

front and hind margins meeting mandible at same level. Pronotum with ante-

basal pubescent band widely broken. Horn without pits beneath, 3-8 teeth on

margin. Crest margin often crenulate, apex pointed, usually indistinct and

merging with horn. First and second protarsomeres with strong spur on

posterior apical margin.

Males with fifth sternite slightly emarginate, circular impression before

apex.

Females with fifth sternite rounded-triangular. Elytral apices slightly

pointed.

62 Contrib. Amer. Ent. Inet., vol. 15, no. 3, 1977

Redescribed “: Junction Highways 190 & 195, Chiapas, MEXICO (DSC).

Length 3.65. Head 0.80 long, eye 0.30 long, raised setae over eyes average

0.11 long; antennomere ratios: 12.32, 111.57, II 2.20, IV 1.56, V 1.70,

VIII 1.25, X 1.14, XI 2.33; fourth maxillary palpus 0.22 long, 0.13 wide;

mandibular explanation 0.11 long, 0.04 wide. Pronotum 1.20 long, length from

base to point of greatest width 0.38, break in antebasal pubescent band 0.30

wide; horn 0.52 long, 0.32 wide at base, 3 teeth on margin; crest 0.29 long,

0.10 wide at base, margin crenulate, apex indistinct and merging with horn;

crest apex 0.04 over horn, horn apex 0.22 from crest apex; pronotal pit 0.12

long. Raised elytral setae 0.12 long, average angle 41°. Emargination of

fifth sternite 0.15 wide, 0.01 deep.

Specimens examined, 18: MEXICO: Chiapas: Junction Highways 190 &

195, 20 May 1969, J. M. Campbell (CNCI); same locality, 6 June 1969, H. F.

Howden & J. M. Campbell (CNCI); same locality, 24 June 1973, G. Ekis

(DSC, USNM); 5 mi W San Cristobal, 7500', 23 May 1969, J. M. Campbell

(CNCI); Zinacantlan, 1900 meters, 27 June 1973, G. Ekis (DSC). GUATE-

MALA: San Geronimo, Champion, Holotype female of N. crucifer (BMNH).

Distribution: Chiapas and Guatemala. Present from May through June.

Discussion: It is most similar to opacus by the usually crenulate crest

margin, the moderately raised setae on the elytra, and the apical emargina-

tion of the male fifth sternite. It is separated by the shallow emargination

and circular impression of the male fifth sternite, and the cylindrical first

metatarsomere of the male.

Notoxus opacus Champion

(Figs. 40, 62)

Notoxus opacus Champion 1890: 207, Table IX, fig. 16. Type locality: Cerro

de Plumas, Veracruz, MEXICO. Holotype ~ in BMNH. Pic 1894: 47;

1911: 10. Not Pallister 1955: 6.

Length 3.45-3.85. Head and pronotum red-brown to brown, elytra red-

brown to brown with dark brown markings: oval mark on omoplates often

expanded laterally to form postbasal band, broken along elytral suture, often

extending posteriorly along elytral suture to postmedian band, mark on apices

extending anteriorly along elytral suture. Pubescence brownish when arising

from marks, whitish over rest of body. Raised setae moderately long and

erect on pronotum and elytra, average angle on elytra 40°. Mandibular ex-

planation narrow, rarely with front and hind margins meeting mandible at

same level. Pronotum with antebasal pubescent band widely broken. Horn

without pits beneath 4-7 teeth on margin. Crest margin usually smooth, apex

pointed, rarely with apex indistinct and merging with horn. First and second

protarsomeres with strong spur on posterior apical margin. :

Males with fifth sternite deeply emarginate, transversely impressed im-

mediately before apex, first metatarsomeres enlarged and flattened.

Females with fifth sternite rounded-triangular.

Redescribed “: Puebla, Puebla, MEXICO (CASC). Length 3.38. Head

0.74 long, eye 0.26 long, raised setae over eyes average 0.13 long; antenno-

mere ratios: 12.50, 111.25, HE2.50, 1V 2.25, V 1.80, VU ft, 45, X71, 34,

XI 2.26; fourth maxillary palpus 0.20 long, 0.11 wide; mandibular explanation

0.10 long, 0.04 wide. Pronotum 1.04 long, length from base to point of great-

est width 0.32, break in antebasal pubescent band 0.25 wide; horn 0.44 long,

Chandler: Notoxus of Central and South America 63

0.27 wide at base, 5 teeth on margin; crest 0.28 long, 0.08 wide at base,

apex distinct and smooth; crest apex 0.04 above horn margin; horn apex 0.16

from crest apex; pronotal pit 0.12 long. Raised elytral setae 0.11 long, aver-

age angle 349. Emargination of fifth sternite 0.15 wide, 0.02 deep.

Specimens examined, 6: MEXICO: Puebla: Puebla, May, A. Fenyes coll.

(CASC); 5 mi SW Chipilo, 26 June 1975, D. S. Chandler, sweeping low vege-

tation (DSC). Oaxaca: 3.2 miS Ocotlan, 20 July 1974, Clark, Murray, Ashe,

Schaffner (TAMU); 30 mi NW Oaxaca, 6500', 27 June 1965, Burke, Meyer,

Schaffner (DSC). The holotype record is from Veracruz.

Distribution: South-central Mexico. At 5 mi SW Chipilo two females were

collected sweeping, while none were attracted to either cantharidin/or drying

meloid beetles which had been placed within thirty meters of the collection

site. Present from May through July.

Discussion: It is most similar to crucifer. It is separated by the deep

'emargination and transverse impression of the male fifth sternite. The first

metatarsomeres of the male are enlarged and flattened.

Notoxus sp.

(Figs. 17, 58)

2: Foz do Iguacu, Parana, BRAZIL (MGF). Body brown, no markings on

elytra. Pubescence whitish. Raised setae long and erect on pronotum and

elytra, average angle on elytra 47°. Mandibular explanation moderately wide,

front and hind margins on same level at mandible. Pronotum with antebasal

pubescent band widely separated. Horn without pits beneath, 6 teeth on mar-

gin. Crest margin serrate, apex pointed and indistinct. Carina extending

from horn base to anterior margin of pronotal pits. First and second protarso-

meres lacking large spur on posterior apical margin.

Male unknown.

Female with fifth sternite triangular.

Length 2.60. Head 0.58 long, eye 0.17 long, raised setae over eyes

average 0.16 long; antennomere ratios: 12.00, I1 1.90, III 2.38, IV 2.25,

V 2.25, antennomeres VII-XI missing; fourth maxillary palpus 0.16 long,

0.08 wide; mandibular explanation 0.08 long, 0.05 wide. Pronotum 0.91 long,

length from base to point of greatest width 0.32, break in antebasal pubescent

band 0.24 wide; horn 0.37 long, 0.29 wide at base; crest 0.24 long, 0.12 wide

at base; crest apex merging into horn, horn apex 0.16 from crest apex; pro-

notal pit 0.11 long. Raised elytral setae 0.15 long.

2: Foz do Iguacu, Parana, BRAZIL, November 1955, G. B. Frey (MGF).

Distribution: Southern inland Brazil.

Discussion: This species is very distinct in the possession of lateral

carinae between the horn base and the cervix. This character is found only

in the African and Indian Notoxus with which this species has a high degree of

resemblance. It is probable that this species has been introduced from the

Old World and it will require the collection of the male in order to decide if

the species has been described.

64 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Incertae sedis

Notoxus dugesi Pic

Notoxus Dugesi (sic) Pic 1901: 89. Type locality: Guanajuato, Guanajuato,

MEXICO. Type in MNHN. Pic 1911: 7.

Pic's description of this species is typical in its brevity and gives no

clues as to which species it could be.

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New species of Coleoptera collected by the expeditions for

geographical surveys west of one hundredth meridian, in charge

of Lieut. Geo. M. Wheeler, United States Engineers, U. S.

Army, 1876, Appendix JJ, p. 516-520.

Leopold, A. S. 1950. Vegetation zones of Mexico. Ecology 31: 507-18.

Leng, C. W.

Loan, CC...

1920. Catalogue of the Coleoptera of America, North of

Mexico. John D. Sherman, Jr., Mount Vernon, N. Y.

x + 470 p.

1972. Parasitism of adult Notoxus anchora Henz.

(Coleoptera: Anthicidae) by Syrrhizus agilis (Cress. )

(Hymenoptera: Braconidae). Proc. Entomol. Soc. Ontario

103-76.

Orphanides, G. M., D. Gonzalez and B. R. Bartlett. 1971. Identification

Pallister, J.

Peterson, A.

and evaluation of pink bollworm predators in southern California.

Jour. Econ. Entomol. 64: 421-424.

C. 1955. The ant-like flower beetles of north-central

Mexico... . American Museum Novitates, #1720, 18 p.

1951. Larvae of insects, an introduction to Nearctic species.

Part II, Coleoptera, Diptera, Neuroptera, Siphonaptera,

Mecoptera, Trichoptera. Edwards Brothers, Inc., Ann Arbor,

Michigan. 416 p.

Pic, M. 1894. Liste des Anthicides décrits postérieurment au catalogus de

1897,

1901a.

MM. Gemminger et Harold (1870-1893). Ann. Soc. Entomol.

Belgium 41: 212-224.

Premier supplement a ma liste des Anthicides. Ann. Soc.

Entomol. Belgium 41: 2.2-224.

Diagnoses d'Anthicidae exotiques. Ann. Soc. Entomol.

Belgium 45: 89-91.

68 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

1901b. Descriptions de Coleopteres nouveaux. Le Naturaliste 23: 227-8.

1904. Description de deux ptinides et d'un Notoxus faisant partie des

collections du Museum de Paris. Bull. Mus. Hist. Nat.

10: 226-8.

1911. Anthicidae, Part 36. In: Coleopterorum catalogus. Ed. S.

Schenkling. W. Junk, Berlin. 102 p.

1912. Coleopteres nouveaux de diverses familes originares de la

republique Argentine. Anal. Mus. Nac. Buenos Aires 22: 455-8.

1913. Descriptions de 29 Especes. Melanges Exotico-Entomol.

dD: 8-9.

1914a. Coleopteres nouveaux du genre Notoxus Geofr. (Hétéromeéres).

Bull. Soc. Zool. France 39: 250-1.

1914b. Coleopteres exotiques en partie nouveaux. L'Echange 30: 61-2.

1916. Diagnoses specifiques. Melanges Exotico-Entomol. 17: 11.

1918. Anthicides exotiques nouveaux (Col. Heteromera). Bull. Soc.

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Entomol. Zeit. 30: 119-128.

Rosen, D. E. 1975. A vicariance model of Caribbean biogeography. Syst.

Zool. 24: 431-464.

Say, T. 1817. American Entomology or descriptions of the insects of

North America. S. A. Mitchell and Ames. Unnumbered pages.

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van Hille, J. C. 1954. Cantharidin and Anticidae (sic). South African Jour.

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Life. Eds. Hanstrom, Brinck, Rudebeck. Almaqvist and Wiksells,

Uppsala. Vol. 8, 557 p.

Werner, E. G. 1964. A revision of the North American species of Anthicus,

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America 4: 195: 242.

Chandler: Notoxus of Central and South America 69

1965a. Family Anthicidae, p. 122-29. In: The Beetles of the Pacific

Northwest. Ed. M. H. Hatch. University of Washington Press,

Seattle. Vol. IV, vii + 268 p.

1965b. A preliminary account of the Anthicidae of Venezuela. Rev.

Facul. Agronomia (Maracay) 3: 10-23.

70 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

penis

—— paramere

phallobase

horn

mandibular

explanation

tooth

crest

longitudinal

ridge

antebasal impression

Figures 1-6. 1. Hindwing of N. arizonensis, 2. Ventral view of gener-

alized Notoxus genitalia. 3. Left lateral view genitalia of Plesionotoxus

lebasi. 4. Ventral view genitalia of Plesionotoxus lebasi. 5. Dorsal view

of generalized Notoxus pronotum. 6. Posterior view mandibular explanation

N. haustrus.

Chandler: Notoxus of Central and South America

0.5mm

D E

7 pueblensis 8 (aia

leonensis 9 gelidus

D E

10 hirsutus

12 occidentalis

16 toltecorum

15 nubilus

i nsi 3

13 arizonensis 14 durangoensis

Figures 7-16. Notoxus spp. A. Elytral color pattern. B. Male fifth

sternite. C. Ventral view mandibular explanation. D. Ventral view male

genitalia. E. Left lateral view male genitalia.

1977

Io A

18 eximius 20 + solus

19 lateralis

21° impressus

S le =

: { )

} \ ie

( ( ) D E

. : : ‘ e : 26 bipunctatus

22 acuminatus os

cumanensis

23 desertus

D E

24 celatus

Figures 17-26. Notoxus spp. A. Elytral color pattern. B. Male fifth

sternite. C. Ventral view mandibular explanation. D. Ventral view male

genitalia. E. Left lateral view male genitalia.

27 jamaicus

Chandler: Notoxus of Central and South America 15

~~)

29 ruficollis 30 peruvianus

A>.

VAY Oe

32 nuperus

34 postictus

D E 35 orientalis

33 haustrus

36 aztecorum

Figures 27-36. Notoxus spp. A. Elytral color pattern. B. Male fifth

sternite.

genitalia.

C. Ventral view mandibular explanation.

E. Left lateral view male genitalia.

D. Ventral view male

74 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

37 truncatipennis

D E D E

D E D E 41. crucifer

40 opacus

38 desperatus

39 stephani

42 talpa

pygidialis

44 photus

Figures 37-44. Notoxus spp. A. Elytral color pattern. B. Male fifth

sternite. C. Ventral view mandibular explanation. D. Ventral view male

genitalia. E. Left lateral view male genitalia. F. Female elytra.

Chandler: Notoxus of Central and South America 15

D E

| ) f 46 zapotecorum

o Yt (

45 fraternus D E

49 mexicanus

: j ‘ D E

. :

47 apicalis E

48 marginatus

50 anchora

t c

53 conformis

52. sparsus

D E

54 bajae

Figures 45-54. Notoxus spp. A. Elytral color pattern. B. Male fifth

sternite. C. Ventral view mandibular explanation. D. Ventral view male

genitalia. E. Left lateral view male genitalia.

51. serratus

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

®calcaratus

@pueblensis

*hirsutus

Woccidentalis

@gelidus

@leonensis

@lateralis

Simpressus

keximius

Wacuminatus

Gsolus

Figures 55-56. Distribution maps. 55. Pueblensis -group and

occidentalis. 56. Eximius-group.

Notoxus of Central and South America

Chandler

®celatus

Bcumanensis

kdesertus

Wruficollis

@jamaicus

Sbipunctatus

®murinipennis

Figure 57. Distribution map of monodon-group.

78 Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Figure 58.

®peruvianus

*Notoxus sp.

Distribution map of peruvianus and Notoxus sp.

Notoxus of Central and South America

Chandler

@®arizonensis

Snubilus

* durangoensi

®toltecorum

®postictus

@haustrus

xk orientalis

nuperus

tZONENSTS- Group.

09.

Distribution maps.

Figures 59-60.

Nuperus-group.

60.

Figures 61-62.

62.

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

ED Aer

®stephani

@fraternus

* truncatipennis

Waztecorum

Odesperatus

™4,

fps SE fe

aA:

®zapotecorum

@ campus

*crucifer

Wopacus eS

Distribution maps of species not placed in groups.

61. Stephani, fraternus, truncatipennis, aztecorum, and desperatus.

Zapotecorum, campus, crucifer, and opacus.,

Chandler: Notoxus of Central and South America

a8 La

aot ) ars ‘f

@ apicalis &) Lr 2 ae |

\- oe pus as, i Vi,

BS al \, , ? :

@ mexicanus oe as oe Peay,

( } BD Sagan NN Ce irae ark

; é y ‘) Sea en onda een pee

* marginatus He ®) 7 Aj eas

f / by

®talpa 5

*pygidialis | AOR

@photus mee

' !

Le Bd atl

63 : Beet

t t

i '

. / we Ava (

® bajae S ‘ bod te se

woot iN re ae cay i

&conformis \ ° te ee if . a [0

PST SR 9 / 4, SOA, faa Ms =

ane Teak lat pS a

* sparsus Cua ee. ian

AT at S : :

Wserratus iene 2 ig S ve VePRD

\ a Se itis eee £3 Wp

64 : ) \s oe 2Ay Cy.

2 tes

We ee ee ar

, ee

Figures 63-64. Distribution maps. 63. Apicalis- and talpa-groups.

64. Sparsus- and monoceros-groups.

Contrib. Amer. Ent. Inst., vol. 15, no. 3, 1977

Correlation

calcaratus

gelidus

hirsutus

caudatus

pueblensis

leonensis

arizonensis

durangoensis

nubilus

tolfecorum

balteatus

subtilis

bifasciatus

montanus

parvidens

schwarzi

solus

n. sp. - U.S.

desertus

cumanensis

monodon

celatus

jamaicus

murinipennis

bipunctatus

ruficollis

peruvianus

planicornis

filicornis

campus

nuperus

postictus

orientalis

haustrus

aztecorum

brevicornis

talpa

pygidialis

photus

crucifer

truncatipennis

stephani

desperatus

marginatus

apicalis

mexicanus

pallidus

zapotecorum

fraternus

n. sp.-U.S.

opacus

conformis

sparsus

n. sp.- U.S.

spatulifer

denudatus

breviusculus

cavicornis

n. sp.- U.S.

bajae

lustrellus

anchora

robustus

pictus

serratus

nevadensis

Figure 65. Phenogram of New World Notoxus produced by numerical

analysis using correlation.

Chandler: Notoxus of Central and South America

INDEX TO GENERIC AND SPECIFIC NAMES

USED IN KEY AND DESCRIPTIONS

acuminatus Champion, 12, 30

anchora Hentz, 13, 50

apicalis LeConte, 17, 44

argentinus (Pic), 8

arizonensis Fall, 17, 23

atripennis Champion, 38

aztecorum n. sp., 17, 57

bajae n. sp., 13, 53

balteatus Casey, 15, 23

bicolor Say, 37

bipunctatus Chevrolat, 13, 35

boliviensis (Pic), 9

brevicornis Fall, 17, 55

calcaratus Horn, 12, 22

campus n. sp., 15, 60

caudatus Fall, 16, 18

celatus n. sp., 13, 32

conformis LeConte, 13, 54

constrictus Casey, 34

cristatus Champion, 21

crucifer Champion, 17, 61

cumanensis LaFerte, 13, 31

desertus Casey, 13, 33

desperatus n. sp., 17, 58

digitatus LeConte, 51

dugesi Pic, 64

durangoensis n. sp., 15, 24

elegantulus LaFerte, 47

eximius Champion, 14, 27

fraternus Champion, 14, 56

gelidus n. sp. 16, 20

germaini (Pic), 9

gounellei (Pic), 9

haustrus n. sp., 15, 40

hirsutus Champion, 12, 21

informicornis (Krekich), 9

innotatipennis (Pic), 9

impressus Champion, 14, 28

jamaicus Pic, 13, 36

krugi Quedenfeldt, 35

lateralis n. sp., 13, 28

lebasi (LaFerte), 8

leonensis n. sp., 16, 19

marginatus LeConte, 16, 45

mexicanus Champion, 17, 46

Monocerus Faldermann, 9

montanus Casey, 16, 23

murinipennis (LeConte), 13, 37

Notoxus Fabricius, 9

nubilus n. sp., 15, 25

nuperus Horn, 15, 39

nuperoides Fall, 44

occidentalis n. sp., 16, 55

Opacus Champion, 17, 62

orientalis n. sp., 15, 42

pallidus Fall, 12, 44

peruvianus Pic, 12, 38

photus n. sp., 14, 49

Plesionotoxus n. gen., 8

postictus n. sp., 15, 41

pueblensis Champion, 16, 18

pygidialis n. sp., 14, 48

quinquemaculatus Pic, 36

ruficollis Champion, 12, 38

serratus (LeConte), 16, 51

solarii Pic, 27

solus n. sp., 13, 29

sparsus LeConte, 13, 52

stephani n. sp., 17, 57

talpa LeFerte, 14, 47

toltecorum n. sp., 15, 26

truncatipennis Champion, 17, 59

vandykei Blaisdell, 51

ventralis Champion, 47

venustus (Pic), 9

zapotecorum n. sp., 17, 60

Contributions

of the

American Entomological Institute

Volume 15, Number 4, 1978

A REVIEW OF THE NEW WORLD GENUS

ATHYREUS MACLEAY

(SCARABAEIDAE, GEOTRUPINAE, ATHYREINI)

Henry F. Howden and Antonio Martinez

A REVIEW OF THE NEW WORLD GENUS ATHYREUS MACLEAY

(SCARABAEIDAE, GEOTRUPINAE, ATHYREINI)

Henry F. Howden! and Antonio Martfnez”

ABSTRACT

Thirty-one species are included in the genus Athyreus Macleay. Seven-

teen previously proposed names are considered valid, six are considered

Synonyms, and three names, billbergi Gray, subarmatus Westwood and

vicinus Laporte are listed as incertae sedis. Fourteen new species are

described: acuticornis from Brazil, alvarengai from Brazil, anneae from

Brazil and Paraguay, biarmatus from Trinidad, bilobus from Brazil,

brasilicus from Brazil, capricornis from Brazil, conspicuus from Argentina,

hypocritus tte Argentina and Paraguay, juanae from Brazil, nitidus from

Bolivia, parvus from Brazil, pyriformis from Peru, and unicornis rnis from

Colombia. The introduction and keys are given in both Spanish and English.

Following the incertae sedis a list of the species included in the genera

Neoathyreus and Parathyreus is given.

CONTENTS

INCNOOUCHION . 6 Us ye oe ee ee Se ee ee 2

ReSWHCN) 2656. 6 Oe See ee ie - eee Cee es =

Collections Sudied . 5°... 6 kes CR ae eee Ce is eos 5

Key to the species of Athyreus Macleay ....... ea Bb ees 5

Clave para Athyreus Macleay ..... od amines oO ees ; 12

Species descriptions

i. ee scene lie. 6 fe ee ek ae eS 20

Ai CAPCOM GA SRG sa kk ek eke 4 ee ees oe 21

Bs ACUEUS GIS. 2. we eke oe ee 6 en ene ee a 22

fn MOAI BOUPOMNOM Ada Re a ee ee Pe is Oe ee 23

BR. MVDOCC Me Ne SR. Se ee Wg a OR a eS ese ee eee 8 ae

6. BEAeimOus Ne Cie Aire Wh eee Een a ae 25

T. BiObUS B. 80s. Gon Gras Oe ee ee ue a ee 26

thepartment of Biology, Carleton University, Ottawa, Ontario, Canada.

“tnvestigador de Carrera. Consejo Nacional de Investigaciones

Cientificas y Técnicas. Buenos Aires, C.C. 21, 1642 San Isidro, Prov. Bs.

As., Argentina.

2 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

8. hemrspnsericus DOUuCOMmONE Core “ais ceeds OR en vee 28

Oo OMLETOUS Ws BO Oe ee seater 2) eee hy Susu n ogee tad ee 29

POs OMe de Be SOs es a a ee ee oo BO, ae ee ty tg gs qth 30

Tl .. JOVGipalus BOUCeINOnt spree er Fe a ER 31

LA e cee, ee ee ie he ee 31

bg sO Os ee ee ee ee ee ea en ew cas a2

es UO ES The (SO a ee a OR te ea ee a Ne a ss 33

iD. “armatus Wesmvood-s:. ta ee ree ra Gy 34

VG Ra oss nes ec ee een wae ee sles Generale se ce 35

PT Bie ae ese O0G oe se. Se Pe ON Ns 25S, 36

US a eek ea eee os ae bec, eels ee oe ee 6 37

19. biutcatus Macleay os ar Oe, eGo Wee EER? ge oegi a ee 38

OO. CGR ie ECC NOs se at GS eae Sas e Kine cet aac ees cee 39

Gils Weds We BOs ee ea re eee ete gi 41

22.2 Devator Weeiwo0d 2. 6k SS. Sea ee et 42

Doe Eee Gore. a a ee ee ke ae ele 42

Za POO ROM abe s 6 eee es Gee ee ee ee wes 43

Zoe PCULICOLRIS Tie SD. ees es POR Ee Ed ee ee ee ee lee 44

7G oo SEI Mis FeloChe 6 2) ee a ee ew we ee ts 46

OW gape Maines ke PS ea eee 47

28. martinezi Howden ... 1. ee eee ee ee ee ee ee eee 48

20>. Waster Folecne 6. i Se kee oe ea eee on 49

Oe Gel eed PoIriMaire | a ee eae pe ce ee kw 50

Ole COMSDICUUS TD. SPs. . be Fie See Ree ee we ee aed ec a1

eS SO re a Ne ge Ghat eo ee aoa a owiehe pe a, 53

List of species in Neoathyreus Howden and NMeptime: ie 5a

List of species in Parathyreus Howden and MARCCe fe eee we 54

Acknowledgments Soe oe oe es Eg eee eae Ome ane eps a as 54

POPC OCTUTe Cle oh ak ee ee en ew eri ee es Va we bes 54

PACU CS os ee ee ee kali aye OR ee ee aa 57

INTRODUCTION

In 1819 Macleay proposed the name Athyreus. Until 1963 the genus included

a heterogeneous mixture of New and Old World forms with some 61 names being

listed in the genus by Boucomont (1912). In 1963 Howden and Martinez created

the tribe Athyreini and restricted the genus Athyreus to include only a group

Howden and Martinez: The Genus Athyreus 3

of relatively large, sexually dimorphic species with at least four, often six,

relatively distinct intervals on each elytron. Athyreus bifurcatus Macleay

was designated as type for the genus but no complete list of included species

was attempted at that time as no key or revision for any of the groups had

been attempted.

Both authors have slowly been accumulating information and specimens,

and many of the types of Athyreus have been studied. We now consider 26 old

names referable to Athyreus as presently constituted. Three names are

based on females seemingly belonging to the bifurcatus complex of species

and we consider them unassignable, listing them as incertae sedis. The

other 23 names we believe can be assigned to 17 species with reasonable

accuracy. However, there is some uncertainty and when the lack of specimens

in series is also considered, we do not consider that a revision of the group

is feasible. Hence, we have merely tried to assign the earlier names,

describe fourteen new species and to present keys, brief descriptions, and

illustrations. The keys, unfortunately, are based largely on males except

for the chalybeatus group. Unless males and females can be associated in

the field their association is usually questionable.

In Athyreus, well developed males often have a well developed clypeal

horn and one or more pronotal horns or strongly elevated lobes. In contrast,

females, with one exception (tuberifer), have a transverse carina across the

anter ior face of the clypeus and a second transverse carina posteriorly at the

clypeal-frontal junction. This posterior carina is normally trituberculate,

with a distinct median tubercle and a smaller one on each side at the lateral

margin. In addition, female pronota may have several conspicuous carinae,

particularly in the chalybeatus group, and in the other groups low central

tubercles, but in only one species (tuberifer) do females have what could be

termed a pronotal "horn". If there is any doubt about the sex it is best to

check the genitalia.

Neither the use of the term "groups" in the keys or text nor the sequence

used in numbering the species is meant to imply phylogenetic relationships.

Rather atypical species, such as gigas Westw., armatus Westw., biarmatus

n. sp., unicornis n. sp., and pyriformis n. sp., are arbitrarily placed and

pyriformis at least could be assigned to either the aeneus or bifurcatus

groups. The divisions were established, at least in part, as a convenient

method of proportioning the descriptive work evenly between the authors.

Little can be said about the habits of Athyreus. Almost all specimens

have been, as far as we can ascertain, collected at light. A few have been

caught flying at dusk (crepuscular) but most are apparently nocturnal. The

genus seemingly occurs mainly below 1500 m in elevation in a variety of

habitats varying from relatively undisturbed lowland rainforest to seasonally

wet, semiarid sandy habitats. Unfortunately, attraction to light seems to be

an infrequent occurrence, and the rarity or lack of material in collections

has seriously hindered our studies. |

We have attempted to avoid special terms but, even so, our attempts to

describe the complexities of the pronotal carinae or male genitalia can easily

cause some confusion. Hopefully, the numerous illustrations will alleviate

this problem to a major degree.

4. Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

RESUMEN

Los autores presentan una tentativa de revisién del género Athyreus,

propuesto por Macleay en 1819. En 1912 Boucomont cataloga unas 60 especies

para el mismo, incluyendo formas heterogéneas del viejo y nuevo mundo. En

razon de tal anédmala situacion, Howden y Martinez en 1963 crean la tribu

Athyreini, restringiendo el genero Athyreus para el grupo de especies

relativamente grandes, con dimorfismo sexual y como maximo con 4 a 6

claros interespacios en cada elitro, designando como tipo a Athyreus

bifurcatus Macleay, pero sin incluir una lista completa de especies, hasta no

efectuar una clave o revision del mismo o de los otros grupo-género por

ellos creados.

Durante el tiempo transcurrido desde ése estudio, ambos autores

estuvieron acumulando informacion y material, habiendo sido estudiados en

ese lapso muchos de los tipos de Athyreus. En la actualidad han considerado

26 nombres referidos a especies del género; 3 de ésos nombres basados en

hembras asignadas al complejo "bifurcatus" y las que han considerado como

"incertae sedis''’. Los otros 23 nombres han podido ser asignados con

relativa seguridad a 17 especies. Sin embargo en alguno de los casos, la

falta de series de algunas de las especies, ha hecho no poder considerar a

este trabajo como una revisién en el sentido estricto y si, como lo sefalan

precedentemente como una primera tentativa. Eso ya se nota en la clave,

que dasafortunadamente ha tenido que ser basada en machos bien desarrollados,

con excepcidn hecha del grupo chalybeatus, vya que no siempre | machos y

hembras han podido ser asociados, siendo ésa una investigacion de campo.

En los Athyreus bien desarrollados, los machos tienen cuerno clipeal

y uno mas cuernos 0 l6bulos pronotales prominentes. Las hembras, con la

sola excepcidn tuberifer, tienen quilla transversal en la regién dorso-anterior

del clipeo y una segunda quilla, también transversal, en el limite clipeo-

frontal, siendo ésta ultima normalmente tridentada, con el diente o tubérculo

mediano mas desarrollado y uno menor a cada lado sobre el borde lateral.

Sin embargo, el pronoto de las hembras puede tener algunas quillas

conspicuas, especialmente en el grupo chalybeatus; en los otros grupos puede

haber éstas o tuberculitos centrales, exceptuando una sola especie, tuberifer,

en las que las hembras tienen un cuerno pronotal, necesitando, a veces, para

salir de dudas acerca del sexo, la verificacion del genital.

Referente a habitos y comportamiento de Athyreus es muy poco lo que

se sabe. Muchos de los ejemplares han sido colectados a la luz. De unos

pocos se conoce que vuelan durante el crepusculo, pero una gran mayoria

aparentemente son nocturnos. No son conocidos ejemplares del género

capturados por arriba de los 1500 m de altura y su "habitat" son aparentemente

los bosques relativamente no perturbados de las regiones bajas en la estacion

huméda o regiones arenosas semiaridas. Desafortunadamente la atraccion por

la luz no es muy frecuente y la rareza o falta de material en las colecciones

obstruye muy seriamente los estudios. Es esa falta de series razonables de

material en las colecciones,es lo que a hecho hacer un intento He agrupar y no

revisar el genero.

Como se nota, en éste intento se ha evitado una ieeeninilects especial,

por que ello puede llegar a causar alguna confusién cuando son descritos los

complejos cuernos, quillas o genitales; por ello son las numerosas

ilustraciones que se dan pensando que ése problema puede ser asi solucionado

en mayor o menor grado.

Para la mejor comprension de las personas de lengua espanola 0

portuguesa, es que ademas de Ja clave en inglés, se d& otra, poco mas

ampliada, en idioma espanol.

Howden and Martinez: The Genus Athyreus 5

COLLECTIONS STUDIED

In citing the collections studied we utilize the name of the city in which

the museum is situated, and in a private collection, we cite the individual's

last name. The major collections studied and the curators assisting are as

follows:

Buenos Aires Museo Argentino de Ciencias Naturales

"Bernardino Rivadavia". Dr. M.J. Viana.

Cambridge Museum of Comparative Zoology, Harvard

University. Dr. A. Newton.

Chicago Field Museum of Natural History. Mr. Henry Dybas.

Leiden Rijksmuseum van Natuurlijke Historie. Dr. J.

Krikken.

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

Ottawa Canadian National Collection, Biosystematics

Research Institute. Dr. A. Smetana.

Oxford University Museum, Hope Department of Entomology.

Mr. E. Taylor.

Paris Museum National d'Histoire Naturelle. Dr. A.

Descarpentries.

Washington | United States National Museum. Dr. R. Gordon.

Hardy Dr. A. Hardy, Sacramento, California.

The personal collections of the authors.

Deposition of types cited as 'Howden" are deposited on loan in the

Canadian National Collection, Biosystematics Research Institute, Canadian

Department of Agriculture, Ottawa, Canada.

Key to the species of Athyreus Macleay

i. Pronotum in male never with a complete median longitudinal

furrow, often with central median horn or tubercle or with

nearly cylindrical horn on each side; female pronotum

with central horn or with two small tubercles on disc, one

on either side of midline, or with transverse carina across

midline in anterior third of disc. ..... ee ee a ILS 2

Pronotum in male with median longitudinal furrow or shallow

depression narrow or wide, variable, essentially dividing

pronotal prominences (these not cylindrical); female

pronotum without tubercles on disc either side of midline

or without anterior transverse carina... . I.aeneus Group (p. 6)

3(2).

4(3).

5(3).

6(5).

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Pronotum of male never with lateral carina extending from

edge of a central depression to lateral edge; female pro-

notum with central horn or with two small tubercles near

center of disc, one on either side of midline....... .

GOR eo org ok ee ge Gon es Bate te eae II. bifurcatus Group (p. 8)

Pronotum of male and female with a lateral carina extending

from edge of discal depression to lateral edge, carina

sometimes briefly interrupted; female with transverse

carina in anterior third of discal region. .........-.

Cee Cee ee oe ee Td ehalypeatis Group (p. 11)

I. aeneus Group

Posterior border of pronotum on each side, anterior to

humerus, interrupted. Brazil..... 1. cyanescens Klug (p. ©)

Posterior marginal bead or border of pronotum entire .....

- Median anterior margin of pronotum more or less angular

CUNG) POLES igs ah vet gt sa van oe eciee wee A ve Sh wh ea ww eo 3

Median anterior margin of pronotum transversely keeled or

not, but-never unidentate -. 00.) 6 4 Bee wea oe re 7

Acute tubercle of anterior pronotal margin well marked,

SOMOtINES SMAil, 410 CCUG. oie Sa ee ee ek ee e's 4

Acute tubercle of anterior pronotal margin poorly developed,

Obeuse or obsolete. 2. secs ey ERS ee Gis te eae we see 9)

Pronotal disc with anterior transverse carina reaching

lateral edge; color dark bluish-green with more or less

bronze-caste. Brazil... «00% 66 aw oe 3. aeneus Klug (p. 22)

Pronotal disc with lateral carina obsolete, short, situated

on posterior half; color largely testaceous or castaneous.

Brazil Ge sa eee oe ee 4. vavini Boucomont (p. 23)

Pronotum of male with anterior depression broadly cordi-

form; longitudinal carina each side of midline costiform,

raised and not dentate; first (most anterior) transverse

carina reaching lateral edge; second (lateromedial)

carina broad, short, and shiny; third (posterior) carina

fine; color of dorsum shiny black with slight steely

reflections; pronotum of female with the three lateral

transverse carinae asin male. Brazil...... ‘

« hg GN son ao SRR HS Sa ps eke As capricornis n Nn. sp. “(p. 21)

Pronotum of male with medial depression circular or oval,

with longitudinal furrow impressed (distinct); transverse

carina, if present, not attaining lateral margin; discal

horn or carina on each side of midline dentate; pronotum

of female without transverse anterior carina or carina

extending to lateral margin; both sexes brown to dark

brOWR COrSally ius nace See we Ee ee ace ce ee 6

Male with horn or keel on either side of depressed midline

near middle of pronotum laminiform, the anterior and

posterior edges dentate; prescutellar region concave and

Howden and Martinez: The Genus Athyreus 7

with small punctiform impression. Brazil, Paraguay ;

bie. 6) DC etree Sie no ee ere 10. anneae n. sp. (p. 30)

Male with carina on each side of medial depression anteriorly

unidentate, feebly elevated; prescutellar region with median

furrow narrow, well impressed, each side elevated and

posteriorly dentiform. Brazil, E. Paraguay and N.

APOONUINA: ok we ve sees 8. hemisphaericus Boucomont (p. 28)

7(2). Anterior margin of pronotum in median fourth distinctly

elevated, carinate; carina dentate or abruptly arcuate on

GACH Stile. 5 ta bee eae ee ee ee ee es alien ae 8

Anterior edge of pronotum in median fourth arcuately

elevated; obsolete denticle often present at midline ..... EL

8(7). Pronotum of male on each side with lateral border of medial

depression elevated into a laminate, bidentate horn or

keel. “BOlNVid 2s ee ae a ee 9. nitidus n. sp. (p. 29)

Pronotum of male on each side with lateral border of

depression elevated; elevated carina or horn with apex

blunt to abruptly rounded, never bidentate. ...... ee 9

9(8). Male with clypeal horn robust, wider than long in cross

section, posterior face excavated or flat; pronotum on

each side with keel or horn which borders medial depression

wide and not angular, apically rounded, sometimes more so

posteriorly, terminating in front of and on each side of wide

prescutellar depression; depression large, subquadrate,

usually wider than long. Brazil, Paraguay. 7. bilobus n. sp. (p- 26)

Male with clypeal horn slender, as long as wide or longer

in cross section, posterior face narrow; pronotum with

keel or horn on each side of median depression lobate,

with apex unevenly rounded, blunt to acute; prescutellar

depression small, scarcely wider than scutellum. ...... 10

10(9). Male with prescutellar impression very small, normally

elongate and situated in median angle of posterior margin;

central depression without longitudinal line of median

granulations; medial carina of anterior margin slightly

wider than half width of prescutellar region, high, with

edge centrally slightly concave and laterally denticulate.

N.E. Argentina, Paraguay...... 5. hypocritus n. sp. (p. 24)

Male with prescutellar impression in form of a medial

longitudinal furrow; central depression with longitudinal

band of granulations which connects anterior and

posterior regions; medial carina of anterior margin as

wide as width of prescutellar region, low, with edge

centrally sinuate, not denticulate laterally. Brazil,

rae rere Rt aA HE AMM: -... 6. brasilicus n. sp. (p. 25)

11(7). Pronotal depression anterior to horns mostly smooth on

either side of median granulate area; setose granules

in elytral intervals distinctly separated, third interval

with moderately even double row of granules. Bolivia.

oe ee Rae eae ek a ee 11. forcipatus Boucomont (p. 31)

201).

(2).

4(2).

5(4).

6(5).

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Pronotal depression anterior to horns almost entirely

granulate, briefly smooth in narrow band anterior to

each horn; setose granules in elytral intervals close,

often almost contiguous, granules in third interval in

three irregular rows. Brazil....12. alvarengain. sp. (p. 31)

II. bifurcatus Group

Prescutellar area of pronotum with deep longitudinal

furrow behind horn; declivous sides of furrow posteriorly

more or less dentiform; pygidium broadly rounded

apically - female (only species with female having

median anterior clypeal horn). Brazil, E. Paraguay

and N.E. Argentina. ... 20. tuberifer Felsche (in part) (p. 39)

Prescutellar area of pronotum lacking deep longitudinal

furrow behind horn, depression if present not posteriorly

dentiform; apex of pygidium narrowly rounded, truncate

or otherwise modified - males (females not included in

subsequent key — often identifiable only by association

WIG WAGE) eo sa es ew eels Se ee ee 2

Pronotum with slightly to distinctly overhanging latero-

discal expansions on either side of central horn or

tubercle(s); anterior margin medially elevated, with

feeble median denticle. . ...+.seecsese+- Be as 3

Pronotum without overhanging latero-discal expansions;

anterior margin regular, sometimes medially almost

angular and obsoletely unidentate. ......-seeccesee 4

Pronotum with central horn slender, cylindrical,

anteriorly inclined. (probably Brazil) .......se+e+4-.

SS Gis oS ca OR ee es 15, armatus Westwood (p. 34)

Pronotum with horn (tubercles) bifurcate, base only

slightly elevated. Trinidad..... 16. biarmatus n. sp. (p..35)

Pronotum on each side with an oval or circular, slight to

distinct depression (not lateral fovea) anteriorly or

antero-laterally, situated adjacent to or in front of

horns; midline anterior to horn or horns not longitudinally

Teg CSOT NRT RE ES ES eee ee er ee ee 5

Pronotum without anterior depression either side of midline,

at most with shallow medial depression or fine longitud-

inal furrow anteriorly; pronotal horn single, medial;

simple, bidentate or tridentate distally............ 10

Disc of pronotum with two horns or triangular corniform

lamellae CH Set Soot Yaa e ® e ® e e e @ OO OB OA Be @ 8 e e ® e e@ e e e @ 6

Disc of pronoun: with three horns: ic6 26s es Sowa 2 ew 9

Clypeal armature consisting of three small but distinct

HOMMsS OF PLOMMIENE FUDCECIES . 56 oa. 6 ce is Gh as tie ce oe 7

Clypeal armature consisting of an elongate medial

anterior horn, laterally clypeus with or without small

GCONMCICS 3.0 6k. este. re ar ee ees eer ae eee 3 ee 8

7(6).

8(6).

9(5).

10(4).

11(10).

Howden and Martinez: The Genus Athyreus

Pronotal armature consisting of a low, central bifurcate

horn, each bifurcation broadly triangular; pronotum

anterior and lateral to horn with much of surface

ae not granulate or setose. Brazil. .........-.

pee Bie, Logan Agar ad Seas aaa ay as gigas Westwood (p.

Prout armature ‘consisting of a moderately developed

central bifurcate horn, each bifurcation tapered to

acute tip; pronotum anterior and lateral to horn with

surface granular or punctate, setose. Brazil. .......

a ey r e il kia GPL el OU TES Darvas. Sp. (p-

Clypeal horn inclined anteriorly, posterior face with a

very distinct medial carina, latero-posterior carina

denticulate in front of canthus; pronotum with lateral

concavity large, elongate-oval, shallow, almost

reaching posterior edge, horns contiguous or approx-

imate at base and obliquely directed upward and

forward. Brazil, E. Makpiticg 3 and N. E. Argentina.

Sarat, Sas ging Ue talely uid eee _bifurcatus Macleay (p.

posterior face and latero-posterior carina not den-

ticulate near ocular canthus; pronotum on each side

with deep circular fovea situated in front of horn and

almost on anterior edge; horns widely separated at

base, somewhat compressed and vertical. Peru. .....

o Soha eee igs CER EER es 26. tribuliformis Felsche (p.

Pronotum with anterior impressions and horns well sepa-

rated from each other, each impression large and with

lateral edge outside ocular canthus; prescutellar horn

conical; fore tibia with six teeth on outer margin.

Bolivia, Brazil, :Peru. uw eo 27. zischkai Martinez (p.

Pronotum with anterior impressions well marked and nar-

rowly separated from each other; lateral edge of im-

pression inside lateral edge of eye canthus; two anterior

horns narrowly separated, third prescutellar horn com-

pressed; fore tibia with five teeth on outer margin.

Peru, Cuadere ag ule a gt 28. martinezi Howden (p.

Middle of pronotal disc with upright longitudinal, laterally

compressed horn or keel; in lateral view edge of horn

dentate on angles, depressed between. Brazil. ......

gitiapt tarts" iran Saran ie Ye rir cruel ke COM ene FUBNAS Ns SPs (p.

Disc of pronotum with central, nearly cylindrical horn;

never with a central longitudinal, laterally compressed

median Hoth or ESL iy ie ee ee a ee

Disc of pronotum with horn simple, acute apically. ......

Dise of pronotum with horn bi- or tri-dentate at its

ADEN ard tla. Mea ue Lee tae soa 4 Oe eRe ery ego ee

36)

37)

38)

46)

47)

48)

41)

2h):

13(11).

14(13).

15(13).

16(15).

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Head with clypeal horn, latero-posterior carina on

each side subdentate; pronotum with horn conical,

vertical; pronotal fossae of anterior edge behind

head, small, searcely evident. Brazil. .....°. 2.4%

Meee nae Se ea oe ay oh ce 25. acuticornis n. sp. (p. 44)

Head unarmed, without horn, convex anteriorly;

pronotum with horn long, directed forward and

somewhat ventrally arcuate; pronotal surface in

anterior half medially concave, pronotal fossae

of anterior margin behind head well marked.

COlOMNOIS. | 96 we 6 5s oe sw ss. bs, UCOrNIS n. Ssp..(p. 33)

Pronotal armature apically tridentate ...... 6. 6s 8% 14

Pronotal armature apically bifurcate ..... pe eal Les ‘ 15

Pronotum opaque, except anterior edge of horn which

is smooth, rest completely punctate to granulate;

color piceous or black. Costa Rica, Panama to

Western HCundOr. 2. 2.6 is 2 ae ce 24. championi Bates (p. 43)

Pronotum with medial or antero-lateral area or areas

impunctate and shiny, sides punctate and granulate;

color yellowish to reddish castaneous. S. E. Brazil,

Paraguay, Es Bolivia and N. Ea Argentina... 16s otis

s eis wes Rao CR es os «se -. « 20. tridens Laporte (p. 42)

Clypeal horn lacking distinct posterior median carina,

horn noticeably wider than long in cross section,

apex broadly truncate; pronotal horn vertical, slender,

apex narrowly divided into two longitudinally plate-

like projections, apex of each in lateral view broadly,

bluntly rounded. Peru. ..... 13. pyriformis n. sp. (p. 32)

Clypeal horn with carina or tubercle on posterior median

edge, horn not noticeably wider than long in cross

section, apex rounded or acute; bifurcation of pronotal

horn consisting of divergent, somewhat cylindrical,

ACULEIY POIMLEC DFOCESSES . te ni she eos ole es el et fe 8s 16

Pronotal horn slightly transversely compressed and

vertically bifurcate at apex; each side of bifurcation

nearly cylindrical, tip acute; horn anteriorly with

fine longitudinal median furrow and with punctiform

median depression posterior to anterior margin;

clypeal horn inclined anteriorly and with posterior

median carina. BrazilandN.. 22. bellator Westwood (p. 42)

Pronotal horn consisting of a large protuberance, the

apex of which slants anteriorly in two sharp, divergent

dentiform processes, surface anterior to horn feebly

transversely concave, smooth and shiny, with median

longitudinal furrow obsolete and without distinct

median impression; head with the clypeal horn strong-

ly inclined anteriorly, posterior edge of horn basally

with slightly compressed tooth. Brazil, E. Paraguay

and N. E. Argentina. .. 20. tuberifer Felsche (in part) (p. 39)

Howden and Martinez: The Genus Athyreus 11

III. chalybeatus Group

is Males, pronotum with horn projecting anteriorly, on each

side or below horn with a very conspicuous fovea; pro-

notal disc never with transverse carina at anterior

third or fourth; clypeus with anterior median horn. ..... 2

Females, pronotum without horn or antero-lateral fovea;

pronotal disc with transverse carina at anterior third

or fourth; clypeus without distinct anterior horn. ...... 3

2(1). Pronotal horn situated above wide fovea or medial con-

cavity, central depression often continuous laterally

with deep lateral, oval fovea, carina on each side above

(posterior to) concavities conspicuous, continuous and

not interrupted; clypeus anteriorly with small, slightly

recurved horn; posteriorly with transverse carina

strongly tridentate; color black or reddish testaceous

with elytra black and light metallic bluish green. S. E.

Brazil,. Nw: E «Argentina and S.-E. Paracuaye ee cas

Ae eee . -.... 29. hastifer Felsche (in part) (p. 49)

Pronotal horn dividing anterior depression, depression

forming at each side a very deep oval fovea; adjacent

carina irregularly marked and laterally obsolete;

clypeus anteriorly with small horn with base poster-

iorly denticulate; transverse carina lacking or

obsolete, a tubercle present on each side near gena;

color black with metallic bluish-green reflections.

Argentina, Uruguay... 30. chalybeatus Fairmaire (in part) (p. 50)

3(1). Pronotum medially deeply impressed, cordiform, de-

pression delimited on all sides by carinae, anterior

transverse carina slightly dentate on either side of

midline; midline of impressed area with two distinct

foveae, one anterior, one posterior, united by a fine

longitudinal line; color almost uniformly testaceous.

(Male unknown). Argentina. ... 31. conspicuusn. sp. (p. 51)

Pronotum medially shallowly to moderately impressed;

anterior transverse carina not tuberculate either

side of midline; elytra black with bluish or steely

POTIOCHIONGs 2666 een Ha Ne a re ce ee eC. 4

4(3). Anterior median clypeal tubercle distinct, united to

posterior median tubercle of transverse carina by a

fine, longitudinal carina; pronotum on either end of

anterior median transverse carina with short carina

at right angle to transverse carina extending anter-

iorly about half the distance to anterior margin; fore

tibia usually with six distinct teeth on outer margin;

color and distribution as described for male.

Argentina, Uruguay. . 30. chalybeatus Fairmaire (in part) (p. 50)

Anterior median clypeal tubercle obtuse, not carinate

posteriorly; pronotum on each side of anterior median

transverse carina lacking distinct anterior carinate

extensions; fore tibia usually with five distinct teeth

1).

3(2).

4(3).

5(3).

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

on outer margin; color and distribution as described

for male. S. E. Brazil, N. E. Argentina and 8S. E.

Paraguay. “nisi Se, 29. hastifer Felsche (in part) (p. 49)

Clave para Athyreus Macleay

Pronoto en el macho sin surco o depresion longitudinal

medial completa; en la hembra con dos pequenos

tuberculitos discales paramedios, o con quilla o

tubérculo medio anterior, limitando al disco por

delante< 4-4 (a3. Pe RS a, 2

Pronoto en el fagehe Vv hembra con surco o depresién

longitudinal medial mas o menos ensanchada marcada

y completa; en la hembra sin tuberculitos, quilla o

tubereulo: diseales.c 1.06 Wea ee I. Grupo aeneus (p. 12)

Pronoto del macho nunca con quilla lateral que desde el

borde de la depresién llegue al borde lateral; en la

hembra con dos pequefios tuberculitos discales para-

medios o, tuberifer, con tubérculo medio anterior. ....

ee a - .. II. Grupo bifurcatus (p. 15)

Pronoto del miechip y hembra con quilla lateral que desde

el borde de la depresion discal llega al borde lateral;

hembra con quilla medial anterior que limita la region

discal. 2... kt ge toe ee. III. Grupo chalybeatus (p. 18)

I. Grupo aeneus

Borde posterior del pronoto, préximo al himero, in-

terrumpido: Brasil. o.:. 40 .>. -.. 1. cyanescens pe (p- an

Borde posterior del pronoto entero. .....s+e-+s-eees

. Margen anterior del pronoto en la mitad mas o menos

ansuladoy WRitemtadOs ie Gos ee 3

Margen anterior del pronoto en la mitad aquillado o no,

PCKO MINCA UNIGENtAdO o25:.0. 2 ee et Oe Oe EE 7

Denticulo del borde anterior del pronoto bien marcado. ... 4

Denticulo del borde anterior del pronoto poco marcado,

a.veeis Solo como antulosidad << .6:.0sete ar ea MO eS 5

Pronoto con quilla transversal anterior que llega al

borde lateral; color verde azulado oscuro con visos

mas o menos bronceados. Brasil. .... 3. aeneus Klug (p. 22)

Pronoto con quilla lateral obsoleta, corta, situadaen

la mitad posterior; color testaceo castafio,

generalmente con los lados del pronoto y bordes

laterales de los élitros aclarados. Brasil. .......-.

oie eee ee Ue See GP ae ee avis Boueomont (p. 23)

Pronoto del macho con quilla transversal anterior que

llega al borde lateral; depresién anterior anchamente

cordiforme; surco longitudinal medial poco marcado,

sin fosa anterior notable; quilla discal costiforme,

6(5).

1D).

Bt7).

9(8).

Howden and Martinez: The Genus Athyreus 13

realzada y no dentada; quilla l4tero-medial ancha y

brillante; quilla posterior fina y poco conspicua;

cuerno clipeal erecto, pequenho, la base con fina

quilla conectada al borde lateral, que es elevado en

diente alargado y comprimido, la cara posterior

finamente aquillada. Color negro algo brillante con

leves matices acerados. Hembra en el pronoto con

quilla transversal anterior que llega al borde lateral.

Brasil. ss eisay e's 6 8) es es Be Capricormie Nn. sp. (p. 21)

Pronoto del macho sin quilla transversal anterior; la

depresién medial circularforme u oval, con surco

longitudinal impreso y fosa anterior marcada; quilla

discal siempre dentada; sin quilla a4ntero-medial o

ésta angosta; falta la quilla péstero-lateral; cuerno

clipeal sin quilla en la cara posterior. Hembra en

el pronoto sin quilla transversal anterior. .....-..ee-. 6

Macho con la quilla que margina la depresion discal

mas o menos elevada, laminiforme, alargada y con

los bordes anterior y posterior dentados; regidn pre-

escutelar deprimida y con impresion puntiforme |

marcada. Hembra desconocida. Brasil, Paraguay. :

sw ws Oe hem eee ee weet ae a re se 10. anneae n. sp. (p. 30)

Macho con la carena que margina la depresion discal

dentiforme, poco elevada, aproximada a la opuesta

y unidentada; regioén preescutelar con SurCO medio

bien impreso, de borde elevado, por detras denticulado

sobre la region escutelar y sin impresidn puntiforme.

Hembra con la quilla que margina la depresién discal

del pronoto poco marcada, subdentada, posteriormente

rasa y no formando denticulo por detras. Brasil, E.

Paraguay y N. Argentina. . 8. hemisphaericus Boucomont (p. 28)

Borde anterior del pronoto elevado medialmente en

quilla transversal a cada lado subdentada o ligeramente

OURCOC ake gk Gb OES Se eo ee es 8

Borde anterior del pronoto regular, no elevado en

qualia ni denticulado.: <a 816) eee oc ees Gi eR 4G 11

Macho en el pronoto con el borde lateral de la de-

presién medial elevado en alto proceso laminiforme

bidentado; quilla del borde anterior estrecha, su

ancho menor que el ancho de la depresién. Hembra :

desconocida. Bolivian sayy. 40sec. 9. nitidus n. sp. (p. 29)

Macho en el pronoto con el borde lateral de la depresion

poco elevado, su 4pice romo y nunca dentado; quilla

del borde anterior subigual - Oo apenas mas angosta

que la depresién en la region preescutelar. ...... or 9

Macho con el cuerno clipeal robusto, la cara posterior

excavada o plana, ancha y limitada lateralmente por

quilla bien marcada, arqueado-dentada en la base y

que alcanza al borde lateral; pronoto con la quilla

que bordea la depresién medial poco elevada, a veces

10(9).

1147).

Contrib. Amer. Ent. Inst., vol.

15; no. 4;° 1978

un poco levantada por detras, pero ancha y no

angulosa, terminando por delante y a cada lado de

la depresion preescutelar, ésta amplia, impresa y

subcuadrada, o mds ancha que larga. Hembra:

pronoto con el borde anterior medialmente elevado

en arista baja, fina y marcada, con su ancho igual

al ancho de la — preescutelar.

Brasil, Paraguay.

- 7. bilobus n. sp. (p. 26)

Macho con el cuerno clipeal ‘erdcil, la cara posterior —

angosta, limitada a cada lado por fin

a quilla que

puede o no perderse hacia la base; pronoto con la

quilla que bordea la depresion en 16b

ulo de Apice

convergente y romo; depresion preescutelar no

ocupando toda la regidn. ......

Macho con la impresi6én preescutelar m

he ee yee ee 10

uy pequefa,

normalmente alargada y situada sobre el Angulo

medial del borde posterior; depresi6:

zona longitudinal de escultura granul

n central sin

osa que la

divida en dos partes; quilla medial del borde

anterior poco mAs ancha que la mitad del ancho

de la region preescutelar, alta, con

la arista

ligeramente céncava y denticulada lateralmente;

cuerno clipeal con la quilla péstero-lateral poco

evidente, perdiéndose hacia la base.

Argentina, Paraguay. ee

N. E.

-5. hypocritus n. sp. (p. 24)

Macho con ancha impresién preescutelar en forma

de surco longitudinal medial; depresién central

con ancha zona longitudinal de escult

ura granulosa,

que conecta las regiones anterior y posterior;

quilla medial del borde anterior tan ancha como

el ancho de la regién preescutelar, baja, de arista

sinuosa y no denticulada lateralmente; cuerno

clipeal con la quilla péstero-lateral fina, entera,

llegando al borde lateral de la cabeza. Brasil.

Pronoto con la depresién anterior, por

6. brasilicus n. sp. (p. 25)

delante de la

carena, con area semicircular lisa y englobando

el area granulosa antero-medial; granulos setiferos

de los interespacios elitrales claramente separados,

ser. interespacio con dos hileras de

Bolivia.acn eo eee ee LE

eranulos.

forcipatus Boucomont (p. 31)

Pronoto con la depresién anterior casi enteramente

granulosa y con banda lisa corta y angosta por

delante del cuerno; granulos setifero

s de los

interespacios elitrales frecuentemente contiguos

y en el 3er. interespacio ordenados en 3 hileras

inreculoves.: Brasil, i... wks 12. alvarengai n. sp. (p. 31)

Howden and Martinez: The Genus Athyreus

II. Grupo bifurcatus

1, Hembra: pronoto en la regién preescutelar con surco

longitudinal mas o menos impreso y dentiforme por

detras; en el disco con tubérculo pequefio y los

cuernos dentiformes poco desarrollados, en la parte

dorsal y por detras con un denticulito; mitad anterior

con profunda depresion transversal y la superficie

centralmente granulosa; cuerno clipeal corto, algo

oblicuo y en la cara posterior y basalmente inerme.

Brasil, E. Paraguay y N. BH. Argentinas). BO. 6 er.

oa en EE PE el eats 20. tuberifer Felsche (parte) (p.

Machos nunca con la regidn preescutelar surcada longi-

tudinalmente €¢4herme sn. sce) Pe ea

2(1). Macho en el pronoto con expansiones alares latero-

discales; el borde anterior medialmente elevado en

quilla subdentadaw so Paine ee a Oe

Macho en el pronoto sin expansiones alares lAtero-

discales; el borde anterior regular, a lo sumo

medialmente algo anguloso y obsoletamente ,

denticuiado. 748.8 rain a as pea oe elie

3(2). Pronoto con cuerno delgado y dirigido hacia adelante.

(Braeil?) 6S ee ae aS 15. armatus Westwood (p.

Pronoto con dos laminas triangulares corniformes.

TRU 1 i ol is ie eo 16. biarmatus n. sp. (p.

4(2). Macho en el pronoto, a cada lado, con una depresioén

oval o circulariforme anterior o antero-lateral al

cuerno discal, estando éste mas o menos separado

del opuestoe en 1a base interna... 0.6 ee

Macho en el pronoto sin depresién, a lo sumo con fino

surco medial longitudinal anterior; armadura discal

unica, medial, simple, bidentada o tridentada

Cistatmente. 946 Vee oe eee Ee ae Ng

5(4). Macho en el disco del pronoto con dos cuernos o laminas

LPIEGNOUIATOS COrniItOrniCs 5k ie ee BE oil a REN

Macho en el disco del pronoto con tres procesos corni-

TOV ie a ea TE a Lea eh cr pete a are ea

6(5). Macho con la armadura clipeal consistente en tres

pequefios pero evidentes dientes o cuernos; armadura

pronotal formada por dos laminas triangulares diver-

POULS So ela a gama. a We ete GNIS Cala ee ae ie

Macho con la armadura clipeal consistente en un cuerno

medial anterior mas o menos largo, lateralmente a

este con o sin denticulo; armadura pronotal constituida

por un cuerno mAs o menos cilindroide o algo aplanado,

pero més lareomue anche. sss fe a gets te

39)

35)

7(6).

8(6).

9(5).

10(4).

11(10).

12(11).

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Pronoto con areas Antero-laterales impunteadas Vy

Orillantes., Brasil... ..5 5. 6] 17. gigas Westwood (p. 36)

Pronoto con areas laterales totalmente con puntos

granulosos,. Bragile: « t2caceds 3 eo 18. parvus n. sp. (p. 37)

Macho con la armadura clipeal formada por un

cuerno oblicuamente dirigido hacia arriba y

adelante, que tiene en la cara posterior una

quilla medial bien evidente, la quilla l4tero-

posterior denticulada sobre el canto ocular;

pronoto con la impresién grande, alargado-oval

bien lateral, casi alcanzando al borde posterior,

el cuerno algo aproximado del opuesto en su base

interna y oblicuamente dirigido hacia arriba y

adelante. Brasil, E. Paraguay y N. E. Argentina. :

een wes ech? «obifurcatus Macleay (p. 38)

Macho con la armadura clipeal consistente en un

cuerno m&s o menos vertical y algo arqueado, sin

quilla medial en la cara posterior y la quilla

l4tero-posterior no denticulada sobre el canto

ocular; pronoto con févea circulariforme profunda,

situada por delante del cuerno y casi sobre el

borde anterior, éste ampliamente separado del

‘opuesto en la base, algo comprimido y vertical.

Pevas 6. 26. tribuliformis Felsche (p. 46)

Macho en el pronoto con la impresién antero-lateral

y cuernos bien separados entre si, aquella grande

y con el borde lateral por fuera del canto ocular,

cuerno preescutelar cénico. Bolivia, Brasil. ......

Per. . ce ee sg eieta @ aie: 27. zischkai Martinez (p. 47)

Macho en el pronoto con la impresién antero-medial

muy marcada y conjuntamente con los cuernos

subcontiguos, el borde lateral de aquella por

dentro del borde lateral del canto ocular; cuerno

preescutelar comprimido. Pert. . 28. martinezi Howden (p. 48)

Macho: pronoto en el disco con quilla longitudinal

medial, la arista escotada y dentada sobre los

Anelodi Basile << 2 ak cas a eras 21. juanae n. sp. (p. 41)

Macho: pronoto en el disco nunca con quilla

LONCICUCING! MICdIAl. | woes ce ee a a Al

Macho: pronoto en el disco con cuerno simple. ....... 12

Macho: pronoto en el disco con cuerno bi o

tridentado, en el,apice:. .. .. « Le a ee 13

Macho en la cabeza con cuerno clipeal y con la

quilla la4tero-posterior a cada lado de éste

subdentada; pronoto con el proceso delgado,

coniforme, vertical, la superficie lateral y

anteriormente granulosa, la fosa del borde

anterior, por détras de la cabeza, pequefia,

poco evidente. Brasil. .... 3. 25. acuticornis n. sp. (p. 44)

Howden and Martinez: The Genus Athyreus 17

Macho con la cabeza inerme, sin cuerno, convexa

anteriormente; pronoto con proceso dirigido hacia

adelante, largo y algo arqueado, la superficie en

la mitad anterior muy brillante y con sdélo algunos

puntitos obsoletos lateralmente, fosa del borde

anterior, por detrds del borde lateral de la cabeza,

bien marcada....Colombia.) .:i..5... 14. unicornis n. sp. (p. 33)

13(11). Macho con la armadura pronotal apicalmente

EV EGOICA CA. io sie otis: Ayla eelh craigs oR stk eas ban, eae wy aan ied 6 14

Macho con la armadura pronotal apicalmente

DHPC ACA ass. Se axieid ea S: abecel area eiiet au ee ered kee. LS

14(13). Macho con el pronoto opaco, excepto el borde anterior

del proceso discal que es liso, totalmente punteado

tornandose los puntos granulosos y bien notables

hacia los lados; parte posterior de la frente y

vértice densa y claramente punteada; color piceo a

negro. Costa Rica, Panama y Eeuador: 4... 406 shou .

ee ee ee 24. championi Bates (p. 43)

Macho en el pronoto con 4rea o Areas medial o antero-

laterales impunteadas y brillantes, hacia los lados

punteado o punteado granuloso, los granulos

pequefios; parte posterior de la frente y vértice

superficial y poco claramente punteada; color

castano amarillento a castano rojizo. S. E. Brasil,

Paraguay, i... bolivia y N. BE. Argentina. «20... 66 «'s

es Bige he, Shas «5 6.4 3804s & 6 aos. tridens Laporte (p. 42)

15(13). Cuerno clipeal sin quilla ni diente en la cara posterior,

ensanchado y truncado en el Apice; pronoto con la

regién preescutelar en surco longitudinal subpiriforme

de borde elevado; borde lateral, un poco por delante

del angulo medio, brevemente quebrado; la regién

lisa central no sobrepasando lateralmente el borde

lateral del canto ocular: la region lateral con densa

granulacién pilosa; proceso pronotal regularmente

angostado distalmente y allf estrechamente hendido

longitudinalmente, formando dos dientecitos

comprimidos, la cara posterior claramente surcada

: el surco continuandose con el surco preescutelar;

élitro con la elevacion costiforme I apareciendo por

detras del escudete; el interespacio II en la mitad

basal como el doble de ancho que elIII. Perd. .....

ih Sl sas aha oa ie aan a 13. pyriformis n. sp. (p. 32)

Cuerno clipeal con quilla o diente en la cara posterior,

aguzado hacia el apice; pronoto con la regién pre-

escutelar plana o ligeramente deprimida, nunca en

surco profundo de borde elevado; borde lateral

entero; la region lisa central sobrepasando amplia-

mente el borde lateral del canto ocular; la regién

lateral lisa o solo con algunos granulos pilosos

dispersos; proceso pronotal en el Apice expandido

y bifurcado en horqueta, formando dos claros y

18 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

evidentes dientes y sin ningtin surco en la cara

posterior; elitro con la elevacion costiforme I

completa desde la base; el interespacio II subigual

ON WNCHO te ee a eS Se ee 16

16(15). Macho con la armadura del pronoto consistente en un

cuerno alto, un tanto comprimido Anteroposterior-

mente y bifurcado en el 4pice; por delante de este

la superficie casi vertical, con fino surco longitudinal

medio, que un poco por detras del borde anterior

tiene una fosa pequeha, a los lados de ése lisa o casi

lisa, tornAndose punteada hacia los bordes laterales;

cabeza con el cuerno clipeal algo oblicuo, en la cara

posterior con quilla que lo recorre totalmente y sin

diente basal. Brasil y N. 3... 22. bellator Westwood (p. 42)

Macho con la armadura del pronoto consistente en una

protuberancia grande, cuyo A4pice superior es

proyectado hacia adelante en dos cuernos dentiformes

aguzados y algo divergentes, por delante de éste la

superficie algo céncava transversalmente, lisa y

brillante, con surco longitudinal medial obsoleto y

sin impresidén puntiforme; cabeza con el cuerno

clipeal muy oblicuamente dirigido hacia adelante,

en la cara posterior y sobre la base con diente

comprimido ligeramente aquillado. Brasil, E.

Paraguay 7 Nei. A eO@eniInas ee ae ee ee ak

ee pe ee 20. tuberifer Felsche (parte) (p. 39)

III. Grupo chalybeatus

ee Machos en el pronoto con cuerno proyectado anterior-

mente, al costado o por debajo de éste con fdvea

muy marcada; disco nunca con quilla transversal

medial anterior; clipeo con cuerno mAs o menos

desarrollado. ..... a er a Feb eee pale. ; 2

Hembras en el pronoto sin cuerno ni féveas antero-

laterales; con quilla medial anterior que limita

el iieeo- clipes an cucino. 2. Se os 3

2(1). Cuerno pronotal situado por encima de Ja févea o

depresion medial, ésta muy ancha y corta,

continuAndose lJateralmente con la profunda fovea

lateral ovaliforme, la quilla anterior que la

margina superiormente, marcada, continua y no

interrumpida; clfipeo anteriormente con pequeno

diente corniforme y por detrds de éste con quilla

transversal ligeramente angulosa cefalad y

fuertemente tridentada; color negro o testaceo

rojizo con los élitros negros y visos verde azulados

met&licos leves. S. E. Brasil, N. E. Argentina y

Db. 2. Parasay, oe oe ee 29. hastifer Felsche (parte) (p. 49)

Cuerno pronotal interrumpiendo _ la depresion anterior

y formando a cada lado una f6vea ovaliforme muy

profunda, que no sobrepasa lateralmente a los cantos

oculares, la quilla anterior irregularmente marcada

Howden and Martinez: The Genus Athyreus 9

y lateralmente algo obsoleta; clipeo anteriormente

con pequeno diente corniforme, en cuya base

posterior hay un denticulo, sin quilla transversal

y sobre los cantos oculares con otro denticulo

triangulariforme algo comprimido. Color negro

con leves reflejos verdoso azulados metalicos.

Argentina, Uruguay. .. 30. chalybeatus Fairmaire (parte) (p. 50)

3(1). Pronoto con la region discal cordiforme, muy impresa,

marginada por quilla divergente y algo arqueada, que

en la region anterior se acoda fuertemente, para

confluir con la opuesta en una pequefiisima angulosidad

medial, que puede ser ligeramente biimpresa; a cado

lado yen la mitad denticulada y convergiendo por

detras con la quilla transversal posterior, que es

larga, muy marcada y finaliza sobre la rasa ps poco

visible févea lateral; la superficie de la regién discal

con dos fosas marcadas, unidas por fino surco

longitudinal medial; a cada lado con 4 quillas bien

evidentes, siendo la anterior arqueada; cabeza en la

frente con dos quillitas paramedias algo arqueadas

y notables, el vértice claramente deprimido. Color

general testaceo. Macho desconocido. Argentina. ...

Pah aia, eT ; o 6 + +e « ». Ol» CONSPICUUS ND. (p- 51)

Pronoto con la renige discal rombiforme alargada,

marginada por quilla divergente y recta, no

denticulada en el medio y que se pierde a cada lado

de la region preescutelar; carena transversal antero-

medial inerme; Ja superficie rasa y sin fosas

marcadas, pero con surco medial longitudinal

impreso; a cada lado con 3 o 4 quillas, en éste

tltimo caso rectas y cortas; la quilla posterior

nunca unida a la quilla longitudinal que limita al

disco; cabeza en la frente con dos tuberculitos

pequefos, obsoletos, a veces un tanto aquillados,

el VOICe DIABO.. ol ss ear a ae gy aaa ae ad cian a : 4

4(3). Diente clipeal marcado, unido posteriormente al diente

medio de la quilla transversal por fina, pero neta

quillita. Pronoto con la quilla transversal medial

recta; la dntero-lateral corta, recta; la medial an-

terior obsoleta sobre el Angulo 4ntero-superior de

la fovea lateral; la quilla posterior marcada desde

el Angulo posterior hasta por detras de la misma

fovea; pilosidad del élitro corta. Color y distribucién

geografica similar a la del macho. Argentina,

PUG ac) ae epee ee chalybeatus Fairmaire (parte) (p. 50)

Diente clipeal marcadu como una pequena angulosidad,

no unido por detrAs al diente medio de la quilla

transversal. Pronoto con la quilla transversal

medial un tanto arqueada; la 4ntero-lateral falta;

la medial anterior bien marcada en toda su longitud;

la quilla posterior fina, irregular; pilosidad de la

superficie del élitro larga. Color y distribucién

20 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

geografica similar a la del macho. S. E. Brasil,

N. E. Argentina yo. E. Paraguay. 2. 6 Se ee ees ae

BA ae eG es cier. 20e “etuicr Pelsche (parte) (p. 49)

1. Athyreus cyanescens Klug

Figures 1-3, Map 1

Athyreus cyanescens Klug, 1845, p. 30.

TYPE. Male (Berlin ?), not examined.

MALES. Length16.0to19.0mm, greatest width 9.5t010.0 mm.

Dorsally black with bluish tinge. Labrum on anterior median surface with a

distinct, anteriorly arcuate, upright tubercle; a character not noted in any

other Athyreus. Clypeal horn (Fig. 1) in well developed males approximately

as long as length of frons and vertex combined; horn in apical half in cross

section wider than long, apex rounded; anterior lateral carina present basal-

ly; posterior lateral carina on each side distinct for most of length of horn

and curving posteriorly on frons to feebly elevated junction at gena; posterior

- median carina absent, posterior surface of horn slightly concave. Frons

medially moderately, longitudinally convex; surface with scattered punctures;

vertex nearly flat, moderately setose-granulate. Pronotum (Fig. 1) with

anterior margin between large fossae upwardly and anteriorly angulate,

dentate at median angle, a faint raised ridge extending posteriorly from

denticle for about 1mm. Pronotal midline feebly impressed, prescutellar

depression lacking; in posterior third on each side raised carina or horn with

acute anterior and posterior lobes (see fig. 1), posterior lobes much closer

to each other than anterior lobes; on each side anterior to and parallel with

pronotal margin, a short carina present anterior to humerus; pronotal margin

broken, deeply excised in this area; pronotal surface except for carinae

setose-punctate (median third) or granulate. Elytron with margin dentate at

anterior-lateral angle. Fore tibia with five teeth on outer margin. Genitalia

as in fig. 2.

FEMALES. Length15.0to18.0mm, greatest width 9.0 to10.0 mm.

Labrum with unusual anterior median tubercle as in male. Clypeal carinae

varying from most species in that, in some specimens, the anterior and

posterior carinae medially with a longitudinal carina between, in most cases

joining both carinae but always contiguous with median tubercle of posterior

carina. Pronotum (Fig. 3) with low arcuate carina on either side of indented

midline, anterior margin not as elevated as in male but with larger median

denticle (tubercle). Pronotal margin near humerus and adjacent short carina

as described in male.

MATERIAL EXAMINED: 10 males, 10 females.

BRAZIL (type locality, no other data). Parana, Curitiba, X1.1940, F.

S. Pereira; X.1940, Lange. Santa Catarina, Pinhal, XII.1951, A. Maller;

Cauna, X11.1945, A. Maller. Sdo Paulo, Jundiaf, XII.1956; Campos do

Jorda0, Lane; XII1.1945, Pereira. Brasilia, Laferte, ex Musaeo Sharp.

Guanabara, Rio de Janeiro.

Specimens are in: London, Ottawa, Paris, Howden, Martinez.

REMARKS. The black color with a distinct bluish caste, the odd

tubercle on the anterior dorsal midline of the labrum, and the interrupted

pronotal margin adjacent to the elytral humerus are all characters that

readily distinguish both males and females of cyanescens from other

Athyreus.

Howden and Martinez: The Genus Athyreus mask

2. Athyreus capricornis n. sp.

Figures 4-6, Map 1

DIAGNOSIS. Black with steely hue; males with clypeal horn erect,

anterior lateral carina interrupted near base, posterior lateral carina

present apically and laterally, posterior median carina pronounced, extend- —

ing to middle of frons; frons and vertex of both sexes closely, irregularly

granulate or punctate; pronotum medially posterior to horns with wide cordi-

form depression in males; median pronotal area concave anteriorly, horn of

males on each side low, irregularly rounded or sinuate, with anterior lateral

branch extending to lateral pronotal margin; females with same carina not

greatly elevated; two other short carinae present in both sexes on each side,

one wide, one narrow, in posterior lateral half of pronotum paralleling pos-

terior margin.

HOLOTYPE. Male, length15.0 mm, greatest width 9.5mm. Overall

black with steely luster, feebly shining. Labrum short, wide (8:1), anterior

median edge with face strongly impressed, concave, impunctate; dorsal sur-

face of labrum confluently coarsely punctate, coriaceous. Clypeus (Fig. 4)

anteriorly truncate, the truncate area bordered by a fine carina which is bow-

shaped and with apex of upward angle on anterior base of clypeal horn; horn

vertical, anterior lateral carina present on each side but not joining bow-

shaped carina; posterior lateral carina pronounced on horn, becoming fine

basally, laterally carina lobed; posterior median carina pronounced, extend-

ing down horn onto middle of frons, there becoming obsolete. Frons and

vertex closely, often confluently punctate, posteriorly granulate. Pronotum

(Fig. 4) with anterior margin obtusely, feebly tuberculate medially, lateral

margins arcuate; posteriorly median depression posterior to horns deeply

cordiform, internally smooth with scattered punctures; depression elevated

between low rounded horns posterior to pronotal center, midline indicated as

impressed line in broader depression; area anterior to horns with scattered

punctures, then near anterior margin with large discrete granules; lateral

thirds of pronotum excluding carinae closely granulate, setose. Pronotal

horn on each side of midline low, cariniform, edge in lateral view rounded

anteriorly, downwardly angulately truncate posteriorly; horn continuing

anteriorly as carina, dividing at base of horn, pronounced lateral branch

extending as sinuous carina of varying width to lateral margin; posterior to

this carina and midway to posterior margin a short wide carina and more

posteriorly a feebly developed carina. Elytra with six raised, polished inter-

vals between suture and umbone, striae relatively narrow for genus; finely,

closely granulate. Fore tibia with six teeth on outer margin. Genitalia (Fig.

5) with parameres apically thickened and lobed, the apical protrusions fili-

form and medially strongly arcuate.

ALLOTYPE. Female, length15.0 mm, greatest width 9.0 mm.

Clypeus lacking horn, with low anterior transverse carina and trituberculate

posterior transverse carina (as is typical for most Athyreus females).

Clypeus, frons and vertex all heavily granulate or confluently punctate.

Pronotum (Fig. 6) not differing greatly from male; central longitudinal carina

on either side of midline not elevated, very wide, posterior end narrowing

and curving inward to midline, joining carina from opposite side just anterior

to posterior margin; in posterior half midline impressed and smooth, granu-

late on either side, most deeply impressed posteriorly; pronotal surface

eranulate except for carinae and midline posteriorly.

TYPE MATERIAL: Holotype, male, Brazil, Estado de SAo Paulo,

22 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Batatais, V.1945, F.S. Pereira (Martinez).

Allotype, female, same data as holotype (Martinez).

REMARKS. This species, which has been confused with cyanescens,

can be easily distinguished by its uninterrupted lateral pronotal margin,

which is broken before the humerus in cyanescens. Also cyanescens has a

denticulate labrum. The form of the labrum and anterior lateral carinae will

distinguish capricornis from other related species, the closest of which is

probably hemisphaericus.

3. Athyreus aeneus Klug

Figures 7-9, Map 1

Athyreus aeneus Klug, 1845, p. 31.

TYPE. Male (Berlin ?), not examined.

MALES. Length16.0to17.0mm, greatest width11.0to12.0 mm.

Dorsally black with distinct greenish reflections, particularly central third

of pronotum and elytra. Clypeus (Fig. 7) with stout vertical horn, apex

truncate, horn in cross section wider than long; anterior lateral clypeal

carina evident only basally; posterior lateral carina well developed for length

of horn, basally fine to lateral angulate junction at gena; posterior medial

carina absent; posterior surface of horn in apical half shallowly concave.

Frons medially distinctly, longitudinally convex; surface of frons and vertex

with scattered punctures or granules. Pronotum (Fig. 7) with anterior

margin between large fossae strongly elevated, acutely angulate at midline;

median angle with faint ridge extending posteriorly almost to small deep

fovea at midline, fovea of same diameter as marginal fossae. Midline de-

pressed behind anterior fovea; prescutellar depression posteriorly wider,

smooth, extending to pronotal margin. On each side of midline at posterior

third an abruptly elevated plate-like horn with truncated apex sloping anter-

iorly; horns narrowly separated basally. Pronotum on each side lateral to

horn with three irregular, obliquely transverse carinae; anterior carina

joining lateral margin midway between anterior and median angles, at junc-

tion forming small tubercle; median of three carinae short, posterior carina

nearly twice as long; all carinae distinct; surface between carinae and in

lateral thirds of pronotum with unusually large, coarse granules; granular

area setose but setae inconspicuous. Elytra with surface unusually polished,

shining, striae feebly depressed, indicated by one or two irregular rows of

punctures. Fore tibia with six teeth on outer margin. Genitalia as in fig. 8.

FEMALES. Length17.0to18.0mm, greatest width 11.0to12.0 mm.

Head with anterior and posterior clypeal carina, the carinae somewhat

atypical; anterior carina arcuate either side of dorsal median denticle, a

longitudinal carina extending from denticle posteriorly to median tubercle of

posterior, tridentate carina; median tubercle of posterior carina approximate-

ly twice size of lateral ones. Dorsal surface of clypeus rugose; frons and

vertex granulate. Pronotum (Fig. 9) similar to male but with inner carina

not elevated, feebly arcuate either side of deeply indented midline; depression

not greatly widened in prescutellar region.

MATERIAL EXAMINED: 2 males, 4 females.

BRAZIL (type locality, no other data): Parana, Curitiba, II .1935,

X1I.1936, X.1940, and 1.1946.

Specimens are in: Howden, Martinez.

REMARKS. Athyreus aeneus is very distinctive; the black color with a

definite greenish luster, relatively shining elytron and the three obliquely

Howden and Martinez: The Genus Athyreus 23

transverse carinae on each side in the posterior half of the pronotum of both

sexes should distinguish it from any other species in the genus.

<S oF eee ro ee pe

\ carn \ Ke te A

, ) a > A @

\ ‘ ‘ ‘ A ey

oat Y SEES LBS ONAL Uist terest

( & A + x ne rE A

3 ey vv

/ r ‘ A ii fe

‘ 1 o

| ewe 5 ‘ ‘ ‘

; js y, f : We y

A =1. cyanescens Klug @ = 4. vavini Bouc.

@ =2. capricornis n. sp. * = 5. hypocritus n. sp.

@=3. aeneus Klug @=6. brasilicus n. sp.

A=7. bilobus n. sp.

4. Athyreus vavini Boucomont

Figures 10-12, Map 2

Athyreus vavini Boucomont, 1902a, p. 184; 1902b, Dp. B80.

TYPE. Male (Paris), examined. 2 |

MALES. Length14.0to17.0 mm, greatest width 10.0 to12.0 mm.

Dorsally reddish brown, often tan laterally, some specimens dark brown in

center of pronotum and on elytra. Clypeal horn (Fig. 10) erect, stout; verti-

cal portion with four distinct carinae, two anterior lateral and two posterior

lateral carinae, near base posterior lateral carinae divided in some well

developed males, the inner carinae extending onto frons, there becoming

obsolete; usual posterior lateral carina on each side arcuately extending to

slightly raised, rounded terminus above antennal insertion. Frons mostly

smooth, vertex mostly granulate. Pronotum (Fig. 10) with anterior margin

between fossae thickened, abruptly raised into acute tubercle on posterior

edge at midline; lateral pronotal margins arcuate. Pronotum in median third

24 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

concave, concavity deepest along midline, anteriorly near margin coarsely

granulate, mostly smooth in remainder of concavity to base of low, obtusely

rounded horn on each side at posterior fourth; a few widely separated gran-

ules near base of each horn. Median depression posterior to horns variable;

in well developed males depression rapidly deepens and expands to approx-

imately 1 mm anterior to posterior margin, base of horn on each side extend-

ing posteriorly and overhanging depression forming posterior denticles; in

male minors only a broad shallow posterior depression present, in larger

specimens sides become ridged, as in type; however even in male minors

low horns obtusely angulate. Fore tibia with five teeth on outer margin.

Genitalia as in fig. 11.

FEMALES. Length15.0to17.0 mm, greatest width 11.0to12.0 mm.

Clypeus typical of Athyreus females, anterior carina upwardly arcuate, pos-

terior carina trituberculate; posterior carina when viewed from above feebly

anteriorly arcuate, the median tubercle slightly anterior to lateral ones.

Pronotum (Fig. 12) with anterior margin with median tubercle as in male;

pronotal midline impressed, area on either side sloping upward, in posterior

area depression limited by longitudinal carina on each side, carina slightly

divergent anteriorly and becoming obsolete just anterior to middle; a short

lateral carina present on either side anterior to humerus. Pronotal surface

setose, closely granulate, granules sometimes reduced or absent medially in

depression just anterior to middle. The granules and degree of elevation of

carina are useful comparative characters. —

MATERIAL EXAMINED. 10 males, 5 females.

| BRAZIL: (type locality, no other data): Rio Grande do Norte, Natal,

TI, £950, TIT /1V 195d, 11, VEY Vit. £958, VILI/&. 1954, 1 VV. 1957,

IV.1961, I1.1962. Paraiba, Joazeirinho, Soledade, I1V.1957. Ceara,

Paraguru, III1.1953. Bahia, Vitoria da Conquista, IV.1961. Minas Gerais,

Paraizo, I.1962; Collectors: Alvarenga, Machado, Pereira, Sebra.

Specimens are in: Paris, Howden, Martinez.

REMARKS. Boucomont's unique type is a relatively minor male with a

small clypeal horn and only a moderate prescutellar impression on the pro-

notum, the impression lacking the dentiform posterior projection on each

side.

The relatively small size, color, median tubercle on the anterior

margin of the pronotum and the range will separate vavini from related

species.

5. Athyreus hypocritus n. sp.

Figures 13-15, Map 2

DIAGNOSIS. Reddish brown to brown; similar to vavini. Males with

pronotal concavity broad, as wide as or wider than head; pronotal horns pos-

teriorly acutely rounded; anterior pronotal margin between fossae distinctly

elevated, tuberculate on either side of raised margin; pronotal midline pos-

teriorly with small nearly circular depression, depression contiguous to

margin, marginal line obsolete medially; pronotal concavity with most or

part of median area not granulate. Females with posterior clypeal carina

feebly arcuate anteriorly, tridentate; anterior pronotal margin as in male

but less distinctly dentate; longitudinal pronotal carina extending anteriorly

to anterior third, slightly more widely separated than in vavini.

HOLOTYPE. Male, length15.0 mm, greatest width 9.5mm. Dorsal-

ly reddish castaneous, labrum, clypeus, sides of pronotum and elytra, and

Howden and Martinez: The Genus Athyreus 25

legs lighter, yellowish tan to light brown. Most closely related to bilobus,

differing in having more slender clypeal horn with rounded apex and lacking

complete lateral posterior carina; front and vertex with small granulate

punctures extending sparsely to near base of horn. Pronotum (Fig. 13) with

anterior margin between fossae distinctly elevated, the elevation on either

side with a small tubercle, slightly lower at middle than laterally; pronotum

posteriorly with small, deep, cordiform (nearly circular) depression poster-

iorly contiguous with margin, adjacent margin lacking distinct marginal line;

horns just anterior to and on either side of midline lobed, acutely rounded,

delimiting posterior of broad, largely smooth, concavity; pronotal concavity

at anterior base of horns as wide as head, medially in anterior half granulate-

setose, posteriorly between horns with only scattered setose punctures.

Lateral thirds of pronotum lacking distinct short carinae, granulate-setose.

Fore tibia with five teeth on outer margin. Genitalia (Fig. 14) with apical

edges of parameres plain, not bent or lobed, the slender protrusion on each

paramere long, evenly arcuate.

ALLOTYPE. Female, length 16.0 mm, greatest width 11.5 mm.

Carinae (Fig. 15) on head typical for genus, anterior clypeal carina slanted

posteriorly, arcuate in frontal view; posterior clypeal carina slightly anter-

iorly arcuate, trituberculate. Frons and vertex punctate to granulate. Pro-

notum (Fig. 15) with anterior margin similar to holotype, less distinctly

tuberculate laterally; depression between longitudinal carinae anteriorly

approximately as wide as vertex, sparsely granulate near center on either

side of median granulate band; midline slightly more deeply impressed in

scutellar region; longitudinal carina on each side becoming obsolete in anter-

ior half; small transverse lateral carina present on each side of pronotum

anterior to humerus.

TYPE MATERIAL: Holotype, male, Argentina, Formosa, Clorinda,

X1I.1949, Martinez (Martinez).

Allotype, female, Paraguay, Santa Barbara, III.1948, Schade

(Martinez).

PARATYPES: 3 males, 4 females.

ARGENTINA: one male, same data as holotype.

PARAGUAY: Puerto Villeta, I.1945, Tepper. Departamento del

Guayra, Villarrica, III.1941, Schade. Departamento Capital, Trinidad, X.

1948, Asuncién, XI.1945, Podtiaguin.

Paratypes are in: Howden, Martinez.

REMARKS. The type series shows little important variation; one male

measures 14.5 mm and the allotype represents the maximum size seen.

The small male has slightly smaller horns and a deeper indentation in the

pronotal prescutellar area. This type of allometric variation is relatively

minor for the genus.

The combination of characters mentioned in the diagnosis should serve

to identify hypocritus.

6. Athyreus brasilicus n. sp.

Figures 19, 20, Map 2

DIAGNOSIS. Clypeal horn small, slender, erect, similar to bilobus in

vavini group; frons and vertex punctate or granulate, setose. Pronotum with

anterior margin elevated between fossae, margin thickened, posterior edge

Sinuate, slightly indented medially, lacking abrupt, tuberculate angle on each

side near fossa; concave area of pronotum in central third, including inner

26 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

surface of flattened horns mostly smooth, shining, the smooth area divided

by median granulate-setose band extending along midline from anterior to

posterior margin; median area posterior to horns not sharply depressed nor

otherwise modified.

HOLOTYPE. Male, length13.0 mm, greatest width 9.2 mm. Dorsal-

ly brownish black, labrum and lateral portions of pronotum brown, elytral

depressions (striae) dull, not shining. Similar to bilobus and hypocritus,

differing primarily as follows: clypeal horn (Fig. 19) slender, tapered

apically; posterior lateral carina fine, merging with lateral carina above

antennal insertion, the carina subdentate or tuberculate. Frons and vertex

irregularly punctate or granulate, setose. Pronotum with anterior margin

elevated between fossae and thickened, posterior edge of elevated margin

slightly sinuate, feebly indented medially, rounded laterally near fossae.

Pronotal horn or lobe (Fig. 19) on each side of midline in posterior half

laterally compressed, rounded at apex, somewhat longer than high, horn

anteriorly divergent from opposite horn. Posterior of pronotum or pre-

scutellar region feebly, longitudinally depressed, surface setose, granulate;

the depression and granules continuing anteriorly in a band along midline to

base of anterior margin; the granulate band completely dividing smooth areas

of anterior central concavity into two. In other related species the median

granulate band is incomplete or absent in well developed males. Pronotum

laterally granulate, similar to vavini, bilobus and hypocritus in this respect.

Fore tibia with five teeth on outer margin. Genitalia (Fig. 20) with ventral

apex of each paramere reflexed, thickened, the apical protrusions small,

arcuate and acute.

FEMALE. Unknown.

TYPE MATERIAL: Holotype, male, Brazil (Ottawa).

PARATYPES: 2 males.

BRAZIL: Sabara-Bello Horizonte, Rio das Velhao (sic), A.G.N.Chalmers.

Paratypes are in: London, Howden.

REMARKS. Athyreus brasilicus is a member of the vavini complex and

may be distinguished from closely related species by the complete granulate

band that extends along the midline of the pronotum from the anterior to pos-

terior margins. The two paratypes do not vary significantly from the holo-

type.

7. Athyreus bilobus n. sp.

Figures 16-18, Map 2

DIAGNOSIS. Color almost uniform brown to dark brown. Related to

vavini, hypocritus and brasilicus. Males readily distinguishable from these

species by the clypeal horn in cross section wider than long, usually notched

at apex, and with posterior vertical surface concave; anterior pronotal margin

raised between fossae, edge of raised area relatively thin, truncate, lateral-

ly not noticeably tuberculate. Females with anterior pronotal margin between

fossae elevated into a low, thin ridge; width of anterior elevation approx-

imately equal to width of prescutellar depression.

HOLOTYPE. Male, length19.0 mm, greatest width 12.0 mm.

Dorsally brown with apices of horns and tubercles darker. Anterior clypeal

face vertical, delimited dorsally by a semi-circular ridge, slightly dentate

medially; clypeal horn (Fig. 16) immediately posterior to this with anterior

face likewise vertical, horn unusually stout, as wide as long or wider in

cross section near blunt apex; horn with anterior lateral carina on each side

Howden and Martinez: The Genus Athyreus cag

distinct, not extending to arcuate anterior clypeal ridge; posterior lateral

carina on each side extending laterally and slightly arcuately posteriorly to

merge with ridge at anterior edge of gena. Frons posterior to horn smooth,

sparsely punctate, vertex granulate, setose. Pronotum (Fig. 16) with anter-

ior margin distinctly elevated between fossae, raised area truncate at top,

edge relatively thin, not distinctly tuberculate laterally; pronotum posterior-

ly in prescutellar region with broad, almost circular depression; anterior,

strongly declivous margin of depression setose and situated directly between

posterior edges of horns; pronotal midline anterior to depression indicated

by finely depressed line and small fovea midway between anterior margin and

horns; area anterior to broadly lobed horns widely, shallowly concave; con-

cavity largely smooth, punctate near anterior margin and on inner sides of

lobes or horns. Pronotum lateral to horn on each side with feebly developed

carina extending from near base of horn toward median lateral angle of

margin; lateral to horns and concavity pronotum granulate-setose. Fore

tibia with five distinct teeth on outer margin, a sixth minute tooth present

basally (not present in other specimens). Genitalia (Fig. 17) with a distinct

apical lobe on each paramere adjacent to arcuate, long, acute prolongation.

ALLOTYPE. Female, length18.0 mm, greatest width 11.0 mm.

Head with carinae as is typical in genus, the posterior clypeal carina slightly

anteriorly arcuate, trituberculate, median tubercle slightly larger. Frons

and vertex entirely, irregularly granulate, granules never separated by more

than three diameters, usually less. Pronotum (Fig. 18) with raised portion

of anterior margin as in male, a faint denticle medially; pronotum almost

completely granulate except for carinae, part of midline extending between

faint anterior fovea at anterior third and faint fovea at posterior sixth

(=prescutellar depression); on either side of midline between foveae granules

irregular, sometimes widely separated; longitudinal carina on each side

distinct from anterior fifth to posterior sixth; anteriorly separated by approx-

imately width equal to width of clypeus, posteriorly converging to 1/2 anterior

width. Lateral oblique carina feebly developed, similar to male.

TYPE MATERIAL: Holotype, male, Brazil, Estado do Paranda,

Arapongas, XII.1951, A. Maller (Martinez).

Allotype, female, same data as holotype (Martinez).

PARATYPES: 11 males, 6 females.

BRAZIL: Parana, XII.1951, XI.1956, A. Maller, XII.1956,

Dirings; N. Parana. Sd&o Paulo, Rio Claro, X.1940, 1.1952, F.S. Pereira;

Araraquara, Rio Aukumas, XII.1954, Olivastro; ha Comprida, 1V.1969,

J.S. Carvalho; Aguapehy, III.1941, Dufaux. D.F., Rio de Janeiro, 1926,

Moraes-Mello. Minas Gerais, Rio Preto, II.1962, ae Machado; Belo

Horizonte, III.1961, Machado; Varginha, X.1952. Goias, Anapolis, XI.

1930, Zellibor.

PARAGUAY: no other data. Guayra, Villarrica, XI.1941, F. Schade;

Colonia Elisa, 17. XI.1940, B. Podtiaguin.

Paratypes are in: Buenos Aires, Cambridge, London, Ottawa, Howden,

Martinez.

REMARKS. Variation is not great. Some allometric growth is evident

in the variation of male horn length, and the prescutellar depression of the

pronotum in males also shows variation in the size and depth of the depression.

Color also varies, the dark brown specimens we suspect are older as most

are more abraded than lighter colored specimens. Characters needed for

identification are summarized in the diagnosis.

28 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

8. Athyreus hemisphaericus Boucomont

Figures 25-27, Map 3

Athyreus hemisphaericus Boucomont, 1902a, p. 184; 1902b, p. 582; 1928, p.

206.

Athyreus canaliculatus Boucomont, 1928, p. 206. NEW SYNONYMY.

TYPE, hemisphaericus. Lectotype here designated, female originally

described as male by Boucomont, 1902a, p. 184 and 1902b, p. 582 and used

for figure 3, plate 5 of volume 71 (1902b). Type locality "Jatahy" (Prov.

Goias, Brazil). (Paris), examined.

TYPE, canaliculatus. Male (Paris), examined.

MALES. Length15.0to17.0 mm, greatest width 10.5 to1l.5 mm.

Dorsally reddish brown to dark reddish brown, apices of horns, mandibles

and margins often darker. Clypeal horn (Fig. 25) small, laterally compress-

ed, anterior face obliquely slanted anteriorly then curved posteriorly in

apical fourth, apex rounded, longer than wide in cross section; anterior

lateral carina distinct, posterior lateral carina extending from base of horn

in some specimens, obsolete in others, when present low and arcuate to

junction with gena. Frons nearly flat, closely punctate. Vertex posteriorly

concave on each side of convex midline, elevated median area expanding

anteriorly, sparsely to densely punctate-granulate, setose. Anterior lateral

edge of eye canthus nearly right-angled, not acute or more strongly recurved

than remainder of canthus as is typical in related species. Pronotum (Fig.

25) with anterior margin between fossae obtusely angulate and subdentate at

midline; pronotal midline deeply, narrowly impressed for most of length,

almost forming fovea at anterior third with smooth area on either side extend-

ing posteriorly to anterior angles of raised carinae (equivalent to "horns"' of

vavini and relatives). Pronotum centrally with raised, obtusely angled

carina on each side of midline, carina highest anteriorly, low and very

slightly convergent posteriorly, ending in obtusely rounded, posteriorly and

medially directed lobe just anterior to posterior margin and overhanging

deep longitudinal median indentation; indentation 1 to 1. 5 mm in length,

narrowed anteriorly by overhanging sides, interior of indentation distinctly

setose. Angulate pronotal carina anteriorly extending laterally, carina

becoming feeble, obsolete in some specimens, joining lateral margin slightly

posterior to mid point between anterior and median angle; juncture feebly

angulate; a second short carina present on pronotum midway between anterior

carina and posterior margin anterior to humerus, carina usually wide and

well developed. Lateral pronotal surfaces, excluding carinae, closely granu-

late, setose. Fore tibia with five distinct teeth on outer margin, a sixth

minute basal tooth often present on tibia of unabraded specimens. Genitalia

(Fig. 26) with parameres apically simple, moderate in length, protrusion

uniformly arcuate.

FEMALES. Length15.5to17.5 mm, greatest width10.5to12.0 mm.

Head with typical female carinae, the posterior transverse carina often with

median tubercle well developed and slightly anterior to smaller, more

oblique lateral tubercle on each side; frons and vertex granulate, vertex not

noticeably concave posteriorly near eyes. Pronotum (Fig. 27) with anterior

margin and carinae not greatly different from small males; major difference

is in lack of deep prescutellar indentation and overhanging lobes; instead

area unmodified, midline narrow and deeply impressed, setose from just

before posterior margin to anterior fourth; carina on either side almost

parallel to anterior angulation, carina extending laterally from angulation to

Howden and Martinez: The Genus Athyreus 29

pronotal margin poorly delimited, in some specimens absent; short carina

near posterior pronotal margin anterior to humerus usually well developed.

MATERIAL EXAMINED: 17 males, 16 females.

ARGENTINA: Corrientes, Ituzaingé, Isla Apipe Grande, X1.1945,

Martinez. Misionés, Santa Maria, Viana; Loreto (Exp. St.), Ogloblin; San

Ignacio, Villa Lutetia, January, Wagner.

PARAGUAY: Alto Parand, Hohenau, XI.1941, Jakob.

BRAZIL: Sa&o Paulo, Cidade (Ipiranga Santo Amaro), Pastore, Lane,

Martinez. Minas Gerais; Uberaba, Summer 1956/57, Mendoza.

Specimens are in: Buenos Aires, London, Ottawa, Paris, Howden,

Martinez.

REMARKS. The species is readily recognizable by the uniform reddish

brown color, shape of pronotal carinae in both males and females, by the

deep, sulcate indentation in the prescutellar region of males and by the male

genital characters.

Boucomont originally (1902) confused the females of two different species

describing one female (herein chosen as the lectotype of hemisphaericus) as a

male. Subsequently when he described canaliculatus (1928) he stated that the

male canaliculatus might be the same as s hemisphaericus as he had discovered

that two females were represented in his original description, the "male" of

his description being the female of "canaliculatus". In selecting the lectotype

it seemed best to select the female that could be definitely related to males

without any doubt that they represented a single species, hence the establish-

ment of canaliculatus as a synonym of hemisphaericus.

9. Athyreus nitidus n. sp.

Figures 21, 22, Map 3

DIAGNOSIS. Dorsally brown; head, pronotum and elytra unusually

shiny, granules and punctures greatly reduced; clypeal horn erect, anterior

and posterior lateral carinae obsolete at base of horn, distinct on horn;

anterior pronotal margin raised and dentate on either side of midline; pronotal

horn on either side of midline blade-like, erect, apically bidentate, widely

separated; middle third of pronotum broadly, shallowly, concave.

HOLOTYPE. Male, length15.0 mm, greatest width 10.0 mm. Dorsal-

ly shining brown. Clypeal horn (Fig. 21) erect, anterior and posterior lateral

carinae present on horn, anterior carina briefly interrupted near base, pos-

terior lateral carina present only on apical half of horn, absent elsewhere;

posterior median carina lacking. Frons and vertex with scattered, long,

semi-recumbent setae, seta-bearing punctures small, sparse, surface

between shining. Pronotum (Fig. 21) with anterior margin between fossae

raised medially, an obtuse tooth present on either side of midline; lateral

pronotal margin somewhat irregularly arcuate to feebly sinuate near rounded

median angle. Pronotum centrally, longitudinally broadly concave; on either

side at edge of concavity just behind middle of pronotum with a laterally

flattened, erect, blade-like horn having apex bifurcate, the posterior bifur-

cation much larger. Median posterior (prescutellar) area of pronotum not

noticeably depressed, posterior margin at midline very narrow, marginal

line obsolete. Elytra with broad indentations (striae) feebly delimited, much

of surface unusually smooth and shining. Fore tibia with five teeth on outer

margin. Genitalia as in fig. 22, with apical lobes very similar to vavini and

hypocritus k

FEMALE. Unknown.

30 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

TYPE MATERIAL: Holotype, male, Bolivia, Dpto. Sta. Cruz, Pcia.

Cordillera, Rio Seco, II. 1962, Martinez (Martinez).

REMARKS. The rather polished, shining brown appearance of nitidus

is quite characteristic, and this coupled with the widely separated, blade-

like bifurcate pronotal horns should be sufficient for identification.

10. Athyreus anneae n. sp.

Figures 28, 24, Map 3

DIAGNOSIS. Color tan to dark brown. Clypeal horn with anterior

lateral carina complete or only slightly interrupted near base; posterior

lateral carina present on horn, lacking on base; posterior median carina

absent. Pronotum with midline depressed, posterior third with laterally

compressed horn on each side adjacent to and paralleling median depression,

each horn bidentate, the horn arcuately depressed between anterior and pos-

terior denticles; posterior margin of pronotum not interrupted or greatly

modified. Female unknown.

HOLOTYPE. Male, length 21.5 mm, greatest width13.0mm. Dorsal-

ly light brown. Clypeal horn (Fig. 23) vertical, anterior lateral carina com-

plete, slightly interrupted at base of horn; posterior lateral carina present

only on vertical portion of horn, absent elsewhere; posterior median carina

absent. Frons and vertex almost completely impunctate, non-granulate;

vertex with distinctly circular concavity on each side adjacent to eye, median

area longitudinally strongly convex. Pronotum (Fig. 23) with midline indented

except at anterior fifth or sixth; horn either side of and paralleling midline

in posterior third, apex of horn bidentate, arcuately depressed between acute

anterior and posterior denticles; horn separated from one on opposite side by

distance of 3 to 4 mm; areas anterior and posterior to horns to margins most-

ly smooth, impunctate, non-granulate. Outer surface of each horn near base

and lateral thirds of pronotum closely setose-granulate; lateral margin

arcuate, anterior margin between fossae very obtusely angulate, summit of

angulation at midline with minute tubercle; posterior margin at midline

obtusely rounded and extended slightly posteriorly, area just anterior to pos-

terior margin and posterior to horns concave, impunctate, non-granulate.

Fore tibia with six teeth on outer margin. Genitalia as in fig. 24.

FEMALE. Unknown.

TYPE MATERIAL: Holotype, male, Paraguay, Caaguazu District,

Estancia Primera, 28.xii.1931, R.F. Hussey (Howden).

PARATYPES. 2 males.

BRAZIL: Sd&o Paulo, Jundiaf, XI1.1956, Martinez.

PARAGUAY: Cerro Amambay, X.1939, A. Schultze.

Paratypes are in: Washington, Martinez.

REMARKS. Variation is mainly evident in the allometric growth of the

horns, the paratype from Paraguay having the poorest development but with

the carinae and shape as described above. In the small paratype the vertex

and frons are mostly granulate or punctate and the color is tan rather than

brown. The species is named for Anne T. Howden in recognition of her con-

tribution to many aspects of this paper.

Howden and Martinez: The Genus Athyreus 31

11. Athyreus forcipatus Boucomont

Figures 31-33, Map 4

Athyreus forcipatus Boucomont, 1928, p. 207.

TYPE. Male (Paris), examined.

MALES. Length 15.0tol7.0mm, greatest width 10 .5to11.0 mm.

Color dorsally varying from brown centrally, tan laterally, to black central-

ly, brown laterally. Clypeal horn (Fig. 31) almost an equilateral triangle in

median cross section, anterior surface flat, posteriorly with posterior

lateral carinae joining near base to form single carinate posterior edge; near

base of horn anterior and posterior lateral carinae present; posterior median

carina absent. Frons and vertex with widely scattered punctures, otherwise

smooth, vertex shallowly concave near eye either side of slightly convex mid-

line. Pronotum (Fig. 31) with horn on posterior half either side of midline;

viewed laterally horn trilobed, anterior lobe sharp, directed slightly anter-

iorly, median lobe distinct, usually slightly obtuse and nearly upright, pos-

terior lobe extending horizontally over posterior pronotal margin and partially

obscuring scutellum; posterior lobes narrowly, deeply separated; middle

lobes separated by approximately 1mm, anterior lobes by 1.5 to 2.0 mm.

Pronotal depression anterior to horns with center of depression granulate,

the surrounding area laterally and posteriorly largely smooth, with sparse,

scattered punctures, non-granulate; lateral convex portions of pronotum

closely granulate, lateral carinae absent; lateral margin arcuate. Elytral

depressions or wide striae (Fig. 33) with setose granules distinctly separated,

third depression with moderately even double row of granules. Fore tibia

with five teeth on outer margin. Genitalia as in fig. 32.

FEMALES. Unknown or unassociated (granules in elytral intervals

should be useful for identification of females when distribution is also con-

sidered).

MATERIAL EXAMINED: 4 males.

BOLIVIA: La Paz (type locality),Santa Cruz. Cochabamba, Pcia.

Chapare, Km. 14, 800m, Martinez.

Specimens are in: Paris, Washington, Howden, Martinez.

REMARKS. In the four males seen the trilobed horn on each side of the

pronotal midline varied both in height and in the degree of anterior-posterior

separation of the lobes, the distance between the anterior and median lobes

being most noticeable. The widely spaced granules in the elytral intervals

seems to be a fairly unique character in the group and this coupled with the

large, non-granulate area in the anterior portion of the pronotum should

identify the species.

12. Athyreus alvarengai n. sp.

Figures 34-36, Map 4

DIAGNOSIS. Very similar to forcipatus. Males with posterior lateral

carinae obsolete near base of clypeal horn; vertex distinctly concave on each

side near eye, midline of vertex distinctly convex; pronotal depression anter-

ior to horns largely granulate, granules in elytral depression (striae) dense,

third stria with three irregular rows of granules.

HOLOTYPE. Male, length15.8 mm, greatest width 10.5 mm.

Dorsally black with lateral pronotal margins dark brown. Clypeal horn (Fig.

34) more acutely triangular than in forcipatus, anterior-posterior length near

ae Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

base 1/4 more than in forcipatus; posterior lateral carinae obsolete near base,

distinct laterally. Frons mostly punctate; transverse band at junction of

frons and vertex impunctate; concave area of vertex on each side near eye

impunctate; median convex portion of vertex granulate. Pronotum (Fig. 34)

similar to that of forcipatus but with trilobed horn on each side in anterior

half separated from horn on other side by 2.5 to 3 mm; concavity anterior to

horns occupying median third of pronotum, setose-granulate except for vague

non-granular ridge anterior to each horn, the vague ridge delimiting central

concavity; pronotum lateral to ridge and horns convex, closely setose-granu-

late. Elytral striae (Fig. 36) closely setose-granulate, granules in third

stria forming three irregular rows. Fore tibia with five teeth on outer mar-

gin. Genitalia as in fig. 35.

FEMALE. Unknown.

TYPE MATERIAL: Holotype, male, Brazil, Pernambuco, Escada, X.

A. Naller (Martinez).

Paratype, 1 male.

BRAZIL: Para, Belém, Utinga, IX .1956, Martinez (Howden).

REMARKS. The single male paratype measures 16.0 mm in length

and both the clypeal and pronotal horns are reduced in size compared to the

holotype. Except for the allometric difference the two specimens are essen-

tially similar. ,

This species is named for Col. Moacir Alvarenga from Rio de Janeiro

in recognition of his numerous entomological endeavors which have advanced

entomology in general and greatly aided this study in particular by providing

the authors with many rare species of Athyreus.

13. Athyreus pyriformis n. sp.

Figures 37, 38, Map 4

DIAGNOSIS. Males with clypeal horn at apex much wider than long,

horn anteriorly-posteriorly compressed, apex nearly truncate. Pronotum

with anterior median half, smooth, shallowly concave; concavity rising nearly

vertically and narrowing posteriorly, forming an upright horn; horn along

median line narrowly, deeply sulcate, the midline indented posterior to

sulcus, widening and deepening about 1 mm before posterior pronotal margin;

margin posteriorly arcuate at posterior of indentation, marginal line incom-

plete. Pronotum in lateral view approximately three times higher than long.

Females not recognized.

HOLOTYPE. Male, length16.5 mm, greatest width 10.8 mm.

Dorsally black, head and sides of pronotum and elytra very dark brown.

Clypeal horn (Fig. 37) with anterior and posterior lateral carinae feebly

developed, largely obsolete on horn, posterior median carina absent; horn

(in cross section) near base only slightly wider than anterior-posterior

length, in apical half two to three times wider than anterior-posterior length;

apex bluntly rounded to truncate. Frons and vertex setose-punctate except

for small median area in center of vertex smooth, a few scattered punctures

present. Pronotum (Fig. 37) with anterior edge of horn rising nearly vert-

ically from posterior edge of large, shallowly concave area extending from

just posterior to anterior margin onto face of horn; central anterior half of

pronotum smooth. Pronotal horn obliquely blunt in lateral view, in posterior

(or anterior) view deeply sulcate the two plates of the horn narrowly separated

(0.5 mm at apex); indentation continuing posteriorly down horn, deepening

and widening on posterior portion of pronotum, forming oval cavity posteriorly

Howden and Martinez: The Genus Athyreus 33

that is delimited by thin posterior pronotal margin, marginal line broken.

Pronotum in lateral view much higher than long, lateral third on each side

from anterior angles to posterior median depression behind horn closely

setose-granulate; lateral pronotal margin just anterior to median angle, very

feebly interrupted immediately below lateral fovea. Fore tibia with five teeth

on outer margin. Genitalia as in fig. 38, the prolongations of the parameres

relatively short for group.

TYPE MATERIAL: Holotype, male, Peru, Junin, Chanchamayo, 3.X.

1961, J. Schunke L. (Howden).

Paratype, 1 male.

PERU: Loreto , Pucallpa, Hocking (Washington).

REMARKS. The single male paratype does not vary significantly from

the holotype except that the size is slightly less (about 0.5 mm).

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. alvarengai n. sp.

- pyriformis n. sp.

. unicornis n. sp.

14. Athyreus unicornis n. sp.

Figures 28-30, Map 4

DIAGNOSIS. Males lacking clypeal horn, pronotum with unique slender

horn arising from center of pronotum, extending anteriorly and inclined down-

ward over head, terminating near anterior edge of clypeus. Females with

34 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

two distinctly separated central pronotal tubercles, a short carina on each

side delimiting central dark brown to black area that extends anteriorly to

margin; remainder of elytra and sides of pronotum reddish brown. Both

sexes with five teeth on outer margin of fore tibia. Occurring in northern

Colombia.

HOLOTYPE. Male, length17.5 mm, greatest width 12.0 mm. Dis-

tinctly bicolored dorsally; frons, vertex central third of pronotum and all but

lateral and apical fourth or third of elytra very dark brown to black; labrum,

clypeus, lateral portions of pronotum and elytra reddish brown. Clypeus

(Fig. 28) with a single, uniformly low, arcuate carina extending from above

antennal insertion on each side anteriorly to middle of clypeus; clypeus anter-

ior to carina sloping gradually downward to margins; medially, a low tubercle

present in center of frons. Frons evenly, vertex less evenly, punctate.

Pronotum (Fig. 28) with slender, nearly cylindrical horn extending anteriorly

and inclined downward, terminating above anterior margin of clypeus; basal

dorsal half of horn and adjacent midline posterior to horn indented, indenta-

tion punctate on horn, setose granulate behind. Pronotum on each side with

short vague carina near base of horn, second more distinct carina near pos-

terior pronotal margin anterior to elytral umbone. Pronotum anterior to

horn smooth and shallowly concave; lateral and posterior thirds of pronotum

setose punctate or granulate; lateral pronotal margin on each side between

acute anterior angle and median angle bisinuate. Fore tibia with five teeth

on outer margin (basal notches between teeth enlarged by some mechanical

process in type, not natural abrasion; also same is true for notches of

mandibles). Genitalia as in fig. 29, similar to bifurcatus.

ALLOTYPE. Female, length18.0 mm, greatest width 13.0 mm.

Dorsally similar in color to male. Head not differing greatly in shape of

carinae from females of related species. Pronotum (Fig. 30) with two

central tubercles distinct, conical, separated by approximately 1 mm; on

each side at edge of dark area posterior to tubercles a short, poorly developed

carina (better developed in paratypes). A short distinct carina present

paralleling posterior pronotal margin just anterior to elytral umbone. Pro-

notal surface entirely setose-granulate; lateral margin on each side between

anterior and median angles vaguely sinuate; fovea above median angle, small,

oval, moderately deep and shining. Fore tibia with five teeth on outer mar-

cin.

TYPE MATERIAL: Holotype, male, Colombia, Valle Medellin, X.1945,

F.L. Gallego M., U.S.N.M. Ins. no. 176630. Col. 37-XX3 (Washington).

Allotype, female, same data as holotype (Washington).

PARATYPES. 1male, 3 females.

COLOMBIA: Magdalena, Sierra Nevadade Santa Marta, Pueblo Bello,

1100 m, 10-17.VI.1968, B. Malkin. Santander, Socorro, VIII.1940.

Paratypes are in: Chicago, Howden, Martinez.

REMARKS. Color normally is not a useful character in the bifurcatus

gsroup, but coloration is constant in the six specimens seen. At present the

range by itself can be considered diagnostic since no other species of

Athyreus is known from the area of Colombia in which unicornis occurs.

15. Athyreus armatus Westwood

Figure 39

Athyreus armatus Westwood, 1848, p. 386; 1851, p. 459; Boucomont, 1912,

p. 4.

Howden and Martinez: The Genus Athyreus 35

TYPE. Male (Oxford), examined.

MALE. Length18.0mm, greatest width about 11.0 mm. Dorsally

black with mandibles and sides of pronotum brown. Anterior lobe of each

mandible projecting strongly anteriorly, forming an elongate lobe. Clypeus

anteriorly with a small median tubercle; a nearly straight carina on each side

extending from median tubercle to lateral margin above antennal insertion,

there forming a small tubercle. Frons and vertex centrally briefly impunc-

tate, most of frons and vertex setose, punctate or granulate. Pronotum (Fig.

39) very unusual for genus; centrally with a single, anteriorly inclined horn;

on each side anterior to horn a rounded, ''flap-like" dilation or lobe extending

to anterior margin behind eye, the dilation most pronounced near posterior

terminus. Lobes or dilations laterally and horn posteriorly effectively de-

limiting anterior, central half of pronotum; area within shallowly concave and

sparsely granulate. Area posterior to pronotal horn medially depressed,

closely setose-granulate; pronotum laterally, including indentation below

dilations, largely impunctate. Fore tibia with five teeth on outer margin.

MATERIAL EXAMINED: 1 male (holotype).

"AMERICA MERIDIONALI": no other data. |

REMARKS. The single male of armatus differs greatly in the pronotal

armature from other species of Athyreus. The laterally directed "flap-like"

dilations projecting from lateral third of pronotum anterior to transverse line

of central horn is nearly unique. The elongate, anteriorly projecting lobe on

each mandible resembles to a degree the mandibles of gigas. The essentially

hornless clypeus and the anteriorly inclined pronotal horn vaguely resembles

unicornis. The lack of any additional specimens since Westwood's descrip-

tion indicates both rarity and lack of collecting. Boucomont, 1911, in his

catalogue lists the locality of armatus as '"Brazil'' (which probably does equal

"America meridionali'').

16. Athyreus biarmatus n. sp.

Figures 79-81, Map 5

DIAGNOSIS. Male lacking clypeal horn, horn represented anteriorly by

a low transverse tubercle; pronotum in anterior half on either side behind

eye with a laterally directed lobe slightly overhanging pronotal surface;

center of pronotum with horn represented by a low swelling bifurcate apically,

each furcation distinct and acutely pointed. Female with very distinct U -

shaped carina enclosing a short distinctly raised carina on either side of mid-

line, the U -shaped carina incomplete posteriorly; short inner carinae

slightly convergent posteriorly but completely separate.

HOLOTYPE. Male, length 15.0 mm, greatest width 9.9 mm, bi-

colored dorsally; vertex, longitudinal central half of pronotum, and elytra

very dark reddish brown; labrum, lateral fourths of pronotum, and legs light

reddish brown. Clypeus (Fig. 79) with complete transverse anterior carina,

carina feeble; medially and contiguous to carina a small transverse tubercle;

posterior carinae obsolete, evident laterally above antennal insertions; frons

slightly tumid, surface of frons and vertex with scattered setose granules.

Pronotum (Fig. 79) with central third in anterior half lobed on either side

posterior to fossa, the lobe raised, projecting laterally and slightly over-

hanging lateral surface; viewed from above lobes and anterior margin pre-

senting a broadly U -shaped raised area with central portion slightly de-

pressed and with scattered granules. Immediately behind this area at center

of pronotum a feeble swelling present with apex distinctly bifurcate, each

36 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

portion of bifurcation abruptly raised,slightly divergent and acute at apex; on

each side of horn posteriorly a slightly sinuate carina terminating just poster-

ior and lateral to the posterior terminus of lobe; midline posterior to central

horn or swelling shallowly, distinctly indented; surface of pronotum except

for raised areas moderately granulate, setose; lateral pronotal margins

sinuate. Elytron between suture and umbone with intervals strongly convex;

each stria largely represented by irregular, double row of granules. Fore

tibia with five teeth on outer margin. Genitalia as in figure 80.

ALLOTYPE. Female, length17.1mm, greatest width 10.5 mm.

Color, granules, elytra, striae, and fore tibia as described for male. Clypeus

anteriorly with evenly arcuate transverse carinae; posterior clypeal carina

tridentate, median tubercle better developed and very slightly anterior in

position to lateral tubercles. Vertex centrally flat, granules widely spaced.

Pronotum (Fig. 81) with longitudinal central third largely enclosed by strongly

raised U -shaped carina with greatest elevation near anterior margin; a fine

carina extending posteriorly from margin on each side just inside fossa and

merging with U -shaped carina; each terminus of U -shaped carina con-

verging posteriorly, terminating approximately 1mm before posterior mar-

gin; area between broadly, shallowly depressed. Center of pronotum on

either side of midline with strongly raised longitudinal carina, the two carinae

slightly convergent posteriorly.

TYPE MATERIAL: Holotype, male, Trinidad, 92-66 (London). Allo-

type, female, Trinidad, St. Augustine, 22.8.1925, C.L. Withycombe

(London).

REMARKS. The species is closely related to armatus but can be

readily distinguished by the bifurcate pronotal horn of the male; the male of

armatus has a long, slender, anteriorly inclined horn. The lateral pronotal

lobes will distinguish armatus and biarmatus males from all other species.

17. Athyreus gigas Westwood

Figures 40-42, Map 5

Athyreus gigas Westwood, 1848, p. 386; 1851, p. 458.

TYPE. Male (Oxford), examined.

MALES. Length 24.0 to 25.0 mm, greatest width13.5 mm. Dorsally

reddish tan to brown. Each mandible with three lobes in addition to acute

apical tooth; anterior apical lobe directed anteriorly, acutely triangular in

shape, similar in this respect to armatus. Clypeus (Fig. 40) with three low

horns of equal height; one medially at anterior edge of clypeus with vague but

typical anterior lateral, posterior lateral and posterior medial carinae

evident; obsolete posterior lateral carina on each side forming an acutely

triangular horn just anterior to gena, horn projecting slightly outward over

antennal insertion. Clypeus and area between horns with scattered, distinct-

ly separated punctures. Frons largely impunctate, posteriorly with a small,

low tubercle on each side of midline; vertex posterior to tubercles setose-

punctate. Pronotum (Fig. 40) with two divergent, blade-like horns arising

from common base, nearly as in some bifurcatus but with horns stouter and

recurved upward on anterior edge. Anterior face of horns impunctate,

smooth; anterior and posterior surfaces setose-granulate. Pronotum lateral-

ly on each side of horns shallowly, broadly concave, the concavity slightly

irregular and non-granulate, impunctate; area near anterior angles flat, with

minute scattered punctures. Lateral pronotal margin straight from anterior

angle to just anterior to median angle where briefly sinuate. Elytral intervals

Howden and Martinez: The Genus Athyreus 37

between suture and umbone poorly delimited, somewhat irregularly spaced.

Genitalia (Fig. 41) with prolongation of paramere (Fig. 42) slightly expanded

and bluntly rounded at apex.

FEMALE. Unknown.

MATERIAL EXAMINED: 2 males.

BRAZIL: no other data (holotype). Mato Grosso, near Cuiaba, XI. 48.

Specimens are in: Oxford, Martinez.

REMARKS. The two specimens seen are very similar, and are easily

separable from other Athyreus, except possibly for the smaller parvus. The

three low horns on the head are an excellent specific character and this, in

conjunction with the smooth areas of the pronotum, immediately separates

gigas from related species such as parvus and bifurcatus.

m=16. biarmatus n. sp. m= 20. tuberifer Fels.

*=17 . gigas Westw. A= 21. juanae n. sp.

@® =19. bifurcatus Macl. @= 22. bellator Westw.

18. Athyreus parvus n. sp.

Figures 82-84

DIAGNOSIS. Clypeus with three horns or large tubercles; anterior

median horn with anterior and posterior lateral carinae and feeble median

posterior carina; lateral "horn" on each side formed by strongly protruding

38 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

angulation of lateral margin positioned above base of mandible distinctly

anterior to gena. Central horn of pronotum bifurcate, as in some bifurcatus;

a circular concavity present on either side of midline anterior to horn; anter-

ior and lateral surfaces of pronotum granular or punctate, not smooth.

HOLOTYPE. Male, length14.9 mm, greatest width 9.8 mm. Clypeus

(Figs. 82, 83) with three horns or tubercles; anterior central horn most pro-

nounced, occupying most of center of clypeus; anterior face of clypeus nearly

vertical, triangular; face delimited on each side by the anterior lateral carina,

carina merging with opposite one near slender apex of horn; horn in lateral

view broadly triangular, slightly bent posteriorly; posterior median carina

feeble; posterior lateral carina on each side strong, elevated, forming a

large, acute tubercle or small horn overhanging base of mandible. Frons

rising uniformly from vertex to base of clypeal horns or tubercles, inclined

surface closely, often confluently punctate; vertex moderately punctate, nearly

flat medially, with a minute tubercle on either side of midline in line with

posterior margin of eyes. Pronotum (Figs. 82, 83) with central horn strongly

bifurcate nearly from base, each furcation inclined anteriorly, divergent from

opposite side, and tapered to acute tip. Pronotum on each side anterior to

horn with moderately deep, almost circular depression; midline behind horn

shallowly impressed; pronotum on each side posterior to horn with two short

feeble carinae, posterior one anterior to humerus and parallel to posterior

pronotal margin, more anterior and shorter carina positioned midway between

base of horn and posterior carina; pronotal surface except for carinae punc-

tate or granulate, largely setose, lacking smooth areas; lateral pronotal mar-

gin between anterior and median angles distinctly sinuate. Fore tibia with

five teeth on outer margin, a suggestion of a minute basal sixth tooth present

on right tibia. Genitalia (Fig. 84) with prolongations of parameres long,

stout, feebly twisted and with blunt apices.

FEMALE. Unassociated.

TYPE MATERIAL: Holotype, male, Brazil (Howden).

REMARKS. Athyreus parvus was at first included with a series of

bifurcatus because of the shape of the pronotal horn. The small size of parvus

originally caused confusion in regards to the short clypeal horn. However

when the genitalia was examined the species was found to be only distantly

related to bifurcatus. We now consider it to be a well developed male with

the clypeal characters showing affinities with gigas. The pronotal depressions

and large prolongation of each paramere are similar to the tribuliformis -

zischkai - martinezi complex. Athyreus parvus seems to be intermediate

between these species and gigas.

19. Athyreus bifurcatus Macleay

Figures 438-45, Map 5

Athyreus bifurcatus Macleay, 1819, p. 124; Guerin, 1956, p. 262; Martinez,

1950, DP. 2a2-

Athyreus herculeanus Laporte, 1840, p. 102; Boucomont, 1911 (1910), p.

346. NEW SYNONYMY. |

Athyreus rotundus Westwood, 1848, p. 386; 1851, p. 461. NEW SYNONYMY.

Athyreus tuberculatus Westwood, 1848, p. 386;1851, p. 460. NEW

SYNONYMY.

TYPE, bifurcatus. Male (location uncertain), not examined.

Howden and Martinez: The Genus Athyreus 39

TYPE, herculeanus (possible type). Female (Paris), examined.

TYPE, rotundus. Female (Oxford), examined.

TYPE, tuberculatus. Male (Oxford), examined.

MALES. Length 18.0 to 22.0 mm, greatest width 11.0 to13.0 mm.

Dorsally brown to very dark brown, pronotum often lighter in color than

elytra, particularly near margins. Clypeal horn (Fig. 43) strongly inclined

anteriorly; anterior lateral, posterior lateral, and posterior median carinae

all present; the posterior lateral carina on each side forming an abrupt,

sharply pointed low tubercle just anterior to gena. Frons and vertex setose,

punctate or granulate. Pronotum (Fig. 43) in well developed males with two

widely divergent horns arising from a common base; on each side of horns a

large, shallow concavity extending from near posterior pronotal margin 2/3

of distance to anterior margin. Lateral third of pronotal surface usually

impunctate; pronotal margin between anterior and median angles sinuate,

area just inside anterior angle tumid. In poorly developed males pronotal

horns indicated by two slightly separated divergent, longitudinal flaps (as in

type of tuberculatus), surface of pronotum largely granular, pronotal con-

cavities lacking, lateral pronotal margins between anterior and median angles

nearly straight. Fore tibia usually with six distinct teeth on outer margin.

Genitalia as in fig. 44.

FEMALES. Length17.0 to 24.0 mm, greatest width 11.0 to14.0 mm.

Females are either extremely variable or the series seen is a mixture of

species, probably both factors are present. All specimens have six teeth on

the fore tibia as do males of bifurcatus. Pronotum (Fig. 45) with central

swelling single or double, low or distinctly elevated; carina on either side

extending to anterior margin or incomplete laterally, in addition to large

arcuate carina, one or two short carinae present.

MATERIAL EXAMINED: 42 males, 48 females (only data from males

listed).

ARGENTINA: Misiones, Loreto, X1.1939, Ogloblin; Misiones, Oberda,

X.1944, Pastrana; Misionés, San Ignacio, 1V.1945, Réhmer; Misionés,

Puerto Victoria, XI.1940; Misionés, no other data.

BRAZIL: No other data (type locality). Sao Paulo, Barueri, 10.IV.

1956, K. Lenko; SdAo Paulo, CIPO, 25.11.1974, 18 .X.1972, 4.XII.1972,

V.N. Alin; Sao Paulo, Ypiranga, 24.X1.1942, A. Alves. Mato Grosso,

Pocone, XII.1948. Minas Gerais, Varginha, X.1950. Parana, Jaguariaiva,

XI1I.1955; Rio Tijuca, XII.1884, E. Gounelle. Rio de Janeiro.

PARAGUAY: Alto Parana, Hohenau, 1929, 15.X1.1948, H. Jacob;

Hohenau, 23.XI1.1945, G. Angel.

Specimens are in: Buenos Aires, Leiden, London, Ottawa, Oxford,

Paris, Washington, Hardy, Howden, Martinez.

REMARKS. Males of bifurcatus, except minor specimens, can be

distinguished by the distinctly anteriorly inclined clypeal horn, the two ©

laterally divergent pronotal horns with a shallow lateral concavity on each

side, and the fore tibia with six teeth. In male minors it is best to check the

genitalia (Fig. 44) which have the protrusions of the parameres abruptly

widened, almost bifurcate, at the apex. Females can only be determined by

association with males.

20. Athyreus tuberifer Felsche

Figures 46-48, Map 6

Athyreus tuberifer Felsche, 1909, p. 762; Martinez, 1953, p. 233 (the female

40 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

"Allotype" is not tuberifer).

TYPE. Female (Dresden ?), not examined.

MALES. Not distinguishable from female by usual dimorphism in horns

and carinae; males lack median dentate pronotal sulcus just anterior to pos-

terior margin (see female description). Length 20.0 to 24.0 mm, greatest

width 14.0to15.0mm. Dorsally dark brown to black with lateral margins

of pronotum and elytra often lighter. Clypeus (Fig. 46) with elongate, anter-

ior median horn strongly inclined anteriorly, both anterior lateral and pos-

terior lateral carinae extending nearly to apex; posterior median carina

obsolete, represented by a small, abrupt tubercle at base of horn. Frons and

vertex centrally impunctate, smooth, shallowly concave. Pronotum (Fig. 46)

centrally with a nearly cylindrical horn (the base 1.0 to1.5 mm high) having —

a small obtuse denticle at median posterior summit, and with an anterior

extension directed slightly upward; apex of extension deeply (1 to 2 mm)

bifurcate, each bifurcation slender, acute apically. Surface of central pro-

notal horn, except base anteriorly, closely punctate; on each side of midline

from near base of horn to posterior margin a cluster of setose-granules;

remainder of pronotal surface, except near anterior margin, smooth, impunc-

tate, non-granulate; a few small scattered setose punctures present near

anterior margin; surface of pronotum of male minors largely granulate. Pro-

notum either side of horn slightly concave (male majors), margin nearly

straight between anterior and median angles; pronotal midline just anterior to

posterior margin not modified, impunctate, slightly indented. Fore tibia with

six teeth on outer margin. Genitalia as in fig. 47, the apical protrusions of

the parameres small, acute.

FEMALES. Readily confused as males of a different but related species

unless genitalia examined. Clypeus (Fig. 48) with single anterior, moderately

elongate (about 2 mm) horn; anterior base of horn usually arising slightly

behind anterior, transversely cariniform edge of clypeus (in males, base of

horn anteriorly forms edge of clypeus); horn with both anterior and posterior

lateral carinae, but lacking posterior median carina; posterior lateral carina

on each side forming distinct tubercle above antennal insertion. Frons and

vertex relatively evenly setose-granulate. Pronotum (Fig. 48) in anterior

median half with a distinct oval concavity, width approximately equal to width

of vertex; a low horn present posterior to and contiguous with concavity, oval

horn almost truncate apically except for distinct vertical tooth on each side of

anterior edge and vague median tubercle at posterior edge. Pronotum pos-

terior to horn with midline narrowly depressed, an acute, posteriorly directed

denticle present on each side of depression just anterior to posterior margin

(this character readily identifies the females of tuberifer). Pronotum on

either side of horn with one short, distinct and one obsolete carina; pronotal

surface largely granulate except for smooth area on each side of central de-

pression. Fore tibia with six teeth on outer margin.

MATERIAL EXAMINED: 9 males, 7 females.

ARGENTINA: Misiones, Loreto; Misiones, Puerto Victoria.

BRAZIL: Pernambuco (type locality) . Terr. do Acre, Cruzeiro do

Sul, i. 1954. Bahia. Terr. Guaporé, Pérto Velho, III.1944, A. Parko.

Para, Cachimbo, 25.IX-X.1956, Travassos, Oliveira, and Adao. Paranda,

Rolandia, I1.1955, A. Maller. Rio de Janeiro. Sao Paulo, Campinas, XI.

1955, Martinez, at light; Sao Paulo, Lusanvira, 1936, H. Zellibor; Sao Paulo,

Pirassununga, 1.1951, Schubart; SAo Paulo, Anhembi, Faz, Barreiro Rico,

XI.1965, W.D. Edmonds.

Specimens are in: Dresden (?), London, Paris, Howden, Martinez.

REMARKS. In the original description Felsche (1909) did not state the

Howden and Martinez: The Genus Athyreus 41

sex of the unique type, but from his drawing and description the type is a

female. The single horn on the head of the female, normally a male charac-

ter, has subsequently caused confusion, a female of another species of the

bifurcatus group being described as the female of tuberifer (see Martinez,

1953). The species, once the sexual differences are recognized, can be

distinguished by the central, oval, anteriorly bifurcate or bidentate and pos-

teriorly, medially obtusely tuberculate horn (Figs. 46 and 48).

21. Athyreus juanae n. sp.

Figures 61, 62, Map 6

DIAGNOSIS. Pronotal horn of males very distinctive; laterally com-

pressed, extending medially from center of pronotum to posterior fourth;

horn viewed from posterior or anterior acutely elongate, viewed laterally

consisting of anterior and posterior points, apical edge arcuately emarginate

between, deepest point slightly anterior in position; pronotum medially between

horn and anterior margin feebly concave, concave area and adjacent lateral

area on each side not setose-granulate, lateral areas with a few scattered

small punctures. Fore tibia with five teeth on outer margin. Females not

associated.

HOLOTYPE. Male, length 22.5 mm, greatest width12.5 mm. Dorsal-

ly very dark brown to black, lateral edges of pronotum and elytra dark brown.

Clypeal horn (Fig. 61) nearly vertical, inclined slightly forward; anterior

lateral carina on each side only evident from outer angle to base of horn; pos-

terior lateral carina on each side distinct to near apex of horn, feebly arcuate

from base to terminus with gena; posterior median carina distinct from base

of horn nearly to apex, not tuberculate at terminus near middle of frons.

Frons and vertex largely punctate or granulate, surface smooth mainly in

transverse depression approximating junction of frons and vertex. Pronotum

(Fig. 61) with laterally compressed horn extending along midline from center

to posterior fourth, horn highest posteriorly, slightly less so anteriorly, the

edge between arcuately depressed, deepest at anterior third; apical two-

thirds of horn punctate. Area posterior and anterior to horn smooth, non-

granulate; anterior area shallowly concave, a smooth area on either side of

concavity distinctly delimited by lateral setose-granular area extending from

near median angle upward to within 1 mm of base of horn. Side of pronotum

straight to slightly arcuate anteriorly. Fore tibia with five teeth along outer

margin, the seemingly closely related tuberifer and bifurcatus both having

six teeth. Genitalia (Fig. 62) very similar to tuberifer.

FEMALES. Not associated.

TYPE MATERIAL: Holotype, male, Brazil, Guaporé, Pérto Velho,

I1.1945, A. Parko (Martinez). Paratype, 1 male, same data as holotype

(Howden) .

REMARKS. In the bifurcatus group females having five teeth on the

fore tibia have at least two distinct types of pronotal carinae. Since none of

these females were collected near the type locality no attempt was made to

associate one of the forms with juanae.

This species is named for Sefiora Juana de Martinez.

42, Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

22. Athyreus bellator Westwood

Figures 49-51, Map 6

Athyreus bellator Westwood, 1848, p. 387; 1851, p. 461.

TYPE. Male (Oxford), examined.

| MALES. Length17.0to 22.0 mm, greatest width 10.5 to12.0 mm.

Dorsally tan to very dark brown, lateral pronotal margins sometimes lighter.

Clypeal horn (Fig. 49) moderately inclined anteriorly, carinate near anterior

angles; on each side posterior lateral carina extending along length of horn;

posterior median carina extending along length of horn and sometimes onto

frons. Frons and vertex, except laterally, largely impunctate; vertex

shallowly concave. Pronotum (Fig. 49) with slender, upright central horn

laterally bifurcate at apex, the bifurcation V-shaped, anterior midline of horn

and pronotum shallowly impressed; pronotum medially posterior to horn with

cluster of setae. Pronotal margin anteriorly between marginal fossae upward-

ly arcuate; pronotal surface laterally setose-punctate, punctures closer pos-

teriorly, surface usually impunctate anterior to horn. Fore tibia with five

teeth on outer margin. Genitalia (Fig. 50) with prolongation of parameres

relatively short, slightly twisted near apices.

FEMALES. Length 20.0 to 22.0 mm, greatest width 11.0 to12.5 mm.

Pronotum (Fig. 51) with central swelling low, the encompassing carinae often

obsolete anteriorly; setose granules near anterior angles separated by approx-

imately 3 to 4 diameters, near posterior angles close, separated by approx-

imately one diameter. Sometimes not separable from females of tridens or

females in the bifurcatus group.

MATERIAL EXAMINED: 18 males, 8 females.

BRAZIL: No other data (type locality). Amazonas, Maues, H. Zellibor.

Para, Obidos, IX.1955, Martinez; I11.1956, I-III1.1959, F. Oliveira.

FRENCH GUIANA: Cayenne (=Cayenna, type locality); St. Jean du

Maroni, VIII, Moult; St. Laurent du Maroni, V, Moult.

SURINAM: Zanderij, 26.XII.1961, J. Belle.

TRINIDAD: No other data.

Specimens are in: Cambridge, Leiden, London, Oxford, Paris,

Washington, Howden, Martinez.

REMARKS. In all males seen including some relatively "minor'' forms

the pronotal horn is distinctly bifurcate, although the width and degree of

bifurcation varies. In small males most of the pronotum is at least sparsely

punctate. The sparsely placed anterior pronotal punctures (males) or gran-

ules (females) and the closely spaced posterior punctures or granules also

seem characteristic but this, at present, is difficult to assess. Some

females that we consider to be bellator have been identified as 'billbergi Gray"

(London), but the latter name could be referred to any female in the

bifurcatus group.

23. Athyreus tridens Laporte

Figures 55-57, Map 7

Athyreus tridens Laporte, 1840, p. 102; Klug, 1848, p. 25; Westwood, 1851,

p. 457; Boucomont, 1928, p. 207; Martinez, 1953, p. 232.

Athyreus similis Felsche, 1909, p. 760. NEW SYNONYMY.

Athyreus villosus Boucomont, 1902a, p. 184; 1902b, p. 581; 1928, p. 207.

Howden and Martinez: The Genus Athyreus 43

TYPE, tridens. Male (Paris), not examined.

TYPE, similis. Male (Dresden ?), not examined.

TYPE, villosus. Male (Paris), examined.

MALES. Length 16.0 to 22.0 mm; color tan to brown with the center

of pronotum sometimes darker. In well developed males clypeal horn (Fig.

55) either vertical or slightly inclined anteriorly; anterior of horn usually

lacking distinct carinae; posterior lateral carinae present; sometimes a

median, posterior carina present; frons smooth, not punctate or granulate;

vertex shallowly concave, setose-punctate posteriorly. Pronotum (Fig. 55)

centrally with erect, nearly cylindrical (flattened on anterior surface) horn,

apex distinctly tridentate, an acute tooth present on either side anteriorly,

and with single, prominent median posterior tooth; pronotal surface anterior

and posterior to horn minutely setose-punctate (Brazil) or not setose and

impunctate (usual for Argentina, Bolivia, Paraguay); pronotum anteriorly with

small median depression midway between horn and margin; lateral pronotal

margin slightly sinuate near median angle. Male genitalia as in fig. 56, with

apical prolongation of each paramere abruptly bent near apex.

FEMALES. Apparently quite variable and only placed by association

with males taken in same general area. Head and pronotum as in fig. 57;

pronotum setose-granulate instead of setose-punctate as in males; median

swelling feebly V-shaped to distinctly bituberculate; pronotum anteriorly

briefly impunctate; carina on either side poorly developed, arcuate to straight

posteriorly.

MATERIAL EXAMINED. 11 males, 5 females.

ARGENTINA: Corrientes, Alto Paranda, Isla Apipé Grande, XI.1945,

Martinez. Misiones, Concepcion, Sta. Maria.

BOLIVIA: Santa Cruz, Ichilo, Buena Vista, Martinez; Pcia. Sara, XI.

BRAZIL: (Type locality).Parand, Arapoti, XI, Miiller. Goias, Jatahy;

Goias, Monjolinho, Barreto; Goias, Rio Verde, 7.X1.1945, Zellibor. Minas

Gerais, Sertaé de Diamantina, 10. XI.1902, Gounelle. Rio Grande do Norte,

Natal, 2.VIII.1950, 28.V111.1951, Alvarenga.

PARAGUAY: Dept. del Guayra, Col. Independencia, XII. 1950,

Martinez. Asuncion, San Lorenzo.

Specimens are in: Buenos Aires, Cambridge, Paris, Howden,

Martinez.

REMARKS. All males examined had the central pronotal horn at least

feebly tridentate. The only closely related species with a tridentate pronotal

horn is A. championi. In championi, which is allopatric (Map 7), the color

is normally black, the clypeal horn of the males is anteriorly carinate basally

and the pronotum is almost entirely punctate, only the anterior face of the

horn sometimes impunctate.

24, Athyreus championi Bates

Figures 58-60, Map 7

Athyreus championi Bates, 1887, p. 108; Boucomont, 1928, p. 207; Howden,

L964, p. 1431974, p. 567,

TYPE. Male (London), examined.

MALES. Length17.0 to 20.0 mm; greatest width 10.0 to11.0 mm.

Color dorsally very dark brown to dull black, often with faint purplish tinge.

Head (Fig. 58) with erect clypeal horn; anterior edge of horn arising from

anterior clypeal margin; horn triangular in shape, sharp in front and near

anterior base bifurcating into two carinae; carinae extending to proximate

44. Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

anterior angles; posterior flattened surface of clypeal horn faintly carinate;

basally, on each side of horn, a carina extending to gena. Surface of frons

largely impunctate, with only scattered punctures posteriorly and laterally.

Vertex nearly flat and setose-punctate. Disc of pronotum (Fig. 58) with

conspicuous, central, trifurcate horn; posterior prong of horn much higher

than two anterior ones. Surface of pronotum finely setose-punctate, granulate;

area anterior to horn sometimes impunctate. Carina on either side of central

horn indistinct. Lateral pronotal margin in front of median angle slightly

sinuate. Genitalia as in fig. 59.

FEMALES. Length 20.0 to 22.0 mm; greatest width 11.5to12.5 mm.

Clypeus anteriorly with sharp, arcuately raised, transverse carina; clypeal

surface in front of carina largely impunctate. Clypeus posteriorly with

tridentate carina, middle tubercle more anteriorly placed than lateral ones,

normally more elongated. Pronotum with central, shining, shallowly bifurcate

protrusion (Fig. 60); on either side of central protrusion and joining anterior

pronotal margin, a distinct carina forming an inverted ''U" around median

projection; distinct oval impression present medially, just behind anterior

margin; midline shallowly, distinctly impressed near posterior margin; on

each side posteriorly, between U-shaped carina and posterior lateral pronotal

margin, an indistinct, short, slightly elevated carina occupying the same

position as in males. Other characters as described for males.

MATERIAL EXAMINED. 8 males, 6 females.

COSTA RICA: Cartago, Monte Redondo, 15. VI.1893; Turrialba, K.

Cooper. San José, 1000-1200m., F. Nevermann, at light; San Vito, VI.1967,

Robertson.

ECUADOR: Pichincha, Sto. Domingo de los Colorados, I.1976,

Venedictoff.

PANAMA: Canal Zone, Barro Colorado Is., VII.1938, C.E. Williams.

Chiriqui, Volcan de Chiriqui, 3000-4000 ft., Champion (Type); Mte. Lirio

(nr. Sta. Clara), 4000 ft., 5.VI.1977, R. Hartmann; Boquete, 3.I1X.1944,

G.B. Fairchild.

Specimens are in: Cambridge, Chicago, London, Ottawa, Washington,

Howden, Martinez.

REMARKS. Athyreus championi seemingly is allopatric to all other

species in the genus, being the only species so far collected in the wet lowland

forests of northwest South America and in Panama and Costa Rica. Even if

this is incorrect, championi males can be confused only with males of tridens.

Characters separating these species are given under tridens, but the nearly

black color of championi alone would serve to distinguish it.

25. Athyreus acuticornis n. sp.

Figures 52-54, Map 7

DIAGNOSIS. Males with central pronotal horn vertical, tapering to

slender, acute apex; base of horn, at least anteriorly, impunctate; posterior

to horn, a shallow depression present, midway between base and pronotal

margin, the depression with distinct, closely clustered group of setae.

Clypeal horn nearly vertical, with distinct posterior median carina in addition

to anterior and lateral carinae. Females difficult to separate from related ,

forms; pronotum as in fig. 54, with the carina beside the median swelling _

posteriorly sinuate; the lateral margin posterior to median angle arcuate,

curving distinctly inward anterior to elytral humerus.

Howden and Martinez: The Genus Athyreus 45

HOLOTYPE. Male, length 23.0 mm, greatest width13.5 mm. Color

reddish brown; sides of pronotum, labrum, anterior and lateral portion of

clypeus reddish tan. Labrum with median length of anterior vertical face

equal to median horizontal length of dorsal portion of labrum. Clypeus (Fig.

52) with horn vertical, on each side a carina extending from anterior lateral

clypeal angle onto anterior face of horn; posterior lateral carina extending on

each side from base of horn to tubercle above mandibular insertion; median

posterior carina extending from frons almost to vertex of horn. Frons im-

punctate except sparsely so laterally and on basal portions of horn between

carinae. Vertex shallowly concave, median two-thirds impunctate, laterally

with scattered setose punctures or granules. Pronotum (Fig. 52) with anter-

ior margin arcuately raised and thickened between fossae, lateral margins

anteriorly arcuate until slightly inwardly arcuate immediately anterior to

rounded median angle; posteriorly arcuate until distinctly bowed inwardly

anterior to humerus. Pronotum centrally with horn vertical, tapering to

acutely pointed apex; horn anteriorly flattened and broader than posteriorly,

both surfaces with a fine median indentation extending almost to apex; anter-

ior base of horn smooth, remainder of surface setose-granulate, granules

separated by two to three diameters. Pronotum laterally with short carina

parallel to posterior margin present just anterior to humerus, length of carina

approximately equal to that of umbone; lateral fovea with vague carina at

anterior margin. Fore tibia with five teeth on outer margin. Genitalia (Fig.

53) with narrow apical prolongation of each paramere unequally bifurcate.

ALLOTYPE. Female, length 23.5 mm, greatest width 13.6 mm.

Head (Fig. 54) with anterior transverse clypeal carina relatively evenly

arcuate; posterior clypeal carina distinctly trituberculate, the median tubercle

larger and carinate posteriorly; frons and vertex setose-granulate, the gran-

ules separated by two to three diameters. Pronotum (Fig. 54) centrally with

median swelling briefly V-shaped, bituberculate; anterior face of swelling

shining, with scattered punctures, not granulate; median swelling on each

side with carina extending posteriorly almost to margin, carina slightly

sinuate; near posterior margin midline briefly, sharply indented; lateral

carinae, foveae, lateral granulate surface, and elytra similar to male.

TYPE MATERIAL: Holotype, male, Brazil, Villa Braga, XII, 1919

(Howden). Allotype, female, Brazil, Pernambuco, Tapera, VII. 27

(Martinez).

Paratypes; 6 males, 3 females.

BRAZIL: Rio Grande do Norte, Natal, 28.VIII.1951, M. Alvarenga.

Maranhao, Barra do Corda, IV.1957, O. Rego. Para, Belém, XII.1951,

Martinez. Paraiba, 4.V.1953, J. Pessoa.

Paratypes are in: Howden, Martinez.

REMARKS. Variation is only moderate when other species in the genus

are considered. Size varies from 18 to 24 mm and color varies from speci-

mens similar to the holotype to ones uniformly reddish brown to very dark

brown to black. Morphological variation is mainly evident in the development

of the male horn and pronotal horn which may be reduced to large acute

tubercles approximately 1to 2 mm high. In large males a small, distinct

pit is present on the pronotum approximately midway between horn and anter-

ior pronotal margin. Other characters are quite consistent. The slender,

vertical, acutely pointed pronotal horn of the male normally is sufficient for

identification.

4.6 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

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A = 23. tridens Lap. * = 26. tribuliformis Fels.

@ = 24. championi Bates @ = 27. zischkai Mart.

@ = 25. acuticornis n. sp. @ = 28. martinezi Howd.

26. Athyreus tribuliformis Felsche

Figures 64-66, Map 8

Athyreus tribuliformis Felsche, 1909, p. 760; Howden, 1955, p. 671.

TYPE. Male (Dresden ?), not examined.

MALES. Length17.0to 21.0 mm, greatest width 11.0to12.0mm.

Dorsally dark reddish brown. Clypeal horn (Fig. 64) nearly vertical, anter-

ior lateral carina on each side extending from anterior clypeal angle to

median basal third or half of horn anteriorly; posterior lateral carina on each

side extending from apex of horn to junction with gena; posterior median

carina absent. Frons and vertex smooth medially, laterally setose-punctate.

Pronotum (Fig. 64) anteriorly with a large, deep circular fovea on each side

behind eye; fovea and convex area between impunctate, smooth. An upright

horn present posterior to each fovea; pronotum posterior to horns flat to

convex, median horn lacking; median area near posterior margin smooth,

area surrounded by setose-granulate area extending along posterior and

lateral margins. Single short pronotal carina present on each side anterior

to humeral area. Lateral pronotal margin sinuate near median angle; normally

shallow fovea just above median angle small, represented by a single large,

Howden and Martinez: The Genus Athyreus A7

deeply indented, circular puncture. Fore tibia with five or six teeth on outer

margin. Genitalia as in fig. 65.

FEMALES. Length17.0to 20.0 mm, greatest width 11.0 to12.0 mm.

Head and pronotum (Fig. 66) with all carinae relatively low, not greatly

elevated. Pronotum with central swelling low, shining, vaguely V-shaped;

encompassed by an arcuate carina on each side extending from anterior near-

ly to posterior margin; two short carinae present on each side; lateral pro-

notal fovea above median angle same diameter as anterior marginal fossa,

deeply indented. Fore tibia with five teeth on outer margin, a vague sixth

tooth sometimes present basally.

MATERIAL EXAMINED: 3 males, 3 females.

PERU: Iquitos, Rio Cachiyacu (type locality); Pucallpa, 250 m., 16.

VII.1961, 25.1X.1963, 22.X%.1962, J. Schunke L. (Some specimens "at

light").

Specimens are in: Washington, Hardy, Howden, Martinez.

REMARKS. The upright clypeal horn and the two well separated pro-

notal horns with the deep fovea anterior to each pronotal horn immediately

identify males of tribuliformis. If consistent, the unusually deep circular,

small fovea above the lateral median marginal pronotal angle may distinguish

females of tribuliformis from those of related species. Also, generally,

tribuliformis is slightly smaller in average size than is martinezi, zischkai,

and other species in the bifurcatus group.

27. Athyreus zischkai Martinez

Figures 67-69, Map 8

Athyreus zischkai Martinez, 1953, p. 227; Howden, 1955, p. 671.

TYPE. Male (Martinez), examined.

MALES. Length 18.0 to 26.0 mm, greatest width 11.0 to12.5 mm.

Dorsally very dark brown to black, sides of pronotum often brown. Clypeal

horn (Fig. 67) nearly vertical, lacking anterior carinae; posterior lateral

carina on each side sometimes present (lacking in holotype); posterior median

carina absent. Frons entirely or centrally smooth, vertex largely setose-

punctate; flat to shallowly concave. Pronotum (Fig. 67) with three horns,

one on either side at center of disc behind eye inclined anteriorly and sep-

arated by distance equal to distance between eyes; median posterior horn

inclined posteriorly, often bent posteriorly, paralleling surface (holotype),

sometimes laterally compressed. On each side of horns one (major males)

or two (minor males) short carinae present as is normal for genus. Anterior

to lateral horns on each side a deep circular or oval fovea separated from

opposite fovea by one diameter, area between foveae convex, impunctate;

male minors may lack fovea, if so then median surface granulate. Lateral

pronotal margin slightly sinuate near median angle. Fore tibia with six

teeth on outer margin. Genitalia as in fig. 68, prolongations of parameres

evenly curved and narrowed.

FEMALES. Length 20.0 to 27.0 mm, greatest width 11.0 to13.0 mm.

Similar to other females in tribuliformis group but pronotum (Fig. 69) with

central swelling often more distinctly elevated and more narrowly divided,

sometimes with acute apex. Fore tibia with six teeth (related females usually

have five).

MATERIAL EXAMINED: 8 males, 5 females.

BOLIVIA: Cochabamba, Pcia. de Chapare, Villa Gral. Roman, 400 m.,

18 .IX.1952, R. Zischka (type locality). Santa Cruz, Pcia. Sara, Nueva

4.8 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Moka, XII.1960, Martinez; Reys, X.1921, Mulford Exp.

BRAZIL: Amazonas, Moura, VIII.1943, A. Parko.

PERU: No other data. Chanchamayo, 15.V.19%73, J. Schunke L.

Loreto, Pucallpa, 6.1V.1973, X1I.1969, J. Schunke L.; Loreto, Ucayali

Riv., Yarina Cocha, 26.X1.1956, P. Hocking.

Specimens are in: Chicago, Paris, Washington, Howden, Martinez.

REMARKS. The nearly vertical clypeal horn and the three horns on the

pronotum will separate males of zischkai from all known species except

martinezi. From the latter the more widely spaced foveae and lateral pro-

notal horns will distinguish males of zischkai. Unworn specimens of zischkai,

both male and female, have six teeth on the fore tibia and this normally dis-

tinguishes them from the related martinezi.

28. Athyreus martinezi Howden

Figures 70-72, Map 8

Athyreus martinezi Howden, 1955, p. 667.

TYPE. Male (St. Mus. f. Naturkunde, Stuttgart), examined.

MALES. Length 20.0 to 23.0 mm, greatest width 12.0 to13.0 mm.

Chestnut brown to largely black dorsally. Clypeal horn (Fig. 70) vertical,

anterior carina on each side extending from anterior-—lateral angle of clypeus

to center of horn; posterior lateral carina on each side extending down horn

and arcuately from base to anterior edge of gena; posterior median carina

absent. Frons centrally smooth, remainder of frons and vertex punctate or

granulate, setose. Pronotum (Fig. 70) with anterior margin strongly arched,

forming small median point. A deep fovea present behind margin on either

side of midline, diameter of fovea about 2 mm, interior shining. Foveae

separated by less than one fovea diameter. Behind each fovea at summit of

pronotum on either side a nearly vertical horn; width between horns less than

distance between eyes; posterior to upright horns a median posteriorly

inclined, laterally compressed horn projecting over posterior pronotal mar-

gin. Two short carinae present on each side laterad to upright horn, the

carinae anteriorly divergent, inner carina less conspicuous. Surface of pro-

notum setose punctate-granulate, each puncture with upright seta, punctures

laterally separated by 2 to 3 diameters, medially by 3 to 4 diameters, punc-

tures obsolete in foveae and near apices of horns. Genitalia (Fig. 71) with

apical prolongation abruptly bent and widened at middle.

FEMALES. Length 22.0 to 25.0 mm. Very similar to females of

tribuliformis differing slightly in shape of central V-shaped swelling in

middle of pronotum (Fig. 72). Fore tibia with five teeth (both sexes) on outer

margin.

MATERIAL EXAMINED: 3 males, 4 females.

ECUADOR: Loja, Abbé Gaujon. ,

PERU: No other data (type locality). Junin, Chanchamayo, 1200-1500

m., 4.11]. 1976, 15.V.1973, 10 .X11.1968, J. Schunke L.; Loreto, Pucallpa,

200 m., 22.X.1960, J. Schunke L.

Specimens are in: Paris, Stuttgart, Frey, Howden, Martinez.

REMARKS. The narrowly separated lateral horns and foveae on the

male pronotum of martinezi and the five teeth on the fore tibia distinguish

it from zischkai. The presence of a posterior median horn immediately

separates it from tribuliformis. However, in the case of small males and

worn specimens the genitalia should be examined. Females in some cases

cannot be identified with certainty, being very similar to other females in the

bifurcatus group.

Howden and Martinez: The Genus Athyreus 49

29. Athyreus hastifer Felsche

Figures 73-75, Map 9

Athyreus hastifer Felsche, 1909, p. 759; Bruch, 1914, p. 541.

TYPE. Male (Dresden ?), not examined.

MALES. Length16.5to18.0mm, greatest width 12.0to13.0mm.

Color entirely black with bluish caste (original description) or with entire

body generally reddish brown except elytra black, with strong, shiny, steel-

like luster. Setae atypical for genus, reddish in color, very dense ventrally,

unusually sparse on elytra with single long upright setae arising from scatter-

ed strial punctures; entire suture of elytra with dense upright band of reddish

setae. Clypeal horn (Fig. 73) strongly inclined anteriorly and upwardly

arcuate, wider than long in cross section, shorter in height than length of

clypeus, apex rounded; a single lateral carina feebly indicated on each side

or obsolete; clypeus posteriorly with trituberculate carina similar to that of

females but with tubercles three times as large, median tubercle slightly

anterior to lateral ones. Vertex with minute tubercle near anterior third on

either side of midline, vertex posterior to tubercles smooth, vertex (laterally

and anteriorly) and frons granulate. Pronotum (Fig. 73) with anterior mar-

gin only slightly widened medially, anterior angles very broadly rounded,

flared obliquely posteriorly so that median angle only slightly posterior to

anterior angle; pronotum centrally with anteriorly directed, tapered, laterally

compressed horn with acute apex; apex of horn overhanging base of frons;

pronotal ridge above large transverse concavity extending 5/6 of distance

between lateral margins; concavity with four depressed areas within; one

either side of thin, slightly elevated midline, a larger, oval one near each

margin; interior of concavity mostly smooth, shining. Pronotum posterior

to horn with midline deeply, narrowly impressed, prescutellar region un-

modified; pronotum each side with three carinae, anterior one arising either

side of midline,then arcuate laterally extending along dorsal posterior edge

of concavity, sinuous laterally then flared and anteriorly arcuate at junction

with margin; second (middle) carina short, arcuate, anterior to humerus and

umbone; posterior marginal carina arising anterior to humerus and termina-

ting laterally at edge of fovea; dorsal surface of horn and surface between

carinae granulate, mostly setose. In less well developed males transverse

concavity greatly reduced, often forming four feebly concave areas, horn

reduced to a large acute tubercle with an anterior branch of anterior carina

extending forward on each side, becoming obsolete on transverse line with

apex of horn. Fore tibia with five teeth on outer margin. Genitalia as in

fig, 74,

FEMALES. Length16.0to17.0mm, greatest width 11.5to12.0 mm.

Anterior clypeal carina obtusely angulate, obsoletely tuberculate medially;

posterior clypeal carina typically trituberculate; frons and vertex as described

for male. Pronotum (Fig. 75) with sides not flared as in male, pronotum

similar to male minor except acute central tubercle (horn) replaced by ele-

vated transverse carina that merges on each side with anterior branch of

anterior transverse carina; other lateral carinae as described for male;

surface between carinae strongly granulate.

MATERIAL EXAMINED: 5 males, 2 females.

ARGENTINA: Misionés, Loreto, 1.1955, F.H. Walz ; Misiones,

Pindapoy, X1.1945, Martinez ; Misionés, San Pedro, 1.1956; Misionés,no

other data, Bonpland.

PARAGUAY: Itapua, Cantera, 20 m NE Encarnacion, X1.1956.

50 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

BRAZIL: Rio Grande do Sul, Porto Allegro (Type locality from

Felsche).

Specimens are in: Buenos Aires, Dresden (?), Ottawa, Howden,

Martinez.

REMARKS. The dense band of long setae along the elytral suture will

separate hastifer from all other species of Athyreus except chalybeatus which

has a somewhat similar line of setae. Size, color, and the presence of only

five teeth on the fore tibia will separate hastifer from chalybeatus; the distri-

butional patterns are also different.

@ = 29. hastifer Fels.

@ = 30. chalybeatus Fairm.

* =31. conspicuus n. sp.

30. Athyreus chalybeatus Fairmaire

Figures 76-78, Map 9

Athyreus chalybeatus Fairmaire, 1892, p. 242; Bruch, 1928, p. 1.

TYPE. Female (Paris), examined.

MALES. Length15.5to17.0 mm, greatest width 9.0 to10.5 mm.

Dorsally black with faint aeneous or greenish blue luster; ventral setae and

those in elytral suture conspicuous, very dense, yellowish white to tan (in

specimens not collected in fluid). Clypeus (Fig. 76) of male majors with

short stout horn; anteriorly with lateral carina each side extending to base of

horn but not to clypeal margin; other carinae absent, posterior base of horn

with distinct contiguous tubercle; laterally above antennal insertion lateral

carina obtusely angled. Frons and vertex mostly granulate, vertex anteriorly

with minute tubercle on each side of feebly depressed, smooth midline. Pro-

notum (Fig. 76) with anterior margin between small fossae slightly thickened

Howden and Martinez: The Genus Athyreus 51

but not elevated or tuberculate; anterior angles obtusely rounded; lateral and

posterior margins sinuate. Pronotum centrally with acute, anteriorly directed,

laterally compressed horn; contiguous to horn on each side in male majors a

deep, oblique, oval concavity, interior of each concavity smooth and polished.

Posterior to concavity on each side three carinae present nearly paralleling

posterior margin. Anterior of three carinae extending from side of deeply

indented midline posterior to horn then above (posterior to) concavity becom-

ing obsolete and sinuous anterior to shining, shallow lateral fovea, then dis-

tinct, raised abruptly, arcuate anteriorly to overhang, then join lateral mar-

gin; second (middle) of three carinae short, feebly arcuate, anterior to hum-

erus in position; third (posterior) carina close to and parallel with posterior

pronotal margin, extending from just below humerus to 0.5 mm from lateral

margin. Surface of pronotum, except for smooth indentation, concavities and

carinae granulate, inconspicuously setose. Elytral intervals shining, striae

punctate. Fore tibia with six teeth on outer margin. Genitalia as in fig. 77.

FEMALES. Length16.0to17.0 mm, greatest width 11.0 to11.5 mm.

Anterior face of clypeus delimited dorsally by obtusely triangular carina,

feebly tuberculate medially, a longitudinal carina extending posteriorly to

strongly developed median tubercle of posterior clypeal carina; frons and

vertex as described for males. Pronotum (Fig. 78) laterally similar to males,

differing centrally; at anterior third a strongly raised transverse carina

extending a distance equal to width of vertex, terminus on each side with

carina extending both anteriorly and posteriorly; anterior branch becoming

obsolete in 0.5 mm, posterior branch merging with most anterior of the

three oblique carinae (as described in males). Midline posterior to trans-

verse carina narrowly, distinctly indented, surface on each side rising to

carina. Pronotal surface granulate except for midline, particularly anterior

to transverse carina, and near lateral foveae.

MATERIAL EXAMINED: 5 males, 4 females.

ARGENTINA: ''Plata orientale" (type locality); Buenos Aires, General

Madariaga, San Clemente del Tuyu, XII.1952, II1.1957, Kormilev; Buenos

Aires, General Lavalle, Punta Norte, I.1936; Buenos Aires, Playa Juancho.

URUGUAY: Montevideo, XII.1946, Ruffinelli, Malvin.

Specimens are in: Buenos Aires, Paris, Howden, Martinez.

REMARKS. The black dorsal color, with faint greenish blue luster and

tan ventral surface, odd clypeal armatures of males and females (see descrip-

tion), unmodified anterior pronotal margin and three lateral carinae on each

side of pronotum should serve to identify chalybeatus. Male minors lack the

deep pronotal concavities and central horn but the characters mentioned as

useful for identification seem reasonably consistent.

31. Athyreus conspicuus n. sp.

Figure 63, Map 9

DIAGNOSIS. Color reddish brown to brown. Clypeal characters

similar to those of hastifer. Pronotum on each side with four oblique carinae

either reaching or extending toward lateral or anterior margins; female

(males not known) with cordiform concavity in center of pronotum sharply de-

limited by surrounding carina, midline of concavity with one anterior and one

posterior fossa or small deep fovea; posterior transverse carina of pronotum

almost straight; extremely close to posterior pronotal margin from humeral

area to midline, this constricted area with distinct, erect band of setae.

HOLOTYPE. Female, length16.0 mm, greatest width10.5 mm.

52 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Color reddish brown with carinae on head, central pronotal carinae, margins

of mandibles and parts of legs brownish black. Clypeus with anterior carina

obtusely angulate, the median angulation with posterior longitudinal carina

extending to median tubercle on posterior transverse clypeal carina; poster-

ior clypeal carina nearly straight, trituberculate, central tubercle slightly

more elevated. Vertex anteriorly with a short, fine arcuate carina on each

side of midline; carina laterally anteriorly arcuate, becoming obsolete near

lateral margin. Dorsal surface of clypeus, frons and lateral portion of vertex

closely, often confluently punctate. Pronotum (Fig. 63) with anterior margin

unmodified, lateral margin angulate at middle where joined by an oblique

carina. Pronotum centrally from anterior fourth or third to posterior mar-

gin with large cordiform depression, deepest anteriorly, at bottom of depre-

ssion anteriorly a small deep fossa or fovea, midline faintly impressed pos-

teriorly, extending to second fossa at posterior edge of depression just anter-

ior to carina adjacent to posterior margin; depression widest anteriorly, ele-

vated and subdenticulate on anterior carinate margin on either side of midline,

carina then posteriorly arcuate to obtuse, inwardly directed angles on either

side near middle of depression, then converging to fuse just behind posterior

fossa. Four carinae present on either side of cordiform carina, anteriorly a

short carina arising from edge of cordiform carina and extending half distance

to anterior margin; second carina arising from just behind obtuse angulation

of cordiform carina and extending to and joining lateral margin midway

between anterior and median angles, carina sinuous in lateral half, obtusely

angled at junction with margin; third carina (adjacent to posterior carina)

relatively short, feebly arcuate, relative position anterior to humeral umbone;

fourth or posterior carina long and atypically nearly straight, extending from

posterior midline to faint lateral fovea near lateral margin; area between

carina and posterior pronotal margin and anterior to base of elytron with

dense band of erect setae. Surface of pronotum except for carina punctate

(in central depression) or granulate, setose. Each elytron with six very con-

vex intervals between suture and umbone. Fore tibia with five teeth on outer

margin.

MALE. Unknown.

TYPE MATERIAL: Holotype, female, Argentina, Santiago del Estero,

Rio Salado, Icafio (Martinez).

Paratypes: 5 females.

ARGENTINA: one, same data as holotype; Santiago del Estero, Afiatuya,

1.1944. La Rioja, Liar. Tucuman.

Paratypes are in: Buenos Aires, Tucuman (Instituto Miguel Lillo),

Howden, Martinez.

REMARKS. Structural variation occurs primarily in the degree of

development of the fine carinae and denticles on the frons and vertex, which

in one case is large, arcuate and extends to the margin at the mandibular

insertion. Some minor variation occurs in the pronotal carinae,and size

varies from 12.5 to 16.0 mm in length.

The reddish brown color, lack of a dense band of long, upright setae

along the elytral suture, and the deep central depression of the pronotum

will identify female conspicuus. The species is undoubtedly related to (

hastifer and chalybeatus and males will probably show similar sexual modifi-

cations.

Howden and Martinez: The Genus Athyreus 53

Incertae sedis

(Species known only from females not associated with males.)

Athyreus billbergi Gray, 1832, p. 508; Westwood, 1851, p. 462.

Athyreus subarmatus Westwood, 1848, p. 386; 1851, p. 460.

Athyreus vicinus Laporte, 1840, p. 102.

List of species, many described in Athyreus,

assigned to Neoathyreus Howden and Martinez, 1963.

(In this and the following genus the numerous new combinations are marked

by an asterisk (*) preceding the species name.)

*angulatus (Klug), 1843.

*anthracinus (Klug), 1843.

*arribalzagai (Martinez), 1951.

*biceps (Felsche), 1909.

*bicolor (Laporte), 1840.

*bidentatus (Macleay), 1819.

*castaneus (Guerin), 1829.

*catharinae (Bates), 1887.

*centralis (Westwood), 1848.

*centromaculatus (Felsche), 1909.

*corinthius (Klug), 1843.

*corniculatus (Felsche), 1909.

*excavatus (Laporte), 1840.

fissicornis (Harold), 1880.

*flavithorax (L.-Arribalzaga), 1880.

*fossulatus (Westwood), 1851.

*fulvescens (Blanchard), 1837.

*soyasensis (Boucomont), 1902.

granulicollis Howden, 1964.

hamifer (Boucomont), 1902.

interruptus Howden, 1964.

lanei (Martinez), 1952.

*latecavatus (Boucomont), 1902.

mexicanus (Klug), 1845.

mixtus Howden, 1964.

panamensis (Robinson), 1946.

54 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

*pholas (Westwood), 1848.

_ planatus Howden, 1964.

*purpureipennis (Westwood), 1848.

quadridentatus Howden, 1964.

*reichei (Westwood), 1851.

*sexdentatus (Laporte), 1840.

tridentatus (Macleay), 1819.

tridenticeps (Bates), 1887.

*tweedianus (Westwood), 1848.

*violaceus (Klug), 1843.

*viridis (Boucomont), 1902.

*vulpinus (Harold), 1880.

List of species originally described in Athyreus,

here assigned to Parathyreus Howden and Martinez, 1963.

*bahiae (Arrow), 1913.

*fissicollis (Arrow), 1913.

trituberculatus (Klug), 1843.

ACKNOWLEDGMENTS

This study was greatly expedited by an operating grant to H. Howden

from the National Research Council of Canada. Support from this grant made

it possible for A. Martinez to visit Ottawa to complete the project. Laboratory

support was also derived from the same source, and we are both greatly

appreciative of this support. We are also indebted to the Consejo Nacional de

Investigaciones Cientificas y Técnicas, Buenos Aires, for granting to A.

Martinez a temporary transfer to Ottawa in order to complete this work.

Several individuals very kindly donated a number of specimens used in

this study. Coronel Moacir Alvarenga and Dr. Carlos Alberto Campos

Seabra, Rio de Janeiro; P. Francisco S. Pereira, S&o0 Paulo; Manuel J. Viana

and P. Gregorio J. Williner, Buenos Aires, all were of great assistance

through their energetic collecting and we acknowledge with thanks their gen-

erosity.

We would also like to thank Anne T. Howden for assistance with trans-

lation of the key into English. Mr. L.E.C. Ling, Carleton University, took

the Scanning Electron Micrographs used herein and Mrs. Betty Perry typed

the manuscript. Museum curators who assisted us are listed at the end of

the introductory section along with their institutions. We thank all of these

people and institutions for their support.

LITERATURE CITED

Bates, H.W.

1889. Pectinicornia and Lamellicornia. In Biologia Centrali-Americana,

Insecta, Coleoptera, I1(2): 1-381; Supplement: 382-432.

Howden and Martinez: The Genus Athyreus 55

Blackwelder, R.

1944. Checklist of the Coleopterous Insects of Mexico, Central

America, the West Indies, and South America. United States

National Museum, Bulletin 185(2): 189-341.

Boucomont, A.

1902a. Description sommaire de quelques Bolbocerini (Col.). Bulletin

de la Société entomologique de France, 184-185.

1902b. Description de quelques Bolbocerini nouveaux. Annales de la

Société entomologique de France 71: 581-587.

1911 (1910). Contribution 4 la classification des Geotrypidae (Col.).

Annales de la Société entomologique de France 79: 333-350.

1912. Scarabaeidae: Taurocerastinae, Geotrupinae. In Junk,

Coleopterorum Catalogus, Berlin 19 (46): 3-47.

1928. Coprophages d' Amérique du Sud nouveaux ou peu connus suite.

Bulletin de la Société entomologique de France: 202-207.

Bruch,.c..

1911. Catalogo Sistemdtico de los Coledpteros de la Reptblica

Argentina. Revista del Museo de La Plata17: 181-225.

1914. Suplemento al Catalogo Sistematico de los Coleopteros de la

Republica Argentina. Revista del Museo de La Plata19: 538-573.

1928. Athyreus chalybeatus Fairm. descripcién del macho. Revista

de la Sociedad Entomologica Argentina 2(1):1-4.

Feltsehe, <¢:

1909. Neue und alte coprophage Scarabaeiden (Col.). Deutsche

Entomologische Zeitschrift: 751-765.

Gray, “Gave

1832. In Griffith and Pidgeon. The animal kingdom arranged in

conformity with its organization by the Baron Cuvier. Vol. 14

(Insecta, vol. 1). London. 570 pp.

Guerin, J.

1956. Coledpteros do Brasil. Dept. de Zoologia, Univ. de SAo Paulo,

S40 Paulo, Brasil: 356 pp.

Howden, H.F.

1955. Description of a new Peruvian Athyreus with notes on the Method

of Illustration. Entomologischen Arbeiten aus dem Museum G.

Frey 6(2): 667-671.

1964. The Geotrupinae of North and Central America. Memoirs of the

Entomological Society of Canada 39:1-91.

1974. Additional records and descriptions of North and Central

American Geotrupinae (Coleoptera, Scarabaeidae). Canadian

Journal of Zoology 52(5): 567-578.

Howden, H.F., and A. Martinez. |

1963. The new tribe Athyreini and its included genera (Coleoptera:

Scarabaeidae, Geotrupinae). The Canadian Entomologist 95(4):

345-352.

Klug, H.

1845 (1843). Die Coleopteren-Gattungen: Athyreus und Bolboceras,

dargestellt nach den in der Sammlung hiesiger Koenig] Universitaet

davon vorhandenen Arten. Akademie der Wissenschaften Berlin

Physikalische Abhandlungen: 21-56.

Laporte, F. i.

1840. Histoire naturelle des animaux articulés. Paris, vol. 2. 540

Pp-

56 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Martinez, A.

1953. Una nueva especie de Athyreus (Col. Scarab. Geotrup.).

Mitteilungen der Muenchener Entomologische Gesellschaft e.v.

43: 227-233.

Westwood, J.O.

1848. Description of some new species of Athyreus, MacL., a genus

of lamellicorn beetles. Annals and Magazine Natural History, ser.

2, 1: 386-387.

1851. Descriptions of some new species of Athyreus, a genus of

lamellicorn beetles. Transactions Linnean Society 20: 453-467.

Howden and Martinez: The Genus Athyreus 57

Figs. 1-3. Athyreus cyanescens Klug. 1. Male, head and pronotum;

2. Male genitalia, dorsal view; 3. Female, head and pronotum. Figs. 4-6.

Athyreus capricornis n. sp. 4. Male, head and pronotum; 5. Male genitalia,

ventral view; 6. Female, head and pronotum.

Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

a — Te

Figs. 7-9. Athyreus aeneus Klug. 7. Male, head and pronotum;

8. Male genitalia, dorsal view; 9. Female, head and pronotum. Figs.

10-12. Athyreus vavini Boucomont. 10. Male, head and pronotum;

dae.

Male genitalia, dorsal view;12. Female, head and pronotum.

Howden and Martinez: The Genus Athyreus 59

Figs. 13-15. Athyreus hypocritus n. sp. 13. Male, head and pro-

notum; 14. Male genitalia, dorsal view;15. Female, head and pronotum.

Figs. 16-18. Athyreus bilobus n. sp. 16. Male, head and pronotum;

17. Male genitalia, dorsal view; 18. Female, head and pronotum.

60 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Figs. 19, 20. Athyreus brasilicus n. sp. 19. Male, head and pro-

notum; 20. Male genitalia, ventral view. Figs. 21, 22. Athyreus nitidus

n. sp. 21. Male, head and pronotum; 22. Male genitalia, oblique ventral

view. Figs. 23, 24. Athyreus anneae n. sp. 23. Male, head and pronotum;

24. Male genitalia, dorsal view.

Howden and Martinez: The Genus Athyreus 61

—

‘ ‘

— ”

Figs. 25-27. Athyreus hemisphaericus Boucomont. 25. Male, head

and pronotum; 26. Male genitalia, dorsal view; 27. Female, head and pro-

notum. Figs. 28-30. Athyreus unicornis n. sp. 28. Male, head and pro-

notum; 29. Male genitalia, dorsal view; 30. Female, head and pronotum.

62 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Figs. 31-33. Athyreus forcipatus Boucomont. 31. Male, head and

pronotum; 32. Male genitalia, dorsal view; 33. Left elytron, inner basal

portion showing striae and intervals. Figs. 34-36. Athyreus alvarengai n.

sp. 34. Male, head and pronotum; 35. Male genitalia, dorsal view;

36. Left elytron, inner basal portion showing striae and intervals.

wad

en and Martinez: The Genus Athyreus 63

: Moll

Figs. 37, 38. Athyreus pyriformis n. sp. 37. Male, head and pro-

notum; 38. Male genitalia, dorsal view. Fig. 39. Athyreus armatus

Westwood. 39. Male, head and pronotum (reproduced from 35 mm color

slide of type). Figs. 40-42. Athyreus gigas Westwood. 40. Male, head and

pronotum; 41. Male genitalia and genital capsule; 42. Male genitalia, apex

of left paramere.

64 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

Figs. 48-45. Athyreus bifurcatus Macleay. 43. Male, head and pro-

notum; 44. Male genitalia, dorsal view; 45. Female, head and pronotum.

Figs. 46-48. Athyreus tuberifer Felsche. 46. Male, head and pronotum;

47. Male genitalia, dorsal view; 48. Female, head and pronotum.

Howden and Martinez: The Genus Athyreus 65

Figs. 49-51. Athyreus bellator Westwood. 49. Male, head and pro-

notum; 50. Male genitalia, dorsal view; 51. Female, head and pronotum.

Figs. 52-54. Athyreus acuticornis n. sp. 52. Male, head and pronotum;

538. Male genitalia, dorsal view; 54. Female, head and pronotum.

66 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

iS .

Figs. 55-57. Athyreus tridens Laporte. 55. Male, head and pro-

notum; 56. Male genitalia, dorsal view; 57. Female, head and pronotum.

Figs. 58-60. Athyreus championi Bates. 58. Male, head and pronotum;

59. Male genitalia, dorsal view; 60. Female, head and pronotum.

Howden and Martinez: The Genus Athyreus 67

Figs. 61, 62. Athyreus juanae n. sp. 61. Male, head and pronotum;

62. Male genitalia, dorsal view. Fig. 63. Athyreus conspicuus n. sp.

63. Female, head and pronotum. Figs. 64-66. Athyreus tribuliformis

Felsche. 64. Male, head and pronotum; 65. Male genitalia, dorsal view;

66. Female, head and pronotum.

68 Contribs Amer int. Insts. Vols 1a, noe 4. 1978

Figs. 67-69. Athyreus zischkai Martinez. 67. Male, head and pro-

notum; 68. Male genitalia, dorsal view; 69. Female, head and pronotum.

Figs. 70-72. Athyreus martinezi Howden. 70. Male, head and pronotum;

71. Male genitalia, dorsal view; 72. Female, head and pronotum.

Howden and Martinez: The Genus Athyreus 69

» \

Figs. 73-75. Athyreus hastifer Felsche. 73. Male, head and pro-

notum; 74. Male genitalia, dorsal view; 75. Female, head and pronotum.

Figs. 76-78. Athyreus chalybeatus Fairmaire. 76. Male, head and pro-

notum; 77. Male genitalia, dorsal view; 78. Female, head and pronotum.

70 Contrib. Amer. Ent. Inst., vol. 15, no. 4, 1978

| _

a WAS

: Q ]

‘ i

Figs. 79-81. Athyreus biarmatus n. sp. 79. Male, head and pro-

notum; 80. Male genitalia, dorsal view; 81. Female, head and pronotum.

Figs. 82-84. Athyreus parvus n. sp. 82. Male, head and pronotum,

lateral view; 83. Male, head and pronotum, dorsal view; 84. Male genitalia,

dorsal view.

Contributions

of the

American Entomological Institute

Volume 15, Number 5, 1978

A REVISION OF THE NEARCTIC SPECIES OF THE GENUS

ONTHOPHILUS LEACH (COLEOPTERA: HISTERIDAE)

Jussi Helava

- ate ere h

“5 : i

a, en pe ee ee

A REVISION OF THE NEARCTIC SPECIES OF THE GENUS

ONTHOPHILUS LEACH (COLEOPTERA: HISTERIDAE)

Jussi Wel

ABSTRACT

The Nearctic species of the histerid genus Onthophilus Leach are

revised. Fourteen species are recognized, of which six are described as

new: O. deflectus, O. intermixtus, O. wenzeli, O. cynomysi, O. thomomysi

and O. giganteus.

All species are described and illustrated with scanning electron micro-

scope pictures. Drawings were made of male genitalia for species in which

males are known, of mouth parts for alternatus, and of the hind wing for

alternatus and flohri. A key and diagnoses are provided for separating

species. The distributions of species are indicated on maps. A brief discus-

sion of the distribution, ecology and taxonomic affinities and evolution is in-

cluded.

CONTENTS

PRUISOCICENGI, To Mur dase, sec wheeen tt ah MEG MR oe Lier ye Seg leek Smit ull td

a StOLECAICREVACW Le thee hts. thee tine coh, dos CAS eT ee:

[DiS Bigs oF Ui Kerr el aaa ape ele ae eee a Ka ee, oe er

Heolos va Nitra We ).w oe eee: we (ee re Oe eRe) hres

TAXOnOmMic AllINItIES ANG CVOIIPON 5 seta ono: fe codes oo Sample aed antes ues

Niethods?*..8. winreds 5 hh OA ee Sk ee, Be Sd 2m: 8,

A BROM OIA a PeeAe eke LUNewes. Rel ce isin a ewley SNe Lorc, Tole ote siceat «seabed Yontid weno.

CHICO MEMOS UENCE Rr) Cle ee iy Se hee AB

O-=- 43 cau -@ Erg: CON ory

KevaLO- NCARCHICESDECLES UF.) Jitu il | Whee Aire ae Cee eee

Species descriptions

ee LEN ey 8 Se Seer Saar ch naire OME ReneS OMS ee Aer i

LY)

2; - Gehlechis mew Species. ..°% Ge eats HA. ee GP eS

ROE MERC i i en woe he Tee eee. ee

Poe

Oo o

A= ARCCPIMIRIUSIDEW ASPOCTCS! 43 are Esa Gi @ eee oo eee ee

B: valternatiuis (Say): ~ =<: elle. deelece) pues Gb cere abies

—

ee)

nodatus Leconte «. 5 i234 lee... ae eee eee

tpepartment of Biology, Carleton University, Ottawa, Ontario, Canada.

2 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

(. PhuIicosravus Heeconte ss ts. Ge et ied ave Reh) 2 ae Behe OLY ean 4 20

Wenzeli NeW Species “4 2%... ™ Se 22

Je IGYMOMNSENEW SPECIES suis Se Bae & gs ek Ges Side Pe 6 23

ON) MSo@luatn ASE tt. Sart: oom aie eee ee ORS eel a a leerte Ge ee ang 25

eis WE COMUCTS EOIN. ve, elt te, See lenet BEARS ae! He ee a we 26

12s: ROMO SEIEW SDCCICS 2) met Joes 'snsee A & Bie ea ee Be ee 27

3. ASEPRIGROSS* i es 2 a Be crys, Ge Mie Pence, OM TE hk Seg gw 29

da (STS AMPCMIS MEW SCCICS ice Si sins 66S Mee a 8 saa ; 30

cen Gnile denen merenedten . 8 Se ets Sh, eats kerermeat let). ag BPO a PU onl

re inate Clee Oe toler Fy de Wwe coe teeta fh terelnldege Seay oe Us ee ors 32

ECGS! Sue. becaees ges bors i ele PO. lk Sey 34

IOS "hes ste cee Siege es te SEMMTUN MRO sgh Sen mat es at S. Ko a 7 slicey *s 42

INTRODUCTION

Onthophilus Leach is a small genus of the family Histeridae. The

strong sculpturing of its dorsal surface makes this a readily identifiable

genus, but because it is rarely encountered by general collectors, Onthophilus

is never represented by many specimens in most collections. The Palearctic

species of Onthophilus were revised most recently by Reichardt in 1933. The

Nearctic species have not been revised since Horn's revision of 1873, in which

he recognized four species. Since then, four more species have been described

and substantially more new material has been accumulated.

This study offers a taxonomic treatment of Nearctic Onthophilus Leach,

and consolidates existing information. Fourteen species are recognized.

Six are described as new, and previously named species are redefined. An

artificial key and diagnostic characters are provided for identifying the

Species. Brief discussions are included on the distribution, on what is known

about the habits of the species, and on the taxonomic affinities of the species.

HISTORICAL REVIEW

Leach erected the genus Onthophilus in 1817 to accommodate two pre-

viously described species, Hister striatus Forst. and H. sulcatus Fab.

Leach provided a key, based mainly on antennal characters, to separate this

genus from others that were extant at that time, but he did not include a

description. The genus was better defined by Leconte in 1845, on the basis

of three North American species, alternatus Say, nodatus Leconte and

pluricostatus Leconte. A more complete description is found in Marseul's

Monograph of 1856. Both Leconte's and Marseul's concept of the genus

Onthophilus was broad enough to include three species, hispidus Paykull,

costipennis Fahrs., and novemcostatus Marseul. These species differ from

the other species of Onthophilus in several marked ways. They are covered

Histeridae: Nearctic Onthophilus 3

dorsally by dense setae or scales which often obscure the elytral sculpture,

the prosternum is bicarinate, and they have a pronounced fovea in the front

angles of the metasternum. Thirty-two names were proposed before 1891.

At that time, Lewis erected the genus Epiechinus to accommodate seven

hispid species of Onthophilus. Reichardt's concept of the genus in 1933, com-

prises those species which were left in Onthophilus after Lewis erected

Epiechinus, and is followed here.

The first Nearctic species to be placed in Onthophilus was alternatus,

described by Say in 1825 from the Melsheimer collection. Leconte (1845) in

his revision of the North American Histeridae, added nodatus and pluricostatus.

In addition to these three species, Horn recognized a new species from

California, in 1870; in his 1873 revision of the North American species of

Histeridae. Horn considered nodatus and pluricostatus as varieties of

alternatus. Subsequently various catalogues, for example Bickhardt (1910),

have treated the Say and the Leconte species as synonyms, although Lewis

(1905) and Reichardt (1933) retained their specific status. They are recog-

nized as valid species in the present paper and types are designated for them.

Lewis described two species from Mexico, flohri and julii, in 1888 and in

1892 respectively. In 1893, Casey described a species from Colorado and

named it in honour of the collector H. Soltau. Ross (1944b) named a species

kirni from Texas in honour of A.J. Kirn with whom he was collecting arthro-

pods associated with Geomys. The numerous species added since 1856, make

Marseul's key inadequate for identifying Nearctic material.

The higher classification of the Histeridae is very unclear and is in

need of much work. Onthophilus was previously placed close to Abreus in

the subfamily Abreinae, for reasons which are difficult to explain. Wenzel

(1944) moved Onthophilus to the subfamily Tribalinae, but this genus and

other closely related genera, Peploglyptus, Glymma, Stictostix and Epiechinus,

can not be placed satisfactorily until a revision of the Histeromorphae is com-

pleted.

DISTRIBUTION

At present a total of thirty species of Onthophilus Leach have been

described. There are thirteen species, one subspecies and one variety

recognized from the Palearctic region. Seven of these are extant in Siberia,

China, Formosa and Japan, and six in western Europe, the Near East and

northern Africa. All these species are confined to either the eastern or the

western part of the region. Two species are recognized from Burma. One

was collected from 7000'; no data are provided with the other description.

One species, australis, was collected by H. F. Howden in Australia. This is

the first record of Onthophilus in that continent.

Fourteen species are recognized from the Nearctic region. Two species,

deflectus and pluricostatus, range from Michigan and New York in the north,

south along the Appalachians and the Mississippi valley to the Gulf states.

Onthophilus alternatus and nodatus are found in the coastal lowlands of the

southeastern states; nodatus has been recorded as far west as eastern Texas;

giganteus is known only from Archer, Florida. Five species occur in the

central states, each recorded from only one locality -- soltaui from Denver,

Colorado; cynomysi from Grady Co, Oklahoma; kirni from Somerset, Texas;

wenzeli from Nebraska and intermixtus from Graham Co., Arizona. One

4. Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

species, lecontei, has been collected along the coast of California, from San

Francisco to San Diego. Three species are from Mexico; thomomysi from

Durango and also Eagle, New Mexico; flohri from Durango; and julii from

Mexico and Oaxaca. All three species occur above 8000' in Mexico.

From the distribution patterns noted above, it is evident that Onthophilus

is largely confined to temperate climates. Even those species that occur

farthest south, in Mexico and Burma, are confined to high elevations.

ECOLOGY

Virtually nothing is known about the biology of Onthophilus. The only

source of biological data available is collecting records. Collecting records,

however, provide no information that would indicate whether these are the

usual habitats in which the species occur, or whether they are only occasional

visitors to these habitats.

Adult beetles of most Nearctic species of Onthophilus have been collected

in traps baited with malt, carrion or dung. O. pluricostatus has been found

under a dog carcass. O. deflectus has been recorded from detritus in tree

holes and between tree buttresses, from forest leaf litter, from dead

Polypore, from mouse nests, from chicken decay, from under a pig carcass

and from cow dung. QO. kirni, cynomysi and thomomysi are recorded from

prairie dog burrows. O. lecontei has been found in gopher burrows, ina

Vespula pennsylvanica nest, and on dandelion.

Most species of histerids are thought to be predators of fly larvae and

of other small arthropods, although there are very few literature references

that confirm this generalization by direct observation. The structure of the

mouth parts of Onthophilus suggests that the species are predators. It is not

unlikely, therefore to assume that the adults of this genus, are general

predators of fly larvae or of other small arthropods in leaf litter, in dung, in

carrion and in the nest and fecal chambers of some rodents and other small

mammals. It is their secretive habits and the relatively specialized techniques

required to collect them, that undoubtedly account for the poor representation

of the various species of Onthophilus in most collections.

No records or biological observations of the larval stages of Onthophilus

exist. Because dung and carrion disappear very rapidly in the summer, it is

likely that the larvae will be found in some more stable niche, such as rodent

burrows in the case of some species, or in leaf litter in others. The larvae

of a few other histerid genera are known to be predators on the eggs and

larvae of small arthropods. Onthophilus larvae will probably conform to this

predatory trend in the Histeridae.

TAXONOMIC AFFINITIES AND EVOLUTION

The family Histeridae is evidently an old one. Onthophilus and the

genera related to it, represent an unusual and divergent branch of this family.

One fossil species, O. intermedius Handschin (1944) establishes that this

genus dates from at least the Oligocene. It may be assumed that in the past

this genus had a much more continuous distribution. The present distribution

of the species indicates a progressive series of radiations and of subsequent

fragmentations.

Histeridae: Nearctic Onthophilus 5)

The species now extant in the Nearctic region are apparently polyphyletic

in origin. Those ten species for which males are known can be placed into

three groups on the basis of the structure of the male genitalia. These groups

may be conveniently referred to by the name of the oldest named species

within that group. The composition of the groups is as follows:

(1) lecontei group: lecontei, cynomysi, thomomysi and kirni

(2) alternatus group: alternatus, nodatus and pluricostatus

(3) flohri group: flohri, deflectus and intermixtus

The median lobe of the male aedeagus of the lecontei group, is extruded ven-

trally between the bifid apex of the parameres. The median lobe of the

alternatus group and the flohri group, is extruded dorsally. The flohri group

differs from the alternatus group in the form of the basal piece. The basal

piece of the alternatus group is long and tubular. The basal piece of the flohri

eroup is short and ovoid. The basal opening is rotated sinistrally 45 degrees

from the longitudinal midline and is subapical. These groups share external

characteristics within groups, which allows the placement, with some confi-

dence, of wenzeli and soltaui into the lecontei group. The only species which

cannot be placed with any confidence is julii. No other species of Onthophilus,

- anywhere, has such greatly interrupted elytral costae. The only other species

which approaches julii in this characteristic, is tuberculatus from Burma.

The general outline, punctation and so forth, of julii suggests that it is closer

to the flohri group than to either of the other two Nearctic groups.

All three Nearctic groups have close taxonomic affinities to species in

the eastern and/or western Palearctic region, but the exact phylogenetic

relationships are obscure. For example, the head and thorax of silvae from

Japan resemble those of alternatus very much, but its elytral costae are

equally well developed. Similarly, the pronotum of wenzeli resembles that

found in the alternatus group, but in the rest of its characters, it is very close

to the other species of the lecontei group.

The general east-west division of the Nearctic species, is probably an

expression of the Tertiary orogeny and the subsequent development of the

recent xerophytic vegetation (Axelrod, 1958). Apparently the species of the

lecontei group have survived the xeric conditions by utilizing the more amen-

able conditions found in mammal burrows and similar habitats. It is not clear,

whether this is a primitive adaptation of the group, or whether these species

invaded these niches and speciated within them after the onset of xeric condi-

tions. Certainly this mode of behaviour is not confined to the lecontei group,

but is found in other species of Onthophilus, in the Nearctic and the Palearctic

regions (Bedwell, 1907), and in other genera of both the Histerinae and

Saprininae.

A more recent distribution pattern that is the result of the Pleistocene

glacial cycles is superimposed on the older one. QO. deflectus and flohri are

two morphologically very similar species. The former is found in eastern

United States and the latter in Mexico. This distribution pattern suggests

that these species arose from an ancestral population by prolonged isolation

during a glacial cycle. Interestingly, this species separation is accompanied

by a reduction of the inner wings of flohri. The specimens of flohri that I

have seen are from the western edge of the Sierra Madre Occidental at high

elevations. Very likely, suitable habitats are sparsely and irregularly distrib-

uted in this area and are subjected to strong prevailing winds from the ocean.

Under these circumstances the reduction and the consequent loss of flight

would have high survival value and would be selected for in order to minimize

the loss of individuals. Whether this is also true for other populations of this

species, is difficult to say, until further collecting reveals the complete

6 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

range of this species. Pleistocene isolation is expressed in other species by

pockets of variability within their range. For example, specimens of nodatus

from eastern Texas have larger punctures than specimens from the eastern

portion of its range. Possibly these specimens represent a population which

was separated from the main body of the species during the last glacial period.

METHODS

This study was based on an examination of approximately 500 museum

specimens of Onthophilus.

Cleaning: Most specimens of Onthophilus are heavily covered by a waxy

exudate to which soil particles and other detritus have adhered. This accre-

tion forms a hard coating which obscures many important taxonomic charac-

ters. Many different compounds were tested in attempts to clean specimens.

The most successful of these was a 10 % (w/v) solution of sodium phosphate,

tribasic (TSP, NagPO4). This solution softened the waxy exudate sufficiently

that it could be peeled away with a camel hair brush. In addition this solution

relaxes specimens very efficiently.

Dissection Techniques: Male genitalia were extracted through the

pygidium of specimens relaxed in hot water or TSP. The genitalia were

cleared in a hot 10 % solution (w/v) of potassium hydroxide, prior to the

separation of the terminalia and the aedeagus. For further study and illus-

tration, microscope slides were prepared using a glycerin gel medium .

Mouth parts, wings and other structures were handled in a similar manner,

except that the mouth parts were stained with a 10 % solution of basic fuchsin

in 95% ethyl alcohol.

Illustrations: A net reticule was used to make all the measurements

and as an aid in making the drawings. The length was measured from the tip

of the front angles to the hind margin of the elytra. The width was measured

at the widest point. At least one specimen of each species was photographed

uncoated, using a JEOL JSM-U3 scanning electron microscope. Magnifica-

tions did not exceed 10 0X.

Terminology: Striae and other surface features are important charac-

ters for defining and identifying species of Histeridae. For the terminology

of the sclerites and striae, I have followed the useage of Wenzel and Dybas

(1941). The following is a definition of the abbreviations used in the text for

important costae and carinae that have no prior names.

Head (Fig. 21)

VFC verto-frontal carina, is located along the longitudinal midline reaching

from the occipital carina to between the antennae; may be variously

reduced.

LFC latero-frontal carina runs from the antennal pit, obliquely across the

face to, or just above the clypeus.

Pronotum (Figs. 42, 43, 44)

These costae are bilaterally symmetrical.

PC 1 pronotal costa one is the costa closest to the lateral edge.

PC 2 pronotal costa two is the costa immediately mesad of PC 1.

PC 3 pronotal costa three is the costa closest to the longitudinal midline.

Elytron (Figs. 42, 48, 44)

There are eight striae on the elytron of Onthophilus. Beginning from the

Histeridae: Nearctic Onthophilus 7

suture and going towards the lateral margin these are as follows: the closest

to the suture is the sutural stria; lying to the outside of this are striae five to

one, stria five being the closest to the sutural stria; then the internal sub-

humeral stria; and on the inflexed lateral area of the elytron is a sinuous

external subhumeral stria. The striae give their names to the major elytral

costae that lie immediately mesad of them. The minor carinae between the

striae and costae are not named. The strial numbering in the Histeridae is

opposite to that found in most other beetles. This convention is followed

because the striae closest to the suture are more frequently obsolete than the

striae farthest from the suture.

SC sutural costa lies mesad of the sutural stria, between it and the suture.

EC 5 elytral costa five, lies mesad of stria five.

EC 4 elytral costa four lies mesad of stria four.

EC 3 elytral costa three lies mesad of stria three.

EC 2 elytral costa two lies mesad of stria two.

EC 1 elytral costa one lies mesad of stria one.

ISC internal subhumeral costa lies mesad of the internal subhumeral stria.

ESC external subhumeral costa lies mesad of the external subhumeral

stria, and because it is straight and not sinuous it is separated from

the stria at its front end by a semicircular area.

Pygidium (Fig. 35)

TC transverse carina runs across the pygidium in the front half; it may be

nearly straight or W-shaped.

LC longitudinal carina runs along the longitudinal midline; it may be vari-

ously reduced.

Propygidium (Fig. 35)

PMC carina that runs along the longitudinal midline; if present usually tear-

shaped. 7

PLC carina that runs along the lateral margin.

TAXONOMY

Onthophilus Menene

Onthophilus Leach 1817: 78; Leconte 1845: 50; Marseul 1856: 549; Horn

1873; 138.

TYPE SPECIES. Hister sulcatus F. designated by Westwood 1840: 22.

(Onthophilus misspelled Orthophilus on pg. 22, corrected on pg. 157.)

The generic characters which are diagnostic for Onthophilus are the

following: the prosternal lobe is present but narrow; the antennal scape is

not expanded or strongly angulate; the labrum has at least two setiferous

punctures; the antennal cavities are in the anterior prothoracic angles and

partly covered by the prosternal lobe; the pronotum has two to six costae;

the elytron has eight striae and eight costae and appears strongly sculptured;

the prosternum is without striae; the metasternum is without pronounced

fovea; setae if present on the dorsal and ventral surface are very short and

not prominent.

sony literature dealing with Nearctic species at the revisional level is

cited. For literature dealing with Palearctic species refer to Bickhardt

PO LOs 72,

8 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

This combination of characters (based in part on Wenzel, 1962) serves

to distinguish Onthophilus from other genera in the family Histeridae and in

the subfamily Tribalinae. |

The Nearctic species of Onthophilus are dark brown to reddish brown

and are under 6.0 mm in length. Their shape is oval and they are moderately

convex transversely and longitudinally.

The head is deflexed. The antennae are inserted in front of the eyes,

are eleven segmented, the last three are fused into an oval club with the

sutures indistinct, and are hidden under the thorax in fossae that are in the

anterior angles of the prothorax and are partly covered by lateral extensions

of the prosternal lobe. The mouth parts are illustrated in Figs. 1-3.

Pronotum has two or six costae. The marginal and lateral striae of the

pronotum are obsolete. Elytron has eight striae with strong punctures and

eight strong costae. External subhumeral stria sinuous and subcariniform.

Area between this stria and lateral margin of elytron with irregular punctures.

Scutellum small and triangular. Propygidium and pygidium strongly deflexed,

with or without carinae.

Prosternum without striae. Hind margin emarginate. Mesosternum

short, front margin lobed to fit prosternal emargination. Metasternum with-

out fovea. Meso and metaepisterna with large contiguous punctures. Clearly

defined by sutures. Legs are short and retractile; femora slightly swollen

with minute, elongate punctures; protibia expanded apically and spinulose;

meso- and metatibia narrow with few rows of spines, tibial spurs unequal and

distinct; tarsi 5-5-5, short with simple claws.

Male aedeagus is surrounded by terminalia composed of the sclerites

of the eighth, ninth and tenth abdominal segments. The tegmen is composed

of a tubular or ovoid basal piece that is free moving and articulated to the

parameres. The parameres are fused for most of their length, forming a

tube that surrounds the penis and median lobe; their apex is bifid and strongly

curved. The penis is flat, its apex is curved and surrounds the median lobe;

its basal end is formed into two apodemes. The median lobe is extruded

dorsally or ventrally between the bifid apex of the parameres. Female geni-

talia with hemisternite narrow and articulated to stylus; no sclerotized

spermatheca was observed. The terms follow usage by Lindroth and Palmen

(1956) and the male and female genitalia are illustrated in Figs. 7-19.

ARTIFICIAL KEY TO THE NEARCTIC SPECIES OF ONTHOPHILUS LEACH

1. Length from front angles of pronotum to hind margin of

elyika. oreaver than A. 91. uses 6 ote and 14. giganteus n. sp. (p. 30)

Length from front angles of pronotum to hind margin of

te STATE A ANN accra doth boat SuSE Remailhs Bic seeks) wisn, iets loon 2

2. Elytral costae one to five interrupted at least three times,

consequently costae are reduced to series of elongate

PUDERCILCS Hie BENey mc x) vidi de he e Sw 4 HOR 1. julii Lewis (p. 10)

Elytral costae one to five entire or costa one or three

interrupted NO-MOre than ONCE, — csses, pls Se Gudens aotsee | s 3

3. Elytral costae two and four much more strongly developed

than costae one, three and five (figs. 40, 42). ...... 10

Histeridae: Nearctic Onthophilus a

Elytral costae one to five equally, or very nearly equally,

strongly developed (figs. 41, 45). .......4.2+4ese-s 4

Pronotum with costae two and three absent, or very

faintly indicated by weakly raised, shining areas

(figS.). 49. 50)enicees 1 seem hepeal dais Et ee Gee a 5)

Pronotum with costae two and three clearly indicated

and entire (figs. 45, 46, 47, 48)... 6 2. eee se ee wes 6

Elytral costa three interrupted before the front margin

by a short, deep, transverse fossa (fig. 50). Meta-

sternum with shallow, fine punctures on the central

pmed (liek Gils <i skies. castade aban inbhaad 13. kirni Ross (p- 29)

Elytral costa three entire to the front margin, without

short, deep, transverse fossa. Metasternum with

deep, medium sized punctures along front and hind

margins; two large, triangular, discal areas with

only minutespunctuses.d Me mo2)\cubars «wations iis aycepesls ae

(ok a cas, $n baebdremd abesceersar i 12. thomomysin. sp. (p- 27)

Median longitudinal carina of pygidium strong. Pronotal

punctures elongate on discal area; interstices arranged

in a strigose pattern (fig. 41). ... 4. intermixtus n. sp. (p.15)

Median longitudinal carina of pygidium absent. Pronotal

punctures round or oval on discal area; interstices not

arrangeciiies Sstrvigese DatlCEn? comiaecwils cow Pome oh ive ai

Second pronotal costa strong and complete from front to

hind margin (fig. 45). Mesosternum with two rows

of punctures only in the middle (fig. 58). ..........

ens ste beep bod senda test tects 8. wenzelin. sp. (p. 22)

Second pronotal costa clear but abbreviated before front

and hind margin; the ends sometimes covered with

punctures that are nearly as large as those in the

intercostal areas (figs. 46, 47, 48). Mesosternum

with at least two complete rows of punctures across

te Gee ee SPUOOSes wie wlarsa those is tad preee std 8

Second pronotal costae convergent behind; these costae

higher and broader at the hind ends (fig. 46). Meta-

sternum rather densely punctate; without two distinct,

triangular discal areas that are punctulate (fig. 59).

oy tp ee arpa Bs Missa Heute sy toeegee ie 9. cynomysi n. sp. (p. 23)

Second pronotal costae parallel or divergent behind; these

costae not distinctly higher and broader in the hind

half. Metasternum with two distinct, triangular discal

areas that are punctulate; punctures along front and

hind margin large, dish-like, sometimes with smaller,

deep punctures inside their leading edge (figs. 60, 61).

Lateral edge of pronotum smoothly arcuate or weakly

angulate, never parallel in hind 0.5 (fig. 48). First

abdominal sternum with large punctures along front

10.

ies

12s

13.

Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

margin and a distinct, semi-circular median area

thathicepunctuiate (is olljen Rite eke 2k 11. lecontei Horn (p. 26)

Lateral edge of pronotum strongly angulate at mid-

point, sides parallel in hind 0.5 (fig. 47). First

abdominal sternum rather densely punctate; punc-

tures becoming gradually smaller from front to

backlogs. 160 emer see Cee 10. soltaui Casey (p. 25)

Metasternum with a common or two triangular discal

areas that are smooth or minutely punctured; punc-

tures elsewhere small or large and dish-like with a

small, deep puncture inside the leading edge (figs.

SONS ome aiid chy ICEL Jha BIN Booed. Hea: BE es 11

Metasternum coarsely, rather uniformly punctate

throughout; punctures large and shallow, without

small punctures inside their leading edge (figs. 52,

Hoye tes eA COVE JOOR <UL) WER Oke gusto 13

Metasternum with a strongly impressed, longitudinal,

broad channel; triangular, minutely punctured discal

areas elevated and irregularly oval (fig. 57). .......

ike, DERMOT Reale Sey he. 7. pluricostatus Leconte (p. 20)

Metasternum without strongly impressed, longitudinal,

broad channel; triangular, minutely punctured discal

areas fused or not elevated (figs55, 56)... «2% 2.0. w% 12

Metasternum with no coarse punctures along the longi-

tudinal midline, or with punctures along the longitud-

inal midline and on the front 0.3 that are not signifi-

cantly larger than the punctures on the prosternum

and that are separated by two to three times their

GAMeLE RS f1ShV55) 2 see 5. alternatus (Say) (p. 16)

Metasternum with coarse, crateriform punctures with

small deep punctures inside their leading edge, along

the longitudinal midline and on the front 0.3 (fig. 56). ...

o) Ais oray =a; te, GUS Chain MRE aa HD, By te 6. nodatus Leconte (p. 18)

Lateral edges of pronotum straight and strongly conver-

gent towards the front, and not reflexed in hind 0.5

(iG. (SO SENG. oles. BID) Bins fied eaLk 2. deflectus n. sp. (p-. 12)

Lateral edges of pronotum arcuate or feebly angulate at

midpoints, and distinctly reflexed in hind 0.5 (fig.

ADs MS LEME DY ol, ARM ee gg a 3. flohri Lewis (p.14)

1. Onthophilus julii Lewis

Figures 20, 31, 38, 51, Map 2

Onthophilus julii Lewis, 1892:124. TYPE. Holotype, female, from Mexico:

Amecameca. Sacromonte, X.1891, collected by Julius Flohr from

under decaying plants, alt. 8,300 ft (BMNH, London), not seen.

DIAGNOSIS. This species is readily distinguished from all other New

World species of Onthophilus by the greatly interrupted elytral costae.

Histeridae: Nearctic Onthophilus 11

Elytral costae one, three and five are low and interrupted five times. Elytral

costae two and four and the internal subhumeral costa are much higher and

are interrupted three times. The general outline of the body and the puncta-

tion, especially on the meso and metasternum, suggest that this species is

most closely allied to flohri, deflectus and intermixtus.

FEMALES. Length: 2.4 mm; width: 2.0 mm. Black with brassy tinge

on head, pronotum and elytra. Head with VFC strong and complete. LFC

strong and curved inward; carinae meeting above clypeus, forming a short

single carina. Punctures on vertex, frons and genae large, round to irregu-

lar, and contiguous; on clypeus and vertices of LFC much smaller. Labrum

with setiferous punctures in lateral 0.2; front margin straight. Pronotum

transverse. Lateral margin broadly, obtusely angulate, somewhat rounded;

lateral bead weak but distinct; surface along lateral edge reflexed. Disk 6-

costate. PC 1 strong, very short and slightly curved inward. PC 2 strong,

reaching from front 0.3 to just before the hind margin and curved gently out-

ward. PC 3 strong; reaching from front margin to hind 0.1, but may be

completely interrupted in the middle third of its length; curved slightly out-

ward. Punctures large, round to irregular, contiguous, except smaller

along hind margin; vertices of elevations with double row of minute punctures.

Surface between PC 1 and PC 2 strongly concave, forming a broad trench

that curves to the front angle; trench deepest just behind front angle. Elytron

wider than pronotum. EC 1, EC 3 and EC 5 much lower than EC 2, EC 4 and

ISC; interrupted five times; elongate tubercles more prominent in hind 0.3.

EC 2, EC 4 and ISC very prominent; interrupted three times; tubercles behind

front margin longest. ESC low and interrupted only once in the middle. SC

very much attenuated in front 0.3. All costae with two rows of irregular

punctures along their base; punctures prominent between tubercles. Inter-

spaces between costae with clear stria that is bordered by two low carinae;

strial punctures small and round, deepest at front margin, with two small

lateral tubercles that interrupt the low carinae. Surface depressed behind

front margin, forming shallow, transverse trench in front of costae.

Propygidium with PMC strong; slightly abbreviated in front and behind. PLC

strong, about 0.3 times as long as PMC. Punctures large, shallow, irregu-

lar and contiguous; on vertices of elevations very fine. Pygidium with LC

strong, bifurcating in the middle of the pygidium, arms reaching hind 0.2.

TC strong and straight; intersecting LC infront 0.2. Surface depressed

before and behind TC. Punctures similar to those on the propygidium, except

smaller and more pronounced on outer margins. Prosternum slightly con-

cave between procoxae, convex before. Hind margin broadly, roundly emar-

ginate. Punctures coarse, irregular, shallow and contiguous; about 0.5

times as large and more rounded on prosternal lobe. Mesosternum short and

transverse. Front margin lobed to fit prosternal emargination. Punctures

similar to those on the prosternum. Metasternum separated from mesoster-

num by a suture which is irregular due to the punctures encroaching on it.

Median longitudinal suture fine and obscured in part by punctures. Surface

slightly depressed behind the front margin and along the median longitudinal

suture; elevated triangular area between meso- and metacoxae. Punctures

intermixed large and small, irregular and contiguous, except small on tri-

angular area. First abdominal sternum punctate throughout; punctures

similar to those on prosternum. Surface strongly reflexed behind front mar-

gin. Protibia slightly expanded distad, with eight short spicules each on a

weakly elevated base.

MALE. Unknown.

SPECIMENS. Mexico. Oaxaca. km 154 on Route 175 south of Oaxaca,

£2 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

8,000 ft, 2-3. VI.1971, H.F. Howden, 1female. 20 km n. Oaxaca, 9,500

ft, 7.VI.1971, S. Peck, under carrion, 1 female.

REMARKS. I have seen only two specimens of this species, both

females. The variation between these two specimens from Oaxaca is very

slight. In specimen collected south of Oaxaca both PC 3 carinae are com-

pletely obsolete in the middle 0.3, but no other significant differences were

detected. P.M. Hammond kindly compared my specimens with the holotype,

which is a little smaller and the elytral costae are not quite so raised.

2. Onthophilus deflectus new species

Pigurés 11, 39, 52, Map 1

Onthophilus deflectus n. sp. TYPE. Holotype, male, Trelease Woods,

Urbana, I111., 28.IV.1951, R.K. Benjamin, (FMNH, Chicago), col-

lected on dead Polypore. Allotype, female, with the same data as the

type, (FMNH, Chicago). Paratypes, all material listed in the section

on specimens examined.

DIAGNOSIS. The densely punctate meso- and metasternum separates

this species from other species, with the exception of flohri, in which the

major elytral costae are unequally developed. This species may be disting-

uished from flohri, which it resembles closely, by the following characters.

The lateral pronotal edges of deflectus are nearly straight and strongly con-

vergent in the hind 0.5; the front angles of the pronotum are broadly round;

and the meso- and metasternum are convex. The lateral pronotal edges of

flohri are arcuate or weakly, broadly angulate and subparallel in the hind

0.5; the front angles of the pronotum are not broadly round; the meso- and

metasternum are concave in the males and flat or weakly concave in the

females. In its general outline and punctation deflectus also resembles

intermixtus but may be separated from the latter by the almost equally well

developed elytral costae, smoothly arcuate lateral edge of the pronotum, and

the long pronotal costa one, of intermixtus.

MALES. Length: 1.8 to 2.0 mm; width: 1.6 to1.8 mm. Head with

VFC strong. LFC strong and meeting just above clypeus. Punctures on

frons, vertex and genae large, roundish, deep and contiguous; punctures

much smaller on clypeus and on the vertices of elevations. Labrum with

setiferous punctures in lateral 0.2. Pronotum 0.5 times as long as wide.

Lateral margins nearly straight in hind 0.5 and strongly convergent front-

wards; lateral bead poorly developed; hind angles not reflexed, surface

smoothly concave to lateral edges; front angles broadly rounded. Disk 6-

costate. PC 1 short and straight; reaching from just before the hind margin

to front 0.5. PC 2 long and straight; reaching from the hind margin to just

before the front margin. PC 3 long and straight; reaching from the front

margin to hind 0.1. Surface not deeply concave between PC 1 and PC 2.

Punctures between PC 1 and lateral edge and along hind margin large, round,

and contiguous; punctures between costae large, elongate and contiguous,

with interspaces arranged in strigose pattern; punctures on vertices of ele-

vations minute and sparse. Elytron wider than pronotum. EC 1, EC 8, EC 5

and ESC much lower in the front 0.75 than EC 2, EC 4 and ISC. ISC, EC 2,

EC 4 and SC with a clear row of large quadrate punctures along their bases.

EC 1 and EC 3 abbreviated before front margin. SC strongly attenuated in

front 0.2. Vertices of costae with two rows of minute punctures. Spaces

between costae with strong stria bordered by two low carinae; strial punc-

Histeridae: Nearctic Onthophilus 13

tures strong, oval and deep with small but clear tubercles on their lateral

edges that interrupt the low carinae. Surface with slight transverse depression

near the middle running from ISC to EC 3. Propygidium with PMC strong;

reaching from front 0.3 to hind 0.1. PLC short, reduced to round tubercle.

Punctures large, irregular deep and contiguous. Vertices of elevations with

minute punctures. Pygidium with LC strong; reaching from front margin to

hind 0.1. TC strong, obtuse V-shaped, meeting LC in front 0.3. Punctures

similar tothose on the propygidium. Prosternum flat between procoxae, con-

vex in front. Hind margin broadly, roundly, shallowly emarginate. Punc-

tures round, large and small intermixed, and contiguous; unctures smaller

on prosternal lobe. Mesosternum short and transverse. Front margin lobed

to fit the prosternal emargination. Punctures similar to those on the pro-

sternum. Metasternum separated from the mesosternum by a fine suture.

Median longitudinal suture distinct. Surface with a broad, oval discal ele-

vation between meso- and metasternum. Punctures round to irregular, larger

than on mesosternum and contiguous; punctures slightly smaller on elevations,

with minute punctures on interstices. First abdominal sternum punctate

throughout; punctures similar to those on prosternum. Protibia with 12-14

small spicules on only slightly elevated bases. Male genitalia nitalia shown in fig.

ik.

FEMALES. Length: 1.9 to 2.2 mm; width: 1.6 to 2.0 mm. Very

similar to males. They can usually be distinguished from males by the meta-

sternum, which is in females only very slightly depressed along the longitud-

inal median suture.

SPECIMENS. CANADA: ONTARIO. Essex Co., Lasalle, 19-20.V.

1967, K. Stephan, 1 male, 3 females. UNITED STATES: ALABAMA.

Jackson Co., Scottsboro, 4-8.IV.1967, S. Peck, forest litter, 1 female.

ILLINOIS. Cook Co., Joliet, 18 .IV.1927, O. Park, cow dung, 1. Urbana,

Trelease Woods, 17.111.1944, W.E. Snow, ex tree hole, 1 female; 28.IV.

1951, R.K. Benjamin, from dead Polypore, 1 male, 3 females. INDIANA.

Porter Co., 1 mi. S.W. of McCool, 4.X.1952, H.S. Dybas, 1 female,

berlese leaf litter. MARYLAND. Takoma Pk., 16.V.1948, E.S. Ross, 2

males. MICHIGAN. Berrien Co., Lakeside, Warren Woods, 25. VI.1966,

W. Suter, buttress debris beech, 1 male. MISSISSIPPI. Meridian, 8.II.,

H. Soltau, 1. MISSOURI. Boonville, 4.V.1964, G.G. Kennon, ex mouse

nest under split oak log, 1 male, 1female. NEBRASKA. F.C. Bowditch, 1.

NORTH CAROLINA. Black Mts., VI.1902, Van Dyke, 1. Round Knob, 25.

VI., Hubbard and Schwartz, 17. SOUTH CAROLINA. Clemson, 28.I1.1962,

J.-A. Payne, chicken decay, 1; 27.11.1967, J.A. Payne, dead pig remains, 6.

TENNESSEE. Blount Co., Great Smoky Mts. Nat. Pk., Cedar Cove, 8.VI.

1960, W. Suter and J. Wagner, alt. 1700 ft, leaf litter, 1. Specimens are in

the following collections: Cal. Acad., San Francisco; FMNH, Chicago;

USNM, Washington; Howden, Ottawa.

REMARKS. This species is named deflectus because unlike any other

species, the lateral edge of the pronotum is not reflexed in the hind 0.5.

The hind angles of the pronotum appear to bend down.

Most of the specimens of this species that I have seen are quite uniform

in appearance, except those from Black Mountain and Round Knob, North

Carolina and the specimen from Nebraska. The PC 3 costae of these speci-

mens may be weakly to strongly, or not at all attenuated at their mid-points,

and the EC 2 costa is weakly to rather strongly attenuated in the front 0.3.

The strial puncture in stria 3 at this point is sometimes strong and reinforces

the appearance that there is a transverse fossa across the elytron in the front

0.3. There is a spectrum of variability from specimens in which only the EC

14 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

2 is slightly attenuated to specimens in which the PC 3 is virtually interrupted

and the EC 2 is strongly attenuated. No differences were noted in the male

genitalia of the specimens. At this time, I would like only to note that this

variability exists. Further studies and collecting will determine whether

these forms should receive formal recognition as subspecies or whether they

should be elevated to specific status.

3. Onthophilus flohri Lewis

Figures 6, 9, 12, 21, 32, 40, 53, Map 2

Onthophilus flohri Lewis, 1888: 235. TYPE. Holotype, sex unknown, labelled

'type''’, Mexico. Durango. 8.VIII.1887, J. Flohr, (BMNH, London),

not seen.

DIAGNOSIS. O. flohri resembles deflectus and intermixtus in general

outline and the densely punctate meso- and metasternum. Characters that

are useful for separating flohri from the former, are discussed in the diag-

nosis for deflectus. Intermixtus has almost equally well developed elytral

costae and the pronotal costa one is long. In flohri, elytral costae one, three

and five are much lower than elytra costae two and four, and the pronotal

costa one is short. This species is readily distinguished from other species

with unequally well developed elytral costae by the densely punctate meso-

and metasternum.

MALES. Length: 2.0 to 2.5 mm; width: 1.6to1.8 mm. Head with

VFC strong, reaching occipital carina. LFC strong above antennal pit,

becoming obsolete above clypeus; curved inward slightly. Punctures large,

shallow, irregular, and confluent on frons and genae; smaller and elongate

above antennal pit; interstices arranged in strigose pattern; smaller and

rounded along front margin of clypeus; vertices of elevations with small,

round punctures. Labrum with setiferous punctures in lateral 0.2; front mar-

gin arcuate. Pronotum transverse, 0.50 as long as wide. Lateral margin

arcuate, or weakly angulate at mid-point; slightly reflexed. Front margin

slightly emarginate; front angles obtuse. Disk 6-costate. PC 1 short,

straight; reaching from front 0.5, hind 0.10 or 0.05. PC 2 curved inward;

abbreviated only slightly in front and behind. PC 3 straight; abbreviated

behind, reaching from front margin to hind 0.20. PC 2 and PC 3 convergent

towards the front. The two PC 3 almost parallel. Punctures at edges small

and round; between costae elongate, interstices arranged in strigose pattern.

Elytron wider than pronotum. ESC, ISC, EC 2, EC 4, and SC abruptly ele-

vated and prominent; EC 1, EC 3 and EC 5 much lower and abbreviated just

behind front margin. All costae abbreviated just before hind margin. SC

attenuated in front 0.3; EC 1 slightly attenuated in the middle. All costae

with a row of minute punctures on each side just below the vertex; EC 2, EC

4, SC and ISC with two basal rows of large quadrate punctures along each

side. Spaces between costae with a strong stria bordered on each side by a

low carina; strial punctures oval and deep, deepest in front, with a clear

tubercle on each side that interrupts the carinae. Propygidium with front

margin reflexed. PMC strong; reaching from just behind front margin to

hind 0.2 to 0.1. PLC 0.5 times as long as PMC. Punctures large, round

and separated by 0.5 times their diameters; vertices of elevations with

minute punctures. Pygidium with LC strong; reaching from front margin to

hind 0.4 to 0.3. TC strong, obtuse V-shaped, meeting TC in front 0.35.

Punctures similar to those on propygidium. Prosternum with distinct depre-

ssion between procoxae. Hind margin obtusely, roundly emarginate; some-

Histeridae: Nearctic Onthophilus 15)

what sinuous. Punctures large, round, separated by their diameters; much

smaller punctures between larger punctures, more dense towards the lateral

margin. Prosternal lobe with punctures much smaller. Emargiuation pre-

sent between prosternal lobe and proepisternum. Mesosternum short and

transverse. Front margin lobed to fit prosternal emargination. Punctures

smaller and sparser than those on prosternum. Metasternum separated from

mesosternum by an indistinct suture. Median longitudinal suture very fine

and indistinct. Surface strongly depressed in the central area. Punctures

large, round, separated by their diameters; punctures on discal area and

along midline 0.50 times as large. First abdominal sternum punctate through-

out with punctures similar to those on metasternum, but becoming smaller

towards the hind margin. Protibia with eight weak spicules each on a slightly

elevated base. Male genitalia illustrated in fig. 12.

FEMALES. Length: 2.6 to 2.8 mm; width: 1.6to 2.1mm. The

females are very similar to the males. The metasternum of females is flat

or only weakly depressed.

SPECIMENS. MEXICO: Durango. EI Salto, 5.VI.1937, E.S. Ross,

44, (Cal. Acad.); 10 mi. W. El Salto, 16.V1.-12.VII.1964, H.F. Howden,

317, (CNC, Howden); 11-15.VII.1964, 9000 ft, J.E.H. Martin, 18, (CNC);

1. VII.1964, Linguist, oak, fir duff, dung, 7 (CNC); 11 mi. W. El Salto,

14-17.VI.1964, S. Peck, dung and carrion, 27, (CNC).

REMARKS. This species is at present known only from El] Salto,

Durango and has been collected in great numbers in baited traps. The vari-

ability expressed is quite low. The PC 1 costa of the males is slightly vari-

able in length, and some males are slightly, but noticeably, smaller than

most. The hind wing of at least the females is very much reduced. P.M.

Hammond has kindly compared some of my specimens with the type.

4. Onthophilus intermixtus new species

Figures 14, 41, 54, Map 4

Onthophilus intermixtus n. sp. TYPE. Holotype, male, ARIZ. Graham Co.

Pinaleno Mts. Wet Cn. 6100 ft, 29.VII.1969, A. Smetana, (CNC,

Ottawa). Paratype, 1 male with the same data as the holotype (Howden,

Ottawa).

DIAGNOSIS. This species resembles most closely flohri and deflectus

in its general outline and punctation. It may be distinguished from them

most readily by the almost equally well developed elytral costae. Other

characters that are useful in separating these species are included under the

specific description of deflectus. Although intermixtus has almost equally

well developed elytral costae, it differs from those species in which the

elytral costae are equally well developed in many significant ways. The dense,

round to irregular punctation of the meso- and metasternum will immediately

separate it from those species. The elytra are more convex, more elongate

and the elytral costae are narrower and more sharply elevated than in the

other two species.

MALES. Length: 2.5 mm; width: 1.8 mm. Head with VFC strong and

complete. LFC strong, sharply elevated and meeting above clypeus. Punc-

tures on vertex large and irregular, yielding strigose pattern; on frons and

genae large, round to irregular, shallow and contiguous; on vertices of ele-

vations small to minute. Labrum with front margin weakly arcuate; setiferous

puncture in lateral 0.2; very short setae interspersed irregularly in front

0.5, separated by 3.0 to 5.0 times their length. Pronotum 0.5 times as long

16 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

as wide. Lateral margin smoothly arcuate; lateral bead strongly developed;

surface along lateral edge reflexed from hind angles to front margin. Disk

6-costate. PC ilong, straight and well developed; reaching from front 0.25

to hind margin. PC 2 strong, curved inward in front 0.25, straight in hind

0.75; reaching from just before front margin to hind margin. PC 3 strong

and straight; reaching from front margin to just before hind margin. Surface

smoothly convex between costae. Punctures large, round and shallow between

lateral edge and PC 1; elongate, with interstices arranged in strigose pattern

between costae, becoming sparser towards the middle; small to minute on

vertices of elevations. Elytron wider than pronotum. EC 3, EC 5 and SC

slightly lower than other costae in front 0.3. All costae with two rows of

minute punctures just below the vertex; short yellow setae arising from them

in the hind 0.5. AIJl costae with a row of small quadrate punctures, which

form a fine stria along the base, cutting off a fine and irregular carina. EC

3 slightly attenuated just before the front margin. Spaces between costae

with a strong stria between two low carinae; strial punctures strong, oval

and deep, with clear tubercles on their lateral edge, that interrupt the low

carinae. Propygidium with PMC strong; reaching from front 0.1 to hind 0.1.

PLC strong, 0.3 times as long as PMC. Punctures large, deep, irregular

and contiguous; smaller on vertices of PLC. Pygidium with LC strong, com-

plete from front to hind margin. TC very obtuse V-shaped, indistinct with

punctures encroaching on its course, meeting LC in front 0.25. Punctures

similar to those on propygidium, except smaller on two indistinct lateral

tubercles and much smaller on vertex of LC. Prosternum flat between pro-

coxae, convex in front. Hind margin broadly, roundly, shallowly emarginate.

Punctures large, round, deep and separated by 0.5 times their diameters,

with much smaller punctures on the interstices; punctures smaller on pro-

sternal lobe. Mesosternum short andtransverse. Front margin lobed to fit

prosternal emargination. Punctures similar to those on prosternum, but

slightly smaller. Metasternum separated from mesosternum by a fine suture.

Median longitudinal suture fine and distinct; surface weakly concave along it.

Punctures intermingled large and medium sized, round, deep and nearly

contiguous, with small punctures on interstices;.small punctures dense on

small oval area behind mesocoxa. First abdominal sternum with punctures

along the front margin similar to those on metasternum, becoming progress-

ively smaller towards the hind margin. Protibia with six small spicules on

elevated bases in apical 0.5. Male genitalia illustrated in fig. 14.

FEMALE. Unknown.

SPECIMENS. This species is known only from the type material.

REMARKS. This species is named intermixtus because it apparently

shares characters with both the flohri group and the lecontei group. It

resembles flohri in most respects but the elytral costae of intermixtus are

equally well developed and in this character it resembles the lecontei group.

5. Onthophilus alternatus (Say)

Pemees doy. 2.1 3514, D».0.20; LOe 22.08, 42,200,004, Map-s

Hister alternatus Say, 1825: 46. TYPE. Neotype here designated, female,

bearing the following labels -- "orange", "'O. alternatus (Say) Er.",

red "MCZ TYPE 32249", red "NEOTYPE Helava '75"". (Say stated

that this species belonged in the genus Onthophilus Leach.)

Onthophilus alternatus (Say) Leconte, 1845: 52. Marseul, 1856: 558. Horn,

Histeridae: Nearctic Onthophilus 17

1873: 303. |

DIAGNOSIS. QO. alternatus is very similar to nodatus and pluricostatus

in its general outline, in the shape and disposition of its pronotal costae and

in the sculpture of its elytra. This species is most readily distinguished

from the latter two species by the sparse punctation of the metasternum and

by the shape of the punctures. Male specimens of alternatus have no large

punctures along the longitudinal midline, only very fine punctures; females

have few large punctures along the midline and have two large oval-triangular

discal areas that are punctulate. The metasternal punctures on the central

area are not ocellate in alternatus, but rather small and deep, with an abrupt

leading edge and a less abrupt and fan-like trailing edge. If the ventral sur-

face is not visible, alternatus may be distinguished with some confidence by

the bituberculate verto-frontal carina and the fine, deep punctation of the

pygidium.

MALES. Length: 1.6 to1.9 mm; width: 1.4to1.7 mm. Head with VFC

bituberculate. LFC strong, curved inwards; the two LFC meeting above the

clypeus. Punctures on frons and genae small, deep and round; separated by

3.0 to 4.0 times their diameters; punctures much smaller, shallower and

denser on vertices of elevations and above antennal sockets. Labrum with

setiferous punctures in lateral 0.25. Pronotum 0.55 times as long as wide.

Lateral margin feebly angulate at midpoint; lateral edge reflexed; lateral

bead not prominent. Disk 6-costate. PC 1 slightly sinuous, strong; complete

from front 0.3 to hind margin. PC 2 subsinuous, strong; complete from

front margin to just before hind margin. PC 8 straight, strong; complete

from front margin to hind 0.2. Surface much higher between inner four

costae, than between PC 2 and lateral margin. Punctures, except on costae,

small, deep, slightly elongate and separated by their diameters; punctures

on vertices of elevations minute. Elytron wider than pronotum. ESC, ISC,

EC 2, and EC 4 much higher and more pronounced than EC 1, EC 3, EC 5

and SC. EC 1 interrupted in front 0.3; EC 3 interrupted before front margin

by a short transverse fossa between stria three and stria four; SC attenuated

just behind the scutellum. Spaces between costae with a strong stria bordered

on each side by a low carina; strial punctures deep and oval, especially deep

just behind the front margin and in the front 0.3 of stria one to stria four,

these giving the effect of a transverse depression; strial punctures with a

tubercle on each side that interrupts the carina. All costae with a row of

minute punctures on each side below the vertex. Propygidium with front mar-

gin strongly reflexed. PMC strong, complete from front 0.2 to hind 0.1.

PLC 0.5 times as long as PMC. Punctures small, deep, round and separated

by 2.0 to 3.0 times their diameters; punctures on vertices of elevations

minute. Pygidium with front margin slightly reflexed. LC obsolete except

in hind 0.2 or 0.1. TC strong and shaped like two contiguous shallow U's.

Punctures similar to those on propygidium. Prosternum slightly depressed

or flat between procoxae. Hind margin broadly, roundly emarginate. Punc-

tures small, deep, round and separated by 2.0 to 3.0 times their diameters;

prosternal lobe and interstices of punctures punctulate. Mesosternum short

and transverse. Front margin lobed to fit prosternal emargination. Punc-

tured like prosternum. Metasternum separated from mesosternum by a fine,

indistinct suture. Median longitudinal suture a fine, distinct, impressed

line. Surface slightly to rather strongly depressed in front 0.5. Punctures

along front margin small, deep and round, spaced irregularly; punctures

along hind margin and on lateral areas large, irregular and sometimes

indistinct, shallow and dish-like, with small, deep, round punctures near

the leading edge. Central areas and interstices of punctures punctulate.

18 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

First abdominal sternum with front margin reflexed. A single row of punc-

tures along front margin that are like those on the hind area of metasternum;

punctures dense and much smaller on lateral areas; punctulate on central

area. Protibia with seven to eight spicules, each on a weakly elevated base.

Male genitalia illustrated in fig. 10.

FEMALES. Length: 2.0 to 2.5 mm; width: 1.6to 2.0 mm. The

females are very similar to males, except that the metasternum is not

depressed and there is a double row of punctures along the longitudinal mid-

line, leaving two large oval-triangular, punctulate, discal areas. The

pygidial carina LC is longer; it is obsolete except in the hind 0.4 or 0.5.

SPECIMENS. UNITED STATES: ALABAMA. Seale, 21.III.1954,

H.F. Howden, 9. FLORIDA. Clarksville, 21.I11.1954, H.F. Howden, 20.

DeFuniak Spr., 21.1I11.1954, H.F. Howden, 5. Levy Co., P. Laurent, 1.

Mossy Head, 21.II1.1954, H.F. Howden, ex skunk dung, 1 female.

LOUISIANA. 10 mi. S.W. Alexandria, 1.IV.1960, J.G. Chillcott, 1 male,

4 females. NORTH CAROLINA. Raleigh, 18.X.1952, H. & A. Howden,

ex Rhizopogon piceus, 25. Southern Pines, 6.X.1949, H. & A. Howden, ex

rotten watermelon rind, 14. SOUTH CAROLINA. Columbia, 3.I1II.1932,

O.L. Cartwright, 1. Windsor, X1I.1933, E.S. Ross, ex dung, 31. Speci-

mens are in the following collections: Cal. Acad., San Francisco; USNM,

Washington; FMNH, Chicago; Howden, Ottawa; CNC, Ottawa; AMNH, New

York; MCZ, Cambridge; ANSP, Philadelphia.

REMARKS. There is not a great deal of variability between the speci-

ments examined. The pygidial carina LC is somewhat variable in length,

both in males and females. Also the metasternal punctation is reduced in

some males; the punctures are smaller and more irregular, often fused to-

gether behind the mesocoxae.

Say described alternatus from the Melsheimer collection under the

name Hister alternatus, but stated that this species belonged to the genus

Onthophilus Leach. It is very difficult to determine from Say's description

to which species this description corresponds, and unfortunately Say's types,

if extant in the Melsheimer collection, were not labelled. Leconte redescribed

alternatus in 1845, but did not provide any useful diagnostic characters to

separate alternatus from nodatus and pluricostatus which he described at the

same time. However, Leconte did send a Zimmerman specimen from

Georgia to Marseul, who redescribed alternatus in 1856. In the key to the

species of Onthophilus, Marseul described the metasternum as "imponctué

et impressioné au milieu" and in the description he stated that the meta-

sternum is "fovéolé au milieu finement pointillé et entouré de quelque gros

points €pars". For these reasons, I have decided to attach the name

alternatus to the species in which the males have no large punctures along

the longitudinal midline, and in which the females have two large oval-tri-

angular, punctulate discal areas. Although the distribution of this species _

appears to be centered in the southeastern states, and it may be unlikely that

Say had a specimen which corresponds to my useage, I have decided to follow

the useage of Leconte and Marseul, and have chosen a Leconte specimen as

neotype.

6. Onthophilus nodatus Leconte

Figures 16, 23, 34, 48, 56, Map 2

Onthophilus nodatus Leconte, 1845: 53. TYPE. Holotype, (MNHN, Paris)

not seen. Marseul, 1856: 556. Horn, 1873: 303.

Histeridae: Nearctic Onthophilus 19

DIAGNOSIS. O. nodatus is very nearly allied with alternatus and

pluricostatus. This species may be separated from the former by the charac-

ters given in the diagnosis for alternatus. O. nodatus may be distinguished

from pluricostatus by the following characters. The metasternum of

pluricostatus is strongly depressed along the longitudinal midline, forming a

broad, rounded trench from the front to the hind margin. This trench is

stronger in the males than in females. The metasternum of nodatus is not

depressed along the midline, although the large crateriform punctures along

the midline sometimes give that impression. The tubercles on each side of

the strial punctures are much stronger and clearer in nodatus, especially in

striae three, four and five, than the tubercles in pluricostatus. The pronotal

costae three of nodatus are, in general, parallel in the front 0.5 and are

closer to each other than to pronotal costae two. The pronotal costae three

of pluricostatus are divergent in the front 0.5 and are closer to pronotal

costae two than to each other. The lateral margin of the pronotum of nodatus

is more strongly reflexed than that of pluricostatus.

MALES. Length: 2.3 to 2.6 mm; width: 1.8to 2.3 mm. Head with

VFC strong and entire, reaching from occipital carina to frons between

antennae. LFC strong, the two convergent, meeting above clypeus forming

a short single carina. Punctures large, round, deep and separated by their

diameters; interspaces of large punctures and vertices of elevations with

minute punctures. Labrum with small, shallow punctures; setiferous punc-

tures in lateral 0.25. Pronotum transverse, 0.5 times as long as wide.

Lateral margins angulate in middle and very strongly reflexed; lateral bead

prominent. Front margin weakly emarginate; front angles obtuse; front mar-

gin strongly reflexed from front angles to PC 2. Disk 6-costate. PC 1 long,

straight or curved inwards; reaching from front 0.25 to hind margin. PC 2

subsinuate; reaching from front margin to just before hind margin. PC 3

curved feebly outward; reaching from front margin to slightly before hind

margin. Surface between the four median costae higher than between PC 2

and lateral margin. Punctures medium, oval or round and deep; separated

by their diameters. Elytron wider than pronotum. ESC, ISC, EC 2, EC 4,

and SC abrupt and more strongly developed than EC 1, EC 3, and EC 5. EC

1 interrupted in front 0.3-0.5; EC 3 interrupted before front margin by a

short transverse fossa between stria three and four, and interrupted in front

0.5; SC attenuated in front 0.2. All costae with a row of minute punctures on

each side just below vertex. Spaces between costae with a strong stria bord-

ered on each side by a low carina; strial punctures large, oval and deep;

strial punctures strongest just behind front margin and in the front third of

stria one to stria four; strial punctures with very strong tubercles on each

side that interrupt the low carinae. EC 1, EC 2 and EC 8 attenuated before

hind margin. Propygidium with front margin strongly beaded and reflexed.

PMC strong; reaching from front 0.2 to hind 0.2. PLC slightly shorter than

PMC. Punctures large, round and deep; separated by their diameters;

vertices of elevations with minute punctures. Pygidium as long as wide. LC

briefly interrupted behind transverse carina. TC bisinuate; with two short

branches extending backwards in lateral 0.25 that are sometimes joined to

LC. Punctures like those on propygidium. Prosternum flat between procoxae,

strongly convex and higher in front. Hind margin broadly, roundly emargin-

ate. Punctures large, round and deep; separated by their diameters; pro-

sternal lobe with small, round, deep punctures. Mesosternum transverse,

short. Front margin lobed to fit prosternal emargination. Punctures large,

round and deep and contiguous. Metasternum separated from mesosternum

by a fine suture. Median longitudinal suture fine. Surface flat or slightly

20 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

depressed. Punctures on narrow, triangular discal area minute, round and

shallow; punctures elsewhere very large, shallow dish-like depressions with

large, deep, round punctures within. First abdominal sternum with a row of

large punctures like those on the hind margin of the metasternum, behind the

front margin, becoming smaller and denser on the lateral areas; central area

punctulate. Protibia with seven to eight spicules, the two most distad on

weakly elevated bases. Male genitalia illustrated in Fig. 16.

FEMALES. Length: 2.6 to 3.0 mm; width: 2.3 to 2.5mm. The

females are very similar to males. The metasternum is slightly convex and

the metasternal punctures are generally larger and are denser along the mid-

line.

SPECIMENS. UNITED STATES: ALABAMA. 17 mi. N. Abbeville,

22.I1I1.1954, H.F. Howden, ex malt trap, 1. FLORIDA. 2 mi. W. Archer,

24.1I1I.1953, Howden & Dozier, ex propionic acid trap, 1. St. Augustine, 3.

GEORGIA. 10 mi. N. Ft. Valley, Rt. 341, 17.11I.1956, Howden & Howell,

ex malt, 3. Tifon, 6.X.1968, J.A. Payne, ex gopher tortoise dung trap out-

side burrow, 1. NORTH CAROLINA. Southern Pines, 9.I1.1952, H. & A.

Howden, 1. OKLAHOMA. Payne Co., 20.I1II.1926, W.J. Brown, 1. SOUTH

CAROLINA. Florence, 18.II.1938, 2.. TEXAS. Nacogdoches, 14.V.1974,

V. Nealis, ex human dung trap, pine-oak, 4. Specimens are in the following

collections: FMNH, Chicago; USNM, Washington; CNC, Ottawa; Howden,

Ottawa; MCZ, Cambridge.

REMARKS. There is very little variability between the specimens

examined of this species, other than that already noted. Marseul's descrip-

tion of this species in 1856 corresponds closely to the specimens labelled O.

nodatus in the Leconte Collection.

7. Onthophilus pluricostatus Leconte

Figures 13, 24, 35, 44, 57, Map 5

Onthophilus pluricostatus Leconte, 1845:51. TYPE. Lectotype, here

designated, sex not determined, bearing the following labels -- "orange",

"937", "O. pluricostatus", red "MCZ TYPE 32251", red "lectotype

Helava '75"". Marseul, 1856: 565. Horn, 18738: 303.

DIAGNOSIS. This species is very closely allied with alternatus and

nodatus. QO. pluricostatus may be distinguished from these species by the

broad, rounded, trench-like depression along the longitudinal midline of the

metasternum. Further characters that separate this species from the latter

two are dealt with under the diagnoses for alternatus and nodatus.

MALES. Length: 2.3 to 2.5 mm; width: 1.7to 2.1mm. Head with

VFC entire. LFC strong, meeting above clypeus, and forming a short, single

carina. Punctures large, round and deep; separated by their diameters on

frons, genae and vertex; smaller above antennae; vertices of elevations with

minute punctures. Labrum with small, shallow and round punctures; seti-

ferous punctures in lateral 0.25. Pronotum 0.5 times as long as wide.

Lateral margin strongly angulate at mid-point, sides in hind 0.5 parallel and

convergent in front 0.5; lateral edges strongly reflexed; lateral bead indis-

tinct. Front margin reflexed from front angles to PC 2; front angles 90°.

Disk 6-costate. PC 1 strong, slightly curved outwards; complete from front

0.25-0.3 to hind margin. PC 2 sinuous; complete from front to hind margin.

PC 3 straight, slightly curved outward in hind 0.1; complete from front mar-

gin to just before hind margin. Surface of pronotum elevated between four

Histeridae: Nearctic Onthophilus agi

median costae. Punctures medium, round and deep, separated by their dia-

meters; sparser and slightly elongate between the PC 3; denser and slightly

larger along lateral margin. Elytron wider in front than pronotum. ESC,

ISC, EC 2, EC 4 and SC more abrupt and higher than EC 1, EC 3 and EC 5.

EC 1 weakly attenuated in front 0.35 to 0.5; EC 3 interrupted before front mar-

gin by a short, transverse fossa between stria three and four; SC attenuated

behind front margin. All costae with a row of very minute punctures on each

side just below the vertex. All costae weakly attenuated before hind margin.

Spaces between costae with a strong stria bordered on each side by a low

carina; strial punctures large, oval and deep, deepest just behind front mar-

gin, and with small tubercles on each side; tubercles clearest in stria one and

two, and interrupt the low carinae. Propygidium with front margin smoothly

arcuate and strongly reflexed. PMC strong and complete from front 0.2 to

0.25 to hind 0.1. PLC 0.5 times as long as PMC. Punctures medium, round

and deep; separated by their diameters. Pygidium with front margin slightly

reflexed. TC very obtuse V-shaped. LC complete from front to hind margin

except sometimes briefly interrupted just behind TC. Punctures similar to

those on propygidium. Prosternum flat between procoxae, convex in front.

Hind margin roundly, quite deeply emarginate. Punctures medium, round,

deep and separated by 1.0 to 2.0 times their diameters; punctures on pro-

sternal lobe much smaller. Mesosternum short and transverse. Front mar-

gin lobed to fit prosternal emargination. Punctures much larger. than those

on prosternum; dish-like with medium, round and deep punctures in their

centers, and nearly contiguous. Metasternum separated from mesosternum

by a suture and an irregular impunctate line. Median longitudinal suture fine.

Surface with broad, rounded, trench-like, longitudinal depression along the

midline from the front to the hind margin. Punctures on two triangular, some-

what oval, weakly elevated discal areas small, round and shallow; punctures

in front, behind and on lateral areas large, irregular, dish-like with medium,

round, deep punctures inside them, and contiguous; punctures in depression

smaller. First abdominal sternum with punctures along front margin like

those on the lateral areas of the metasternum, except fan-like at their trailing

edges; punctures smaller on lateral areas, becoming much smaller and

sparser on central area. Protibia with eight small spicules, each on a weakly

elevated base. Male genitalia illustrated in fig. 13.

FEMALES. Length: 2.3 to 2.7 mm; width: 2.1to 2.3 mm. The

females are very similar to the males in most respects, except that the

median depression of the metasternum is not well developed and clear in some

specimens. These females may be identified with some confidence by the

complete VFC and the carinae of the pygidium.

SPECIMENS. UNITED STATES: ALABAMA. Mobile, 24. Springhill,

Zeer, lose, W.P.” Loding, 1: DC... 1." GHRORGIA. "os mit We Doccea,y s.

IX.1967, J.M. & B.A. Campbell, under dog carcass, 5. KENTUCKY.

Slade, 25.VIII.1967, J.M. & B.A. Campbell, 1. MARYLAND. Oakland,

Hubbard & Schwarz, 1. MASSACHUSETTS. Framingham, 30-31.VII.,

C.A. Frost, ex fungus, 1. Natick, 13.IX.1913, C.A. Frost, sweeping in

woods, 1. Springfield, 1.VIII.1927, D.H. Blake, under Lactarius, 1; 1954,

Chapin, 1. MICHIGAN. Detroit, Hubbard & Schwarz, 1. NEW JERSEY.

Lakehurst, 16. VIII.1912, 1. NEW YORK. Peck, 1. NORTH CAROLINA.

Pisgah Forest, Looking Glass Mt., 21, 27. VIII.1940, O.L. Cartwright, 2;

Pisgah Forest, 26. VIII.1941, O.L. Cartwright, 16. PENNSYLVANIA.

Jefferson Co., 10.1X.1950, O.L. Cartwright, 1. SOUTH CAROLINA.

Mountain Rest, 7.IX.1944, O.L. Cartwright, under cow dung, 1. Oconee

22 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

Com Lich watchery,.o+bX oso, is CCC Camp be, 2o.VETL.1937, 15. VIL.

1947, O.L. Cartwright, 3. Sassafras Mtn, Rocky Bottom, 4.IX.1934, O.L.

Cartwright, 1. TENNESSEE. Burrville, 16.X.1954, H.F. Howden, ex malt,

3. Chilhowee Mts., 3.X.1953, H. & A. Howden, ex malt and propionic acid

trap, 8. Great Smokey Mt. Nat. Pk., 6.VIII.1955, H. & A. Howden, ex

malt, 1. VIRGINIA. Church Falls, 19. VII.1919, E.A. Chapin, ex

Lactarius piperans, 1. Lee Co., Jone's Cove, Hubbard & Schwarz, 2. Speci-

mens are in the following collections: FMNH, Chicago; USNM, Washington;

CNC, Ottawa; Howden, Ottawa; MCZ, Cambridge; AMNH, New York; O.L.

Cartwright.

REMARKS. There is little variability between the specimens examined

of this species, other than for the depth and density of punctation of the longi-

tudinal trench of the metasternum. The only specimen in the Leconte Collec-

tion identified as O. pluricostatus is not in very good condition, but appears

to fit Marseul's description (1856) and that of Leconte (1845), and was there-

fore chosen as lectotype.

8. Onthophilus wenzeli new species

Figures 25, 45, 58, Map 3

Onthophilus wenzelin. sp. TYPE. Holotype, female, Nebraska (salt basin),

27.X.1923, Owen Bryant, (USNM, Washington).

DIAGNOSIS. This species may be distinguished from other species with

equally well developed elytral costae, by the well developed pronotal costae

PC 2 and PC 3, and by the fact that the pronotal costa PC lis not strongly and

entirely undercut along its inner edge. The pronotal costae of intermixtus are

well developed, but it is smaller in size and the meso- and metasternum are

densely punctate. The metasternum of wenzeli has two large discal areas

that are punctulate.

FEMALES. Length: 3.4 mm; width: 2.4mm. Head with VFC reduced

to a tubercle between antennae. LFC strong at antennae; attenuated in the

middle; meeting above clypeus, forming an elevation that reaches the clypeal

margin. Punctures round and irregular in size; much finer and denser above

antennae and on vertices of elevations. Labrum finely punctate; setiferous

punctures in lateral 0.25. Pronotum 0.50 times as long as wide. Lateral

margins strongly angulate and reflexed; parallel in hind 0.50, convergent in

front 0.50; lateral bead prominent; front angles acute. Disk 6-costate. PC1

long and straight; reaching from front 0.3 to hind margin. PC 2 subsinuate

and entire from front to hind margin. PC 3 slightly curved inward; reaching

from front margin to slightly before hind margin. PC 2 and PC 3 closer to-

gether than the two PC 3. Surface between PC 1 and PC 2 depressed. Punc-

tures medium size; separated by their diameters; slightly elongate on disk,

rounder along lateral bead. Elytron broader than pronotum, narrower behind.

All elytral costae equally strong and entire. SC attenuated before front mar-

gin. Costae with two rows of minute punctures on vertices. Interspaces

‘between with strong stria bordered by two low carinae; strial punctures

strong, more prominent towards front margin. EC 1 slightly attenuated in

front 0.30 to 0.45. ESC strongly sinuate; punctate in hind 0.75. Propygidium

with PMC strong in hind 0.75. PLC 0.65 times as long; vertex smoothly,

broadly rounded. Punctures medium sized and deep along front margin,

becoming smaller to minute and sparser towards hind margin and along

vertices of elevations. Pygidium with weak, bisinuate TC in front 0.30.

Surface depressed between front margin and TC. Punctures medium size,

Histeridae: Nearctic Onthophilus ZB

round, separated by their diameters along front margin and on discal area,

becoming small to minute along lateral margins and on vertex of TC. Pro-

sternum slightly depressed between procoxae. Hind margin deeply, sinuously

emarginate. Punctures medium size, round and separated by their diameters,

with minute punctures on interspaces and on prosternal lobe. Mesosternum

short and transverse. Front margin lobed to fit prosternal emargination;

strong, bisinuous. Metasternum separated from mesosternum by clear suture.

Median longitudinal suture faint. Surface depressed along midline in front

0.10. Punctures along midline and front margin dish-like with medium size,

round, deep punctures inside front edge; separated by their diameters; behind

mesocoxae and lateral areas irregular and confluent; on interspaces and on

two large triangular discal areas minute. First abdominal sternum with

punctures similar to those on front 0.2 of metasternum along front edge and

lateral areas; large semicircular median area that is punctulate. Protibia

with fifteen slender spicules, each on a weakly elevated base.

MALES. Unknown.

SPECIMENS. Known only from the holotype.

REMARKS. The pronotum of this species is very similar to that found

in the alternatus group. The lateral edge is strongly reflexed and the surface

is depressed between PC 1 and PC 2, and the pronotal costae are very strongly

developed. At present the range of this species is restricted but further

investigations of small mammal burrows will undoubtedly reveal that it has a

much wider range. This species is named for R.L. Wenzel in recognition of

his assistance to me and other students of the Histeridae.

9. Onthophilus cynomysi new species

Figures 15, 26, 37, 46, 59, Map 3

Onthophilus cynomysin. sp. TYPE. Holotype, male, Oklahoma, Grady Co.,

4 mi. N. Blanchard, 17. VIII.1953, M.J. Wilcomb, (FMNH, Chicago).

Allotype, female, with the same data as the holotype, (FMNH, Chicago).

Paratypes, 5 males and 5 females with the same data as the holotype,

(FMNH, Chicago; Howden, Ottawa).

DIAGNOSIS. This species resembles wenzeli, lecontei and soltaui in

general outline. All four species have distinct pronotal costae two and pro-

notal costae three. The pronotal costa one in cynomysi is undercut along the

entire length of the inner edge, while it is undercut only in the hind 0.2 in

wenzeli. The pronotal costae two of cynomysi are convergent behind while

they are straight or divergent behind in the other three species. The strongly

and abruptly depressed surface between pronotal costa one and the lateral

margin of cynomysi serves to separate this species from lecontei and soltaui.

MALES. Length: 2.5 to 3.1mm; width: 2.1lto 2.5mm. Head with

VFC short, indistinct; often obscured by encroaching punctures. LFC meeting

above clypeus; attenuated above clypeus, often indistinct. Surface depressed

between the two LFC. Punctures round, medium sized, and separated by

0.5 to1.0 times their diameters, except finer and setiferous on the vertices

of the elevations. Labrum with two setiferous punctures in lateral 0.25, the

setae long and prominent; the rest of the surface with much shorter setae

that are separated by their lengths. Pronotum transverse, 0.5 times as long

as wide. Lateral margin broadly angulate at midpoint; lateral bead prominent

with inner groove. Front margin shallowly, roundly, emarginate; front angles

slightly obtuse. Disk 6-costate. PC 1 slightly subsinuous; reaching from

front 0.25 to hind margin; surface along outer edge smoothly, quite abruptly

24. Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

depressed, then abruptly reflexed along lateral margin; inner edge strongly

undercut along its entire length; surface between PC 1 and PC 2 depressed,

forming a shallow trench that narrows strongly towards the front. PC 2 sub-

sinuous; reaching from front 0.1 to just before hind margin; attenuated in

front 0.5, higher and broader behind; obscured in front and behind by punc-

tures. PC 3 curved weakly outward; reaching from just behind front margin

to hind 0.25; obscured before and behind by punctures; the two PC 3 conver-

gent behind. Punctures medium sized and separated by their diameters;

round to oval on lateral and hind areas, more elongate in trench and on

central areas. Elytron wider than pronotum. All costae equally well develop-

ed. SC slightly attenuated in front 0.3; two confluent and irregular rows of

punctures just below the vertex, that are more prominent in hind 0.5. All

other costae with two rows of minute punctures that bear short setae in hind

0.5. Interspaces between costae with a strong stria that is bordered by two

low carinae; strial punctures much deeper at front margin. Propygidium

with PMC broad, rounded, strongest just before hind margin; surface depress-

ed behind hind end of PMC. PLC indistinct, broad swelling. Punctures

medium size, deep, round and separated by their diameters, except on vertex

of median elevation, punctures about 0.5 as large and elongate. Pygidium

surface without carinae. Front margin reflexed; discal area convex. Punc-

tate throughout; punctures similar to those on propygidium. Prosternum

generally convex, with slight concavity between procoxae. Hind margin with

deep, cuspidal emargination. Punctures round, medium size, deep and

separated by 0.5 to 1.0 times their diameters. Mesosternum short and trans-

verse. Front margin with a cusp to fit prosternal emargination. Punctures

twice as large as those on prosternum, contiguous or separated by 0.5 times

their diameters. Metasternum separated from mesosternum by clear suture.

Median longitudinal suture fine and clear. Median area weakly concave.

Punctures along front, hind and lateral margins like those on mesosternum;

becoming finer on median discal areas. First abdominal sternum with punc-

tures along front margin and on lateral areas similar to those on front third

of metasternum; semicircular central area punctulate. Protibia with a row

of ten to twelve spicules, on elevated bases. Male genitalia illustrated in fig.

eS.

FEMALES. Length: 3.0 to 3.6 mm; width: 2.5to3.0 mm. Except

for sexual characters the females differ very little from the males. On the

average females are slightly larger than males; the outer edge of the lateral

pronotal trench reaches the front 0.3; the metasternum is not depressed along

the midline and the protibial spicules are stronger.

SPECIMENS. This species is known only from the type material.

REMARKS. There is very little variability within this species. The

facial carina, pronotal costae and the PMC of the propygidium are less dis-

tinct in some specimens than in others. The protibial spicules are broken or

worn on older specimens.

Virtually nothing is known about the biology or habits of this species.

All the material seen was collected by M.J. Wilcomb during his study of

prairie dog burrows. The conditioned soil was scraped from the tunnels and

small arthropods were extracted with Berlese funnels. The name cynomysi

does not mean to imply that this species is characteristic of, or confined to

Cynomys burrows, but merely that this is the niche from which it was taken.

Histeridae: Nearctic Onthophilus 25

10. Onthophilus soltaui Casey

Figures 27, 47, 60, Map 1

Onthophilus soltaui Casey, 1893: 557. TYPE. Holotype, sex undetermined,

bearing the following labels -- ''Col.'', 'Denver 10/1890", "Casey

bequest 1925"", "soltaui Csy", red "TYPE USNM 38463"', (USNM,

Washington), not seen.

DIAGNOSIS. This species is very similar to lecontei and cynomysi.

Characters that separate soltaui from them are dealt with in the diagnoses for

lecontei and cynomysi.

FEMALES. Length: 3.5 mm; width: 2.5mm. Head with VFC faintly

indicated. LFC faint, barely discernable above clypeus. Punctures between

the two LFC, and between LFC and lateral margin, large, round, shallow

and separated by 0.5 to 1.0 times their diameters; above clypeus, on vertex,

and on vertices of elevations strong, fine and dense. Labrum with setiferous

puncture in lateral 0.25; shorter setae along front 0.5 separated by their

lengths. Pronotum 0.6 times as long as wide. Lateral margins straight and

subparallel to middle, then abruptly angulate, strongly convergent to front

angles; lateral bead prominent, abruptly reflexed; front angles slightly obtuse,

scarcely rounded. Disk 6-costate. PC 1 prominent and smoothly arcuate

outwards; reaching from front 0.25 to hind margin; outer edge indistinct, not

abrupt, surface smoothly continuous from vertex of PC 1 to lateral bead;

inner edge very abrupt, deeply undercut and shining; surface between PCl1 and

PC 2 depressed, forming a broad trench. PC 2 straight and faint; reaching

from front 0.25 to hind 0.25. PC 3 faint; weakly curved inward in front 0.1,

then straight; reaching from front margin to hind 0.3. Punctures large,

shallow and separated by approximately their diameters; round between

median costae; elongate between PC 1 and PC 2 forming strigose pattern; less

dense to lacking along lateral margin. Elytron wider at base than pronotum.

Costae strong and even. SC attenuated in front 0.3. Vertices of costae with

double row of fine setiferous punctures. Interspaces between costae with two

fine, lateral carinae bordering a clear stria; strial punctures shallow, strong-

est at front margin. External subhumeral stria more coarsely punctate; area

between it and lateral side coarsely punctate. Marginal stria indistinctly

defined. Propygidium with PMC complete from front 0.2 to hind margin; in-

distinct in front; forming distinct tuberculate elevation behind. PLC indistinct.

Punctures medium, round, shallow and separated by 1.0 to 1.5 times their

diameters; punctures more distinct along their front edges. Pygidium with

surface strongly convex. Front edge reflexed with distinct depression in

front angles. Punctures similar to those on propygidium, but separated by

0.25 to 1.0 times their diameters; denser along hind margin. Prosternum

strongly convex before procoxae, flat behind. Hind margin quite deeply,

obtusely emarginate. Punctures medium, round, shallow and separated by

their diameters; interspaces, hind margin and lateral margin behind procoxae

with minute, round punctures. Prosternal lobe with small and minute punc-

tures densest along midline, becoming sparser laterally. Mesosternum short

and transverse. Front margin bisinuate; somewhat cuspid lobe fitting into

prosternal emargination. Punctures medium, round, shallow and separated

by 0.5 times their diameters. Metasternum separated from mesosternum by

a clear suture. Median longitudinal suture fine. Punctures in front 0.3 and

along hind margin large, round, shallow and separated by 0.5 times their

diameters; becoming irregular and contiguous along lateral margin; inter-

spaces and central areas with dense, minute punctures. First abdominal

26 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

sternum covered throughout with punctures similar to those on front 0.3 of the

metasternum. Protibia with nine spicules mounted on elevated bases.

MALES. Unknown.

SPECIMENS. UNITED STATES. COLORADO. Denver, X.1890, H.

Soltau, 1female, (USNM, Washington).

REMARKS. Although soltaui is very similar to cynomysi and lecontei

there appear to be sufficient differences to retain it as a good species. Since

soltaui is known from only two specimens the ultimate decision as to whether

these differences are species differences, or merely represent extremes of

a clinal variation, must be reserved until more material has been collected.

S. Peck has kindly compared my specimen with the type. No significant

differences were perceived.

11. Onthophilus lecontei Horn

Figures 17, 28, 48, 61, Map 4

Onthophilus lecontei Horn, 1870: 138. TYPE. Holotype, sex undetermined,

bearing the following labels -- "Ca."', red "HOLOTYPE 3237",

"Onthophilus lecontei Horn", type locality Fort Tejon, California, (MCZ,

Cambridge).

DIAGNOSIS. QO. lecontei is very similar in its genera] outline to

soltaui, wenzeli and cynomysi. For characters that separate the last two

from lecontei refer to the diagnoses for these two species. O. lecontei may

be separated from soltaui by the characters given in the key. There are other

comparative characters which separate these species. The lateral bead is

not as robust in soltaui, the lateral margin is more strongly reflexed, PC 1

is more strongly undercut before the hind margin, the trench between PC 1 and

PC 2 is deeper, and the sutural costa has an irregular double row of small

punctures separating it from the suture. These punctures are less dense and

often quite indistinct in lecontei.

MALES. Length: 2.5 to 3.0 mm; width: 1.9 to 2.1mm. Head with

VFC small, low tubercle. LFC attenuated before clypeus; the LFC conver-

gent but not meeting. Surface between antennae to occipital carina depressed;

clypeo-frontal surface elevated along midline. Punctures large, round and

contiguous in frontal area and on genae; punctures much smaller on vertex

and clypeo-frontal elevation. Labrum with setiferous punctures in lateral

0.25. Pronotum 0.5 times as long as wide. Lateral margin smoothly arcuate

or with slight angles at midpoint; lateral bead very strong with inner depre-

ssion deep and extending along front margin. Front margin smooth, deep,

round emargination; front angles 90°. Disk 6-costate. PC 1 slightly curved

outward; reaching from front 0.25 to hind margin; outer edge indistinct, not

abrupt, surface continuous from vertex to lateral margin; inner edge abrupt,

strongly undercut along its entire length; surface between PC 1 and PC 2

depressed, forming a shallow trench that reaches the front angles. PC 2.

very weakly subsinuate; attenuated before hind and front margin. PC 3

straight; attenuated before hind margin; strongest at front margin. Punctures

medium, round along lateral edge and between PC 2 and PC 3; more elongate

in depression between PC 1 and PC 2 and between the two P@3; punctures

encroaching on PC 2 and PC 3 in hind 0.25. Elytron wider than pronotum.

All costae equally well developed and entire. SC attenuated in front 0.2.

All costae with a row of minute punctures on each side just below the vertex,

that bear minute setae in the hind 0.3. Spaces between costae with a strong

stria that is bordered on each side by a low carina; strial punctures oval and

Histeridae: Nearctic Onthophilus 27

deep, with small tubercle on each side that interrupts the carinae; first punc-

ture behind the front margin especially deep; punctures in the mid-section of

stria one, two and three slightly deeper; minute punctures between costae and

carinae. Propygidium with lateral margin angulate in front 0.2. PMC indis-

tinct; reaching from front margin to hind 0.3; higher and broader in hind 0.5.

Surface weakly depressed behind PMC. Punctures medium size, and elongate;

interstices arranged in strigose pattern. Pygidium with no carinae; surface

convex with transverse depression behind front margin. Punctures medium,

round and separated by their diameters; becoming finer towards hind margin.

Prosternum weakly depressed between procoxae, convex in front. Hind mar-

gin with obtuse, somewhat rounded, deep emargination. Punctures medium,

round and separated by 1.0 to 2.0 times their diameters; much smaller punc-

tures on interspaces. Prosternal lobe with small punctures; surface strigu-

late. Mesosternum short and transverse. Front margin lobed to fit pro-

sternal emargination. Punctures similar to those on prosternum with at least

two complete rows across. Metasternum separated from mesosternum by a

fine suture. Median longitudinal suture very fine. Surface distinctly concave

behind front margin. Large dish-like punctures with small deep punctures

inside their leading edge, in front 0.5, along midline, along hind margin and

on lateral areas; interspaces of large punctures and discal areas with minute

punctures. First abdominal sternum with large punctures along front margin

and on lateral areas; large semicircular median area punctulate. Protibia

with six to eight strong spicules each on an elevated base. Male genitalia

illustrated in fig. 17.

FEMALES. Length: 3.2 to 3.7 mm; width: 2.4to2.7 mm. The

females differ from males very little except for sexual characters. They are

on the average larger, the metasternum is not depressed and the protibial

spicules are slightly stronger.

SPECIMENS. UNITED STATES. CALIFORNIA. Atascadero, 8.II.

1940, E.S. Ross, ex gopher burrow, 1 male, 1 female. Contra Costa Co.,

Danville, 5-8.XII.1951, F.X. Williams, ex Vespula pennsylvanica nest or

debris; 1 male, Ifemale; Oeil. 1952, 28=30.111. 1952, 17.bV.1952) FX.

Williams, ex gopher burrow, 15. Monterey Co., Carmel, 5.III.1914, 27.

[12915909 <1 ot9y Aorta l 7 aS.L. Slevin, 4; Monterey, 20. XL. 1925,

S.L. Slevin, 1. Pasadena, XII, A. Fenyes Coll., 1 male, 1female; II, A.

Fenyes Coll., 1female. Salada Beach, 21.I1V.1928, F.E. Blaisdell, 1

male. San Diego Co., Poway, F.E. Blaisdell, 2. Santa Barbara, 22.XII.

1928, F.E. Blaisdell, 1female. Santa Cruz, 21.V.1921, S.L. Slevin, on

dandelion, l1male. Tracy, 5.1I1.1939, 1female. (Cal. Acad., San

Francisco; Howden, Ottawa)

REMARKS. There is not very much variation expressed in the speci-

mens of this species. The trench between PC 1 and PC 2 is less well develop-

ed and the lateral margin is weakly angulate in some specimens. These

differences do not appear to be clinal, but are expressed irregularly through-

out the known range. O. lecontei has been recorded only from the coastal

areas of California, and is apparently well isolated today from other related

species.

12. Onthophilus thomomysi new species

Figures 18, 29, 36, 49, 62, Map 4

Onthophilus thomomysin. sp. TYPE. Holotype, male, MEXICO. Durango.

10 mi. W. El Salto, 4. V1.1964, H.F. Howden, ex Thomomys burrow,

28 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

(CNC, Ottawa). Allotype, female, with the same data as holotype,

(CNC, Ottawa). Paratypes, 11 males and 7 females with the same data

as holotype, (CNC, Ottawa; Howden, Ottawa).

DIAGNOSIS. This species may be distinguished from the other species

with equally well developed elytral costae, except kirni, by the absent or

almost indistinguishable pronotal costae PC 2 and PC 8. For characters that

separate thomomysi from kirni, refer to the diagnosis for the latter. Some

specimens of thomomysi may be confused with very worn specimens of

lecontei in which the pronotal costae PC 2 and PC 3 may be very low. These

species may be separated by the pronotal punctures. The punctures of

thomomysi are small, round and separated by several times their diameters.

The punctures of lecontei are medium sized, oval and separated by their dia-

meters. |

MALES. Length: 3.2 to 3.4 mm; width: 2.6to2.7 mm. Head with

VFC and LFC obsolete; indicated by broad, rounded elevations. Punctures

on frons, vertex and above antennal sockets small, round and separated by

several times their diameters; larger and sparser on genae. Labrum with

setiferous punctures in lateral 0.25; surface densely punctulate. Pronotum

0.5 times as long as wide. Lateral margin weakly, broadly angulate; lateral

bead prominent, with shallow inner groove, strongest in hind 0.5. Front

margin deeply roundly emarginate; front angles 90 . Disk 2-costate. PC 1

strong, arcuate outward; reaching from front 0.25 to hind margin; outer edge

smoothly continuous with surface between it and lateral margin; inner edge

abrupt, strongly undercut and shining; surface mesad of PC 1 depressed,

forming a shallow trench that runs the length of PC 1 and curves to the front

angles. PC 2 and PC 3 entirely absent or sometimes faintly indicated by

more shining and very weakly elevated areas. Punctures round, small on

disk, larger along lateral margin, in lateral trench and along the hind margin;

separated by two or more times their diameters. Elytron wider than prono-

tum. All costae equally well developed. SC attenuated in front 0.1. EC 3

slightly lower just behind front margin. All costae with two irregular rows

of minute punctures, one on each side just below the vertex, that bear short

setae in the hind 0.4. Spaces between costae with two low carinae and a

strong, clear stria between them; strial punctures oval, deep and with small

lateral tubercles that interrupt the carinae; strial punctures much stronger

and deeper at the front margin, and slightly deeper in the midsection of stria

one and stria two. Propygidium with lateral margin obtusely angulate in

front 0.3. PMC and PLC faint, broad and low; more prominent in hind 0.6.

Punctures along front margin small, round and separated by 2.0 to 5.0 times

their diameters, gradually becoming smaller and sparser, almost absent

from middle to hind margin. Pygidium with TC faint, broad, low and sinuate,

its midpoint almost reaching front margin, its lateral ends in front 0.25.

Punctures throughout small and round; separated by 2.0 to 4.0 times their

diameters. Prosternum depressed between procoxae. Hind margin with

deep, cuspid emargination. Punctures small, round and sparse; punctulate

in front of hind margin. Prosternal lobe impunctate. Mesosternum short

and transverse. Front margin lobed to fit prosternal emargination. Two

transverse rows of small, round punctures separated by 2.0 times their dia-

meters. Metasternum separated from mesosternum by a fine suture. Median

longitudinal suture fine. Midline and large discal areas punctulate. Punc-

tures in front and hind 0.3 small and round, separated by 2.0 to 4.0 times

their diameters; punctures in lateral 0.5 large, shallow, somewhat irregular

and dish-like, with small deep punctures within their front edges, confluent

to separated by their diameters. Surface depressed behind front margin.

Histeridae: Nearctic Onthophilus Zo

First abdominal sternum with punctures along front margin and on lateral

areas similar to those along front margin of metasternum; large, semicircu-

lar central area that is punctulate. Protibia with eight spicules each on an

elevated base. Male genitalia illustrated in fig. 18.

FEMALES. Length: 3.5 to 3.6 mm; width: 2.7to2.8 mm. The

females are very similar to the males. The metasternum is not depressed

and there are two rows of punctures similar to those along the front margin,

along the longitudinal midline.

SPECIMENS. Type series. NEW MEXICO. Engle, 1(MCZ, Cambridge).

REMARKS. Very little variability was noted, other than for the slight

differences in the development of the pronotal costae. All specimens were

heavily covered by a hard accretion of waxy exudate and adhered soil particles.

As with cynomysi, the name thomomysi does not intend to imply that

this species is characteristic of, or confined to the burrows of Thomomys,

but it is proposed merely to indicate the niche from which this species was

first collected.

13. Onthophilus kirni Ross

Figures 19, 30, 50, 68, Mapl

Onthophilus kirni Ross, 1944b:115. TYPE. Holotype and allotype, Texas,

seven and one-half miles S.E. Somerset, 18.11.1943, A.J. Kirn,

collected in a bait trap in a Geomys burrow, (Cal. Acad., San

Francisco). Not seen. Paratypes, numerous males and females

collected at the above locality in baited traps and by excavating Geomys

burrows by A.J. Kirn and E.S. Ross, (deposited in numerous collec-

tions).

DIAGNOSIS: The males and females of this species can be readily dis-

tinguished from other species in which the elytral costae are equally well

developed by the following combination of characters. The pronotal costae

PC 2 and PC 3 are entirely obsolete. The third elytral costa is abbreviated

in front by a short, deep, transverse trench that joins stria three and stria

four. The metasternum is densely and finely punctate.

MALES. Length: 2.8 to 3.0 mm; width: 1.9 to 2.2 mm. Head with

VFC and LFC poorly indicated by broad, low elevations. Punctures round,

medium sized, slightly larger on frons and genae than on vertex, and separ-

ated by 0.5 to 1.0 times their diameters. Labrum with setiferous punctures

along front margin; one long seta in lateral 0.25; other setae much shorter

and separated by approximately their lengths. Pronotum 0.75 times as long

as wide. Lateral margin very broadly, obtusely angulate, somewhat rounded;

lateral bead prominent with a deep inner groove along the entire length of

side, deepest at front angles. Front margin shallowly, roundly emarginate;

front angles almost 90°. Disk with two broad trenches in lateral 0.25;

trenches shallow, broader at their hind end, narrowing towards the front,

reaching front 0.3; inner edge poorly indicated; outer edge equivalent to PC 1,

abrupt, undercut, deepest at the hind margin, curved and shining. Punctures

medium sized and separated by 0.5 to 1.0 times their diameters; round and

densest between lateral margin and trench; elongate in trench and along

longitudinal midline. Elytron with all costae approximately equally well

developed, broad and low; EC 3 abbreviated before front margin by a short,

deep, transverse fossa that joins stria three and stria four; EC 2 and EC 4

slightly broader and higher at front margin than other costae; SC slightly

lower than other costae, separated from suture by a well defined, irregular

30 Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

row of small punctures; all costae with a double row of minute punctures that

are indistinct in the front 0.5 and have minute yellow setae in the hind 0.3.

Interspaces between costae with two lower carinae that border a clear stria;

strial punctures not distinct except along front margin and in hind 0.3.

Propygidium with front margin reflexed. PMC reaching from front margin to

just before hind margin; tear-shaped, narrow and low in front, broader and

higher behind. PLC low and indistinct. Punctures oval and medium sized,

separated by 0.5 to 1.0 times their diameters; leading edge of punctures

better defined than trailing edge; punctures finer and sparser on vertex of

PMC. Pygidium as long as wide. Surface convex; front edge most abrupt,

forming a broad, transverse channel just behind the front margin. Punctures

round and the same size as those on the propygidium; deeper and larger in

the transverse channel. Prosternum very slightly concave between procoxae.

Hind margin roundly emarginate. Punctures similar to those on the head,

but with minute yellow setae arising from them in the hind 0.5; punctures on

prosternal lobe smaller. Mesosternum short and transverse. Front margin

lobed to fit the prosternal emargination. Punctures similar to those on hind

0.5 of prosternum. Metasternum separated from mesosternum by a clearly

defined suture. Median longitudinal suture strong and fine. Surface shallowly

depressed behind front margin. Punctures on the central area like those on

hind 0.5 of prosternum; larger and very irregular on lateral 0.25. First

abdominal sternum punctate throughout with punctures similar to those on the

central area of metasternum. Protibia with eight spicules, each on an ele-

vated base. Male genitalia illustrated in fig. 19.

- FEMALES. Length: 3.0 to 3.6 mm; width: 2.4to 2.6 mm. The

females are very similar to the males, except that the metasternum is not

depressed, and the protibial denticules are larger, more numerous and

extend almost to the base.

SPECIMENS. I have seen a series of forty paratypes from California

Academy of Sciences, San Francisco.

REMARKS. E.S. Ross and A.J. Kirn encountered kirni in large

numbers in the nesting chambers of Geomys breviceps attwateri. As many

as nineteen were found in one nest. In total they collected 115 specimens by

digging out burrows and by bait traps (Ross, 1944a, b).

14. QOnthophilus giganteus new species

Figures 65, 66, 67, 68, Map 5

Onthophilus giganteus n. sp. TYPE. Holotype, male, FLORIDA. Alachua

Co. 1mi. W. Archer, 15-19 .1.1976, J.R. Diley, ex malt trap

(FDACS, Gainesville).

DIAGNOSIS. This species may be separated from all other species by

the following combination of characters: size over 4 mm; very strongly

broadened front ends of elytral costae EC 2 and EC 4; very large and deep

strial punctures just behind the front margin of the elytra.

MALES. Length: 5.6 mm; width: 4.1mm. Head with VFC well

developed, reaching from occipital carina to frons between antennae. LFC

present near antennal sockets, obsolete elsewhere. Punctures minute, deep,

separated by several times their diameters; denser on vertex and above

clypeus. Labrum with minute, dense punctures; setiferous punctures in

lateral 0.25. Pronotum transverse, 0.5 times as long as wide. Lateral

margin arcuate, very strongly reflexed; lateral bead prominent. Front mar-

gin broadly, roundly emarginate; reflexed from front angles to PC 2. Disk

Histeridae: Nearctic Onthophilus on

6-costate. PC 1long, strongly angulate in front 0.2; reaching from front 0.2

to hind margin. PC 2 sinuate, broader behind; reaching from front margin

to just before hind margin. PC 3 weakly indicated by broad elevations,

slightly curved outward; reaching from front margin to hind 0.2. Surface

very convex, very deeply depressed between PC 1 and PC 2; depression broad

behind, narrowing towards front margin. Punctures on disk small to minute

intermixed, sparser on costae and along lateral margin, depression impunc-

tate. Elytron wider than pronotum. EC 2 and EC 4 slightly higher, much

more robust at front margin than ESC, ISC, EC 1, EC 3, EC 5andSC. EC 3

interrupted before front margin by short, deep, transverse fossa between EC

2 and EC 4; SC attenuated in front 0.5. Strial punctures oval, deeper, more

pronounced towards hind margin; stria two, three, four and five with extreme-

ly large, deep punctures just behind front margin. Punctures on vertices of

costae minute, oval, sparse. Propygidium with PMC strong, tear-shaped;

reaching from front 0.5 to hind margin. PLC as long as, and converging

towards PMC behind. Punctures medium, round, deep, separated by 2 to 3

times their diameters; contiguous just behind front margin; punctures on

vertices of elevations minute. Pygidium with LC obsolete in middle 0.3;

strong in hind 0.3; low, bicarinate in front 0.3. TC indicated by broad, low

elevations. Punctures similar to those on propygidium. Prosternum slightly

depressed between procoxae. Hind margin broadly, roundly emarginate.

Punctures similar to those on propygidium, denser along lateral margin and

behind; interstices of large punctures and prosternal lobe punctulate. Meso-

sternum short, transverse. Front margin lobed to fit prosternal emargina-

tion. Punctures slightly larger than those on prosternum; contiguous or fused

laterally. Metasternum separated from mesosternum by a fine, distinct

suture. Median longitudinal suture fine, distinctly impressed. Surface with

broad, median depression in front 0.3. Punctures along hind margin like

those on mesosternum; laterally large, irregular, shallow, confluent; fused

into broad, deep, punctate fossa along inner margin of lateral metasternal

stria; punctures on disk and interstices of large punctures sparse and minute.

First abdominal sternum with front margin reflexed. Punctures medium,

deep, sparse behind front margin; forming short fossa behind metacoxae;

large, shallow, irregular, contiguous laterally; minute elsewhere. Protibia

with fourteen spicules, each on a weakly elevated base; apical three contigu-

ous. Male genitalia illustrated in fig. 68.

FEMALES. Unknown.

SPECIMENS. This species is known only from the holotype.

REMARKS. OQ. giganteus is by far the largest species of Onthophilus

yet described. It is nearly 1.5 mm longer than the next largest and nearly

four times longer than the smallest. Apparently it belongs near nodatus,

but the elytral costae are not as unequally developed.

ACKNOWLEDGMENTS

Specimens were borrowed from a number of institutions in Canada and

United States. I would like to thank the following institutions for allowing me

to examine their material, and the following individuals who are responsible

for the loans: J.E.H. Martin, Canadian National Collection of Insects (CNC),

Ottawa; L.H. Herman, Jr., American Museum of Natural History (AMNH),

New York; J.M. Kingsolver, United States National Museum (USNM),

Washington; D.H. Kavanaugh, California Academy of Science (Cal. Acad.),

San Francisco; J. F. Lawrence, Museum of Comparative Zoology, Harvard

University (MCZ), Cambridge; D.C. Rentz, Academy of Natural Science of

ac Contrib » Amend Aunts Instse voleid;- nods; 1978

Philadelphia (ANSP), Philadelphia; R.L. Wenzel, Field Museum of Natural

History (FMNH), Chicago.

| I thank Dr. G. Carmody and Dr. H.H.J. Nesbitt for reading the pre-

liminary manuscript, for their advice and criticism. I would like especially

to thank Dr. R.L. Wenzel for his time and his assistance, and Dr. H.F.

Howden for allowing me to use specimens from his private collection, for his

guidance, criticism and forbearance.

I am indebted to Mr. L.E.C. Ling for the scanning electron microscope

pictures, and to my wife, Maureen, for her assistance with the typing. This

work was partially supported by an operating grant from the National Research

Council of Canada to H. F. Howden.

LITERATURE CITED

1958. Evolution of the Madro-Tertiary geoflora. Bot. Rev. 24: 433-

509:

Balduf, W.V.

1935. The Bionomics of Entomophagous Coleoptera. 8. Histeridae.

John S$. Swift Co., Inc. 220 pp.

Bedwell, E.C.

1907. Onthophilus sulcatus F., ina moles nest. Ent. Monthly Mag.

43: 62.

Bickhardt, H.

1910. Coleopterorum Catalogus, 8, Pars 24, Histeridae. ed. S.

Schenkling. W. Junk, Berlin. 137 pp.

Casey, -T...L.

1893. Coleopterological Notices V. Ann. N.Y. Acad. Sci. 7: 553-578.

Handschin, E.

1944. Insekten aus den Phosphoriten des Quercy. Verlag Birkhauser,

AG, Basel.

Horn, ;Gai.

1870. Description of new species of Histeridae of the United States.

Trans. Amer. Ent. Soc. 3: 134-142.

1873. Synopsis of the Histeridae of the United States. Proc. Amer.

Philos. Soc. 13: 273-360.

Leach, G. |

1817. XXI. A sketch of the characters of the stirpes and genera of

the family Histeridae. Zool. Miscell. 3. London. 151pp.

Leconte, J.E. .

1845. A monograph of the North American Histeroides. Bost. Jour.

Nat. Hist. 5: 32-86.

Lewis, G.

1888. Histeridae: in Godman and Slavin, Biol. Cent.-Amer. Ins.

Coleoptera. 2 (1): 182-244.

1891. A new genus of Histeridae. Ent. Monthly Mag. 11: 319.

1892. Ona new Onthophilus from Mexico. Ent. Monthly Mag. 28:124.

1905. A Systematic Catalogue of Histeridae. Taylor and Francis,

London. 81 pp.

Lindroth, C.H. and Palmen, E.

1959. Coleoptera. in Taxonomist's Glossary of Genitalia in Insects,

ed. S.L. Tuxen. Ejnar Munksgaard, Copenhagen. 284 pp.

Histeridae: Nearctic Onthophilus 33

Marseul, S.A. de

1856. Essai monographique sur la famille des Histérides. XLC Genre,

Onthophilus. Ann. Soc. Ent. Fr. (3) 4: 549-566.

Reichardt, A.

1933. Uebersicht der Palaearktischen Arten der Gattung Onthophilus

Leach (Col. Hister.). Sbornik. Entom. odd. Nar. Musea v. Praze

11: 137-144.

Ross, E.S.

1944a. Arthropod collecting in the burrows of Texas pocket-gopher.

Ent. News 55: 57-61.

1944b. Onthophilus kirni new species and two other noteworthy

Histeridae from burrows of a Texas pocket-gopher. Ent. News

55: 115-117.

say, T.

1825. Descriptions of new species of Hister and Hololepta inhabiting

the United States. Jour. Acad. Nat. Sci. Phila. 5: 32-47.

Wenzel, R.L.

1944. On the classification of the histerid beetles. Field Mus. Nat.

Hist. Zool. Ser. 28: 51-151.

Wenzel, R.L. and Dybas, H.S.

1941. New and little known neotropical Histeridae (Col.). Field Mus.

Nat. Hist. Zool. Ser. 22 (7): 483-472.

Westwood, J.O.

1840. Introduction to Modern Classification of Insects... . VolIIT,

Synopsis of the Genera of British Insects. London.

34 Contrib. Amer: Ent. Inst.; vols 15, no. 5, 1978

0.2 mm

2 8

1.0 mm

Figs. 1-9. Onthophilus spp. 1-5. alternatus, male. 1. anterior

view of left maxilla; 2. anterior view of left mandible; 3. ventral view of

labium; 4. antenna; 5. right hind wing; 6. flohri, female right hind wing;

7-8. alternatus, male. 7. penis; 8. median lobe of penis; 9. flohri,

female styli of ovipositor.

Histeridae: Nearctic Onthophilus

=D >,

0.5 mm

Figs. 10-19. Onthophilus spp. male genitalia (lateral view of

aedeagus, ventral view of 9th sternite, dorsal view of aedeagus). 10.

alternatus; 11. deflectus new species; 12. flohri; 13. pluricostatus; 14.

intermixtus new species; 15. cynomysi new species; 16. nodatus; 17.

lecontei; 18. thomomysi new species; 19. kirni.

Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

1 new

j Ulli;

wenzel

20.

view of head

pluricostatus;

28. leconte

soltau

ior

24.

S spp., anter

nodatus

ies; 27

ilu

Onthoph

20-28.

22. alternatus

- cynomysi new species

igs

flohr

species

Histeridae: Nearctic Onthophilus oT

Figs. 29-37. Onthophilus spp. 29-30. anterior view of head. 29.

thomomysi new species; 30. kirni. 31-37. posterior view of propygidium

and pygidium. 31. julii; 32. flohri; 33. alternatus; 34. nodatus; 35.

pluricostatus; 36. thomomysi new species; 37. cynomysi new species.

38 Contrib..<Amer..bnt.nsts, vols 15,6n0. 5, 1978

Figs. 38-46. Onthophilus spp., dorsal view. 38. julii; 39. deflectus

new species; 40. flohri; 41. intermixtus new species; 42. alternatus;

43. nodatus; 44. pluricostatus; 45. wenzeli new species; 46. cynomysi new

species.

Histeridae: Nearctic Onthophilus 3g

47. soltaui;

48. lecontei; 49. thomomysi new species; 50. kirni; 51-55. ventral view.

Figs. 47-55. Onthophilus spp. 47-50. dorsal view.

51. ju julii; 52. deflectus new species; 53. flohri; 54. intermixtus new species;

99. alternatus.

Contrib. Amer. Ent. Inst., vol. 15, no. 5, 1978

4.0

nodatus;

<<!

€) 6

D ..

. 2'o

Lo) a

Io EB:

Els

3 ao

MH 3)

e Slo

® »- Os

Aes

60S

O oO

nae

Be &

aod

Sic)

Sole

al ee

al2)a138

|. 21

Ole 615

Poa) |

e Vio,

dar

iol 1s

- Bl.

MN Ol Si a

OD =

mt ol SlB

Fy Siral

rt} Ol

Qul Dy ey

<* 3 10

tS nay

LD <On cy

Histeridae: Nearctic Onthophilus

LEW

ior v

(lateral

anter

1 65

male genita

species.

V1EW

gig teus new

ventral vi

ilu

OT;

Onthoph

View

dorsal

Figs. 65

of head; 66.

LEW

, dorsal v

VIEW

4.2, Contribs, Amer Hut. mists, volki15;me.05, 1978

Map 1-2. Distribution of Onthophilus spp.: 1. deflectus new species

(@) (O- state record); soltaui (a); kirni (*). 2. nodatus (@); julii (a);

flohri (yx).

Histeridae: Nearctic Onthophilus 43

Map 3-5. Distribution of Onthophilus spp.: 3. alternatus (@);

wenzeli new species (A- state record); cynomysi new species (%).

4. lecontei (@); intermixtus new species (A); thomomysi new species (x).

5. pluricostatus (@) (O - state record); giganteus new species (4).

ae SUPGENUS STEGDMYTA OF. AEDES

os. Pate papselaktead ee WITH .

Contributions

of the

American Entomological Institute

Volume 15, Number 6, 1979

—"

MEDICAL ENTOMOLOGY STUDIES - XI.

THE SUBGENUS STEGOMYIA OF AEDES

IN THE ORIENTAL REGION WITH

KEYS TO THE SPECIES

(DIPTERA: CULICIDAE).

Yiau-Min Huang

CONTENTS

AB STR eee le Se a ie eG eo aM cee Gk Oe ea eS 1

EN PRODUCTION oo. e oe a ee a a LE ge OR ee 1

SUBGENUS STEGOMY)A THEOBALD ais wee lee SR ee 3

CHARA Rte io es he ae a ee eee ew ee je 4

ie ee te cas rat at ree ee ee a ee a wis ie else wae &

PUN te oe eek OR iN ae RM go ab ae eee ee se eo 6

DESCRIPTIONS OF THE GROUPS OCCURRING IN THE ORIENTAL

PC ee ictal el ae away ek ee ea ek ee Se icas, Sonics 6

GOS PT SEOs, iu es, oe Be ee i AE ee, Ee OH, 6

TOO Te OL ON GA. ce a ee es ee ak a ee ee 7

CU OE ye ye) es a Gee A ay a ke ee ee 7

SCV POU 2 ce sa es ie ee Se he we ke es y

WOOO an a oe a, 6 6 hs ea we Rl Swe 8

DIS PRIBU TION heii o5 5 aetee rtrd nennnreauane OMe eh a et ieee ae Se ere 8

Oe cece ie le ee ee ee el 8

NIP DIC AD INIPOR TANG... sie gow ee ee ee eR ee Ne eee ees 9

KEYS TO THE GROUPS, SUBGROUPS AND SPECIES OF THE SUBGENUS

tee ele ce ea ea a) era 2k 6 kk a wR ea Be 12

ADULTS (NALESAND FEMALES) 36. Ad & aed E eek ee 12

Wit PEN da eG ea ew ws he WN ae ee we 16

ee se eee maar ee ae ea es om Se Wah ac es BE ae we we Ch 20

POS Pen a ey Ie or ek ee Tk we a ee es es 23

DO ee ey ae ata re gl SN yee A Ca GS OR ee eee a Oe Le 27

PONG) Wie PG RIE IN ies gene we a el ee a ee ee we ww 27

LITERATURE’ CITED. 3.0.5. sus Sear ees we ee Oa ee a 28

APPENDIX I. PRESENT STATUS OF THE SPECIES OF AEDES

OLTEGOMYVIA) IN THE ORIENTAL REGION. 6.060562 bis ee ee ee 36

APPENDIX II. DISTRIBUTION LIST OF THE ORIENTAL SPECIES OF

Pe OA es ig ee oe a a ee ECE ewe ee 38

APPENDIX III. SUMMARY OF THE IMMATURE HABITATS OF THE

SPECIES OF AEDES STEGOMYIA) IN THE ORIENTAL REGION. ... 40

APPENDIX IV. Aedes Stegomyia) aegypti (Linnaeus). .........-. 41

APPENDIX V. Aedes (Stegomyia) platylepidus Knight and Hull. ..... 44

MAP OF THE GRIER NTAL REGION 3 5 faa oe SY TASTE PRR. we 45

ite GO er a a ee Ss eae ew Ra ee 46

PAG URE gee es ee cealy ey e's p a h aR a aa ge we ee Se 47

MEDICAL ENTOMOLOGY STUDIES - XI.

THE SUBGENUS STEGOMYIA OF AEDES

IN THE ORIENTAL REGION WITH

KEYS TO THE SPECIES

(DIPTERA: CULICIDAE).!

Yiau-Min Huang

ABSTRACT

The subgenus Stegomyia Theobald of the genus Aedes Meigen and the 5

species groups recognized in the Oriental region are characterized. The

taxonomy, medical importance, immature habitats and distribution of the

species of the region are discussed and summarized. Keys and illustrations

are provided for the identification of adults (males and females), male ter-

minalia, pupae and 4th stage larvae of the 5 species groups and 37 species and

subspecies known to occur in this region. In addition, the detailed descrip-

tion of the male, female, pupa and larva of Aedes aegypti (Linnaeus) and a

redescription of the male terminalia of Aedes platylepidus Knight and Hull are

given.

INTRODUCTION

The subgenus Stegomyia. Theobald is medically very important since it

includes vectors of human filariasis and a number of virus diseases. It is one

of the most dominant subgenera of the genus Aedes Meigen in the Oriental

region, as indicated by the number of species groups and species. At present,

37 species and subspecies of this subgenus are recognized in this region.

The subgenus Stegomyia has not been properly defined since Edwards'

(1932) classification and this has resulted in the incorrect assignment of some

species to it and exclusion of others. As there is no comprehensive review of

the subgenus of this area, this paper is intended to clarify some of these tax-

onomic problems and also provide a guide for the identification of the species

occurring in the Oriental region.

I This work was supported by Research Contracts No. DA-49-193-MD-2672 and

DAMD-17-74C-4086 from the U. S. Army Medical Research and Development

Command, Office of the Surgeon General, Ft. Dietrich, MD 21701.

2Medical Entomology Project, Department of Entomology, Smithsonian

Institution, Washington, DC 20560.

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

The subgeneric characters of Stegomyia and its affinities to other aedine

subgenera and the classification of the species groups are discussed. The 5

species groups of this subgenus, known as the aegypiz, albolineatus, edwardsi,

scutellaris and w-albus groups, occurring in the Oriental region are also char-

acterized. These 5 species groups with their constituent subgroups and 37

species and subspecies recognized here are outlined in Table 1. The role of

members of the subgenus in the transmission of pathogens is presented in

Table 2. Keys and illustrations for the identification of adults (males and

females), male terminalia, pupae and 4th stage larvae of the 5 species groups

and 37 species and subspecies of this region are provided. However, the full

illustrations of those species which were published previously by Huang (1972,

1977a, 1978a, 1978b) will not be included. Data on the present status, geo-

graphical distribution and immature habitats of the species are summarized

in Appendices I, II, and III.

Aedes aegypti (Linnaeus) is the type-species of the subgenus Stegomyia.

It is one of the most variable of all species of mosquitoes with respect to its

coloration (scale pattern). Mattingly (1957) recognized 3 forms of aegypti:

(1) aegypti sensu strictu, thetypeform, (2) @egypti subspecies formosus

(Walker), (3) aegypti variety queenslandensis (Theobald) and gave a provision-

al diagnosis for each. Both the "type form" and "'variety queenslandensis"

occur in the Oriental region. It is interesting to note that these 2 forms appear

to exist sympatrically throughout this region, but "variety queenslandensis"

occurs at a very low frequency. For this reason, I believe that these ''forms"

of aegypti in the Oriental region are not discrete entities, and therefore con-

sider that "variety queenslandensis" is an infraspecific variation. The

detailed description of the male, female, pupa and larva of this species is

given in Appendix IV. Another form of aegypti, ''subspecies formosus"', is

known only from Africa and will not be discussed here.

Aedes platylepidus Knight and Hull, which was assigned by Reinert (1970)

to the subgenus Diceromyia Theobald, is included here in the @lbolineatus

sroup. This decision was reached after a careful study of the male terminalia

of this species which agree well with the members of the albolineatus group,

as demonstrated in this paper. The male terminalia of platylepidus are very

similar to that of other members of the albolineatus group. Its closest allies

within this group are bambusicola Knight and Rozeboom, hoogstvaali Knight

and Rozeboom and impatibilis (Walker), (see Figs. 12C, 16C, 19C). It should

be noted that the description and the illustration of the male terminalia given

by Reinert (1970: 13 and Fig. 10) are in error, particularly with regards to

the basal mesal lobe (claspette) and is corrected in Appendix V (Fig. 24).

Unfortunately, the larva and pupa of this species are unknown. The transfer

of platylepidus from Diceromyia to Stegomyia may well receive further con-

firmation when the immature stages are known.

Aedes meronephada (Dyar and Shannon), was placed by Edwards (1929) in

the subgenus Stegomyia and was assigned by Edwards (1932) to Group B

(w-albus group), on the basis of the tarsal markings and the thoracic ornamen-

tation. Huang (1978b) removed Aedes meronephada from Group B (w-albus

group) and placed it in the subgenus Diceromyia. This conclusion was reached

after a critical study and consideration of all known stages. Thus, Aedes

meronephada is omitted from the key since it is not a Stegomyia.

Hairiness in Stegomyia larvae was first reported by Rosen and Rozeboom

(1954) in members of the scutellaris group in the South Pacific. In Singapore,

Colless (1956) observed that larvae of Ae. albopictus (Skuse) (also a member

of the scutellaris group) collected from tree holes, frequently had many of the

Huang: Aedes (SStegomyia) in the Oriental Region 3

body setae longer, stouter and more branched than usual. The hairiness

factor was shown to be associated with the presence of debris in the tree

holes utilized as larval habitats.

This study was based primarily on specimens accumulated by the Southeast

Asia Mosquito Project (SEAMP) and its successor, the Medical Entomology

Project (MEP), Department of Entomology, Smithsonian Institution. Addition-

al material was borrowed from the following institutions: Bernice P. Bishop

Museum; United States National Museum; Field Museum of Natural History,

Chicago; University of Utah; Cornell University; Johns Hopkins School of

Hygiene and Public Health; California Academy of Sciences; Academy of

Natural Sciences, Philadelphia; Medical Zoology Laboratory, Institute for

Infectious Disease, University of Tokyo; British Museum (Natural History)

and the Instituut voor Tropische Hygiene, Amsterdam.

All the type-specimens of the included species which are in the British

Museum (Natural History), the United States National Museum and the Medical

Zoology Laboratory, Institute for Infectious Diseases, University of Tokyo,

have been studied.

It has been necessary to rely upon available literature both for information

regarding those species recorded from the Oriental region and for information

regarding medical significance.

The nomenclature chosen for the chaetotaxy of the larva and pupa and the

terminology for the structural parts of the adult used in this paper largely

follows that of Belkin (1962) and Huang (1972, 1977a).

Abbreviations cited for the references to the literature conform to the

BIOSIS List of Serials, Biosciences Information Service of Biological Abstracts,

Philadelphia, 1974.

The information on the immature habitats is entirely based on data

accompanying specimens which I have examined.

The term "Oriental region"' as used here encompasses both the Oriental

and Indomalayan areas of Belkin (1962). It comprises the following area:

the Ryukyu islands (Amami, Okinawa, Miyako, Ishigaki, Iriomote), Taiwan,

the Pescadores, Hainan, China (south of the Yangtze Kiang), the Philippines,

Vietnam, Indonesia (the eastern boundary is essentially that of Lee 1944),

Laos, Cambodia, Thailand, Malaysia, the Andaman islands, the Nicobar

islands, Burma and Assam, Bangladesh, Bhutan, Nepal, India (south of the

Himalayas), Sri Lanka, southeastern part of Pakistan, the Laccadive islands,

the Maldive islands and the Chagos islands. This area falls approximately

within 11 degrees south to 30 degrees north latitude and 65 to 130 degrees

east longitude (Map 1).

SUBGENUS STEGOMYIA THEOBALD

Stegomyia Theobald 1901la (June 1), in Howard 1901: 235; Theobald 1901b

(July 15): 235; Theobald 1901c (Sept.): 4, App. ii; Theobald 1901d (Nov.

23): 283. Type-species: Culex aegypti Linnaeus 1762 (see Mattingly,

Stone and Knight 1962).

Scutomyia Theobald 1904: 77. Type-species: S. albolineatus Theobald 1904,

Ampang Jungle, near Kuala Lumpur, Selangor, Malaya; monobasic.

Quasistegomyia Theobald 1906: 69. Type-species: Q. unilineatus Theobald

1906, Bahr-el-Ghazal, Sudan; monobasic.

Pseudostegomyia Ludlow 1908: 10. Type-species: Stegomyia gardnerii Ludlow

1905, Bulacao, Mindoro Island, Philippines; monobasic. - Apparently an

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

error for Quasistegomyia Theobald 1906 (see Theobald 1910: 135). For

complete synonymy, see Knight and Stone (1977: 154).

CHARACTERISTICS.

The subgenus Stegomyia is characterized by the following combination of

characters: ADULT (both sexes). Head. Vertex with largely broad and flat

decumbent scales, erect forked scales not numerous, restricted to occiput;

& palpus not very short, more than 0.5 length of proboscis, 5-segmented,

segments 4,5 subequal, slender and with only a few short setae, total length of

apical 2 segments not very short, at least 0.4 length of the remaining seg-

ments; in 2 about 0.14-0.25 length of proboscis, 4- or sometimes 5-segmented,

when present segment 5 minute. Thorax. Acrostichal bristles absent; spiracu-

lar bristles absent; postspiracular bristles present; scutellum with broad

scales on all lobes; postnotum bare. Wing. With narrow scales on all veins.

Legs. Hindtarsus with basal white band at least ontarsomere 1. Male

Terminalia, Proctiger without cercal setae; aedeagus strongly toothed;

claspette well developed, with numerous setae; distimere usually simple,

elongate, or sometimes distimere complex, expanded at base apically forked,

spiniform present. Female Terminalia. Insula longer than broad, with

minute setae and 3-10 larger setae on apical 0.25-0.50; cerci short and broad;

3 spermathecae, one larger than the other 2. PUPA. Subgeneric characters

not evident, see under groups. LARVA. Head, Seta 4-C well developed,

branched, closer to 6-C than 5-C, cephalad and mesad of 6-C; 4, 6-C cephalad

of antennal base; 6-C cephalad of 5,7-C. Abdomen. Seta 12-I not developed;

2-VIII distant from 1-VIII; comb scales in a single row; ventral brush with 4,

5 pairs of setae on grid; without precratal tufts. This combination of characters

differs from that of all other subgenera of Aedes.

SYSTEMATICS.

Edwards (1932) divided the subgenus Stegomyia into 4 groups which he

designated A,B,C, and D. Knight and Rozeboom (1946) removed Aedes

albolineatus (Theobald) from Group C and defined a 5th group for that species

and its relatives which later was designated Group E (albolineatus group) by

Knight and Hurlbut (1949). Bohart 1956 (1957) added Group F (pandani group)

and Group G (maehleri group). Belkin (1962) removed Aedes edwardsi (Bar-

raud) from Group B and defined a new group (edwardsi group) for it and its

relatives. Huang (1972) redefined Group C (scutellaris group) and subdivided

the scutellayis group into 2 subgroups known as the albopictus and scutellaris

subgroups. Later, Huang (1977a) redefined Group B (w-albus group) and sub-

divided the w-albus group into 4 subgroups known as the annandalei, desmotes,

mediopunctatus and w-albus subgroups. Huang (1977b) removed Group D

(vittatus group), a monotypic species group, (Aedes) vittatus (Bigot)), from the

subgenus Stegomyia and placed it in the subgenus Aedimorphus Theobald of

the genus Aedes. Thus, the subgenus Stegomyia now consists of the following

7 groups: Group A (aegypti group), Group B (w-albus group), Group C

(scutellaris group), Group E (albolineatus group), Group F (pandani group),

Group G (maehleri group), and Group H (edwardsi group).

Of the 7 recognized groups of Stegomyia mentioned above, 5 are represented

in the Oriental region. Only Groups F and G do not occur inthe Oriental region.

These 5 groups with their constituent subgroups, 37 species and subspecies are

indicated in Table 1.

Huang: Aedes Stegomyia) in the Oriental Region 5

TABLE 1. Classification of the subgenus Stegomyia in the Oriental region.

GROUP SUBGROUP

1. aegypti

2. albolineatus

SPECIES AND SUBSPECIES

1. aegypti (Linnaeus)

2. albolineatus (Theobald)

3. arboricola Knight and

Rozeboom

4. bambusicola Knight and

Rozeboom

boharti Knight and Rozeboom

hoogstraali Knight and

Rozeboom

7. impatibilis (Walker)

8. laffooni Knight and Rozeboom

9. platylepidus Knight and Hull

0. pseudalbolineatus Brug

co) o1

3. edwardsi

4. scutellaris albopictus

scutellaris

5. w-albus annanda lei

11. edwardsi (Barraud)

12. seampi Huang

13. albopictus (Skuse)

14. downsi Bohart and Ingram

15. novalbopictus Barraud

16. patriciae Mattingly

17. pseudalbopictus (Borel)

18. seatoi Huang

19. subalbopictus Barraud

20. unilineatus (Theobald)

21. alcasidi Huang

22. alorensis Bonne-Wepster

23. andrewsi Edwards

24. krombeini Huang

29. malayensis Colless

26. paullusi Stone and Farner

27. viversi Bohart and Ingram

28. annandalei (Theobald)

29. craggi (Barraud)

desmotes 30. desmotes (Giles) 3

31. malikuli Huang

mediopunctatus

w-albus

32. mediopunctatus (Theobald)

33. perplexus (Leicester)

34. rhungkiangensis Chang and

Chang

39. gardnervii gardnerii (Ludlow)

36. gardnerii imitator (Leicester)

37. w-albus (Theobald)

6 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

In the identification of the species of the subgenus Stegomyia, the adult

stages appear to be more useful than the immature stages. However, it must

be remembered that specific differences between the members of this subgenus

tend to be very slight. Some members are highly variable in both adult orna-

mentation and in the immature stages. Although the males of all species can

be recognized on the basis of morphological features, the females and the

immatures are extremely difficult or impossible to distinguish in many

instances. The male terminalia of all species are distinct and the most

diagnostic feature of all is the claspette of the basimere. In dealing with

these, special preparations must be made and care taken to study both lateral

and mesal views of the dissected claspette as well as undissected aspects.

AFFINITY. |

The subgenus Stegomyia possesses some rather important basic characters,

as follows, in common with the subgenera Aedimorphus Theobald, Alanstonea

Mattingly, Ayurakitia Thurman and Diceromyia Theobald of the genus Aedes in

the Oriental region: Adult. Male palpus 5-segmented; proctiger without cer-

cal setae; aedeagus with conspicuous teeth; claspette developed; distimere

with one or more spiniforms; Larva. Seta 12-I not developed; pecten teeth

present. These shared characters indicate the affinity of Stegomyia to the

above 4 subgenera. Of these 4 subgenera, Stegomyia shares more important

characters in both adult and immature stages with Diceromyia than with any

other subgenus, suggesting the strongest affinities with that subgenus. How-

ever, it differs from Diceromyia in the development of the male palpus and in

the position of seta 4-C of the larva. The male palpus of Sftegomyia has the

total length of the apical 2 segments not very short, at least 0.4 the length of

the remaining segments while in Diceromyia, the total length of the apical 2

segments is very short, at most 0.3 the length of the remaining segments, or

segment 5 is much shorter than segment 4. The larva of Stegomyia has seta

4-C cephalomesad of 6-C while in Diceromyia, seta 4-C is caudomesad of 6-C.

DESCRIPTIONS OF THE GROUPS OCCURRING IN THE ORIENTAL REGION.

aegypti group

ADULT. Head. Palpi with white scales. Thorax. Dorsocentral bristles

present; scutum with a lateral crescent-shaped white marking on anterior half

of scutum; subspiracular area with broad white scales; postspiracular area

without scales; paratergite with broad white scales; scutellum with broad white

scales on all lobes. Legs. Hindtarsus with a basal white band on tarsomeres

1-4, tarsomere 5 all white. Male Terminalia, Apical margin of tergum IX

with middle part deeply concave; paraproct with sternal arm, without apical

lobed process; distimere simple, elongate, with an apical spiniform process;

aedeagus strongly toothed. Female Terminalia, Apical margin of sternum

VIII with a deep U-shaped notch at middle.

PUPA. Paddle. Margins with distinct denticles; seta 1-P single.

LARVA. Comb scales in a single row, not arising from a sclerotized plate;

marginal spicules of anal segment very small and inconspicuous; ventral brush

with 5 pairs of setae.

Huang: Aedes (Stegomyia) in the Oriental Region 7

albolineatus group

ADULT. Head. Palpi without white scales. Thovax. Dorsocentral

bristles present or absent; scutum with, or without, a long median longitudinal,

white stripe of narrow scales extending from anterior margin to at least the

level of wing root; subspiracular and postspiracular areas without scales; para-

tergite bare; scutellum with broad scales on all lobes. Legs. Hindtarsus with

basal white band at least on tarsomere 1. Male Terminalia, Apical margin of

tergum IX with middle part shallowly concave, flat, or slightly produced; para-

proct without sternal arm, with or without apical lobed process; distimere

simple, elongate, with a submedian or subbasal spiniform process; aedeagus

with several lateral distal teeth. Female Terminalia. Apical margin of stern-

um VIII with middle part shallowly concave, or apical margin of sternum VIII

with a deep U-shaped notch at middle.

PUPA. Paddle. Margins without fringe of very long hair-like spicules,

or distinct denticles, at most with slight spicules; seta 1-P branched.

LARVA. Comb scales in a single row, arising or not arising from a sclero-

tized plate; marginal spicules of anal segment very small and inconspicuous, or

very long, spine-like and conspicuous; ventral brush with 5 pairs of setae.

edwardsi group

ADULT. Head. Palpi with white scales. Thorax. Dorsocentral bristles

present; scutum with a distinct, small, median oval white spot of narrow scales

on anterior 0.33 of scutum; subspiracular area with or without scales; post-

spiracular area without scales; paratergite with broad white scales; scutellum

with broad white scales on all lobes. Legs. Hindtarsus with a basal white

band on tarsomeres 1-4, tarsomere 5 all white. Male Terminalia. Apical

margin of tergum IX with middle part produced into a large lobe; paraproct

without sternal arm, without apical lobed process; distimere simple, elongate,

with an apical spiniform process; aedeagus apically widened, with 6 or 7 apical

teeth on each side. Female Terminalia. Apical margin of sternum VIII with a

deep, U-shaped notch at middle.

PUPA. Paddle. Margins with fringe of very long hair-like spicules; seta

1-P single.

LARVA. Comb scales ina single row, arising from a sclerotized plate;

marginal spicules of anal segment very long, spine-like and conspicuous; ventral

brush with 4 pairs of setae.

scutellaris group

ADULT. Head, Palpi with white scales. Thorax. Dorsocentral bristles

present; scutum with a long, median longitudinal white stripe of narrow scales

extending from anterior margin to about the level of wing root; subspiracular

and postspiracular areas with or without scales; paratergite with broad white

scales; scutellum with broad white scales on all lobes. Legs. Hindtarsus with

a basal white band on tarsomeres 1-4, tarsomere 5 all white. Male Terminalia.

Apical margin of tergum IX with middle part rounded, truncated, or produced

into a large lobe or a median projection; paraproct without sternal arm, without

apical lobed process; distimere simple, elongate, with an apical or a subapical

8 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

spiniform process; aedeagus with a distinct sclerotized, lateral toothed plate

on each side. Female Terminalia. Apical margin of sternum VIII with a deep

U-shaped notch at middle.

PUPA. Paddle. Margins with fringe of very long hair-like spicules; seta

1-P single.

LARVA. Comb scales in a single row, not arising from a sclerotized

plate; marginal spicules of anal segment very small and inconspicuous; ventral

brush with 4 pairs of setae.

w-albus group

ADULT. Head. Palpi with white scales. Thorax. Dorsocentral bristles

sabsent; scutal markings varied; subspiracular and postspiracular areas with

broad white scales; paratergite with broad white scales; scutellum with broad

white scales at least on one lobe. Legs. Hindtarsus with a basal white band on

tarsomeres 1,2, tarsomere 3 with or without a basal white band. Male Ter-

minalia. Apical margin of tergum IX with middle part concave, nearly flat, or

produced into a large lobe; paraproct with or without sternal arm, without

apical lobed process; distimere simple, elongate, with an apical or a subapical

spiniform process, or distimere complex, expanded at base and apically forked,

with spiniform process; aedeagus with a distinct, sclerotized, lateral toothed

plate on each side. Female Terminalia. Apical margin of sternum VIII with a

deep U-shaped notch at middle.

PUPA. Paddle. Margins with fringe of very long hair-like spicules; seta

1-P single.

LARVA. Comb scales in a single row, arising or not arising from a

sclerotized plate; marginal spicules of anal segment very small and incon-

spicuous, or very long, stout and conspicuous; ventral brush with 4 pairs of

setae.

DISTRIBUTION.

Stegomyia is confined to the Old World, except for Aedes aegypti which has

been introduced through commerce into the New World. This subgenus occurs

chiefly in the tropical and subtropical zones throughout the Old World but is

also represented in the southern part of the Palearctic.

Members of the Oriental Stegomyia are distributed mainly in the Oriental

region, with extensions into the southern part of the Palearctic area in the

north, the Papuan, Western Pacific islands and South Pacific areas in the east

and the Ethiopian area in the west. Aedes albopictus (Skuse) is also known in

the Hawaiian islands (where it has been introduced) and Malagasy. In Stegomyia,

it appears that there are several widely distributed dominant species and a

number of specialized endemic species. The distribution records are given in

Appendix II.

BIONOMICS.

The immature stages have been found mainly in tree holes, bamboo stumps,

artificial containers (tin can, water jar, water bucket, broken bottle and tire),

stump holes (palm stumps, banana stumps and coconut stumps), bamboo inter-

nodes, coconut shells (coconut halves, coconut husks and coconut spathes),

bamboo cups (bamboo pots), rock holes, plant axils (Pandanus, banana, nipa

palm, sago palm, taro and Colocasia), log holes, split bamboos, cut bamboos,

fallen leaves (fallen abaca leaf), palm fronds (coconut fronds), rock pools, foot-

prints, wheel ruts, root holes and shelf fungus - a wide variety of natural and

Huang: Aedes Stegomyia) in the Oriental Region 9

artificial container habitats. Females of 14 species and one subspecies (aegypii,

boharti, albopictus, downsi, seatoi, krombeini, malayensis, rivers,

annandalei, craggi, desmotes, mediopunctatus, perplexus, w-albus and

gardnerii imitator) are known to bite man. The immature habitats are sum-

marized in Appendix III.

MEDICAL IMPORTANCE.

Stegomyia is one of the most important subgenera of mosquitoes from the

standpoint of the transmission of pathogens. Aedes aegypit is the principal

vector of urban yellow fever virus in the African, Central and South American

tropics. This species is likewise the primary vector of dengue viruses

through most of the tropical world. It has also been shown to be a vector of

Chikungunya virus in Southern Province, Tanganyika Territory [Tanzania]

(Lumsden 1955) and Venezuelan equine encephalitis virus in Colombia

(Sanmartin-Barberi et al. 1955) and is a known vector of other viruses. Some

members of the @egypii group are efficient vectors of yellow fever virus in

Africa (Gillett 1972). Aedes scutellaris (Walker) from New Guinea has been

incriminated as a vector of dengue virus (Mackerras 1946). In addition to

viruses, some members of the scutellaris group are also efficient vectors of

subperiodic filaria in the South Pacific (Sasa 1976).

Both Aedes aegyptiand albopictus are important vectors of dengue viruses

in Southeast Asia and have been incriminated in the transmission of dengue

virus during outbreaks of haemorrhagic fever in Singapore (Chan, Ho and Chan

1971) and Koi Samui Island, Thailand (Gould et al. 1968). Aedes albopictus

from India can transmit Chikungunya virus in the laboratory (Rao et al. 1964).

The larvae of Dirofilaria immitis have been found in wild-caught albopictus in

Nagasaki, Japan (Suenaga and Itoh 1973) and 3rd stage larvae of Dirofilaria sp.

have been found in wild-caught albopictus in Thailand (Harinasuta et al. 1970).

Several viruses have been isolated from wild-caught aegypti and albopictus from

the Oriental region. Table 2 shows briefly what is known in this regard.

TABLE 2. Isolation of pathogens of actual or potential medical significance

from Aedes (SStegomyia) species in the Oriental region.

Species Pathogen Location Comments Reference

aegypti Chikungunya virus South India 111 isolations Myers et al.

August- 1965

December, 1964

Chikungunya virus Nagpur City, 5 isolations 1965 Rodrigues

India et al. 1972

Chikungunya virus Bangkok, 7 isolations from Halstead et al.

Thailand 150 pools, 1962 1969

Dengue 1 and Vellore, 3 isolations Carey et al.

Dengue 4 viruses India Dengue 1 and 1964

2 Dengue 4 from

77 pools,

September-

December, 1961

10 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

TABLE 2 (Continued).

Species Pathogen

aegypti Dengue 4 virus

Dengue 1 and

Dengue 3 viruses

Dengue 1 and

Dengue 2 viruses

Dengue 1 virus

Dengue 3 virus

Dengue virus

Dengue 2 virus

Dengue 1-4 viruses

Location

Nagpur City,

India

Ajmer City,

India

Bangalore

City, India

Peninsular

Malaysia

Manila,

Philippines

Singapore

Bangkok,

Thailand

Koh Samui

Island,

Thailand

Bangkok,

Thailand

Comments

9 isolations

1965

isolations

September 1969

32 isolations of

both viruses,

Dengue 2 more

plentiful

1 isolation

1956

1 isolation

1966

3 isolations

1958

8 isolations

from 27 pools

July-September

1966

. isolations

Dengue 1,

6 Dengue 2,

8 Dengue 3 and

1 Dengue 4 from

150 pools, 1966;

1 isolation

Dengue 1, 2

Dengue 2 and

1 Dengue 4 from

90 pools, 1963

Reference

Rodrigues

et al. 1972

Ghosh et al.

1974

George and

Soman 1975

Rudnick (pers.

commun. )

1975

Hammon,

Rudnick and

Sather 1960

Lim, Rudnick

and Chan 1961

Chan, Ho and

Chan 1971

Hammon,

Rudnick and

Sather 1960

Gould et al.

1968

Halstead et al.

1969

Huang: Aedes (Stegomyia) in the Oriental Region 11

TABLE 2 (Continued).

Species Pathogen

aegypti Dengue 4 virus

Dengue 2 virus

Dengue 1-4 viruses

Dengue 3 virus

Zika virus

albopictus Dengue 2 virus

Dengue 1 and

Dengue 2 viruses

Dengue 2 virus

Dengue 4 virus

Reovirus 3

Location

Koh Samui

Island,

Thailand

Saigon,

Vietnam

Saigon area,

Vietnam

Rangoon,

Burma

Bentong,

Malaysia

Singapore

Koh Samui

Island,

Thailand

Koh Samui

Island,

Thailand

Koh Samui

Island,

Thailand

Peninsular

Malaysia

Comments

25 isolations

from 88 pools

May-

November 1968

1 isolation

November 1963

30 dengue

isolations of all

strains from

1,892 pools May-

December 1967

1 isolation July

1969

1 isolation

July 1966

1 isolation

November 1960

1 isolation

Dengue 1 and 4

isolations

Dengue 2 from

pools April-

October 1966

4 isolations

from 141 pools

July-September

1966

10 isolations

from 897 pools

August-

December 1967

38 isolations

from 1, 829

pools May-

November 1968

2 isolations

1974

Reference

Smith et al.

1971

Halstead et al.

1965

Russell, Quy

et al. 1969

MI- MI-KHIN

and KHIN-AYE-

THAN 1976

Marchette,

Garcia and

Rudnick 1969

Rudnick and

Chan 1965

Chan, Ho and

Chan 1971

Gould et al.

1968

Russell, Gould

et al. 1969

Smith et al.

1971

Rudnick (pers. )

commun. ) 1975

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

TABLE 2 (Continued).

Species Pathogen Location Comments Reference

albopictus Tembusu virus Peninsular 1 isolation Rudnick (pers.

Malaysia 1974 commun. ) 1975

Dirofilaria Nagasaki, 3.2% of field Suenaga and

immitis Japan dissected Itoh 1973

adults in urban

areas with

filarial larvae

and 5.5% in

rural areas

with filarial

larvae

Dirofilaria sp. Sangkhlaburi 2/571 dissected Harinasuta

District, with stage 3 et al. 1970

Kanchanaburi filarial larvae

Prov.,

Thailand

KEYS TO THE GROUPS, SUBGROUPS AND SPECIES OF

THE SUBGENUS STEGOMYIA

ADULTS (MALES AND FEMALES)

t. Palpi with waite scales (Fic, 1A): 3 6 se a ke SR ew Ou 2

Palpi without white scales (Fig. 6A).

ALBOLINEATUS GROUP (p. 13)

2(1). Dorsocentral bristles present (Fig. 1A). ............0ee8- 3

Dorsocentral bristles absent (Huang 1977a, Fig. 25A). :

W-ALBUS GROUP (p. 15)

32). Scutum with lyre-shaped white markings (Fig. 1A). |

AEGYPTIGROUP

aegypti (p. 41)

Scutum without lyre-shaped white markings. ....... Re pial 4

4(3). Scutum with a long median longitudinal white stripe of narrow scales

extending from anterior margin to about level of wing root

ie. ZORA Ree oe cs ES SCUTELLARIS GROUP (p. 14)

Scutum with a distinct, small, median oval white spot of narrow scales

on anterior 0.33 of scutum (Huang 1977a, Fig. 1A).

EDWARDSI GROUP (p. 13)

Huang: Aedes (Stegomyia) in the Oriental Region 13

THE ALBOLINEAT US GROUP

Ls Scutum with a median longitudinal white stripe. ........... 2

Scutum without a median longitudinal white stripe.

platylepidus (p. 44)

2(1). Scutelium with broad white scales on midlobe and with broad dark

scales’ onlateral dobe( Pig: 6A) ceo se Pee A as 8 3

Scutellum with broad white scales on all lobes (Fig. 20A)..... 6

3(2). Median scutal stripe extending posteriorly to the scutellum

(ised BO)\ow os aais te Bows Boge igre Roe es old 4

Median scutal stripe extending posteriorly only to the level of wing

root (Pig. GA). eek od yo wee ee, Bee eG 5

4(3). Hindtarsomere 3 with white basal band (Fig. 18A). ... hoogstraali

Hindtarsomere 3 entirely dark (Fig. 18B)......... impatibilis

5(3). Posterior pronotum with narrow dark scales only. ... albolineatus

Posterior pronotum with narrow dark scales dorsally and broad white

SCALCS-V OIL Tally se PS Ge eae eee, igh ye, BLES: boharti

6(2). Median scutal stripe extending posteriorly to the prescutellar

area (Pie 20AI Se See PtSi gO, OGG Ba bambusicola

Median scutal stripe extending posteriorly only to the level of wing

OSE CRIay QOB) Se Wie eet ste RHE ee Ses teed CPE oe 7

7(6). Scutum with a patch of broad white scales on lateral margin just before

the level ot wine root (fie. 200). 1 yk OS PORE 8

Scutum without a patch of broad white scales on lateral margin just

before the level of wing foot. oo ee eee ee, arboricola

8(7). Male hindtarsomere 2 with at least basal 0.25 white band; female

hindtarsomere 3 with at most basal 0.5 white dorsally (Fig. 23A).

la ffooni

Male hindtarsomere 2 with at most basal 0.2 white band; female hind-

tarsomere 3 with at least basal 0.87 white dorsally (Fig. 23B).

pseudalbolineatus

THE EDWARDSI GROUP

Posterior pronotum without narrow dark scales on upper portion;

lower mesepimeron with bristles (Huang 1977a, Fig. 4C).

edwardsi

Posterior pronotum with narrow dark scales on upper portion; lower

mesepimeron without bristles (Huang 1977a, Fig. 4F). . seampi*

*The male of seampi is unknown.

4(3).

5(2).

6(5).

7(6).

8(5).

9(1).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

THE SCUTELLARIS GROUP

Supraalar white line incomplete, not clearly defined and with only

narrow scales over wing root (Fig. 380A). ........80648-. 2

Supraalar white line complete and well developed, with broad flat

scales over wing root and toward scutellum (Fig. 30B)..... 9

Scutum with patch of broad flat white scales on lateral margin just

before level ofavint root (Fie. 300s ote ON ee. 3

SCHUUnY WitihOuL SUCN DALCH. . 6 es 6. Re a oe ke 5)

Scutum with a small patch of white scales on scutal angle area; mid-

femur with some pale scales scattered on anterior surface

(Huns OG2. Fies, 154, fo, 15D) ese BE. oe 4

Scutum without such a white patch on scutal angle area; midfemur

without such scales (Huang 1972, Figs. 1A, 1D, 1E). . .albopictus

Tergum I with a large median patch of white scales; midfemur without

a large white spot on anterior surface (Huang 1972, Figs. 15A,

LSD )uilanend: bhi eae es es Shee Se ee wa. seatoi

Tergum I without median patch of white scales; midfemur with a large,

white spot on anterior surface (Fig. 25B)........ unilineatus

Scutum with patch of narrow curved yellowish scales on lateral margin

just before level of wing root. ......- 2.2 se eee eevee 6

Scutum with patch of narrow curved white scales on lateral margin

just before level of wing root (Fig. 30D). .......-2.-+e+6- 8

Fore- and midfemora with some pale scales scattered on anterior

Surface ek ck es a eae te ee Ae nova lbopictus *

Fore- and midfemora without such scales. .......2+-s2.eee-e 7

Subspiracular area with scales; hindtarsomere 4 with at most basal

0. 66-0. 75 white banded (Huang 1972, Fig. 20M). .... patriciae

Subspiracular area without scales; hindtarsomere 4 with basal 0. 83-

0.90 white banded (Huang 1972, Fig. 20N)......... downsi

Scutum with patch of broad dark scales on each side of prescutellar

space between prescutellar white line and postdorsocentral white

line; postspiracular area with scales (Fig. 314A).

pseudalbopictus

Scutum without such patch of scales; postspiracular area without

Sealee (Picu BIB) See ea ew: mwa SS subalbopictus

Abdomen with lunate, lateral, white spots only; wing without a minute

basal spot of white scales on costa (Huang 1972, Fig. 20D).

andrewsi

*The female of novalbopictus is unknown.

10(9).

11(10).

12(10).

13(12).

14(13).

3(2).

Huang: Aedes (SStegomyia) in the Oriental Region 15

Abdomen with some complete pale bands or with indications of such

bands on terga; wing with a minute basal spot of white scales on

Midfemur with a median white line on anterior surface (Huang 1972,

Ps BOR. Se oe gale ae ea eae aa eae ae 11

Midtemur without Sueh. Line.) so.2 ee sow Gia aes ee a a 12

Scutum with a few narrow white scales on lateral prescutal area and on

scutal angle area (Huang 1972, Fig. 20J)......... paullusi

Scutum without any narrow white scales in these positions.

alorensis

Hindtarsomere 3 with basal 0.4 white; hindtarsomere 4 with at most

bagel Qe Giiwhtben i: setae cies ea ae a es ee ess 13

Hindtarsomere 3 with basal 0.5 white; hindtarsomere 4 with basal

0;'75 white: {Etiam 1972: Bie 2 1 Bs te we G ei res alcasidi

Hindtarsomere 1 with at least basal 0.25 white; hindtarsomere 2 with

basas 0:33 whibewd wists tales? sakes alee ca eee) 5. 14

Hindtarsomere 1 with basal 0.20 white; hindtarsomere 2 with basal

0:25 white: (Hudne 1072 Fiee 31 Bw oti © iain ltl: riversi

Hindtarsomere 1 with basal 0.25 white; hindtarsomere 4 with basal

0.66 white (Huang 1972, Fie. 31D). ... 6.5... es malayensis

Hindtarsomere 1 with basal 0.33 white; hindtarsomere 4 with basal

OO Grae hoo tet eer ngtabe ehite: sen eat Me 4 krombeini

THE W-ALBUS GROUP*

Tibiae with white band (Huang 1977a, Fig. 21A).

(desmotes subgroup) desmotes

Tiiae Wimiont. wWhire Mand. 6 a AS eee a as COTE oe 2

Scutellum with broad dark scales on midlobe and with broad white

scales on lateral lobe (Huang 1977a, Figs. 9B, 9D).

| (annandalei subgroup) 3

Scutellum with broad white scales on midlobe and with broad dark, or

ware. OF Dotheon lateral lobe: 2. 4a ay eae Oe ie

Scutum with a large median oval white patch, extending from anterior

margin, narrows slightly posteriorly, to about the middle of the

scutum; scutal patch posteriorly blunt (Huang 1977a, Fig. 9B).

: annandalei

Scutum with a large median triangular white patch, extending from

anterior margin and about anterior half of the lateral prescutal

area, narrows posteriorly, to the middle of the scutum; scutal

patch posteriorly pointed (Huang 1977a, Fig. 9D). .... craggi

*Aedes rhungkiangensis is not included.

4(2).

6(5).

2(1).

3(1).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

Scutum with a broad median longitudinal white stripe, extending from

anterior margin, tapering posteriorly, to the prescutellar area and

forking at the beginning of prescutellar space; midfemur without a

median white spot on anterior surface (Huang 1977a, Fig. 25A).

(mediopunctatus subgroup)

malikuli*

mediopunctatus

perplexus

Scutum without such a median longitudinal white stripe; midfemur with

a median white spot on anterior surface (Huang 1977a, Fig. 21B).

(w-albus subgroup) 5

Scutum with a broad longitudinal white stripe on each side of midline,

extending from anterior margin to the middle of the scutum and

fusing with the antealar white patch (Huang 1977a, Fig. 17D).

gardnerii gardnerii

Scutum with a large median white patch, or 2 lateral white patches, on

anterior ©. 00 6; SCUlUIN G2 a ae ee

Scutal median white patch broader than long, with narrow white scales

reaching from anterior margin to the lateral prescutal area on each

side; antealar white patch with narrow white scales mainly and with

some broad white scales on the lateral margin just before the level

of the wing root (Huang 1977a, Figs. 32B, 32A).

garvdnerii imitator

Scutal median white patch rather narrow and long; antealar white patch

with all narrow white scales (Huang 1977a, Figs. 32D, 32C).

w-albus

MALE TERMINALIA

Pavaoroct with scarnar arm (Fic. 1C). . . ww ak ke ee ee 2

Paraproct wineutl Sterna armas 2.6 Soe es 3

Apical margin of tergum IX with middle part deeply concave

(Pigs 2 2Cy oe ee es AEGYPTI GROUP

aegypti (p. 41)

Apical margin of tergum IX with middle part nearly flat (Huang 1977a,

Pie 140) ree ee eee a ee W-ALBUS GROUP

(desmotes subgroup)

desmotes

Distimere simple, elongate, with a spiniform process. ....... 4

Distimere complex, expanded at base and apically forked, with spini-

forms processes (Huang 1977a, Figs. 19C, 21C, 25C).

W-ALBUS GROUP

(mediopunctatus subgroup) (p. 20)

*The female of malikuli is unknown.

Huang: Aedes Stegomyia) in the Oriental Region 17

4(3). Spiniform of distimere process submedial or subbasal (Figs. 6C,

120) Hee Oa ee ALBOLINEATUS GROUP (p. 17)

Spiniform of distimere process at apex or subapical. ........ 9)

5(4). Apical margin of tergum [IX with middle part concave (Huang 1977a,

Figs, 6C,:19G)6 bay ee ore eee ee W-ALBUS GROUP

(annandalei subgroup) (p. 20)

Apical margin of tergum IX with middle part rounded, truncated, or

produced into a large lobe, or a median projection. ....... 6

6(5). Aedeagus apically widened, with 6 or 7 apical teeth on each side

(Huane 197 7a, Fics UB). <2 See EDWARDSI GROUP*

edwards

Aedeagus with a distinct sclerotized lateral toothed plate on each

Sider UW eecters, pare GT ee i a a ee ao 7

7(6). Spiniform of distimere long, at least 0.28 length of distimere;

apical margin of tergum IX with large median lobe (Huang 1977a,

Pigs 2OCE 880).. Wiky. Se SKS es tee ee W-ALBUS GROUP

(w-albus subgroup) (p. 20)

Spiniform of distimere rather short, at most, 0.25 length of distimere,

or apical margin of tergum IX without large median lobe (Figs. 25D,

vil Oy Mame igi larg eee Sore wee aaa SCUTELLARIS GROUP (p. 18)

THE ALBOLINEATUS GROUP

1 Paraproct with apical loved process. wow 8) SN Se we 2

Paraproct without apical lobed process. .<.cosi un. oO 8S kw es 8

2(1). Spiniform of distimere process before middle, at most, at about basal

Spaatteiaret UnetancgaNeDeSrmeliaaptenterpoekatiRe. cx. 4

3(2). Apical margin of tergum IX with middle part concave; basimere long,

about 3.0 as long as wide, with numerous short, apically curved

setae along mesal side of dorsal surface (Fig. 16C).

hoogstraali

Apical margin of tergum IX with middle part nearly flat; basimere

rather short and broad, about 2.2 as long as wide, with some long,

stout, apically curved setae along mesad side of dorsal surface

UIE, |, NRE Ge 2 ap aun a RRC age eGo OUR e aah Sime ar ar impatibilis

4(2). Lateral plate of aedeagus with less than 10 apical teeth (Figs. 6B,

GOs eas «Paar has ete se a ee ht ae albolineatus

5(4). Claspette with 3 short, stout, spine-like setae and several shorter

setae on distal part (Fig. 14C)...... by gil cig he ee boharti

*The male of seampi is unknown.

6(5).

7(6).

8(1).

3(2).

4(3).

(4).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

Claspette with 3-5 rather long, stout, spine-like setae and several

long setae with slender curved tips on distal part. ....... 6

Claspette with 3,4 stout, spine-like setae and several long setae with

slender curved tips on distal part; apical margin of tergum IX with

middbe concave (Fie. 00). ous es CS 2 cee arboricola

Claspette with 4,5 stout, spine-like setae and several long, stout setae

with slender curved tips on distal part; apical margin of tergum IX

with middle flat (Wigs. 210, 23Cy 23D )iew Sah soetwries Boe 7

The apices of the strongest spine-like setae of claspette reaching to the

level of apex of basimere (Fig. 21C)..........4.4. laffooni

The apices of the strongest spine-like setae of claspette not reaching

to the level of apex of basimere (Fig. 23D). . . . pseudalbolineatus

Spiniform of distimere process subbasal (Fig. 12C).

bambusicola

Spiniform of distimere process submedial (Fig. 24C). |

platylepidus (p. 44)

THE SCUTELLARIS GROUP

Apical margin of tergum [IX with a median lobe or projection. ... 2

Apical margin of tergum IX with middle broadly rounded or

ARICA OR ee ee ee is a URE ES 8

Apical margin of tergum IX with conspicuous horn-like median

projection (Huane 1972; Fig. 2C). 2.4808 2. albopictus

Apical margin of tergum IX with middle part produced into a lobe. . 3

Apical margin of tergum IX with middle part produced into large lobe

with apex serrated (Huang 1972, Fig. 4C)........ downsi

pen Ol joue NoLserraied. a a Ge eS A ee ee 4

Claspette long, slender, reaching to about 0.75 of basimere,

expanded portion facing mesad (Fig. 25D; Huang 1972,

Ee a eS SY 5

Claspette large, broad, reaching to about 0.5 of basimere, expanded

rome horizontal in position (Huang 1972, Figs. 9C, 11C,

FOC er Suk WARS es ootek DSi Eos a SS 6

Apical margin of tergum IX with large median lobe; claspette with

numerous setae and several widened specialized curved ones on

mesal side of slightly expanded distal part, without any setae on

basal 0.4 area (Huang 1972, Fig. 16C)............ seatoi

Apical margin of tergum IX with rounded median lobe; claspette with

numerous setae and several stouter widened ones on distal part,

with a small, mesally directed projection on basal 0.4 bearing one

large seta and with 3 smaller setae near to it (Figs. 25D, 25C).

unilineatus

6(4).

7(6).

8(1).

9(8).

10(9).

11(10).

12(11).

13(11).

14(13).

Huang: Aedes (SStegomyia) in the Oriental Region 19

Specialized setae on sternal side of expanded portion of claspette

spine-like, curved with sharply pointed tips and varying length

(Huane 1972, Figs. BO, OC}. es a ee owe ae 8 novalbopictus

These specialized setae blade-like or clubbed, without sharply pointed

tip Guanes 1972, Pigs. F9Cy PbO. bein Stew ei oe me a i

Claspette with broad stem and lateral distal angle turned through 90°

in lateral aspect (dissected) (Huang 1972, Fig. 18C).

subalbopictus

Claspette with narrower stem and without a 90° lateral distal angle

(Huang TO72y) Peed iCyicep ba Aaley ope eowy Pate Ss patriciae

Claspette complex with numerous setae on expanded distal part,

each seta on distinct cone, a tergal mesal finger-like process

bearing 6 modified setae at tip (Huang 1972, Fig. 24B). . alorensis

Claspette truncate, with a distinct oval face at apex; with numerous

setae and several long, stout ones on tergal side and with 4 spine-

like setae on sternal side of oval face; apical margin of tergum IX

with middle truncated (Huang 1972, Fig. 29C)....... paullusi

Claspette with apex more or less cylindrical. ........... 10

Claspette long, slender, reaching to 0.7 of basimere, with one widened

specialized spine-like seta and numerous setae distal to it (Huang

ISAs Fig 18s eer cai roy detete Seba pseudalbopictus

Claspette rather short, with several widened specialized setae and

numerous unmodified setaeiiial. TR wee Nw Bee, "4 11

Claspette with distal expanded part rectangular in lateral aspect

(dissected), sternal and tergal sides more or less parallel,

apicosternal angle present (Fic. 2TC).-. 6. ea. 12

Claspette with distal expanded part subtriangular in shape in lateral

aspect (dissected), sternal and tergal sides not parallel but tapering,

wethoulapiegsternal angles: 2.5 Geo ew OE Pe Ee ek. 13

Apicotergal area of claspette with several distinctly long and stout setae

Pang tO 2 Weeks. (alee aM OR ih aa andrewsi

Apicotergal area without any such setae (Fig. 27C). krombeini

Apicotergal area of claspette with several distinctly long setae (Huang

PETE eee BOC CRC is Hi es RS Be he he 8 alcasidi

Apicotergal area without any such setae. ..........220- “14

Claspette with 7-10 modified setae forming a prominant row at center

of sternal side and occupying about 0.5 of it (Huang 1972,

Pager CAD AO) tos hag Showin eo ee eRe malayensis

Claspette with 6-8 modified setae at center of sternal side, closer to

sternal angle area than to apicotergal angle area and occupying

about 0.4 of it (Huang 1972, Figs. 26A, 26B)........ riversi

a(t)

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

THE W-ALBUS GROUP

annandalei subgroup

Claspette short and broad, with 3 widened, leaf-like setae on tergal

side and with 2 stout, spine-like setae on sternal side of expanded

distal part, with several slender setae scattered in-between (Huang

POV ta, Rigs 6C 108) 8 a er eS ee es annandalei

Claspette longer and narrower, reaching to 0.75 of basimere, with 3

widened specialized setae on dorsal basal part and with numerous

long setae ventral and distal to it (Huang 1977a, Figs. 11C, 10D).

craggi

mediopunctatus subgroup*

Claspette simple, with numerous long setae on the slightly expanded

distal part and with few shorter ones on sternal side (Huang 1977a,

Pig 190). ek cata Coes s Po ee ee eee. malikuli

Claspette large and bilobed, with numerous setae and with a distinct

stout spine-like seta on apicosternal angle of expanded distal

Claspette with several distinctly long and stout setae on tergal portion

of expanded distal part (Huang 1977a, Figs. 21C, 21D).

mediopunctatus

Claspette without several distinctly long and stout setae on tergal

portion of expanded distal part (Huang 1977a, Fig. 25C).

perplexus

w-albus subgroup

Claspette with numerous setae on the expanded distal part and occupy-

ing about 0.5 of it (Huang 1977a, Fig. 29C). . gardnerii gardnerii

gardnervii imitator

Claspette with numerous stouter, widened setae on the expanded distal

part and occupying 0.67 or more of it (Huang 1977a, Fig. 33C).

w-albus

PUPAE**

Paddle margins without fringe of very long hair-like spicules

(Piss oA TR oe a a ee oa 2

Paddle margins with fringe of very long hair-like spicules

(Ee OTA Sr a a a 3

*Aedes rhungkiangensis is not included.

** The pupa of rhungkiangensis is unknown.

Huang: Aedes Stegomyia) in the Oriental Region 21

2(1). Paddle margins with distinct denticles; seta 1-P single (Fig. 2A).

AEGYPTI GROUP

aegypti (p. 41)

Paddle margins without distinct denticles, at most with slight spicules

seta 1-P branched (Figs. 7A, 10A, 12A, 14A, 16A, 19A, 21A)

ALBOLINEATUS GROUP (p. 21)

3(1). Seta 2-IV, V directly anteriad of 1-IV, V; seta 5-C strong, usually

GGUS er eee ie LC eae a EDWARDSI GROUP*

Seta 2-IV, V laterad of, or mesad of 1-IV,V........-+.... 4

4(3). Setar Zany, Vitaterad al tetris ee ee is Rs we iw we 5

Seta 2-IV, V mesad of 1-IV, V (Fig. 27A; Huang 1977a, Figs. 29A,

B9A). . sae. SBEQE eae SCUTELLARIS GROUP (p. 22)

W-ALBUS GROUP

(w-albus subgroup)

5(4). Seta 2-IV, V laterad of 3-IV, V; seta 1-II usually with 3-6 branches, not

dendritic (Huang 1977a, Fig. 14A)....... W-ALBUS GROUP

(desmotes subgroup)

desmotes

Seta 2-IV, V mesad of 3-IV, V; seta 1-II usually well developed, with

Many AenOPIlIC DYAVCN ES ix ie os Geceks de loogae ww ca pea ese 6

6(5). Male genital lobe short and broad, about as long as wide (Huang 1977a,

Vics; VA, Gon 20 a ae a es ge eS W-ALBUS GROUP

(mediopunctatus subgroup)

malikuli

mediopunctatus

perplexus

Male genital lobe long and broad, slightly longer than wide (Huang 1977a,

Bice GAs Eta a Wwe ee PO W-A LBUS GROUP

(annandalei subgroup) (p. 23)

THE ALBOLINEA TUS GROUP**

1, Seta 5-C short, about length of 4-C (Figs. 12B, 16B, 19B)..... 2

Seta 5-C strongly developed, very long, much longer than 4-C... 3

2(1). Seta 9-VII single, small and slender, about same magnitude as 9-VI;

1-P well developed, with at least 9 branches (Fig. 124A).

bambusicola

Seta 9-VII single or branched, much longer and stouter than 9-VI;

1-P with at most 5 branches (Figs. 16A, 19A)..... hoogstraali

impatibilis

3(1). Seta 5-VI usually single (1,2), short, not reaching beyond posterior

margin of following segment (Figs. 7A, 14A)........... 4

*Known from unpublished material from Andaman islands.

**The pupae of platylepidus and pseudalbolineatus are unknown.

4(3).

5(3).

2(1).

3(1).

4(3).

5(4).

6(3).

7(6).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

Seta 5-VI single or double, or 2,3 forked, usually long, reaching

beyond posterior margin of following segment. ......... 5)

Seta 9-VII usually with 2,3 branches; 6-VII usually single or double and

smaller than 9-Vib (Figs TA). .a 3. SOP ee oh albolineatus

Seta 9-VII usually with 3-5 branches; 6-VII usually with 2-6 branches

(im a) a a ee a ee boharti

Seta 9-VIII usually with 17 (14-21) branches (Fig. 21A). .. J/affooni

Seta 9-VIII usually with 13 (11-16) branches (Fig. 10A). - arboricola

THE SCUTELLARIS GROUP*

THE W-ALBUS GROUP

w-albus subgroup

Seta 9-III-V strongly developed, thickened, much stouter than 9-II

(Huang 1972. Fie. 16A;. Huang 1077a,. Fie. 839A); fs wisdawe es 2

Seta 9-III-V not strongly developed, slender, about same magnitude

as 9-II ee 6 ee ee ee eS Si ek ewe a ee ecm ae eres & 3

Seta 1-II usually with 8-10 branches (Huang 1972, Fig. 16A). ,

'seatoi

Seta 1-II usually with 2,3 branches (Huang 1977a, Fig. 334A).

w-albus

Seta 9-VI much stouter than 9-V, at least 2.0 length of 9-V (Fig. 274;

Nuance 1072, Figs: 44) 184A) 6 vee ee Ak ees ere ele 4

Seta 9-VI about as thick as 9-V, less than 2.0 length of 9-V..... 6

Seta 9-VI usually single and barbed; 9-VII usually single and barbed

or with 2 apical branches; 9-VIII usually with 2 main stems (1, 2)

reaching beyond fringe of paddle, each stem with lateral branches

of varying length (Huang 1972, Fig. 4A).......... downsi

Seta 9-VI single and simple (Fig. 27A; Huang 1972, Fig. 18A)....5

Seta 9-VIII usually with 2 main stems, barbed, not reaching beyond

fringe of paddle (Huang 1972, Fig. 18A)........ subalbopictus

Seta 9-VIII usually with single main stem (1, 2) reaching beyond fringe

of paddle, with lateral branches of varying length (Fig. 27A).

krombeini

Seta 9-VII single, stout and barbed or split at apex. ........ 7

Seta 9-Vilsincle, simple. ta. bales We es Gee os 9

Seta 6-C much shorter than 7-C (Huang 1972, Fig. 9B).

nova lbopictus

Seta 6-C about leneth Of (Cy. oti Se ARTE ws

*The pupa of wnilineatus is not included. The pupae of alorensis and andrewsi

are unknown.

8(7).

9(6).

10(9).

11(10).

12(11).

13(10).

14(13).

Huang: Aedes (Stegomyia) in the Oriental Region 23

Seta 6-C much stouter than 7-C and usually slightly longer; 1-II usually

with 10,11 branches arising from a common stem at base; 9-VIII

usually with 2 main stems, reaching beyond fringe of paddle, each

stem with lateral branches of varying length; paddle margins fringed

close to base, on more than apical 0.75 of paddle (Huang 1972, Figs.

on Me a ee eM EE UME Ce RAMU Nee imu Nem Met ye beBlusi

Seta 6- Cc usually about length of 7-C; 1-II branched, without a common

stem at base; paddle margins with fringe on less than apical 0.75

of paddle (Huang 1972, Figs. 22B, 22A)..... alcasidi (in part)

Seta 9-VIII usually with 2 (1,2) branches, each barbed, not reaching

beyond fringe of paddle (Huang 1972, Fig. 11A)..... patriciae

Seta 9-VIII reaching beyond fringe of paddle. ....... Sie aan 10

Seta: 6-C’ about: 0, Stengel Tee pi Se ee ey PA es 11

weta-6-C about: 0.75-1.00 lenetn of FO. ee a ee ee ek 13

Seta 9-VIII with 2 branches (Huang 1972, Fig. 134A).

pseudalbopictus

Seta O- Vill meually Single Tie ye ea eee ais foe oC 12

Male genital lobe short and broad, about as long as wide (Huang 1972,

SBR ya ees ee ee Ge es albopicius

Male genital lobe long and broad, longer than wide (Huang 1977a,

BE yg ce ye ye a el ae eg ee em gardnerii gardnerii

gardnerii imitator

Sela OC aout), To benctaee el ws. Givi ele BREESE 14

Seta 6-C about 1.0 length of 7-C (Huang 1972, Fig. 22B).

alcasidi (in part)

Seta 9-VIII usually with single main stem (1,2), with lateral

branches of varying length; 1-II with many primary and

secondary branches (Huang 1972, Fig. 27A)...... malayensis

Seta 9-VIII usually with 2 branches tis 2), barbed; 1-II with very few

secondary branches (Huang 1972, Fig. 32A)......... riversi

annandalei subgroup

Male genital lobe with a triangular-shaped fold on ventral side

mie 807 a, Dig. D1Aleicissceepcelgcgeeeaesg CEM craggi

Male genital lobe without a triangular-shaped he on ventral side

(Hunee 20T sa. Rig: GA ee Vee Pee annandalei

FOURTH STAGE LARVAE*

Ventral brush with 5 pairs of setae (Figs. 3C, 8C).......... 2

Ventral brush with 4 pairs of setae (Fig. 28C)............ 3

*Aedes rhungkiangensis is not included.

2(1).

3(1).

4(3).

5(4).

6(3).

7(6).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

Comb scale with very strong basal denticles (Fig. 3C).

AEGYPTI GROUP

aegypti (p. 41)

Comb scale without very strong basal denticles (Figs. 8C, 11C, 18C,

beC 276 3226) see. ALBOLINEATUS GROUP (p. 25)

Marginal spicules of anal segment well developed and conspicuous. . 4

Marginal spicules of anal segment not well developed, very small

Ale INE ONSDICUOUS. 5 668 ete ae ee SN 6

Seta 1-S inserted at level of, or before last pecten tooth. ...... 5)

Seta 1-S inserted beyond last pecten tooth (Huang 1977a, Figs. 20C,

2a, 270)... eee ee ee W-ALBUS GROUP

(mediopunctatus subgroup)

malikuli

mediopunctatus

perplexus

Marginal spicules of anal segment very long, spine-lLike.

EDWARDSI GROUP*

Marginal spicules of anal segment stout, rather short and bluntly

rounded at apex (Huang 1977a, Figs. 7C, 12C).

| W-ALBUS GROUP

(annandalei subgroup)

annandalei

craggt

Comb scales in a single row, arising from a sclerotized plate

(Hiuane 197 fas Wigs 190). oe he W-ALBUS GROUP

(desmotes subgroup)

desmotes

Comb scales in a single row, not arising from a sclerotized plate. . 7

Ventral brush with 4d-X well developed, single or branched and with

bars (Fig. 28C; Huang 1977a, Fig. 34C).

SCUTELLARIS GROUP (p. 25)

W-ALBUS GROUP

(w-albus subgroup)

w-albus

Ventral brush with 4d-X not well developed, single, very small,

much smaller than 4a, b,c-X and without bars (Huang 1977a,

Bio, 300)... ok ee a se oe W-ALBUS GROUP

(w-albus subgroup)

gardnerii gardnerii

gardnerii imitator

*Known from unpublished material from Andaman islands.

2(1).

3(1):

4(3).

5(4).

Huang: Aedes Stegomyia) in the Oriental Region 29

THE ALBOLINEAT US GROUP*

Marginal spicules of anal segment small and inconspicuous (Figs. 13C,

Rs ch penis: er ee, Bak ye Be eee ea in 2

Marginal spicules of anal segment well developed, very long and

conspicuous. (Kies. BCy 11C, TBC ae oc ie ee RE ee eae 3

Comb scales ina single row, arising from a sclerotized plate

CRs AEC: yogis ton! 4 alee eRe. RR ees bambusicola

Comb scales ina single row, not arising from a sclerotized plate

ie CATON os arg Boat mapa Wise UT ae wea See hoogstraali

Seta 5-P long, about 1.0 length of 7-P (Fig. 8B)... . albolineatus

Seta 5-P short, much shorter than 7-P (Figs. 11B, 15B. 22 B)... 4

Comb scale with free portion widened at base and sharply pointed at

apex, with fine denticles on basal portion of apical spine

CBOE ee TC 1 wcao ts spisiannes amianarcenca nee finan tere have, le Spas ee ole det) Aes boharti

Comb scale with free portion rather slender and long, about 2.0

length of attached portion, without fine denticles on basal portion

of Anieal enine dPigse 1iCe B20). fo Cece es Nee OS we 5

Comb scales ina single row, arising from a sclerotized plate

Rig, DiC he a Gti) aes. Bis, GURU Figs te Lh Ete arboricola

Comb scales ina single row, not arising from a sclerotized plate

OO ie cut a. pec se aris vera mer bs 8 es dal Sarees! '« laffooni

THE SCUTELLARIS GROU P**

THE W-ALBUS GROUP

w-albus subgroup

Comb scale with prominent denticles at base of apical spine; seta _

2-VII usually with 6 (5-8) branches (Fig. 31C)....... seatoi

Comb-scale without euch denticlesy: c.28) «Wile ee Ge ee 8 ee ow 2

Siphon with acus; pecten teeth 3-6, each tooth short and stout; usually

with 3, 4 basal denticles (Huang 1972, Fig. 14C).

pseudalbopictus

Siphon without acus; pecten tooth long, at least 4.0 as long as

Anal segment with complete saddle (Huang 1972, Figs. 10C, 19C). . 4

Anal segment with incomplete saddle. .........+6-24-ee+e8- 5)

*The larvae of impatibilis and platylepidus are unknown. The larva of

pseudalbolineatus is not included.

**The larva of unilineatus is not included. The larvae of alorensis and andrewsi

are unknown.

4(3).

5(3).

6(5).

7(6).

8(7).

9(8).

10(7).

11(10).

12(11).

Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

Seta 2-X 2-branched; pecten tooth usually with 3 (2,3) basal denticles;

1-S inserted beyond last pecten tooth and ventrad of teeth (Huang

tes Fie OC) ea ee ee pe ee ea novalbopictus

Seta 2-X usually single, when 2-branched one much smaller than the

other; pecten tooth usually with 2 (2,3) basal denticles; 1-S inserted

well beyond last pecten tooth and in line with teeth (Huang 1972,

Big 29C es gh a ae ee aes subalbopictus

Siphon short, 1.7 as long as wide at middle (Huang 1977a,

Pie. 240)... 2 os Cook ee eae ae W-ALBUS GROUP

(w-albus subgroup)

w-albus

Siphon about 2.0-2.5 as long as wide at middle. ........... 6

Seta 2-X 3-branched; 2-VII usually with 3 (2,3) branches; pecten tooth

with one large and occasionally 1,2 very small basal denticles

(Hinnecl07e.. Pies 20CeAs. So Ges ke ek See paullusi

Seta ek Senranciied. (3 26s SSO ea a Re Sk ce. i

Seta 1-VII usually with 4 (3,4) branches, short, less than 2.0 length

CEOS a a SS iy 8

Seta 1-VII usually with 2 (2,3) branches, long, at least 2.5 length of

Se ee er ee a ay ee ae he 10

Seta 2-Vil with, 3, 4 DrANCHOS. oe er ac a ee PS 9

Seta 2-VII usually single (1,2) (Huang 1972, Fig. 3C). . . albopictus

Comb scale with free portion widened at base and sharply pointed at

Pi Cie tr) ee ee a ee ae eS eS See downsi

Comb scale with free portion rather slender, nearly parallel-sided

from base and at least length of attached portion (Fig. 31E).

patriciue

Seta 1-VII with 2,3 branches; siphon about 2.0 as long as wide; pecten

teeth 10-21, closely arranged in a line; 1-S usually inserted beyond

middle of siphon; comb scale sometimes with apical spine split at

ii (gane 19y2. Fie. S80). oe ew Be Is riversi

Seta 1-VII usually with 2 long branches (2,3), when 3-branched then

one much smaller than other 2; siphon about 2.5 as long as wide;

pecten teeth 10-19; 1-S inserted at middle or before middle of

SIPHON Goes aS ES a ee. 11

Seta 1-S usually inserted at middle of siphon; pecten teeth 10-14, each

with 2-4 basal denticles (Huang 1972, Fig. 28C). . .. malayensis

Seta 1-S usually inserted before middle of siphon. ......... 12

Pecten tooth with one large and 1-3 small basal denticles (Fig. 31F).

alcasidi

Pecten tooth with 1-3 basal denticles (Fig. 28C)......- krombeini

Huang: Aedes Stegomyia)in the Oriental Region 27

NOMEN DUBIUM

Aedes rimandoi Basio. Philippines. 1971a: 52 (o*, 2). Type-locality: U.P.

College of Forestry Campus, Los Banos, Laguna, Luzon, Philippines

(NE).

TAXONOMIC DISCUSSION. According to Basio (1971la: 53), the original

description was based on 20, 19 and the type of 77mandoi was deposited in the

"RGB", Ruben G. Basio Mosquito Collection, Manila, Philippines (Basio 1971b:

31). However, numerous enquiries addressed to several persons in Manila,

Philippines have failed to show that any type-material is still in existence.

Based on the original description, vimandoi is unrecognizable and should be

considered a nomen dubium.

ACKNOWLEDGEMENTS

I wish to express my sincere appreciation to Dr. Ronald A. Ward,

Dr. Botha de Meillon, Dr. Douglas J. Gould, MAJ Bruce A, Harrison,

Mrs. Rampa Rattanarithikul, CPT Michael E. Faran and Mr. E. L. Peyton for

a critical review of the manuscript and for their valuable comments.

Iam most grateful to Dr. P. F. Mattingly, Department of Entomology,

British Museum (Natural History), London, for the loan of several types and

other material in the British Museum; Dr. M. Sasa, formerly Professor of

Parasitology, the Institute of Medical Science, University of Tokyo, for

Yamada's type-specimens and other material; the Thailand material collected,

reared and/or curated by Mr. Kol Mongkolpanya, Mr. Prajim Boonyakanist,

Mr. Sanit Nakngen, Mr. Sorasak Imvitaya, Mr. Chamnong Noigamol,

Mrs. Nantana Eikarat, Mrs. Suda Ratanawong and Mrs. Prasertsri Rohitaratana

of the U. S. Army Medical Component, Armed Forces Research Institute of

Medical Sciences, Bangkok; the Philippines material from Dr. G. L. Alcasid

and his staff, Department of Education, National Museum, Manila; the Ryukyus

material from LTC R. W. Intermill, formerly Chief, Entomology Division,

U.S. Army Medical Center, Ryukyu Islands; the Malaysian and Indian material

from Dr. S. Ramalingam and his staff, Department of Parasitology, the

University of Malaya, Kuala Lumpur; the Sri Lanka material from Dr. K. V.

Krombein and his collection teams, Department of Entomology, National Museum

of Natural History (USNM), are acknowledged with sincere appreciation. I wish

to thank also the following institutions for the loan of material: Bernice P.

Bishop Museum; USNM; Field Museum of Natural History; University of Utah;

Cornell University; John Hopkins School of Hygiene and Public Health; California

Academy of Sciences; Academy of Natural Science; Medical Zoology Laboratory,

The Institute for Medical Science, University of Tokyo; British Museum (Natural

History) and the Instituut voor Tropische Hygiene, Amsterdam.

I also wish to express my gratitude to Mr. Vichai Malikul and Mr. Young

T. Sohn for preparing the drawings, Mrs. Janet D. Rupp, for typing the manu-

script for offset reproduction, and finally to my parents and friends for their

kind encouragement.

28 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

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Asia. Contrib. Am. Entomol. Inst., (Ann Arbor). 5(4): 1-43.

RODRIGUES, F. M., M. R. PATANKAR, K. BANERHEE et al.

1972.

Etiology of the 1965 epidemic of febrile illness in Nagpur City,

Maharashtra State, India. Bull. WHO 46: 173-9.

ROSEN, L. and L. E. ROZEBOOM.

1954.

Morphologic variations of larvae of the scutellaris group of

Aedes (Diptera, Culicidae) in Polynesia. Am. J. Trop. Med.

Hyg. 3: 529-38.

RUDNICK, A. and Y. C. CHAN.

1965.

Dengue type 2 virus in naturally infected Aedes albopictus

mosquitoes in Singapore. Science 149: 638-9.

34 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

RUSSELL, P. K., D. J. GOULD, T. M. YUILL, A. NISALAK and

P. E. WINTER. |

1969. Recovery of dengue-4 viruses from mosquito vectors and

patients during an epidemic of dengue hemorrhagic fever.

Am. J. Trop. Med. Hyg. 18: 580-3.

RUSSELL, P. K., DO VAN QUY, A. NISALAK, P. SIMASATHIEN,

T. M. YUILL and D. J. GOULD.

1969. Mosquito vectors of dengue viruses in South Vietnam.

Am. J. Trop. Med. Hyg. 18: 499-9.

SANMARTIN-BARBERI, G., H. GROOT and E. OSORNO-MESA.

1955. Human epidemics in Colombia caused by the Venezuelan

equine encephalomyelitis. Am. J. Trop. Med. Hyg. 3:283-93.

SASA, M.

1976. Human filariasis. A global survey of epidemiology and

control. University Park Press, Baltimore, London and

Tokyo. 819p., 3 pl.

SMITH, T. J., P. E. WINTER, A. NISALAK and S. UDOMSAKDI.

1971. Dengue control on an island in the Gulf of Thailand II.

Virological studies. Am. J. Trop. Med. Hyg. 20: 715-9.

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

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

Culicidae). Thomas Say Found., Entomol. Soc. Am. Vol. 6,

398 p.

SUENAGA, O. and T. ITOH.

1973. Studies on the filarial prevalence among dogs and the mosquito

vectors in Nagasaki City, western Japan. 95. On the vector

mosquitoes of Dirofilaria immitis in Nagasaki City. Trop.

Med. 15: 131-40.

TANAKA, K., E. S. SAUGSTAD and K. MIZUSAWA.

1975. Mosquitoes of the Ryukyu Archipelago (Diptera: Culicidae).

Mosq. Syst. 7: 207-33. |

THEOBALD, F. V.

1901a. (June), p. 235, in Howard, L. O., Mosquitoes: how they live,

how they carry disease, how they are classified, how they

may be destroyed, New York. 241 p.

1901b. (July), The classification of mosquitoes. J. Trop. Med.

4: 229-35.

1901c. (Sept. ), Notes on a collection of mosquitoes from west Africa,

and descriptions of new species. Report of the malaria

expedition to Nigeria of the Liverpool School of Tropical

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Mem. Lpool. Sch. Trop. Med. 4 (Ap.) ii: 1-14.

1901d.

1904.

1906.

TOT:

Huang: Aedes Stegomyia) in the Oriental Region 30

(Nov.), A monograph of the Culicidae or mosquitoes.

Vol. 1, Br. Mus. (Nat. Hist.), London. 424 p.

New Culicidae from the Federated Malay States (continued).

Entomologist 37: 77-8.

Report on economic entomology. PartI. Second report on

the mosquitoes of Culicidae of the Sudan. Rep. Wellcome

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36 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

APPENDIX I. PRESENT STATUS OF THE SPECIES OF AEDES STEGOMYIA)

IN THE ORIENTAL REGION

SPECIES STAGES BIONOMICS

A. PP. LE

1. aegypti A a. At x SF Immature habitats known,

female bites man

2. albolineatus XK: KH K* XN. = Immature habitats known

3. arboricola a XR Re Immature habitats known

4. bambusicola 5 ae Immature habitats known

D9. boharti a Od Re Immature habitats known,

female bites man

6. hoogstraali xt X* X* MF cg Immature habitats known

7. impatibilis a Se Immature habitats known

8. laffooni a” &* X* X* Immature habitats known

9. platylepidus Bm ee Immature habitats known

10. pseudalbolineatus ae ok Immature habitats known

11. edwardsi Ga, Ca ae Pe Immature habitats known

12. seampi - XK*¥ - - - Unknown

13. albopictus APR XR A a Immature habitats known,

female bites man

14. downsi > Glee Gale, Sa, Glee Immature habitats known,

female bites man

15. novalbopictus Rte. A a = Immature habitats known

16. patriciae Xt. RX Rt. Immature habitats known

17. pseudalbopictus x ae Ke a Immature habitats known

18. seatoi Ke Immature habitats known,

female bites man

19. subalbopictus AO RT XR Immature habitats known

20. wunilineatus a7 ke OR A Immature habitats known

21. alcasidi Ae ee ee Immature habitats known

22. alorensis > Unknown

23. andrewsi BT RX ee oe os Unknown

24. krombeini RO DK = Immature habitats known,

female bites man

25. malayensis > Sse. gee, Gale. Gea Immature habitats known,

female bites man

26. paullusi 2 ae we. S Immature habitats known,

female bites buffalo

27. viversi a ee Immature habitats known,

female bites man

28. annandalei xt X* A* AF KF Immature habitats known,

female bites man

29. craggi X* X* X* X*F - Immature habitats known,

female bites man

30. desmotes At RE Xe KR Immature habitats known,

female bites man

31. malikuli X* - X* X* - Immature habitats known

Huang: Aedes (SStegomyia) in the Oriental Region 37

APPENDIX I (Continued).

SPECIES STAGES BIONOMICS

A PL &E

32. mediopunctatus X* Xe oa AS Immature habitats known,

female bites man

33. perplexus RY RO a Immature habitats known,

female bites man

34. vhungkiangensis a Xx Se RT Unknown

35. garvdnerii gardnevii Par, See. alee, Su Fam Immature habitats known

36. gardnerii imitator A ee a Immature habitats known,

female bites man

37. w-albus D Sale. Glee. Se. Suma Immature habitats known,

female bites man

X* = Stage or sex described and illustrated.

- = Stage or sex unknown.

X = Stage or sex described.

Contrib. Amer. Ent. Inst.

» vol. 15, no. 6, 1979

APPENDIX II. DISTRIBUTION LIST OF THE ORIENTAL SPECIES OF AEDES (STEGOMYIA)

ORIENTAL REGION

SPECIES

1. aegypti

2. albolineatus

Ryukyu Islands

iwan

ee China (S. of Yangtze Kiang)

3. arboricola

4, bambusicola

Cambodia

@¢

5. boharti

6. hoogstraali

7. impatibilis

Sangi Islands, Taroena

Celebes, Kabaena

Sabah

ox Sarawak

W. Malaysia

Poeloch Weh

ee Singapore

~ @& Sumatra

Flores

ap Kalimantan, Tarakan

[

8. laffooni

9. platylepidus

. pseudalbolineatus

11. edwardsi

. seampi

. albopictus

14. downsi

. novalbopictus

16. patriciae

. pseudalbopictus

. seatoi

. subalbopictus

. unilineatus

. alcasidi

22. alorensis

. andrewsi

24, krombeini

. malayensis

zr rversi_—|®

28. annandalei

29. craggi

30. desmotes

31. malikuli

32, mediopunctatus

33. perplexus

ES) @@

34, rhungkiangensis

35, gavdnerii gardnerii

36. gardnerii imitator 12? @eee@

37. w-albus

ee = Areas from which specimens were available for examination.

: = Species recorded from the area but have not been seen by the author.

Huang: Aedes (SStegomyia) in the Oriental Region 39

APPENDIX II (Continued)

ORIENTAL REGION OTHER AREAS

Ethiopian

SPECIES

Chagos Islands

India

Sri Lanka

La Reunion

Madagascar

Solomon Islands peeerey S

Sudan

Bangladesh

Morotai

Ceram

Hawaiian Islands

N. Rhodesia

& Nigeria

@) Mauritius

& Seychelles

1. aegypti

2. albolineatus pee

lee

3. arboricola

4, bambusicola

5. boharti

6. hoogstraali

| [| @e

7. impatibilis

8. laffooni

9. platylepidus

10. pseudalbolineatus

Blea ee a

11. edwardsi

12. seampti

13. albopictus

14. downsi

15. novalbopictus

16. patriciae

17. pseudalbopictus

18. seatoi

19. subalbopictus

20. unilineatus

21. alcasidi

22, alorensis

23. andrewsi

24, krombeini

25, malayensis

26. paullusi

27. viversi

28, annandalei

29. craggi

30. desmotes

. malikuli

. mediopunctatus

. perplexus

. rhungkiangensis

. gavdnerii gardnerii

. gardnerii imitator

ae = Areas from which specimens were available for examination.

? = Species recorded from the area but have not been seen by the author.

40 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

APPENDIX III.

SUMMARY OF THE IMMATURE HABITATS OF THE SPECIES OF AEDES GSTEGOMYIA) IN THE ORIENTAL REGION*

IMMATURE

HABITATS

SPECIES

1. aegypti

2. albolineatus

3. arboricola

coconut stumps, banana stumps)

coconut husks, coconut spathes)

Coconut shells (coconut halves,

Fallen leaves (fallen abaca leaf)

“-

—

Artificial containers (tin can,

Stump holes (palm stumps,

Bamboo internodes

Bamboo cups (bamboo pots)

nipa palm, sago palm, taro,

Colocasia)

Palm fronds (coconut fronds)

Footprints (wheel ruts)

bottle, tire)

& Rock holes

Log holes

Split bamboos

Cut bamboos

Rock pools

Root holes

Bamboo stumps

@ oo water jar, water bucket, broken

4, bambusicola

5. boharti

6. hoogstraali

7. impatibilis

8. laffooni

9. platylepidus

@@@ eee e

| 10. pseudalbolineatus

11. edwardsi

12. albopictus

13. downsi

| 14. novalbopictus

15. patriciae

16. pseudalbopictus

17. seatoi

18. subalbopictus

@ ecoeeeecoeeooeoe eo

eco ee

‘fe @

19. unilineatus

20. alcasidi

hal

21. krombeini

22. malayensis

23, paullusi

24, rviverst

25. annandalei

26. craggt

27. desmotes

28, malikult

ie © eeecoeooe

29. mediopunctatus

30. perplexus

31. gardnerii gardnerii &

32. gardnerii imitator |\@

shea ao

33. w-albus &

*Immature habitats are unknown for the following species: seampi, alorensis, andrewsi and rhungkiangensis.

Huang: Aedes (Stegomyia) in the Oriental Region 41

APPENDIX IV. Aedes (SStegomyia) aegypti (Linnaeus)

Type species: Aedes (Stegomyia)| aegypti (Linnaeus)

(Figs. 1, 2, 3, 4, 5

Culex aegypti Linnaeus 1762: 470 (A).

Culex argenteus Poiret 1787: 245 (9).

Culex fasciatus Fabricius 1805: 36 (A).

Culex calopus Meigen 1818: 3 (¢, 9).

Stegomyia fasciata var. queenslandensis Theobald 1901d: 297 (£).

Stegomyia fasciata persistans Banks 1906: 996 (c, 9°).

Duttonia alboannulis Ludlow 1911: 132 (&).

For complete synonymy, see Knight and Stone (1977).

Aedes (Stegomyia) aegypti (Linnaeus), Dyar 1920a, 1920b: 181, 204 (taxonomy);

Edwards 1932: 161 (systematics); Barraud 1934: 221 (o*, ¢, L*,

variations); Bohart and Ingram 1946: 6, 11, 22, 27, 37, 66 (o*, 2*, P*,

L*, bionomics, Pacific distribution); Knight and Hull 1952: 167 (¢, ¢, L*);

Bonne-Wepster 1954: 75 (o, 2*, L*); Mattingly 1957: 392 (taxonomy,

bionomics); Belkin 1962: 441 ('*, 2, P*, L*); Mattingly 1965: 23 (*, 2*,

P*, L*; Neotype designated).

Aedes Stegomyia) fasciatus (Fabricius), Bonne-Wepster and Brug 1932: 47

(**, 2, L*, biology).

Aedes Stegomyia) argenteus of authors.

Stegomyia fasciata of authors.

Aedes calopus of authors.

Stegomyia calopus of authors.

MALE (Fig. 1A). Head. Proboscis dark scaled, without pale scales on

ventral side, longer than forefemur; palpus 5-segmented, as long as proboscis,

dark, with a white basal band on each of segments 2-95; those on segments 4, 5

dorsally incomplete; segments 4,5 subequal, slender, upturned, and with only a

few short setae; antenna plumose, shorter than proboscis; torus covered with

white scales except on dorsal and ventral sides; clypeus bare; erect forked

scales pale, not numerous, restricted to occiput; vertex with a median stripe of

broad white scales, with broad dark ones on each side interrupted by a lateral

stripe of broad white scales followed ventrally by a patch of broad white scales.

Thorax. Scutum with narrow dark scales and a distinct, small, median white

spot of narrow scales on anterior border; a narrow submedian longitudinal stripe

of narrow pale yellowish scales on each side of midline, submedian stripes reach

to anterior 0.66 of scutum; a few narrow white scales on the posterior portion of

midline forming a short median white stripe just in front of prescutellar space;

on each side a narrow prescutellar white line present; broad curved lateral

prescutal and posterior fossal lines of crescent-shaped silvery white scales

followed by a narrow line of narrow white scales reaching to the posterior mar-

gin of scutum; a patch of broad white scales on lateral margin just in front of

wing root and a few narrow curved pale scales over wing root; acrostichal

bristles absent; dorsocentral bristles present; scutellum with broad white scales

on all lobes and with a few broad dark scales at apex of midlobe; anterior pro-

notum with broad white scales; posterior pronotum with a patch of broad white

scales and some dark and pale narrow scales dorsally; paratergite with broad

white scales; postspiracular area without scales; patches of broad white scales

on propleuron, on the subspiracular and hypostigial areas, on prealar, on the

42 Contrib. Am. Ent. Inst., vol. 15, no. 6, 1979

upper and lower portions of sternopleuron, and the mesepimeron; upper ster-

nopleural scale patch not reaching to anterior corner of sternopleuron; upper

mesepimeral scale patch not connecting with lower mesepimeral scale patch;

lower mesepimeron without bristles; metameron bare. Wing. With dark scales

on all veins except for a minute basal spot of white scales on costa; cell Ro 1.5

length of R913. Halter. With dark scales. Legs. Coxae with patches of white

scales; knee spots present on all femora; fore- and midfemora with a narrow

white longitudinal stripe on ventral half of anterior surface; hindfemur anteriorly

with a broad, white, longitudinal stripe which widens at base and is separated

from apical white scale patch; all tibiae dark anteriorly; fore- and midtarsi with

a basal white band on tarsomeres 1, 2; hindtarsus with a basal white band on

tarsomeres 1-4, the ratio of length of white band to the total length of tarsomere

is 0.33, 0.33, 0.33 and 0.75; tarsomere 5 all white; foreleg with tarsal claws

unequal, the larger one toothed, the smaller one simple; midleg with tarsal

claws unequal, both simple; hindleg with tarsal claws equal, simple. Abdomen.

Segment I with white scales on laterotergite and a large, median, pale patch;

terga II-VIII with basolateral white spots; terga II-VI each with a basal pale

band which does not connect with the basolateral white spots; sterna I-III largely

covered with pale scales; sterna IV-VI each with a basal pale band and lateral

white spots which do not connect with the basal bands. Terminalia (Figs. 1B,

1C, 1D, 2C). Basimere about 2.2 as long as wide, scales restricted to lateral

and ventral areas, with a few setae on basomesal area of dorsal surface, mesal

surface membranous; claspette large, oval, lobelike, with numerous setae and

with several widened specialized setae, of which 3 or 4 of these setae bent at tip,

on mesal side of expanded distal part; distimere simple, elongate, about 0.63

length of basimere, apical 0.28 rather narrow and curved, with a short spini-

form process at apex and with a few setae on apical 0.25; aedeagus strongly

toothed; paraprocts with sternal arms; cercal setae absent; apical margin of

tergum IX deeply concave medially and with 3 setae on each lateral lobe; ster-

num IX without bristles.

_ FEMALE. Essentially as in the male, differing in the following respects:

Head. Palpus about 0.2 length of proboscis, with white scales on apical half,

4-segmented, or sometimes 5-segmented, when present segment 5 minute;

clypeus with lateral white scale patches. Wing. Cell Ro about 2.0 length of

R9+3. Legs. Hindtarsomere 4 with basal 0.4 white band; fore- and midlegs

with tarsal claws equal, all toothed. Abdomen. Terga II-VI each with a basal

white band and basolateral white spots which do not connect with the basal bands;

tergum VII with lateral white spots only; a row of small white scales along pos-

terior border on II-VII; segment VII largely retracted. Terminalia (Figs. 4,5).

Apical margin of sternum VIII with a deep U-shaped notch at middle and with

conspicuous rounded lateral lobes; insula longer than broad, with minute setae

and with 5 larger setae on apical 0.25; apical margin of tergum IX with well

developed lateral lobes, each with 5 or 6 setae; apical margin of postgenital

plate with a shallow notch; cerci short and broad; 3 spermathecae, one larger

than the other 2.

PUPA (Figs. 2A, 2B). Cephalothorax. Trumpet short, about 2.0 as long

as wide at the middle; setae 1,3-C usually single (1,2), longer than 2-C, 2-C

usually double (1,2), 4-C usually single (1,2), 5-C usually with 2,3 branches,

6-C single, longer than 7-C, 7-C usually double (1,2), 8-C usually double

(2,3), 9-C single, 10-C 2-branched, mesad and caudad of 11-C, 11-C single,

stout, 12-C usually single (1-3). Abdomen. Seta 1-I well developed, with more

than 10 branches, dendritic, 2-I single, 3-I single, long, 2,3-I widely separated,

distance between them about 1.5 of the distance between 4, 5-I; seta 1-II usually

Huang: Aedes (SStegomyia) in the Oriental Region 43

with 3 branches (2-4); 3-II, III usually single (1,2), shorter than segment III;

1-II usually double (1-3); 1-IV usually double (1,2); 2-IV, V slightly mesad of

1-IV, V; 5-IV-VI usually single (1,2), short, not reaching beyond posterior

margin of following segment; seta 9-I-VI small, single, simple, sometimes

9-VI double; 9-VII usually single (1,2) and barbed; 9-VII, VIII much longer and

stouter than preceeding ones; 9-VIII usually with 5 branches (3-7) and barbed.

Paddle. Margins with distinct denticles, without fringe of very long hair-like

spicules; apex rounded; seta 1-P single. Male genital lobe short and broad,

much shorter than wide.

LARVA (Fig. 3). Head. Antenna 0.5 length of head, without spicules;

seta 1-A inserted slightly before middle of shaft, single, small, spine-like;

inner mouth brushes pectinate at tip; seta 4-C well developed, usually with 5

branches (4-7), closer to 6-C than 5-C, cephalad and mesad of 6-C, 5-C usual-

ly single, long, 6-C usually single, 7-C usually single, rarely double, 8-10,

13-C single, 11-C usually with 3 branches (2,3), 12-C usually double (2, 3),

14, 15-C usually with 2,3 branches; mentum usually with 12 (10-14) teeth on

each side. Thorax. Setae 1,7, 14-P usually with 3 branches (2,3), 2,6-P

single, 3-P double, 4,9,11-P usually single (1,2), 5-P usually double; 5, 7-M

single, 6-M usually 3-branched, 8-M usually with 4 branches (3,4), 9-M usually

2-branched (2,3), 10,12-M single, long, stout and barbed, 11-M single, small;

7-T usually with 4 branches (4, 5), 9-T usually 2-branched, 10,11-T similar to

those on mesothorax, 12-T much reduced; basal spine of meso- and metapleural

seta long, stout, pointed at tip and apically hooked. Abdomen. Seta 6-I-III

usually with 3 branches (2-4), 7-I single; 7-II usually double (2, 3); 6-IV, V

usually double (2,3); 6-VI usually single (1,2); 1-VII usually double (1-3),

2-VII single; 2-VIII distant from 1-VIII, 1, 5-VIII usually with 3, 4 branches,

3-VIII usually with 6 branches (5-7), 2,4-VIII single; comb of 8 (6-12) scales in

a row, each scale with very strong denticles at the base of the apical spine; anal

segment with saddle incomplete, marginal spicules very small and inconspicu-

ous; seta 1-X usually double (1,2), short, 2-X usually with 3 branches (2, 3),

3-X usually single (1,2); ventral brush with 5 pairs of setae on grid, each seta

usually 2-branched (2,3), sometimes 4e-X single; no precratal tufts; anal

papillae about 2.5-3.0 length of saddle, sausage-like. Siphon. About 1.8-2.5

as long as wide, acus absent; pecten teeth usually 8-20, evenly spaced, or some-

times with the last pecten tooth widely spaced, each tooth with 1-4 basal denticles;

seta 1-S with 3,4 branches, usually inserted beyond last tooth and beyond middle

of siphon.

MATERIAL EXAMINED. More than 1,070 adults (% and $), 390 individual

rearings (2911, 390 p), 43 “ terminalia, 25 2 terminalia, 210 whole larvae

from the Oriental region.

REMARKS. Adult specimens of "variety queenslandensis"' have more pale

scales present on the vertex, thorax and abdomen than that of typical aegypii.

The range of scaling varies from that almost referable to typical aegypti to

complete covering of the vertex and abdomen with white scales. The main

character used to identify "variety queenslandensis" is the white scales present

on at least the first and 2nd abdominal tergite. Typical aegypti has white scales

present on the first abdominal tergite.

4G Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

APPENDIX V.

Aedes (SStegomyia) platylepidus Knight and Hull

(Fig. 24)

Aedes (?) platylepidus Knight and Hull 1951: 201 (9).

Aedes (Diceromyia) platylepidus Knight and Hull, Mattingly 1959: 43 (9);

Reinert 1970: 12 (o'*, 9*).

MALE. Terminalia. (Figs. 24A, 24B, 24C). Basimere short and broad,

about 2.0 as long as wide, scales restricted to lateral and ventral areas, with

several irregular rows of stout, flattened, long setae along mesal side of dorsal

surface, with several short setae scattered on lateral half of dorsal surface;

claspette with 5 stout, spine-like setae and several long, stout setae with slender

curved tips on distal part; distimere simple, elongate, about 0.8 length of basi-

mere, tapering to a blunt apical point, with a long, stout, spiniform process at

basal 0.4, apex of the spiniform blunt and mesally curved; aedeagus with several

apical teeth on each side; paraproct long, without apical lobe-like process; cer-

cal setae absent; apical margin of tergum IX flat at middle, with 5 setae on each

Side.

REMARKS. The above description was based on the only known terminalia,

(slide #68/1142 from adult #2252, Isabella, Basilon Is., Philippine Is., 27 Sept.

1945) in USNM, which was used by Reinert (1970) in his original description of

SO platylepidus. Figure 24C was drawn from the original undissected view of the

o terminalia.

In order to give a correct and detailed description of the S terminalia, I

have dissected the claspette and remounted the terminalia.

MATERIAL EXAMINED: 1¢, 292, 1% terminalia, 1¢ terminalia.

2Oll te =e

svmisi¥H9 J

ES.

OV 13dIHOWy S09 VHO

SONV1S!

& wveooin til

"S| NYNVGNY

i | svuauw(|

1TV9O NAS

ONVTIVH AS, i

ps IYW ONVIHO>

=) i

Lf (wvssy)

PION inviniigyl

ee ~run_,VSNd

Looe

y BVH ig

“S

ningnnve

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eX Ny 37.9NN"

Contrib. Am. Ent. Inst., vol. 15, no. 6, 1979

LIST OF FIGURES

Aedes Stegomyia) aegypti - A, % adult; B, aedeagus; C, paraproct; D,

tergum IX and sternum IX.

Aedes Stegomyia) aegypti - A, B, pupa; C, % terminalia.

Aedes Stegomyia) aegypti - A, B, C, larva.

Aedes Stegomyia) aegypti - 2 terminalia.

Aedes Stegomyia) aegypti - A, B, C, D, ? terminalia.

Aedes (Stegomyia) albolineatus - A, % adult; B, aedeagus; C, © terminalia.

Aedes (Stegomyia) albolineatus - A, B, pupa; C, ¢ terminalia.

Aedes (Stegomyia) albolineatus - A, B, C, larva.

Aedes (Stegomyia) albolineatus - A, B, C, D, $ terminalia.

Aedes (Stegomyia) arboricola - A, B, pupa; C, % terminalia.

Aedes Stegomyia) arboricola - A, B, C, larva.

Aedes (Stegomyia) bambusicola - A, B, pupa; C, % terminalia.

Aedes (SStegomyia) bambusicola - A, B, C, larva.

Aedes (SStegomyia) boharti - A, B, pupa; C, % terminalia.

Aedes (Stegomyia) boharti - A, B, C, larva.

Aedes (Stegomyia) hoogstraali - A, B, pupa; C, % terminalia.

Aedes (Stegomyia) hoogstraali - A, B, C, larva.

Aedes (SStegomyia) hoogstraali - A, & hindleg.

Aedes (Stegomyia) impatibilis - B, % hindleg; C, D, thorax.

Aedes (Stegomyia) impatibilis - A, B, pupa; C, > terminalia.

Aedes (Stegomyia) bambusicola - A, & adult.

Aedes Stegomyia) laffooni - B, C, thorax.

Aedes (Stegomyia) laffooni - A, B, pupa; C, “ terminalia.

Aedes (SStegomyia) laffooni- A, B, C, larva.

Aedes Stegomyia) laffooni - A, % and 2? hindlegs.

Aedes (Stegomyia) pseudalbolineatus - B, % and 2 hindlegs; C, D,

o terminalia.

Aedes (Stegomyia) platylepidus - &% terminalia.

Aedes (Stegomyia) unilineatus - A, thorax; B, midfemur; C, D,

_ & terminalia.

Aedes Stegomyia) unilineatus - A, B, C, D, 2 terminalia.

Aedes (Stegomyia) krombeini - A, B, pupa; C, % terminalia.

Aedes Stegomyia) krombeini - A, B, C, larva.

Aedes (SStegomyia) krombeini - A, B, C, D, ¢ terminalia.

Aedes (Stegomyia) albopictus - A, C, thorax.

Aedes (Stegomyia) krombeini - B, thorax.

Aedes (SStegomyia) pseudalbopictus - D, thorax.

Aedes (SStegomyia) pseudalbopictus - A, thorax.

Aedes (Stegomyia) subalbopictus - B, thorax.

Aedes (Stegomyia) seatoi - C, comb scale.

Aedes (SStegomyia) downsi - D, comb scale.

Aedes (Stegomyia) patriciae - E, comb scale.

Aedes (Stegomyia) alcasidi - F, pecten tooth.

aedeagus

ig. 1

dorsal ventral

sternal

arm

Loe

ee,

dorsocentra/

bristles

Aedes (Stegomyia) aegypti (Linnaeus)

Sekt hus

Y\3

WK. Paraprocts

: \ aedeagus

0-1

basimere

Aedes (Stegomyia) aegypti (Linnaeus)

yes

Aedes (Stegomyia) aegypti (Linnaeus )

cerc/

postgenita/ plate

IXth tergum

spe athecae

Vill th sternum

b+ 0:2. ——__

Aedes (Stegomyia) aegypti (Linnaeus)

/X th tergum

t— 0+] —4

insula

V/// th tergum

has eth

Vill th sternum

/—— 0:2 ——J

Aedes (Stegomy/a) aegypti (Linnaeus)

Fig. 6

aedeagus

Aedes (Stegomyia) albolineatus (Theobald)

a ole gee

Aedes (Stegomyia) a/bolineatus (Theobald)

p (N

ONY CONN :

CUA TG UL, \S

8 Fr SA

A ga y

Bini Gi el “

iin, tin Ge rt

aeV'5, | hol bY es

rs ‘

nat AG

Ait 7a

a : !

Aedes (Stegomyia) albolineatus (Theobald )

rye ae cerc/

) ‘a

Fig. 9

postgenital plate

/X th tergum

t———— 0:] -—+

insula

; , BSR A "4 i “a 1 ASVENG Rat ‘ Ae i

i Gaeta Ww SAYA | ie

nN “i

\ 1X, \ Ky

WW | ‘il iW

—— ZEB

Villth sternum Vill th tergum

hall

Aedes (Stegomyia) albolineatus (Theobald)

Fig.10

Knight & Rozeboom

Aedes (Stegomyia) arboricola

; te,

Wy,

Uy,

ALAN)

ra)

on (

Knight & Rozeboom

Aedes (Stegomyia) arboricola

Aedes (Stegomyia) bambusicola Knight & Rozeboom

Aedes (Stegomyia) boharti Knight & Rozeboom

i AS

‘

0 Wt S

HEAL AA

Aedes (Stegomyia) boharti Knight & Rozeboom

O-2

Fig. 16

distimere

claspette

basimere

Aedes (Stegomyia) hoogstraali Knight & Rozeboom

dlchaitradihed,

* Figs?

A A

\ ke

Mis

Ey awe ; t/

: \y | fp

j : Y, 13

i= 2)

_—

WN UT fal =>

I i hic a oe

Aedes (Stegomyia) hoogstraali Knight & Rozeboom

hoogstraali

lis

tibil

impa

tibilis

impa

Aedes (Stegomyia) impatibilis ( Walker)

Fig.19 2 \

x = ee PS Ps

vit) wf 4

Want. Y

fpr: Ne

Aedes (Stegomyia) impatibilis (Walker)

laffoont

Aedes (Stegomyia) bambusicola Knight & Rozeboom

Aedes (Stegomyia) laffooni Knight & Rozeboom

SSS

tS \}

y LOG

laffooni

pseuda/bolineatus

Aedes (Stegomyia) pseuda/bolineatus Brug

; : Das a i At; >

f (“ ; a" Ie i ;

/ “A 5 ye

\ 5 Zi Z, Be } i/ (

f by

(Sy;

/ Os 1G J

Y Lge

Aedes (Stegomyia) platylepidus Knight & Hull

Fig. 25

midfemur

Aedes (Stegomyia) unilineatus (Theobald)

0-1

cerc/

VIII th sternum

Aedes (Stegomyia) unilineatus ( Theobald)

postgenita/ plate

IX th tergum

Villth tergum

UehaiMalehul,

lateral

aspect

0-2

My (

{

Aedes (Stegomyia) krombeini Huang hehaMbabebal

Lo RIG 28

aif A

: ON ae

= S4R—1

A oe 6 Cc

Aedes (Stegomyia) krombeini Huang

cerci F ig. 99

postgenital

plate

aN 2

A at ey

[Xtergum

ee Ot oa et

spermathecae

Vill tergum

Vill sternum ——__ 0:2 ————“+

Meh Mpalthl

Aedes (Stegomyia) krombeini Huang

Fig. 30

“~

qq ~~

% ©

8S:

Qaé

DS!

m ©

AS)

“en

\@)

over wing root

rea)

a/bopictus

krombeini

a/bopictus

pseuda/bopictus

pseudalbopictus

Fig. 31

Gs fe

— Siar eee BEE

“xk

~ a

a8 a

iS wn

S VY

ee epee eng cite

CS se a) ee ee sek

i on PTE seats ware? he

SSR

aerate Peg

Ben oat

subalbopictus

haiMabhale

downs! seato!

patriciae

alcasidi

78 Contrib. Amer. Ent. Inst., vol. 15, no. 6, 1979

INDEX

Names of valid taxa are set in roman type; synonyms are in italicized type.

Italicized numerals refer to the principal text references.

Roman numerals

refer to secondary text references; the suffix ''k'"’ indicates mention in a key and

the suffix ''t'' indicates mention in a table.

without a suffix refer to the figures.

Aedes Meigen 1, 4, 6

Aedimorphus Theobald 4, 6

aegypti group 2, 4, 5t, 6, 9, 12k,

16k, 21k, 24k

aegypti (Linnaeus) 1, 2, 3, 5t, 8, 9,

9t, 10t, 11t, 12k, 16k, 21k, 24k,

$6t, S8t, SOt, 40¢, 41, 43;:(1, 2,

3, 4, 5)

Alanstonea Mattingly 6

alboannulis Ludlow 41

albolineatus group 2, 4, 5t, 7, 12k,

13k, 17k, 21k, 24k, 25k

albolineatus (Theobald) 3, 4, 5t, 13k,

17k, 22k, 25k, 36t, 38t, 39t, 40t,

(6, 7, 8, 9)

albopictus (Skuse) 2, 5t, 8, 9, 11t,

12t, 14k, 18k, 23k, 26k, 36t, 38t,

39t, 40t, (30)

albopictus subgroup 4, 5t

alcasidi Huang ot, 15k, 19k, 23k, 26k,

36t, 38t, 39t, 40t, (31)

alorensis Bonne-Wepster ot, 15k,

19k, 22, 25, 36t, 38t, 39t, 40

andrewsi Edwards ot, 14k, 19k, 22,

29, 36t, 38t, 39t, 40

annandalei subgroup 4, 5t, 15k, 17k,

20k, 21k, 23k, 24k

annandalei (Theobald) 5t, 9, 15k, 20k,

23k, 24k, 36t, 38t, 39t, 40t

arboricola Knight and Rozeboom 9t,

13k, 18k, 22k, 25k, 36t, 38t, 39t,

40t, (10, 11)

argenteus Poiret 41

Ayurakitia Thurman 6

bambusicola Knight and Rozeboom 2,

ot, 13k, 18k, 21k, 25k, 36t, 38t,

39t, 40t, (12, 13, 20)

boharti Knight and Rozeboom 5t, 9,

13k, 17k, 22k, 25k, 36t, 38t, 39t,

40t, (14, 15)

calopus Meigen 41

Roman numerals in parentheses

Chikungunya virus 9, 9t

Colocasia 8

craggi (Barraud) 5t, 9, 15k, 20k, 23k,

24k, 36t, 38t, 39t, 40t

dengue viruses 9, 10t

Dengue 1 virus 9t, 10t, 11t

Dengue 2 virus 10t, 11t

Dengue 3 virus 10t, 11t

Dengue 4 virus 9t, 10t, 11t

desmotes (Giles) 5t, 9, 15k, 16k, 21k,

24k, 36t, 38t, 39t, 40t

desmotes subgroup 4, 5t, 15k, 16k,

21k, 24k

Diceromyia Theobald 2, 6

Dirofilaria immitis 9, 12t

Dirofilaria sp. 9, 12t

downsi Bohart and Ingram 5t, 9, 14k,

18k, 22k, 26k, 36t, 38t, 39t, 40t,

(31)

edwardsi (Barraud) 4, 5t, 13k, 17k,

36t, 38t, 39t, 40t

edwardsi group 2, 4, dt, 7, 12k, 13k,

17k, 21k, 24k

fasciatus Fabricius 41

formosus (Walker) 2

gardnerii Ludlow 3

gardnerii gardnerii (Ludlow) 5t, 16k,

20k, 23k, 24k, 37t, 38t, 39t, 40t

gardnerii imitator (Leicester) 5t, 9,

16k, 20k, 23k, 24k, 37t, 38t, 39t,

40t

haemorrhagic fever 9

hoogstraali Knight and Rozeboom 2,

ot, 13k, 17k, 21k, 25k, 36t, 38t,

39t, 40t, (16, 17, 18)

impatibilis (Walker) 2, 5t, 13k, 17k,

21k, 25, 36t, 38t, 39t, 40t, (18, 19)

Huang: Aedes (Stegomyia) in the Oriental Region 79

krombeini Huang 5t, 9, 15k, 19k, 22k,

26k, 36t, 38t, 39t, 40, (27, 26,

29, 30)

laffooni Knight and Rozeboom dt, 13k,

18k, 22k, 25k, 36t, 38t, 39t, 40t,

(20, 21, 22, 23)

maehleri group 4

malayensis Colless ot, 9, 15k, 19k,

23k, 26k, 36t, 38t, 39t, 40t

malikuli Huang 5t, 16k, 20k, 21k, 24k,

36t, 38t, 39t, 40t

mediopunctatus subgroup 4, 5t, 16k,

20k, 21k, 24k

mediopunctatus (Theobald) 5t, 9, 16k,

20k, 21k, 24k, 37t, 38t, 39t, 40t

meronephada (Dyar and Shannon) 2

novalbopictus Barraud 5t, 14k, 19k,

22k, 26k, 36t, 38t, 39t, 40t

pandani group 4

Pandanus 8

patriciae Mattingly 5t, 14k, 19k, 23k,

26k, 36t, 38t, 39t, 40t, (31)

paullusi Stone and Farner dt, 15k, 19k,

23k, 26k, 36t, 38t, 39t, 40t

perplexus (Leicester) 5t, 9, 16k, 20k,

21k, 24k, 37t, 38t, 39t, 40t

persistans Banks 41

platylepidus Knight and Hull 1, 2, ot,

13k, 18k, 21, 25, 36t, 38t, 39t,

40t, 44, (24)

pseudalbolineatus Brug 5t, 13k, 18k,

21, 25, 36t, 38t, 39t, 40t, (23)

pseudalbopictus (Borel) 5t, 14k, 19k,

23k, 25k, 36t, 38t, 39t, 40t, (30,

31)

Pseudostegomyia Ludlow 3

Quasistegomyia Theobald 3, 4

queenslandensis (Theobald) 2, 41, 43

Reovirus 3 11t

rhungkiangensis Chang and Chang ot,

15, 20, 23, 37, Jet, 39t, 40

rimandoi Basio 27

riversi Bohart and Ingram 5t, 9, 15k,

19k, 23k, 26k, 36t, 38t, 39t, 40t

scutellaris group 2, 4, 5t, 7, 9, 12k,

14k, 17k, 18k, 21k, 22k, 24k, 25k

scutellaris subgroup 4, 5t

scutellaris (Walker) 9

Scutomyia Theobald 3

seampi Huang 5t, 13k, 17, 36t, 38t,

39t, 40

seatoi Huang 5t, 9, 14k, 18k, 22k, 25k,

36t, 38t, 39t, 40t, (31)

Stegomyia Theobald 1, 2, 3, 4, St, 6,

8, 9, 12k

subalbopictus Barraud 5t, 14k, 19k,

22k, 26k, 36t, 38t, 39t, 40t, (31)

subperiodic filaria 9

Tembusu virus 12t

unilineatus (Theobald) 3, 5t, 14k, 18k,

22, 25, 36t, 38t, 39t, 40t, (25, 26)

Venezuelan equine encephalitis virus 9

vittatus (Bigot) 4

vittatus group 4

w-albus group 2, 4, 5t, 8, 12k, 15k,

16k, 17k, 20k, 21k, 22k, 24k, 25k,

26k

w-albus subgroup 4, 5t, 16k, 17k,

20k, 21k, 22k, 24k, 25k, 26k

w-albus (Theobald) 5t, 9, 16k, 20k,

22k, 26k, 37t, 38t, 39t, 40t

Yellow fever virus 9

Zika virus 11t

| cere e aan

eae

~. 08

beeps Peeled Bee inter ve mt

a ee Ae ae gers es

ae } sbaigora’ss et

ES Bodie PUBS Dhy

ae ae ee re) ibe

a sine here

iAIRB ORua Tl, BAS NS

ai MB 28.5) Siemon

oe mes: HES pie te *

re et eg |

gk es #

q dibisciloints att

BW — ee

Contributions -

of the

American Entomological Institute

Volume 15, Number 7, 1980

MOSQUITO STUDIES (Diptera, Culicidae) XXXIV.

A revision of the Albimanus Section

of the subgenus Nyssorhynchus of Anopheles

Michael E. Faran

CONTENTS

INTRODUCTION

MATERIAL AND METHODS .

TAXONOMIC HISTORY

TAXONOMIC CHARACTERS

SYSTEMATICS

BIONOMICS he

MEDICAL IMPORTANCE .

TAXONOMIC TREATMENT .

Albimanus Section .

Keys to Groups and Species.

Albimanus Group :

1. Anopheles (Nys.) albimanus

OswaldoiGroup .

Oswaldoi Subgroup .

Oswaldoi Complex .

2. Anopheles (Nys.) oswaldoi .

. Anopheles (Nys.) galvaoi

. Anopheles (Nys.) noroestensis .

. Anopheles (Nys.) aquasalis .

. Anopheles (Nys.) ininii

. Anopheles (Nys.) rangeli .

Oo ON WNW NH HB W

. Anopheles (Nys.) trinkae

10. Anopheles (Nys.) nuneztovari

Strodei Complex .

11. Anopheles (Nys.) strodei

12. Anopheles (Nys.) rondoni .

13. Anopheles (Nys.) benarrochi .

Triannulatus Subgroup

14. Anopheles (Nys.) triannulatus

REFERENCES

FIGURES :

TABLE OF DISTRIBUTIONS |

CONSPECTUS OF TAXONOMIC CHANGES .

INDEX TO SCIENTIFIC NAMES

. Anopheles (Nys.) anomalophyllus .

. 106

loge 1

ne ae

At ees

. 136

. 141

ne ae

. 178

eet

ete

Wes We,

MOSQUITO STUDIES (Diptera, Culicidae)

XXXIV. A REVISION OF THE ALBIMANUS SECTION

OF THE SUBGENUS NYSSORHYNCHUS OF ANOPHELES'

Michael E. Faran?

INTRODUCTION

Zavortink (1973:4), when he was reviewing the subgenus Kerteszia of Anopheles,

stated, “*... the systematics of Nyssorhynchus are too poorly known and hopelessly

confused.” In 1942, L. E. Rozeboom (1942b) used only one couplet in a key to the

adult females for 9 different species in the Albimanus Section. The reason for this,

as the pioneering work of W. H. W. Komp (1942:25-26) and Rozeboom recognized,

is that many of the characters that have been used to distinguish the adult females in

the Albimanus Section are extremely variable and unreliable for species identifica-

tion. On the basis of one character alone it is often impossible to identify with any

confidence an adult female as belonging to a particular species in this section, which

includes several important vectors of malaria. The purpose of this revision is (1) to

describe and illustrate, in detail, all the currently known species in the Albimanus

Section, (2) to reconcile some of the systematic problems that have resulted in this

aura of confusion and (3) to develop effective keys that use more than one character,

when possible, for all stages of the life cycle for the species in this section.

I am subdividing the subgenus Nyssorhynchus into 2 sections, the Albimanus Sec-

tion and the Argyritarsis Section. These sections are very closely allied and form a

tight, well-defined unit. The entire subgenus is restricted to the Neotropics except

for albimanus which extends into the Nearctic. The Albimanus Section is distin-

guished from the Argyritarsis Section in the adults primarily by the basal dark band

on hindtarsal segment 5, and in the male genitalia by the variously developed fused

ventral claspette. Only on the basis of these 2 characters can these sections be readi-

ly differentiated. The Myzorhynchella group has been excluded from consideration

because of the paucity of material available for study and because of its uncertain

taxonomic position.

In the present revision 14 species are recognized in the Albimanus Section. The

nominal species sanctielii is not included because of the lack of material. An. gorgasi

‘This research was supported by the Medical Entomology Project, Smithsonian Institution, U.

S. Army Medical Research and Development Command Research Contract DAMD-17-74C-4086;

the Mosquitoes of Middle America project, University of California, Los Angeles, supported by U.

S. Army Medical Research and Development Command Research Contract DA-49-193-MD-2478

and U. S. Public Health Service Research Grant AI-04379; and U. S. Public Health Service Predoc-

toral Trainee Grant 5T01 AI00070-14.

Captain, Medical Service Corps, U. S. Army, Department of Entomology, Walter Reed Army

Institute of Research, Washington, DC 20012.

2 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

is removed from synonymy with albimanus and relegated to status of anomen du-

bium as is evansi which was formerly considered to be the senior synonym of strodei.

The synonymy of metcalfi is changed from oswaldoi to that of noroestensis. An. tri-

annulatus is treated as a single, although variable, form, not as being composed of 2

separate subspecies, t. triannulatus and t. davisi (or t. bachmanni).

An attempt has been made to assemble the species into distinct monophyletic

groups on the basis of correlated characters in the adults and immatures. The charac-

ter state, ancestral or derived, of the taxonomically important features has been de-

termined whenever possible and evolutionary trends stated in formulating the phylo-

genetic relationships within the section. I am dividing the section into 2 groups, the

monotypic Albimanus Group and the Oswaldoi Group. An. albimanus is the least de-

rived species in the section, possessing several ancestral features which it shares with

some of the ancestral species in the Argyritarsis Section. An. albimanus is easily dit-

ferentiated from the Oswaldoi Group by several correlated unique features in the

adult, male genitalia and larva.

I am separating the Oswaldoi Group into 2 subgroups, the monotypic Triannulatus

Subgroup and the Oswaldoi Subgroup composed of 12 species, further separated into

the Oswaldoi Complex and the Strodei Complex.

The Oswaldoi Complex consists of 9 species. Within the complex, 2 separate phy-

letic lines are discernible on the basis of the structure of the ventral claspette of the

male genitalia. One line is composed of oswaldoi, galvaoi, noroestensis, aquasalis, in-

inii and possibly the relict anomalophyllus, and the other by rangeli, trinkae and nu-

neztovari.

The Strodei Complex contains strodei, rondoni and benarrochi. An. strodei and

rondoni are very closely related, but their relationship to benarrochi is difficult to de-

termine as benarrochi is the most derived species in the section. An. benarrochi has

been placed in the Strodei Complex only because of the similarity of its male genita-

lia with those of strodei and rondoni. |

The larvae and pupae of the Albimanus Section occur primarily in ground water

habitats. All the species breed in fresh water, except for aquasalis and often albima-

nus which are often found in brackish water. The females feed predominantly on

large mammals. The adults are active either crepuscularly or nocturnally; triannula-

tus is the only species reported to bite occasionally during the day.

Two species, albimanus and aquasalis, are major vectors of malaria in Central and

South America and the islands of the Caribbean. An. nuneztovari is the primary vec-

tor of malaria in western Venezuela and northern Colombia and possibly a vector in

Suriname. An. triannulatus, strodei and norestensis have been implicated in the

transmission of malaria, since they have been found naturally infected. An. rangeli

has been suspected of transmitting malaria in Ecuador. Two viruses, Venezuelan

Encephalitis virus and Tlacotalpan virus, have been isolated from aquasalis and albi-

manus respectively. Nothing is known about the medical importance of the remain-

ing species.

I would like to express my deep gratitude to John N. Belkin for his constant sup-

port and guidance. I thank the other members of my doctoral committee, A. Ralph

Barr, George A. Bartholomew, Walter Ebeling and Peter Vaughn, for their encourage-

ment and helpful advice.

I am greatly indebted to Ronald A. Ward of Walter Reed Army Institute of Re-

search and Oliver Flint of the Smithsonian Institution for the loan of material; to P.

Cova Garcia and A. Gabaldon, J. F. Reinert and D. R. Roberts, J. B. Kitzmiller, R. F.

Darsie, Jr., and R. S. Panday for valuable specimens from Venezuela, Brazil, Colom-

bia, El Salvador and Suriname respectively. Also, I would like to thank the following

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 3

individuals and institutions that have so generously loaned specimens for this study:

Paul H. Arnaud, California Academy of Sciences, San Francisco; Henry S. Dybas,

Field Museum of Natural History, Chicago; Peter F. Mattingly and Graham B. White,

the British Museum (Natural History), London; L. L. Pechuman, Cornell University,

Ithaca; Selwyn S. Roback, Academy of Natural Sciences, Philadelphia; Milan Trpis,

Johns Hopkins University, Baltimore; and Pedro W. Wygodzinsky, American Museum

of Natural History, New York.

I am very grateful to my friends and colleagues George K. Bryce and Kenneth Lin-

thicum for reading and criticizing the manuscript and to J. Hal Arnell, O. G. W. Ber-

lin and Thomas J. Zavortink for suggestions, advice and assistance whenever sought.

To Sandra Heinemann, William Powder, Thomas Gaffigan, Ellen Paige, Ruby Sims

and Melvin Stave I wish to express my thanks for the preparation of material and

miscellaneous assistance.

During my field studies I have been assisted by many people. I would especially

like to thank Nelson Papavero of the Museo de Zoologia, Sao Paulo, Oscar de Souza

Lopes and his staff of Instituto Adolfo Lutz, Sao Paulo, and Herbert C. Barnett and

Ricardo Iglesias Rios of Instituto de Microbiologia, Rio de Janeiro.

I wish to extend my sincerest appreciation to Sharon Toalson, Trinka Faran and

Cynthia Lomax for the difficult and frustrating job of typing the preliminary drafts

of the manuscript. I thank Sandra Heinemann for criticizing and editing the manu-

script, as well as preparing the text copy for lithoprinting. Also, I thank Nobuko Ki-

tamura, L. Margaret Kowalczyk, Young Sohn and Vichai Malikul for preparation of

the preliminary and final drawings.

MATERIAL AND METHODS

MATERIALS. This taxonomic revision is based on a study of 14,792 specimens:

2816 males, 774 male genitalia, 6078 females, 2088 pupae and 3036 larvae, including

1682 individual rearings (755 larval, 741 pupal, 186 incomplete) and 30 progeny

rearings. Individually reared specimens were available for 11 of the 14 species (all

but benarrochi, galvaoi and rondoni); progeny rearings were available for 5 species

(benarrochi, oswaldoi, rangeli, triannulatus and trinkae). Except for galvaoi and ron-

doni, larvae and pupae of all species were available for study. For 2 of the medically

more important species (albimanus and aquasalis), abundant individually reared ma-

terial from almost their entire geographic range was examined, permitting the analysis

of intra- and interpopulational variation(s) within these 2 species.

The majority of the material for the present study was collected (or otherwise ac-

quired) by the project ‘“‘Mosquitoes of Middle America’ (MOMA); this material has

recently been transferred from the University of California at Los Angeles to the

“Medical Entomology Project” (MEP) at the U. S. National Museum of Natural His-

tory [USNM]. In the Distribution section for each species, the specimens listed are

all deposited at the USNM, unless otherwise noted by the presence of the abbrevia-

tion for the depository in brackets. In order to conserve space, specimens from the

MOMA collection for which collection data have been published are listed only by

code and collection number under the correct locality (example: ““Humazones, HON

59’). All other specimens are listed individually by number and type, along with lo-

cality, date and collector. Most collection localities are not given in detail, but only

by the nearest town. The number and types of specimens are summarized and listed

directly after the country or major political division. The collection records of the

4 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

“Mosquitoes of Middle America’’ project have been published in the following series

of papers: Belkin, Heinemann and Page 1970; Belkin and Heinemann 1973, 1975a,

1975b, 1976a, 1976b, 1976c; Heinemann and Belkin 1977a, 1977b, 1977c, 1978a,

1978b, 1978c, 1979. These papers should be consulted for complete data for the

following codes: ANT, BAH, BH, BRA, BRB, CAY, COL, COM, CR, CUB, DOM,

ECU, FWI, FWIM, FG, FGC, GG, GR, GRR, GUA, GUY, HAC, HAR, HAT, HON,

JA, LAR, LU, MAR, MEX, MNT, MX, NI, NIC, NVS, PA, PER, PR, PRA, PRX,

RDO, SAL, SUR, VI, VZ.

The only other major sources of material were the USNM and the School of Public

Health, Johns Hopkins University. Additional individually reared material collected

by John F. Reinert and Donald R. Roberts from Para, Brazil has been particularly

important in this study since it included the only specimens of ininii. Other sources

of material include collections by A. Gabaldon, P. Cova Garvia, J. B. Kitzmiller, R. F.

Darsie, Jr., R. S. Panday, W. H. W. Komp, F. M. Root and L. E. Rozeboom. Material

was also borrowed from Paul H. Arnaud, California Academy of Sciences, San Fran-

cisco [CAS]; A. Ralph Barr, University of California, Los Angeles [UCLA] ; Marga-

ret Thayer, Museum of Comparative Zoology, Harvard University, Cambridge [MCZ] ;

Henry S. Dybas, Field Museum of Natural History, Chicago [FMNH]; Peter F. Mat-

tingly and Graham B. White, British Museum (Natural History), London [BM]; L. L.

Pechuman, Cornell University, Ithaca [CU]; Selwyn S. Roback, Academy of Natural

Sciences, Philadelphia [ANS]; Milan Trpis, Johns Hopkins University, Baltimore [JH];

and Pedro W. Wygodzinsky, American Museum of Natural History, New York [AMN

H]. Recently, valuable material was collected and field observations made by John

N. Belkin, J. Hal Arnell, Thomas E. Rogers, Kenneth J. Linthicum and myself in the

state of Sao Paulo, Brazil, January to March 1975, and by John N. Belkin, George K.

Bryce and myself in the state of Rio de Janeiro, December 1975 to March 1976; the

data for these collections are given in Heinemann and Belkin 1979. I have examined

the types of davisi, anomalophyllus, emilianus, dunhami, galvaoi (paratype), goeldii,

gorgasi, metcalfi and strodei [USNM]. J. N. Belkin has examined the types of gal-

vaoi, noroestensis, rondoni, cuyabensis, albimanus and albipes.

TAXONOMIC PROCEDURES. The methods and techniques utilized in this study

are essentially those of comparative morphological taxonomy as described by Belkin,

Schick, Galindo and Aitken (1965:10; 1967:10-11). An attempt has been made to

assemble the species described here into separate monophyletic taxa. In almost all

cases this was done on the basis of the correlation of several derived features in the

larva, adult and male genitalia. The determination of the character states, derived

(apomorph) or ancestral (plesiomorph), was accomplished using the methods previ-

ously described and discussed by Mayr (1969:181-257) and Hennig (1966). Bionom-

ics and distributional data also were employed as taxonomic characters in an attempt

to explore evolutionary trends within the section.

DESCRIPTIONS. The terminology, abbreviations and form of presentation are es-

sentially those of Belkin (1962) with the modifications of Belkin (1968a:49). The

descriptions are all composites based on as many specimens as were available. In

most cases the chaetotaxy of the immatures was based on at least 10 individuals, un-

less otherwise stated, and is listed as 2 sets of figures for all the setae that are consid-

ered to be taxonomically important. The first set of numbers after the hair is the

modal condition, representing the frequency of at least 75% of the observations. The

set of numbers following the modal condition is the entire range of all observations.

A few special terms are used for the wing spots and the male genitalia. These terms

are discussed in the chapter on Taxonomic Characters. Most measurements were

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 5

made from slides with the aid of a calibrated ocular micrometer in the eyepiece of a

compound microscope, except for a few of the adults in which measurements were

made in a similar fashion with a stereoscopic dissecting microscope. Intra- and inter-

populational variations and any anomalous forms are analyzed in the Discussion sec-

tions.

SYSTEMATICS. The taxonomic history, when pertinent, and systematics are dis-

cussed for each species. This includes affinities, evolutionary trends, possible centers

of origin, intrapopulational and geographical variations, and any other information

relevant to the systematics of the species. In the discussions under the groups, sub-

groups and complexes particular attention has been given to the interrelationships of

the species within each taxon and the taxon’s relationships with other related taxa.

BIONOMICS. Much of the information regarding the bionomics of the species for

which there was adequate material collected by the project “Mosquitoes of Middle

America” was extracted from the field collection data forms on file at UCLA. Most

of this data has since been published in the “Collection records of the project “Mos-

quitoes of Middle America’ ”’ series (see p 4). The data from the collection forms are

combined with information condensed from the literature about the natural history

of each species. |

MEDICAL IMPORTANCE. A brief summary of the medical importance of each

species is included following the bionomics. This section is only a summary and does

not in any way attempt to be an inclusive discussion of the literature on this subject.

It pertains primarily to malaria and what is known concerning the vector capacity of

each species.

DISTRIBUTION. The known distribution is given for each species followed by a

list of the material examined (see p 3-4). In the ““Table of Distributions,” under each

species for the different countries, a solid circle is used to indicate that a specimen

(or specimens) of that species was examined from that locality. A starred circle indi-

cates a distribution record taken from what is believed to be a reliable reference. The

distribution for each species is outlined in figures 1-3. On the figures, a solid line bro-

ken by a dotted line implies uncertainty regarding the distribution of that particular

species within the area bordered by the dotted line.

TAXONOMIC HISTORY

Several good reviews have been published on the taxonomic history of the Albima-

nus Section, previously referred to as ‘““Tarsimaculatus Series,’’ ““Tarsimaculatus Com-

plex,” “‘Albimanus Series’’ or ‘‘Albimanus Group” (Root 1926b; Lima 1928; Gabal-

don 1940; Rozeboom and Gabaldon 1941; Komp 1941b; Galvao and Lane 1938:

Galvao 1940, 1943; Floch and Abonnenc 1943a; Causey, Deane and Deane 1945; Be-

jarano 1957). For this reason I will only briefly summarize the history of this group,

indicating some of the major taxonomic problems which have led to considerable

confusion. Most of the problems relating to a particular species will be dealt with in

the Discussion section for the species concerned. The single most perplexing prob-

lem in the past regarding the species in the Albimanus Section has resulted from the

use of the name “tarsimaculatus.’’ This one taxonomically invalid name has hindered

the understanding of this group of mosquitoes more than any other factor. The tax-

onomic history of the Albimanus Section is, in many ways, the history of “‘tarsima-

culatus.”’

The oldest species in the subgenus Nyssorhynchus is albimanus, which was describ-

6 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

ed by C. R. W. Wiedemann in 1820. A few years later, J. B. Robineau-Desvoidy

(1827) described argyritarsis, the first species now recognized in the other major sec-

tion of Nyssorhynchus bearing its name. Theobald (1901) described albipes as a va-

riety of argyrotarsis [sic] which, he stated, differed from argyrotarsis only by the

presence of a black basal band on the last hindtarsal segment. The type locality was

not specified. Theobald (1903:110-113) raised albipes to specific rank, placing it in

his new genus Cellia, which included New World as well as Old World species of Ano-

pheles. Shortly afterward, Goeldi (1905), after studying material in Para, Brazil, pro-

posed the name tarsimaculata as an emendation for Theobald’s albipes in order to

avoid confusion with the name albitarsis. Coquillett (1906:8, 13) synonymized albi-

pes with albimanus. Dyar and Knab (1906b:160-161) also correctly synonymized al-

bipes with albimanus and recognized that tarsimaculatus was proposed as a substitute

for albipes. However, they mistakenly concluded that since the material Goeldi exa-

mined was not albimanus or argyritarsis, Goeldi’s name tarsimaculata was available

for the specimens they had examined from Sao Paulo and Manaus, Brazil, and Trini-

dad. In 1907, Dyar and Knab described a new species, gorgasi, based on a badly

damaged, mutant specimen collected by A. H. Jennings on the Pacific side of the

Panama Canal Zone. The same year Theobald (1907:106-109) synonymized cuben-

sis Agramonte 1900, albipes and tarsimaculata with albimana. Theobald (1910:69)

in a footnote stated, ‘“‘All the long series [of tarsimaculatus| sent me by Professor

Goeldi from Para are the same as those received from many other places, and are un-

doubtedly albimana.”’ Knab (1913), like Theobald, synonymized albipes and cuben-

sis with albimanus; however, he synonymized gorgasi with tarsimaculata. In their

fourth volume of “‘The mosquitoes of North and Central America and the West In-

dies,’ Howard, Dyar and Knab (1917:975-979) interpreted tarsimaculata as being a

valid species even though they realized Goeldi had not proposed a new species and

stated, ““There is therefore no original description, but the species is figured and with

a discussion the new name is published. We have therefore felt justified in recogniz-

ing Goeldi’s name as the first valid name for the species before us.”’ They also recog-

nized 2 other species in Nyssorhynchus, albimanus and argyritarsis.

Subsequently, after Howard, Dyar and Knab accepted tarsimaculatus as being val-

id, almost all the species in the Albimanus Section have at some time been designated

as tarsimaculatus or as synonyms of tarsimaculatus. Root (1924b:460-462) studied

the male genitalia of “oswaldoi,”’ tarsimaculata and albimanus. He stated that tarsi-

maculata is easily distinguished from albimanus; however, he could not find any gen-

italic differences between tarsimaculata and oswaldoi, so he “relegated” the name os-

waldoi as a synonym of tarsimaculata. Root (1926b) described strodei and listed al-

bimanus, tarsimaculatus, bachmanni, and tentatively rondoni, triannulatus and cuya-

bensis, plus a number of other species in the Argyritarsis Section, in the subgenus

Nyssorhynchus. Concerning tarsimaculatus, Root stated that the range of variation

in the black area on the second hindtarsal segment is continuous, ranging from 8 to

35%, and the larvae, pupae and male genitalia of the Cellia oswaldoi type of the Bra-

zilians, ‘“‘seem to be quite identical with those of normal tarsimaculatus.”’ Therefore

he saw no reason for retaining the name oswaldoi.

Christophers (1924:38-41) recognized albimanus and 5 different varieties of the

species tarsimaculatus which included oswaldoi, rondoni, cuyabensis, triannulatus

and a Bonne variety from Surinam (Bonne 1923, and Bonne and Bonne-Wepster

1925, recognized 2 types of tarsimaculatus, one on the coast and one in the interior).

Dyar (1928:434-446) in his monograph ‘““The Mosquitoes of the Americas,”’ as did

Root, considered oswaldoi a synonym of tarsimaculatus, listing 15 “‘species” in Nys-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 7

sorhynchus of which 7, albimanus, bachmanni, rondoni, cuyabensis, triannulatus, ev-

ansi and tarsimaculatus, are now included in the Albimanus Section. Later Curry

(1932) described 2 more varieties of tarsimaculatus, var. aquasalis and var. aquacael-

estis. Curry recognized, however, that his aquacaelestis could be synonymous with

_ oswaldoi or possibly a valid species, and not merely a variety of tarsimaculatus.

To unnecessarily complicate matters, Townsend (1933a:102) stated that the tarsi-

maculata of Howard, Dyar and Knab 1917 was not the tarsimaculata of Goeldi 1905,

and proposed gorgasi as an available name for the former species. In another paper

Townsend (1933b) stated, “It seems strange that the name tarsimaculatus was ever

adopted from Goeldi by Howard, Dyar and Knab.”’ Unfortunately, as Komp (1941b:

793) stated, ““He [Townsend] plunged the subject into confusion by making several

unwarrantable assumptions.” Townsend believed that albitarsis and albimanus (the

tarsimaculatus that Townsend thought Goeldi examined) were both present in Belem

during the time Goeldi collected there, and later these 2 species were replaced by

gorgasi of Townsend and An. (Nys.) darlingi Root 1926. Townsend called specimens

he collected at Rio Tapajos, Boa Vista, Para, Brazil gorgasi. Some of this latter ma-

terial was identified as goeldii by Rozeboom and Gabaldon (1941:97). When Komp

(1941b) examined mounts of the male genitalia from Boa Vista labelled by Town-

send tarsimaculatus Goeldi, he discovered some of them to be triannulatus.

In 1937 Galvao and Lane revalidated oswaldoi and stated that tarsimaculatus of

Root 1926 is really a variety of oswaldoi which they called oswaldoi var. metcalfi.

An. metcalfi was based principally on a description of an egg by Root (1926b:700).

Galvao and Lane, at a meeting of the Sociedade de Biologia de Sao Paulo, 15 Sep-

tember 1937, described the varieties metcalfi and noroestensis, but their published

descriptions did not appear until later (Galvao and Lane 1938:169-178). At this

meeting they also stated that they recognized tarsimaculatus Goeldi 1905 as being

valid because of Article 21 of the International Code of Zoological Nomenclature

which stated, “The author of a scientific name is that person who first publishes the

name in connection with an indication, a definition, or a description, unless it is clear

from the contents of the publication that some other person is responsible for said

name and its indication, definition, or description.”’ So by 1937, Galvao and Lane

recognized tarsimaculatus separately from oswaldoi, the latter composed of 3 varie-

ties, oswaldoi, metcalfi and noroestensis. Lane (1939:27-29), in his ‘“‘Catalogo das

mosquitos neotropicos,” listed gorgasi, evansi, aquasalis and aquacaelestis as syno-

nyms of tarsimaculatus.

Three years later Rozeboom and Gabaldon (1941:96) stated, ‘‘. .. we do not see

how Article 21 applies to this [tarsimaculatus| case. Goeldi did not realize that he

was dealing with a new species, as he proposed ‘tarsimaculata’ simply because albipes

has the same meaning that albitarsis has and might lead to confusion. Therefore,

Goeldi proposed his name illegally, for Article 32 of the International Rules states,

‘A generic or specific name, once published, cannot be rejected, even by its author,

because of inappropriateness.’’? Komp (1941b:793) was in agreement with the lat-

ter authors and proposed a new species, emilianus from Belem, Para, in order to ‘“‘ob-

viate further difficulties in the nomenclature of the species in this group... .” An.

emilianus was supposedly the species Goeldi worked with in 1905. One year later,

Galvao and Damasceno (1942), for exactly the same reasons as given by Komp, syn-

onymized emilianus with tarsimaculatus. Since then tarsimaculatus (=aquasalis) has

continued to be used as recently as 1958 (Rachou 1958; Andrade 1958a, b) and pos-

sibly even more recently in Brazil.

Regarding the higher classification of the group, Theobald described the genus La-

8 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

verania in 1902 with the type argyrotarsis [sic]. At the time Theobald (1902) de-

scribed Laverania he also described the genus Cellia, with pharoensis as type species.

Cellia included also pulcherrimus, squamosus and bigotii. The same year Blanchard

(1902) proposed Nyssorhynchus as a replacement for Laverania Theobald 1902, be-

cause the name was preoccupied by Laverania Grassi and Feletti 1890. In 1903, in

Volume 3 of his ““Monograph of the Culicidae of the World,” Theobald catalogued

the species argyrotarsis and albipes in Cellia, even though he synonymized Laverania

with Nyssorhynchus. By 1910, in Volume 5, Theobald included argyrotarsis and al-

bimana of Nyssorhynchus in the genus Cellia, along with several Old World species.

Unfortunately Theobald based his classification on the adult scalation patterns.

Knab (1913:34), criticizing the classification of Anopheles, wrote, “*. . . the subject

[Anopheles| was made needlessly difficult by hasty work and by the sub-division of

the old genus Anopheles into numerous ill-defined and fancifully differentiated gen-

era. |

Christophers (1915) studied the male genitalia of Anopheles and stated that al-

though he was not at that time concerned with nomenclature, generic subdivision of

anophelines would include the following 3 amended genera, Anopheles, Myzomyia

and Nyssorhynchus. Root (1923:266) agreed in general with Christophers’ classifica-

tion but considered Nyssorhynchus to be of subgeneric rank. Lima (1928) followed

Christophers’ interpretation, but divided the genus Nyssorhynchus into 2 subgenera,

Nyssorhynchus and Kerteszia; he further subdivided the subgenus Nyssorhynchus in-

to 2 groups, Group A (in part Argyritarsis Section) and Group B which included 11

““species’’ now in the Albimanus Section. Edwards (1932:43-46) separated the sub-

genus Nyssorhynchus into 3 groups, Group A being Nyssorhynchus excluding Myzo-

rhynchella, which was divided into 3 series, Argyritarsis, Tarsimaculatus and Rondoni.

The Tarsimaculatus series included species with a basal dark band on the second and

fifth hindtarsal segments, hindtarsal segments 3 and 4 being entirely white. The Ron-

doni series comprised those species which had dark bands on hindtarsal segments 3

and 4 in addition to hindtarsal segments 2 and 5.

It was not until 1940 that Gabaldon divided the Tarsimaculatus series of Edwards,

which he called the Albimanus Series, into 3 subseries based on the presence or ab-

sence of setae on the ventral claspette of the male genitalia. These subseries were the

monotypic Albimanus Subseries, the Triannulatus Subseries (triannulatus and stro-

dei) and the Oswaldoi Subseries (oswaldoi, aquasalis, anomalophyllus and nunezto-

vari). By 1952 (Gabaldon and Cova Garcia 1952:178), benarrochi and rondoni had

been added to the Triannulatus Subseries, and galvaoi, ininii, konderi, noroestensis,

rangeli and sanctielii had been placed in the Oswaldoi Subseries.

In another interpretation of the phylogenetic relationships within the subgenus,

Galvao (1943:141-142) divided Nyssorhynchus into 2 series, the Argyritarsis Series

and the Tarsimaculatus Series corresponding to Lima’s Groups A and B respectively.

He further separated the Tarsimaculatus Series into a monotypic group, albimanus,

and 3 complexes: the tarsimaculatus Complex (tarsimaculatus, oswaldoi, aquasalis,

anomalophyllus, noroestensis, rangeli and nuneztovari), the rondoni Complex (ron-

doni and strodei including subspecies albertoi, arthuri, artigasi and lloydi) and the

triannulatus Complex (triannulatus triannulatus, t. davisi, t. chagasi and benarrochi).

In 1949 Levi-Castillo divided the subgenus Nyssorhynchus into the Albimanus

Group and the Argyritarsis Group. The Albimanus Group was divided into 3 series,

Albimanus, Oswaldoi and Triannulatus which corresponded to the subseries of Gabal-

don. However, in addition to strodei, triannulatus, rondoni and benarrochi, Levi-

Castillo also included galvaoi and nuneztovari in the Triannulatus Series. Until the

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 9

present revision no other major changes have been made in the higher classification

of the Albimanus Section.

TAXONOMIC CHARACTERS

I have attempted, whenever possible, to determine the character state, whether an-

cestral (plesiomorph sensu Hennig 1966) or derived (apomorph sensu Hennig 1966),

of the taxonomically important characters, in order to facilitate the hypothesis of

the probable phylogenetic relationships among the species within the Albimanus Sec-

tion (fig. 4). In some cases it has been very difficult to ascertain the character state

of certain features because of the tremendous number of similarities among the spe-

cies resulting from convergent and parallel evolution, and/or secondary loss of mor-

phological traits.

When possible, I have stated evolutionary trends which may have taken place with-

in the section, regarding individual characters in the adult, male genitalia, pupa and

larva. In developing and interpreting the proposed phylogeny of the Albimanus Sec-

tion I have relied on the correlation of as many derived characters as possible in all

the life stages. Ancestral characters, along with certain derived features, were im-

portant in determining the phylogenetic relationships of the 2 more ancestral mono-

typic taxa, the Albimanus Group and the Triannulatus Subgroup.

The most reliable characters for species identification are in the male genitalia and

the larva. The external morphology of the adult and pupa, particularly in the case of

the Oswaldoi Subgroup, is very similar interspecifically, and usually quite variable

intraspecifically. For this reason, the keys for the adults and pupae are not always

entirely reliable when used by themselves. It is highly recommended that one refer

to the species descriptions and discussions when attempting to identify any species

in this subgroup, and that one correlate those data with the information given on the

bionomics and distribution. As has been emphasized by Belkin (1962), it is best to

examine more than one specimen. And to be certain of an identification, the imma-

tures should be individually reared and slides prepared of their exuviae and of the

genitalia of the corresponding males to permit the correlation of characters in the dif-

ferent life stages. Where morphologically similar species occur sympatrically it is

doubly important to use more than one developmental stage. Lastly, it must be em-

phasized that not all the populations from every locality where each species exists

were examined. Thus, the range of variation reported is conservative, may not be in-

clusive and may be exceeded in some individuals. Also, as indicated by Arnell (1973:

5), the illustrations of the larvae and pupae show only the modal condition of the se-

tal branching and cannot represent the range of variation which occurs in a species.

ADULTS

The important characters for differentiating the females are seen in the (1) penul-

timate segment [4] of the palpus, (2) presence or absence of scales on the mesepimer-

on, (3) banding pattern of the legs, (4) relative length of wingspots and (5) dark cau-

dolateral scale tufts of the abdomen.

HEAD. The only character which varies appreciably among the species is the light

color of the scales vesting the penultimate segment of the palpus and, to a much les-

ser extent, the erect scales on the vertex. The trend in the scale pattern of the fourth

palpal segment appears to be a shift from predominantly dark in the ancestral condi-

tion to mostly light in the derived. In the Albimanus Group palpal segment 4 is dark

10 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

or almost completely dark; whereas, throughout the Oswaldoi Subgroup this segment

is conspicuously light, at least mediolaterally. The Triannulatus Subgroup has an ap-

preciable amount of light scaling on palpal segment 4 but generally less than in the

Oswaldoi Subgroup.

THORAX. Very few differences are found in the thorax. Primitively, there was

probably an upper mesepimeral patch of light scales which has been lost in all species

except rangeli, and occasionally trinkae and noroestensis. The premise of the anti-

quity of these scales is based on an out-group comparison, the fact that several spe-

cies in the Argyritarsis Section also possess upper mesepimeral scales. An. triannula-

tus is distinguished by the key feature, a patch of light scales on the anterior mesepi-

meron, rather than on the upper mesepimeron. This character is present only in one

other species in Nyssorhynchus, darlingi of the Argyritarsis Section.

LEGS. The hindtarsus was consistently studied in the past with little regard for

the banding patterns of the fore- and midtarsus. In the fore- and midtarsus, as in the

hindtarsus, there seems to have been a trend toward the development of a larger and

more extensive light apical band. In albimanus foretarsal segments 4 and 5 are al-

most completely dark. In triannulatus foretarsal segment 5 is all dark, but foretarsal

segment 4 has a large apical light band. Throughout the Oswaldoi Subgroup the api-

cal light band varies in size, but is usually present on foretarsal segments 1, 2, 3 and

5, with segment 4 being largely dark. The extreme condition in seen in ininii where

all the foretarsal segments are largely light, with dark scales occasionally present only

as a narrow dorsobasal stripe. The midtarsal segments are usually similarly banded;

however, the light apical band is much less conspicuous, smaller and usually cream or

golden, rarely white. There is no apical light band on midtarsal segments 3 or 4 in al-

bimanus. Midtarsal segment 4 of the Oswaldoi Subgroup is usually all dark or with a

rather inconspicuous apical light band, except in ininii where a distinct apical light

band is present. Primitively the second hindtarsal segment probably possessed a large

basal dark band, 0.5 or more the length of the segment. In all taxa above the species

level, there is at least one species that retains this character. There has been an inde-

pendent reduction in the size of the dark basal band in the Oswaldoi and Strodei

Complexes, the band being smallest in oswaldoi and ininii. In other species, such as

rondoni and aquasalis, the large dark basal band of hindtarsal segment 2 may be inde-

pendently derived.

WING. The terminology used for the wing spots is modified noun Zavortink

(1973: fig. 5). The trend has been for the reduction in size of the dark spots along

the costal vein and corresponding spots on the more posterior veins. The basal dark

spot of the costal vein is moderately large in albimanus and large in triannulatus. In

the Oswaldoi Subgroup it is small, usually less than the length of the humeral light

spot, except in nuneztovari and some trinkae, where it is large and may be indepen-

dently derived. In general the wings are very similar throughout the Oswaldoi Sub-

group except for rangeli and rondoni. In rangeli there has been a very definite in-

crease in the size of the light spots of veins C, R, Rs, R,, R, and Ry.;. In rondoni

the opposite has occurred and the dark spots are much more extensive. The subbasal,

presectoral and sectoral dark spots on vein C usually are fused into a very long, single

dark spot. Correspondingly, the dark spots on the posterior veins are in general very

large. The preapical dark spot of vein M in rondoni extends unbroken onto vein

1+2:

ABDOMEN. I believe that the dark caudolateral scale tufts were primitively pre-

sent on segments II-VII. In albimanus the scale tufts are absent on II.

MALE GENITALIA. The male genitalia offer the best and most easily identifiable

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 11

group characters. In the Albimanus Section, the male genitalia are complex and high-

ly derived with respect to other anopheline groups. The terms dorsal and ventral in

this study refer to the orientation of structures of the male genitalia prior to rotation,

that is, the morphological tergal and sternal surfaces respectively.

The subgenus Nyssorhynchus is characterized by the fusion of the ventral clasp-

ettes to form a single median structure, and by the single parabasal spine on the side-

piece. The male genitalia of the Albimanus Section have been studied extensively by

previous workers and have been regarded as being of primary importance in species

identification. Characters and structures seldom examined in the past are sternite IX,

the relative length of the parabasal spines, the setal pattern of the sidepiece, and fea-

tures of the dorsal claspette. There are no good group characters on segment VIII.

Sternite IX. In ventral aspect, the sternite appears short and subrectangular in the

Albimanus Group and moderately long to long and subtrapezoidal to subtriangular in

the Oswaldoi Group. Along the anterior margin there is an apodeme (anterior apo-

deme) which is very short and inconspicuous in the Albimanus Group and moderate-

ly narrow and subrectangular, to broad and subtriangular in the Oswaldoi Group.

Sidepiece. There is a trend for a relative shortening of the parabasal spine in sever-

al species within the Oswaldoi Complex. The parabasal spine and tubercle are, in

general, longer in albimanus, triannulatus and the Strodei Complex, and shorter in the

Oswaldoi Complex.

Dorsal Claspette. Although not important as a section feature, the morphology of

the dorsal claspette may aid in identification of species within the section and should

be used in correlation with other characters of specific value.

Ventral Claspette. The fused ventral claspette best delimits the taxa within the

section and has been used most often to divide the section into different groups (Ga-

baldon 1940; Galvao 1943; Levi-Castillo 1949). The various components of the ven-

tral claspette, the (1) basal lobules, (2) preapical plate (Gabaldon 1940), (3) refrin-

gent structure (Gabaldon 1940), (4) mesal cleft, (5) membranous area and (6) me-

dian sulcus, are illustrated in figure 10. The trend has been for the acquisition of se-

tae on the ventral claspette; this appears to have occurred in a more or less stepwise

fashion. The ancestral condition is represented in albimanus, triannulatus and all

members of the Argyritarsis Section, by the complete absence of setae. In the Stro-

dei Complex setae are present; however, they do not extend to the apex. The most

derived condition is represented by the species comprising the Oswaldoi Complex

(except ininii) in which setae are continuous on the lateral and ventral surfaces, ex-

tending to or nearly to the apex. The preapical plate has also tended to increase in

size from an inconspicuous spot in the Albimanus Group to a very large, heavily scler-

otized plate in some species in the Oswaldoi Complex.

Phallosome. Primitively the phallosome in the Albimanus Section probably had a

pair of large, serrated leaflets as seen in several species in the Argyritarsis Section.

These leaflets have been lost except in anomalophyllus, a relict species in the Oswal-

doi Complex. Remnants of the leaflets are still present in many of the species in the

latter complex as unserrated, membranous, basolateral expansions at the apex of the

aedeagus. Whenever width of aedeagus is used, this refers to width of aedeagus at

base of apex (subapex where lateral sclerotizations extend ventromedially to form an

incomplete tube, fig. 10). Length of apex of aedeagus refers to distance from subapi-

cal, collarlike, subtriangular, lateral sclerotizations to apex of aedeagus (fig. 10).

12 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

PUPAE

As in several other groups of mosquitoes, the pupae in the Albimanus Section are

remarkably similar. They exhibit few features that vary consistently throughout the

group, and are of phylogenetic importance only when correlated with the significant

group characters in the other life stages. For this reason I have not included them in

the descriptions of the groups, subgroups and complexes except for the monotypic

taxa. The only group trends that are apparent occur in the trumpet and “caudolater-

al spines”’ (hair 9-IJ-VIII). In the ancestral condition of the trumpet, the pinna is

long, usually greater than 3.0 length of the meatus. The pinna is proportionately

long in triannulatus, benarrochi, strodei, rangeli, trinkae and ininii. The pinna is high-

ly derived and shortest in 2 species in the Oswaldoi Complex, oswaldoi and nunezto-

vari. Likewise, hairs 9-IJ-VIII are shorter in the more derived state. Hairs 9-VII, VIII

are 0.5 or less the length of the corresponding abdominal segment in oswaldoi, ano-

malophyllus, benarrochi, noroestensis and nuneztovari; in the other species, these

hairs are at least 0.5 the length of the segment. There is also a correlation between

the short pinna and the short caudolateral spines in oswaldoi and nuneztovari. The

shape of the pinna is described in lateral aspect (as in figs. 8, 25) from dissected pu-

pal exuviae mounted on microscope slides (Belkin 1962:77). In living specimens

when the trumpet is open at the water surface, the margins of the meatal cleft are

widely separated. This separation is much wider apically in some species than in

others; the wider this opening is, the narrower the sides of the pinna appear to be, so

that in lateral view on a microscope slide the pinna may appear to be tapered toward

the apex:

CHAETOTAXY. The chaetotaxy is of little phylogenetic value, but is important

for species differentiation. On the cephalothorax, the relative lengths of the individ-

ual branches of hair 7-C are significant, since in many species one branch is consider-

ably longer than the other(s). The relative lengths of hairs 10-12-C are also of speci-

fic value. In most species 10-C is subequal to 11-C, whereas in triannulatus 10-C is

much shorter than 11-C. On the abdomen, the length of hair 2-I, the number and the

point of origin of branches, correlated with the relative length of hair 3-I, are impor-

tant distinguishing characters in some species. The relative lengths of hairs 6,7,9-I

and 6,7-II are good secondary key characters. Also the derived feature, insertion of

hair 3-II very near the caudal margin of the segment adjacent to hair 1-II, is diagnos-

tic for benarrochi.

PADDLE. The shape of the paddle and/or the relative length of the buttress do

not show any phyletic relationships. The length of hairs 1,2-P and the distance the

marginal fringe of spicules extends along the inner margin of the paddle are second-

ary diagnostic characters in some species.

TERMINAL SEGMENTS. The male genital lobe and its apical mammilliform pro-

tuberance are more strongly developed in some species, but show no phylogenetic

relationships.

FOURTH INSTAR LARVAE

Many characters in the larvae of the Albimanus Section correlate with characters

in the adult and the male genitalia to clearly define the separate taxa. The chaeto-

taxy is very important for species identification; therefore I have included in the spe-

cies descriptions the mode (at least 75% of the observations) and range for all the se-

tae of taxonomic significance. It is important to consider the amount of intraspecific

variation in larval “hairiness”? when examining any of the species in the section.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 13

HEAD. In the ancestral condition, the clypeal hairs (2,3-C) are widely spaced,

with short to minute barbs, occasionally appearing as simple spiniform hairs (hair

3-C in this condition is shorter than 2-C). In the derived state the inner clypeals (2-

C) may be (1) closely approximated, (2) of subequal length and/or (3) with long,

simple or dendritic branches. The relationship between the approximation of the in-

ner clypeal hairs (2-C) and the outer clypeal hairs (3-C) is given by the clypeal index,

which is the distance between the inner and outer clypeals on one side divided by the

distance separating the inner clypeals. An. anomalophyllus in the Oswaldoi Complex,

and strodei and reportedly rondoni in the Strodei Complex, are characterized by ap-

proximated inner clypeals. This character has supposedly evolved independently in

the 2 complexes. In the Oswaldoi Subgroup, for some species, the trend has been

toward more extensive branching of the clypeal hairs along with longer branches, so

that either or both the inner and outer clypeals may be plumose. The most derived

condition is found in oswaldoi, where the clypeal branching is often dendritic. Hair

4-C is long in the Albimanus Group and short throughout the Oswaldoi Group except

in nuneztovari and trinkae, where it is strongly developed. Hairs 8,9-C do not show

any phylogenetic relationships; however, they are important to characterize species,

being strongly developed in some species and more weakly developed in others. The

collar tends to be wider, dorsomedially, in the more derived species than in the more

ancestral, and is of group importance, although this character is highly variable.

ANTENNA. Hair 1-A in the ancestral condition is small; it is large and indepen-

dently derived in benarrochi and ininii.

THORAX. Primitively, hair 1-P was plumose and multibranched; this condition is

found only in the monotypic Albimanus Group. The trend has been toward the de-

velopment of palmate hairs with fewer, broader, lanceolate leaflets. Hair 1-P in trian-

nulatus may represent the intermediate condition with numerous, weakly lanceolate

branches. The branching of hair 2-P and the presence or absence of a common scler-

otized tubercle for hairs 1,2-P or 1-3-P show some group relationships. However,

these characters are unreliable because of considerable intraspecific variation. The

length of the dorsoventrally flattened shaft, and the number and length of the

branches of hair 14-P are characteristic, to some extent, of triannulatus and some

species in the Oswaldoi Group.

ABDOMEN. Derived group characters include the following: (1) small, usually

numerously branched hair 13-I-IV in Oswaldoi Subgroup, with triannulatus some-

what intermediate between Albimanus Group and Oswaldoi Subgroup and (2) large,

5-7 branched hair 11-I in triannulatus. Important characters for species identification

within the Oswaldoi Group include (1) location and size of hair 5-I,II, (2) size and

branching of hairs 0-II and 13-IV and (3) width of palmate hair 1-II-VII.

SPIRACULAR LOBE. In the ancestral condition the lateral arms of the spiracular

apparatus were moderately short. There has been an independent increase in the size

of the arms in the Triannulatus Subgroup and to a lesser degree in ininii of the Oswal-

doi Subgroup. The dentition of the pecten, although not of group importance, is of

value as a secondary key character in species diagnosis within the Oswaldoi Group.

ANAL SEGMENT. In the Section there appears to be a trend in the migration of

the point of insertion of hair 1-X from a dorsal position, within the saddle, to a posi-

tion on or near the ventral margin of the saddle. In the most derived case (oswaldoi),

hair 1-X is not inserted on the saddle at all, but ventrad of it. Hair 1-X is moderately

long to long in all species except benarrochi. The anal gills are short in aquasalis and

sometimes in albimanus; in all other species they are moderately long to very long.

14 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

SYSTEMATICS

The nominal species sanctielii Senevet and Abonnenc 1938 is not included in this

revision because of the lack of material.

I no longer consider the nominal species gorgasi as a synonym of albimanus but as

anomen dubium. After examination of the damaged adult female holotype of gor-

gasi, collected by A. H. Jennings at La Boca, Panama Canal Zone, I have concluded

that this specimen is probably not a mutant of albimanus. Contrary to the condition

in albimanus, this specimen possesses dark caudolateral scale tufts on abdominal seg-

ments II-VII, and the penultimate segment of the palpus appears to have been white.

It lacks upper mesepimeral scales and is too large to be considered triannulatus. Hind-

tarsal segment 2 of this female is brown in about 0.5, and segment 3 has a brown spot

slightly less than half the length of the segment. This anomalous specimen most like-

ly belongs to either strodei, oswaldoi or aquasalis. Previous workers (Rozeboom

1941; Curry 1932) have rejected aquasalis since it is very rarely ever collected on the

Pacific side of Panama, even though it is probably the most likely candidate based on

morphological characters. In any case, gorgasi is not easily recognizable as any spe-

cies, since mutations of the sort found in gorgasi are now known to occur in many

different species in this section.

: I am dividing the Albimanus Section, composed of those species in the subgenus

Nyssorhynchus characterized by the presence of a basal dark band on hindtarsal seg-

ment 5, into 2 clearly defined groups: a monotypic Albimanus Group and the Oswal-

doi Group consisting of 13 species. The 2 groups are easily separated on the basis of

9 correlated characters in the adults, male genitalia and larvae. An. albimanus is the

least derived species in the Albimanus Section and retains many ancestral characters

that it shares with several species in the Argyritarsis Section. The Albimanus Group

is characterized, primarily, in the female by (1) dark palpal segment 4, (2) absence of

dark caudolateral scale tufts on the second abdominal segment and (3) predominant-

ly dark foretarsal segment 5; in the male genitalia by (1) short sternite IX, (2) ventral

claspette completely without setae and with the ventral surface produced into 2 large,

inflated, ovoid, bulbous lobes and (3) very small circular to oval, weakly sclerotized

preapical plate; and in the larva by (1) hair 1-P plumose with filiform branches, (2)

hair 9-P,T pectinate and (3) hair 13-I, III, [1V moderately large, with few branches.

Most of these characters that distinguish the Albimanus Group from the Oswaldoi

Group are ancestral, and are found nowhere else in the section. Exceptions are the 2

derived features, the short ninth sternite in the male genitalia and the pectinate hair

9-P,T in the larva.

I believe that the evolutionary lineage which has resulted in albimanus separated

from the main phyletic line of the Albimanus Section very early in the evolution of

the section. An. albimanus retains many characters shared by relict species in the Ar-

gyritarsis Section, which indicates that it diverged from the Nyssorhynchus phyloge-

netic tree soon after the separation of the 2 sections, Albimanus and Argyritarsis. It

is even possible that the Albimanus Section is of polyphyletic origin, although not

very probable for the reasons enumerated below. The principal character, the dark

basal band on hindtarsal segment 5, separating the Albimanus Section from the Argy-

ritarsis Section, is extremely stable throughtout the Albimanus Section. K. L. Linthi-

cum (personal communication; who is presently revising the Argyritarsis Section)

states that he has never observed any specimen in the Argyritarsis Section with a dark

band present on hindtarsal segment 5. Conversely, I have never observed a speci-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 15

men in the Albimanus Section with the dark fifth hindtarsal band completely absent,

although it is sometimes considerably reduced. Parallel evolution and the very close

phylogenetic relationship of the Albimanus and Argyritarsis Sections may help to ex-

plain the absence of other consistent characters distinguishing the 2 taxa. For exam-

ple, in both sections there has been a tendency for some members to independently

lose the apical leaflets on the phallosome of the male genitalia and for the larval hair

1-P,I to become palmate. The divergence within the subgenus Nyssorhynchus is

probably of very recent origin.

The Oswaldoi Group is characterized in the female by (1) palpal segment 4 light,

(2) presence of dark caudolateral scale tufts on abdominal segment II and (3) foretar-

sal segment 5 usually about 0.5 to completely light (except in triannulatus and some

strodei); in the male genitalia by (1) moderately long to long, subtriangular to subtra-

pezoidal sternite LX, (2) presence of setae on the ventral claspette or, if bare, apex

expanded laterally into large auriculate lobe and (3) absence of bare, ventral, inflated,

ovoid lobes on the ventral claspette; and in the larva by (1) hair 1-P palmate, (2) hair

9-P,T long and single and (3) hair 13-I,III,[V small to moderate or, if large, hair 1 1-I

large and 5-7 branched.

I am dividing the Oswaldoi Group into 2 subgroups: the monotypic Triannulatus

Subgroup and the Oswaldoi Subgroup. I have separated triannulatus from strodei

and rondoni, as did Galvao (1943), since it differs from all species in the Oswaldoi

Subgroup by several correlated characters in all stages except the pupa. The adults of

triannulatus are unique in possessing a patch of white scales on the anterior mesepi-

meron. Several reliable characters in the larva, particularly the length of the lateral

arms of the spiracular apparatus and the development of hairs 1,14-P, 11-I, 13-LI1U,

IV, clearly separate triannulatus from the Oswaldoi Subgroup. The ventral claspette

of the male genitalia of triannulatus lacks setae, as in albimanus, and the apex is ex-

panded laterally into a pair of large auriculate lobes. The reason for including trian-

nulatus in the Oswaldoi Group is on the basis of (1) presence of caudolateral tufts on

the second segment of the abdomen in the adult, (2) more or less lightly colored pal-

pal segment 4 in the female, (3) hair 1-P palmate in the larva, (4) the intermediate

character of hair 13-IV in the larva and (5) overall general similarity of the male geni-

talia, particularly with the Strodei Complex.

The Oswaldoi Subgroup is characterized in the female by (1) predominantly white

palpal segment 4, (2) absence of anterior mesepimeral scales, (3) absence of a large,

apical, white band on foretarsal segment 4, except for some nuneztovari, trinkae and

ininii and (4) foretarsal segment 5 about 0.5 to completely light, except in some Stro-

dei; in the male genitalia by the presence of setae at least on the basal lobules of the

ventral claspette; and in the larva by (1) palmate hair 1-P usually with fewer than 16

leaflets, (2) 14-P with a short to moderately short shaft, (3) 11-I moderately large, 2-

4 branched, (4) 13-I small to moderately large, usually more than 3 branched, (5) 13-

III small, with numerous branches and (6) lateral arms of the spiracular apparatus

short to moderately long, except in ininii.

The Oswaldoi Subgroup is an assemblage of closely related species, which I believe

is composed of 2 major phyletic lines: the Strodei Complex and the Oswaldoi Com-

plex. As indicated above, on the basis of the male genitalia, the Strodei Complex is

more closely allied to the Triannulatus Subgroup than is the Oswaldoi Complex. The

apex of the ventral claspette is expanded laterally, and the sidepiece possesses a long

parabasal spine and tubercle as in triannulatus. Setae are present on the ventral clasp-

ette but do not extend to the apex. Within the Strodei Complex, 2 of the species,

rondoni and strodei, are very closely related and represent one clearly defined line;

16 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

benarrochi, a highly derived species, represents the other. An. rondoni appears to be

avery recent species that may have evolved from an isolated gravid female or an ano-

malous population of strodei, later becoming reproductively isolated. The relation-

ship of benarrochi to strodei is more difficult to understand, since benarrochi is the

most highly derived species in the Oswaldoi Subgroup, possessing several unique char-

acters; only on the basis of the male genitalia is it possible to see its relationship with

rondoni and strodei.

The Oswaldoi Complex is characterized in the male genitalia by (1) setae on the

lateral margins of the ventral claspette extending to or nearly to the apex (except in

ininii), (2) apex of the ventral claspette not expanded into a pair of lateral lobes and

(3) usually moderately short parabasal spine and tubercle. The Oswaldoi Complex is

composed, I believe, of 2 separate monophyletic lines in addition to one relict spe-

cies, anomalophyllus. An. nuneztovari, trinkae and rangeli compose one phyletic

line, and oswaldoi, galvaoi, aquasalis, noroestensis and ininii form the other clearly

defined line, as based on the male genitalia, larvae and adults. Within each phyletic

line of the Oswaldoi Complex the male genitalia are clearly distinct. Also, in the os-

waldoi phyletic line, there is a tendency in the larvae for the acquisition of long

branches on the clypeal hairs as observed in oswaldoi and aquasalis. The relationship

of anomalophyllus to the rest of the species in the complex is unclear. An. anomalo-

phyllus retains a pair of strongly developed serrated leaflets at the apex of the aedea-

gus. Also, it shares with the Strodei Complex several larval characters such as the

closely approximated inner clypeals. It may possibly be that anomalophyllus repre-

sents an annectent species between the Oswaldoi Complex and Strodei Complex.

Within the complex several other species, nuneztovari, rangeli, ininii and (rarely) os-

waldoi, often have remnants of aedeagal leaflets evident as very small, membranous,

pointed projections. |

BIONOMICS

The immature stages of the Albimanus Section are found predominantly in ground

water. They occur in a variety of habitats, such as ponds, lakes, stream and river mar-

gins, canals, seepage and drainage areas, ditches, flooded meadows or pastures, reser-

voirs, swamps, ground pools, borrow pits, and animal and vehicle tracks. An. albima-

nus, although usually collected from ground pools of various sorts and stream mar-

gins, has been found also in crab holes, tree holes, large artificial containers and

brackish swamps. An. aquasalis is the only species primarily restricted to the coast

and preferentially occurs in brackish water such as in mangrove swamps and coastal

ground pools. However, aquasalis is capable of breeding in fresh water, and often is

collected several kilometers from the coast. An. anomalophyllus may be restricted to

stream margins in Costa Rica and northern Panama. An. triannulatus and ininii are

found commonly in lakes, ponds or large ground pools. An. triannulatus is the only

species clearly shown to be often closely associated with a specific plant; it is usually

collected in or between the rosette crowns of Pistia sp. Some species are fairly habi-

tat specific. An. oswaldoi, ininii and triannulatus adults are usually collected in the

interior of forests, although the larvae may be collected from ground pools in inter-

spersed secondary growth areas. An. noroestensis is usually collected in drier regions

than the forest species, in cultivated areas or areas of secondary vegetation.

Regarding altitudinal distribution, strodei breeds at the highest elevations for any

species in the Albimanus Section (1600m), although it also occurs at lower elevations.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 17

An. rangeli, and to a lesser extent benarrochi, are principally found at intermediate

elevations (200-1000m) such as in the upper Amazon and the lIlano plateau region of

Colombia extending south into Mato Grosso and Bolivia. An. albimanus and aquasa-

lis, on the other hand, normally breed on or near the coast in wet areas, usually at el-

evations less than 400m. An. nuneztovari is found at low elevations usually in or at

the margins of forests. An. trinkae breeds in habitats similar to those of rangeli, of-

ten in areas of secondary growth at intermediate elevations. An. rondoni breeds in

ditches, puddles, flooded meadows, etc., in southern Brazil and northern Argentina,

and is never found farther north than the southern margin of the upper Amazon. An.

ininii has been collected only in French Guiana and in Para, Brazil in the interior of

the forest. :

Host preference studies for several species indicate that these mosquitoes feed pre-

dominantly on large mammals such as dogs, cats, cattle, pigs, goats, donkeys and

man, although some do feed on fowl. In all known cases, the species feed readily on

man when given the opportunity; however, anomalophyllus, ininii, trinkae and gal-

vaoi have not been studied in this respect. The adults are active either crepuscularly

or nocturnally, except triannulatus which is somewhat diurnal (Rozeboom 1935:527).

Most of the species are predominantly exophilic and zoophilic and become a serious

health or pest problem only when occurring in high densities without abundant alter-

nate hosts. However, during the peak seasons certain species, such as albimanus, aqua-

salis, nuneztovari and possibly rondoni and strodei, are commonly found inside

houses. Marked capture and release studies indicate that aquasalis and albimanus are

both strong fliers capable of migrating considerable distances. Flight ranges for the

remaining species have not been studied in detail.

MEDICAL IMPORTANCE

An. albimanus is the major vector of malaria in coastal Central America and north-

ern South America, extending to the Paria Peninsula in Venezuela and into the Great-

er Antilles. It does not seem to be predominantly anthropophilic or endophilic, but,

because it occurs in such tremendous numbers during certain times of the year, it is

the most common anopheline found inside houses feeding on man. An. albimanus

has been found naturally infected with Plasmodium sp in nearly every country in

which it is encountered.

An. aquasalis is a primary vector of malaria in the Lesser Antilles, and in Trinidad

and Tobago. Along the coast of Brazil, the Guianas and possibly Venezuela, it is al-

ways a potential vector but usually only important when it occurs in large numbers.

An. aquasalis feeds readily on man and is commonly collected in houses. In the past

it has been an important vector of malaria in coastal Brazil.

An. nuneztovari is a primary vector of malaria in western Venezuela and northern

Colombia, and is a probable vector in Suriname; in some areas where it occurs, spleen

indices have been close to 100% (Gabaldon and Guerrero 1959). The vector potential

of nuneztovari has been reported to depend on the density of the nearby vegetation

surrounding regions of habitations, which may be correlated with the greater life ex-

pectancy and vector density in the forest (Hamon, Mouchet et al. 1970).

An. triannulatus has been implicated as a vector of malaria in Venezuela, and once

was found to have a natural oocyst infection (Gabaldon and Cova Garcia 1946b).

Several authors have succeeded in experimentally infecting triannulatus with Plasmo-

dium vivax and P. falciparum; however, it is much more refractory to infection than

is albimanus.

18 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

According to Correa (1938), strodei transmitted malaria at the Fazenda Santa

Alice, Sao Paulo, Brazil. He reported a natural infection rate of 1.2%. Other Brazil-

ian workers have experimentally infected strodei with Plasmodium vivax, although

there have not been any other reports implicating strodei.as a vector.

Very little is known regarding the vector potential of rangeli, noroestensis, rondoni

and oswaldoi. An. noroestensis was found naturally infected once in Ribeira, Sao

Paulo, Brazil by Correa and Ramos (1942b:385-386). Lucena (1940b) reported find-

ing oswaldoi var. metcalfi naturally infected in Pontesinha, Brazil; however, Lucena

may have been studing aquasalis rather than noroestensis. An. oswaldoi has been ex-

perimentally infected with P. vivax and P. falciparum (Fonseca and Fonseca 1942;

Rozeboom 1942a). An. rangeli has been suspected of transmitting malaria in Ecua-

dor (Forattini 1962), but it has never been shown to be naturally infected. An. ron-

doni was investigated in Jujuy, Argentina during the malaria season by Davis and

Shannon (1928), and it was not found to be naturally infected nor was it possible to

experimentally infect it with P. falciparum, P. vivax or P. malariae in 3 different ex-

periments. Nevertheless, Shannon and Del Ponte (1927) stated that Davis was able to

infect rondoni in other experiments. Nothing is known about the vector potential of

ininii, galvaoi, trinkae or anomalophyllus.

Two viruses have been isolated from 2 species in the Albimanus Section. An. aqua-

salis was found naturally infected with Venezuelan Encephalitis virus (VE) during the

1962-1964 outbreaks in Venezuela (Sellers, Bergold et al. 1965). Tlacotalpan virus,

in the Bunyamwera group, was isolated from albimanus in Tlacotalpan, Mexico

(Scherer, Campillo-Sainz et al. 1967:79-91). ;

TAXONOMIC TREATMENT

ALBIMANUS SECTION

FEMALES (fig. 5). Small to large, wing 2.5-4.0 mm. Head, thorax, abdomen and

legs predominantly dark scaled with distinct markings of light scales and pruinose in-

tegumentary patterns. Head: Integument light brown to dark brown, usually some-

what pruinose. Interocular space moderately wide, 2-7 ommatidial diameters. Ver-

tex and occiput with many erect, moderately long to long, cuneate to lanceolate or

~ weakly forked scales, white anteriorly on vertex, becoming progressively darker cau-

dad and laterad on occiput and postgena. Vertex with a few decumbent light scales.

Interorbital and upper orbital lines with white decumbent scales. Strong, long,

curved, dark orbital setae present. Frontal tuft conspicuous, composed of 10-20

long, slender, white setiform scales. Clypeus prominent, bare, pruinose. Proboscis

much longer than forefemur, entirely dark scaled; labella slightly lighter than labium;

with 3-6 basal bristles. Palpus subequal in length to proboscis; predominantly dark

scaled; appearing weakly metallic when viewed at an angle; with an apical light band

on segments 2,3; band on 3 usually broader than that on 2; segment 4 either light or

dark scaled, when light scaled, a basal and an apical dark, narrow band present, and

occasionally a darker stripe on ventral surface; segment 5 white, occasionally with a

small basal dark band; segments 2,3 with varying amounts of dorsal and dorsolateral

speckling of light scales; segment | less than 0.1 of palpal length; segment 2 about

0.7-0.9 of segment 3; combined length of segments 4 and 5 slightly greater than

length of segment 2 and less than that of segment 3; segments 1,2 and basal 0.3-0.5

of 3 with erect scales, progressively less outstanding from base of segment | to seg-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 19

ment 3; remainder of palpus with decumbent scales. Antenna: 0.60-0.75 length of

proboscis. Torus with dorsolateral row of light scales and a few short lateral and ven-

tral setae. Lateral margin of proximal flagellar segments (1-5) with small, light scales;

flagellar segment 1 subequal to combined length of segment 2 and 3, with long bris-

tles on all surfaces except midventral, bristles most abundant on dorsolateral surface,

and with long, oblanceolate to setiform, white scales on dorsomesal surface and

shorter scales basally; segments 2-13 each with basal whorl of 6-10 long, curved bris-

tles and with short, silver setae distally; segment 13 pointed at apex and slightly long-

er than 2-12. Thorax: Scutum small to large, length on midline 0.9-1.4 mm; integu-

ment generally pruinose; with 2 pairs of bare, light pruinose, longitudinal stripes,

when viewed at an angle from the front as follows: (1) submedian stripe just laterad

of acrostichal bristles, extending from scales of anterior promontory to about middle

of scutum, becoming much less distinct in posterior half of scutum, (2) broad, sub-

dorsal stripe laterad of posterior dorsocentral and lateral prescutellar bristles extend-

ing caudad from a pair of prominent, bare, nonpruinose rectangular or subtriangular

areas at caudal margin of posterior fossa, ending at small nonpruinose area anterior to

scutellum; acrostichal and posterior dorsocentral lines nonpruinose or distinctly less

pruinose than remainder of scutum; fossal region slightly less pruinose than general

surface of scutum. Scutum with small, lanceolate to obovate, silver or yellow to

white, decumbent scales in (1) acrostichal line, extending from anterior promontory

to prescutellar bare space, and anterior and posterior dorsocentral lines, the scales be-

coming less defined into distinct regions posteriorly, (2) fossal area and (3) supraalar

and antealar areas; with elongate, narrow erect, lanceolate scales on median anterior

promontory and along lateral margin of antealar area extending posteriorly onto su-

praalar area; humeral tuft with numerous moderately short to long, light scales above

and numerous, short, obovate or cuneate, dark scales below. Scutum with numerous

moderately long to long, dark setae on acrostichal line, dorsocentral line, posterior

median scutal area and supraalar area; with scattered short, light to dark setae on fos-

sa, antealar area and lateral prescutum. Parascutellum with a single long seta. Pre-

scutellar bare space appearing dark from above, weakly pruinose, triangular or horse-

shoe shaped. Scutellum rounded, with obovate light scales, very long, dark setae and

shorter, light setae along posterior border; posteromedian border usually with a few

long, light setiform scales. Postnotum bare. Pleural integument dark, with gray or

silver to reddish pruinose pattern. Apn reniform, with moderately long setae on dor-

dal surface along ventral margin, and with a dorsal patch of erect, obovate, dark

scales, usually with a few light scales; pp! with 2,3 (1-4) setae; sp with 1-9 short to

moderately long setae and a few elongate light scales; pra with 4-10 light setae and a

patch of elongate light scales; upper stp with 2-8 setae and a patch of 4-18 light scales;

lower stp with 1-3 setae and 2-10 light scales oriented either in a horizontal arc or di-

agonal patch; upper mep with 3-10 setae in horizontal or diagonal row and occasion-

ally 1-4 light scales; anterior mep bare or with a cream to white scale patch in trian-

nulatus, remainder of pleuron without scales or setae. Legs: Forecoxa larger than

mid- or hindcoxa. Integument of coxae and trochanters dark. Forecoxa with a row

of long, strong, dark setae anteriorly and a cluster of 1-4 long, dark setae caudoven-

trally; a patch of light scales dorsolaterally; caudoventral surface with a row of light

and dark scales, light scales usually somewhat cephalad of dark scales. Midcoxa later-

ally with 1-3 long, dark setae 0.3 from base; anteriorly with a small patch of short

spiniform setae and a horizontal line of yellow to white scales; antero- and postero-

apical borders with 1,2 long, dark setae and usually a light scale patch; anteroapically

also with a row of short, dark setae. Hindcoxa with 1,2 long, dark setae and dorso-

20... Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

caudally with a patch of light scales; posteroapical margin with moderately short se-

tae, occasionally 1,2 long, dark setae and a row of light scales; anteroapical border

with a long, curved, dark seta directed ventrad. Trochanter with short setae and yel-

_low to white scales. Forefemur shorter than midfemur; latter subequal to hindfemur.

Forefemur largely dark scaled, with a band of light scales at base; anterior surface

mottled and occasionally with an indistinct, longitudinal streak of light scales; basal,

dorsal and anterior surfaces speckled with light scales; distal 0.3-0.5 of dorsal, poster-

ior and ventral surfaces with cream to yellow scales. Midfemur with basal light band

followed by a narrow dark band; posterior surface light; anterior surface predomi-

nantly dark scaled, with a distal, variously shaped, light (anteroapical) spot just anter-

ior and basad of distinct dorsal knee spot and with a longitudinal streak extending

from near base to near anteroapical spot; apex with row of moderately strong setae.

Hindfemur predominantly dark, with a basal light band, speckled with scales, and

with a light spot of variable length in distal 0.25; knee spot present or absent; ventral

surface light; apex with moderately long setae and whitish scales. Foretibia subequal

to midtibia; hindtibia longest. Tibiae predominantly light on ventral and posterior

surfaces; fore-, mid- and occasionally hindtibia with a yellowish longitudinal streak

on anterior and dorsoanterior surfaces broadening at apex, occasionally appearing as

speckling on foretibia; apices of tibiae yellow to white, usually light scaling weakly

developed on foretibia, progressively more extensive on mid- and hindtibia; dorsal

surface of apex with strongly developed apical spines. Hindtarsal segment | longer

than hindtibia. Dorsal surface of fore- and midtarsal segment | predominantly dark,

speckled with light scales, and with a light apical band usually less than 0.1 length of

segment. Foretarsal segments 1 and 2 with double row of spiniform setae on plantar

surface. Ventral surfaces of fore- and midtarsal segments 1,2 light; segments 3-5 of-

ten speckled with dark scales, varying from completely light to dark. Foretarsal seg-

ment | with an apical band, when present, white to golden; segment 2 with a narrow

to wide, light scaled band in apical 0.15-0.95; segment 3 with a broad, light scaled

band in apical 0.2-0.9; segment 4 from dark to predominantly light; segment 5 from

dark to light. Midtarsal markings generally as in foretarsus except that ventral sur-

face usually darker and apical light bands, when present, white to golden and usually

darker than on foretarsus; dorsal surfaces of segments 2,3 from completely dark to

light in apical 0.4; segment 4 usually dark, occasionally with apical light band (ininii);

segment 5 from completely dark (albimanus and triannulatus) to completely light.

Hindtarsal segment | predominantly dark with speckling of light scales on ventral sur-

face and a longitudinal light streak on anterior surface, apex with or without yellow

to white band; segment 2 with highly variable, white band in apical 0.10-0.95; seg-

ments 3,4 completely white or with dark basal band present on either one (mutants)

or both segments (rondoni, mutants); segment 5 with white to pale yellow band in

about apical 0.5. Pretarsi simple, claw of foreleg largest; claw of midleg larger than

that of hindleg. Wing (fig. 5): As figured but extremely variable. Vein C with basal,

humeral, subbasal, presectoral, sectoral, subcostal and preapical light scaled spots us-

ually present; subbasal, presectoral and subcostal light spots sometimes absent. Inte-

gument of wing usually darkened in region of scales on C and R. Rs-R,.3 variable,

more or less predominantly dark, with 3 or 4 dark spots and 2 or 3 light spots; with

or without an extra subcostal light spot. R, with 2 light spots of varying lengths. R3

usually with 3, occasionally only 2, light spots. R4,; with 2 small to moderate size

dark spots, one subcostal, the other preapical. M with or without sectoral dark spot

of variable length and subcostal dark spot that may reach furcation. M,.. with 2 dark

spots and M3,4 with one dark spot. Base of Cu light with a small to medium sectoral

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 21

dark spot usually not reaching furcation. Cu, largely light, with 3 small, dark spots,

2 toward base in sectoral-subcostal region and one preapical. Cu, all light except for

small to moderate preapical dark spot. Vein A largely light, with 2 dark spots, 1 sub-

basal and 1 subcostal. Apical light fringe spot conspicuous, small to large; fringe of

remainder of wing largely dark with light areas where veins intersect wing margin ex-

cept for R,; usually additional moderately long, light fringe spot present between

base of wing and A. Haltere: Stem pale; apex of knob with small patch of dark

brown scales, surrounded by subapical light scales, particularly abundant laterally.

Abdomen: Integument of tergites predominantly light to dark brown, with light

mottling occasionally present on II-VII, when present usually more extensive on V-

VII. Long setae present along lateral and apical margins of tergites I-VII; and short,

scattered setae medially. Tergites II-VII with patches of light scales; patches triangu-

lar, with bases toward apex of segment on proximal segments, and subrectangular

and longer on more distal segments. Caudolateral margins of tergites and sternites of

segments II-VII (III-VII in albimanus) with conspicuous, outstanding, dark scale

tufts. Sternite I with or without setae. Moderately long setae present on sternites II-

VII in submedian row and along caudal border; sternite I usually with a few incon-

spicuous light scales; sternites II-VII with a submedian, longitudinal patch of light

scales, followed posteriorly by tuft of dark scales which extend laterally along caudal

border of sternite to meet caudolateral tufts of tergite.

FEMALE GENITALIA. Segment VIII: About as long as wide, slightly narrower

than segment II-VII; with scattered moderately long, dark setae; tergite densely cov-

ered with light scales; sternite usually with median longitudinal tuft of dark scales

along caudal border; remainder of sternite with light scales. Tergite IX: Ribbonlike

dorsally, expanding laterally into curved plate projecting slightly ventromesad. Ter-

gite X: With curved subtriangular lateral plates directly caudad of tergite IX, narrow-

ing abruptly medially, and not continuous dorsally. Tergites IX and X densely cov-

ered with short, fine spicules. Cercus: Large, subcylindrical, curved dorsally; dense-

ly covered with lanceolate scales and moderately long, dark setae; scales darker at

base, cream at apex. Postgenital Plate: Moderately long, subtriangular, rounded api-

cally, covered with short fine spicules, with 2 long, strong, subapical setae. Upper

Vaginal Lip: Strongly sclerotized. Insula: Discoid, without distinct margin, usually

with 20 or more short spiniform setae originating from large conspicuous alveoli.

Atrial Plates: Strongly sclerotized, triangular, laterally articulating with tergite IX.

Lower Vaginal Lip: Membranous, spiculose. Spermatheca: One, spherical, strongly

sclerotized, dotted with numerous small, circular, membranous regions.

MALES (fig. 5). Essentially similar to females except for sexual characters. Color-

ation and scale patterns as in females. Head: Clypeus smaller than in females. Pro-

boscis longer than in females, usually apically retrorse, light reddish brown to dark

brown. Palpus 5 segmented, subequal to or slightly longer than proboscis, segment 1

short, 0.06-0.08 of palpal length; segment 2 long, shorter than segment 3, 0.25-0.30

of palpal length; segment 3 long, 0.32-0.39 of palpal length; segments 4 and 5 moder-

ately short, each 0.10-0.16 of palpal length; segments 1, 2 and 3 ankylosed; 3 ex-

panded dorsoventrally at apex and with segments 4 and 5 forming a laterally com-

pressed, dorsoventrally expanded, conspicuous club; palpus bent near apex of seg-

ment 3 so that segments 4 and 5 are directed anterolaterally; segments 1 and 2 with

dark, erect scales, becoming decumbent toward apex of 2; apex of segments 2 and 3,

and base of segment 3, each with a white band; segment 3 and occasionally 2 with a

few dorsal, light scales; base and occasionally ventral surface and apex of segment 4

dark, remainder of lateral surface of segment 4 cream or white, somewhat darker in

a Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

albimanus; lateral surface of segment 5 dark in about basal 0.33 of segment and on

ventral surface, remainder of segment 5 white; dorsal and ventral surfaces of apices

of palpal segments 2 and 3 with numerous, long, apically directed setae. Antenna:

Length 0.65-0.85 length of proboscis; segment 1 slightly longer than segments 2-11,

with long, light scales on inner margin; segment 12 very long, about 0.20-0.25 of an-

tennal length; segment 13 about 0.5 length of segment 12; flagellar whorls on seg-

ments 1-12 strongly developed, with numerous, long setae; setae on inner margins of

whorls 1-5 occasionally white. Legs: Basal plantar surface of foretarsal segment 5

with about 6-9 short to moderately long, spiniform setae; setae longer toward apex;

shorter spiniform setae medially between longer setae. Single claw on foreleg long,

slightly curved, acuminate, with a blunt to acuminate submedian tooth and a curved,

usually blunt, short to moderately short, external basal tooth. Mid- and hindleg with

2 claws; claws on midleg larger than on hindleg. Empodium spinulose, about 0.75

length of claw. Wing: Narrower than in female.

MALE GENITALIA. Segment VIII: Not retracted into segment VII; as narrow as

or slightly narrower than segment VII; densely covered with lanceolate to truncate,

obovate scales except on proximal border and with moderately long, dark setae scat-

tered over surface. Tergite often with a median longitudinal patch of dark scales sur-

rounded by light scales, and with long, dark setae along distal and lateral margins.

Sternite covered with light scales, with or without median subtriangular, dark scale

patch. Segment IX: Spiculose. Tergite small, membranous, without lateral lobes,

articulated caudally with proctiger and laterally with sidepiece. Sternite well devel-

oped, sclerotized; narrowly subrectangular to broadly subtriangular, curved and tap-

ered dorsolaterally, extending a short distance dorsad as a small, sclerotized, subrec-

tangular lobe; articulating with proctiger and sidepiece; with ribbonlike to triangular,

darker, sclerotized, nonspiculose, apodeme projecting along anteromedian border;

apodeme occasionally with a median protuberance. Sidepiece: Sclerotized, spicu-

lose, cylindrical to subconical, moderately narrow, curved mesally. Tergal surface

with a submedian longitudinal row of 4-6 long (about 0.67 length of sidepiece),

strong (tergomedial) bristles arising from large, conspicuous alveoli; basal 3 or 4 bris-

tles separated from more apical bristles; rows of weaker moderately long setae, one

row mesad, and usually one laterad, of long bristles. Tergolateral surface with large,

spatulate scales extending onto sternal surface, usually darker laterally. Basal tergo-

mesal margin with a single, large, very stout, blunt, retrorsely hooked, parabasal spine

inserted on a moderately short to long, protruding tubercle, directed tergomesally.

Basomesal margin of sidepiece cephalad of parabasal spine modified into a long (0.16-

0.25 length of sidepiece), heavily sclerotized, blunt apodeme articulating mesally

with basal piece of phallosome; apodeme equal to or slightly less than length of para-

basal spine. Tergomesal surface of sidepiece with 2 long, apically flattened, blunt-

hooked, accessory spines inserted on 2 adjacent, prominent tubercles that arise 0.40-

0.52 from base of sidepiece; the more dorsal spine about 0.5 length of sidepiece;

shorter more ventral spine 0.58-0.80 length of dorsal spine. Mesal surface with a

long, curved, internal spine arising 0.45-0.60 from base, subequal in length to more

ventral accessory spine. Apicomesal border usually with one long, subapical seta at

base of clasper. Sternomesal and mesal margins with moderately short, weak setae.

Clasper: Subequal in length to sidepiece, curved mesally, slender beyond base. Ven-

tromesal margin with a row of about 10-14 short spinules, from near base to apex.

Spiniform thin to thick, blunt to acuminate. Seta b weak, moderately short to mod-

_ erately long; subequal to slightly longer than spiniform, arising from small crest im-

mediately basolaterad of spiniform. Claspette: Divided into a dorsolateral claspette

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 23

and an apically fused, membranous, mesoventral claspette. Dorsal Claspette: Base

continuous ventrally with phallosome and ventral claspette, and laterally with side-

piece; basal portion modified into sclerotized pedicel with a rounded semispherical

base usually curved mesad, occasionally with a conspicuous internal apodeme; distal-

ly developed into 3 long, broad, membranous, lanceolate leaflets, longest about 0.3-

0.5 length of claspette, each curving mesally, with a medial supporting rib; ventral

leaflet subequal to middle leaflet; dorsal leaflet shortest, with a variously developed,

mesally directed, basal projection (basomesal projection). Ventral Claspette: Highly

variable, supported mesally by 2 sclerotized “rods” arising from a sclerotized bridge

which extends mesad from sidepieces; basally produced into 2 small to large basal

lobules interconnected by an inconspicuous spiculose membrane, area ventrad of

membrane between 2 lobules referred to as mesal cleft; setae present or absent on

basal lobules. A thickened rooflike refractile structure, known as refringent struc-

ture, extends across vertex of fusion of basal lobules and basad a short distance along

inner margin of each lobule, occasionally with lateral extensions or arms. Fused api-

cal portion of ventral claspette differentiated into (1) a hairless dorsal lobe which ap-

poses the aedeagus (possibly to form the external gonopore) and (2) a variously

shaped, ventral lobe with or without setae. Subapicomedial region of ventral clasp-

ette with a small to large, variously shaped, sclerotized, preapical plate. Area imme-

diately basad of preapical plate transparent and membranous, in shape of an inverted

U or V, basal margins being formed by refringent structure. Phallosome: Composed

of a central aedeagus, a pair of parameres and a pair of basal pieces, all fused into one

sclerotized structure. Sides and most of apex of aedeagus sclerotized; apex with or

without leaflets; subapically with a pair of subtriangular, lateral sclerotizations ex-

tending ventromedially to form an incomplete tube; outer border of apex membran-

ous; aedeagus weakly to moderately curved dorsally toward proctiger. Aedeagus ba-

sally contiguous with triangular, highly sclerotized parameres. Basal piece laterally

connected to paramere by a narrow, sclerotized bridge. A dorsocaudal extension of

basal piece articulates with dorsal claspette; a lateral arm of basal piece extending la-

terally, basad of sidepiece, and appearing to be attached to sidepiece by a tendinous

apodeme. Phallosome usually equal to or longer than ventral claspette. Proctiger:

Conical, apex hooked ventrally, spiculose, with sclerotized sides and triangular base,

articulating laterally with sidepiece and tergite IX.

PUPAE. Cephalothorax: Weakly to strongly pigmented; often with longitudinal

pigmented stripes on wing cases. Vertical plate about as long as wide, more sclero-

tized along anterior border, and produced apicomedially into small protuberance.

Hairs 1-3-C 2,3 branched (1-4), moderately developed: 1,2-C adjacent. Hairs 4-7-C

usually few branched, moderately to strongly developed; 4,5-C shorter than 6,7-C; 7-

C laterad or caudolaterad of 6-C. Hair 8-C single (1-3 branched), moderately long,

inserted near lateral ridge. Hair 9-C usually forked, weak, moderately long, inserted

caudolaterad of trumpet base. Trumpet: Length and shape highly variable. Moder-

ately to strongly pigmented. Inserted very Close to wing base. Pinna angusticorn;

strongly tuberculate, varying from short, flared and truncate to long, narrow and tap-

ered; meatal cleft short to long. Tracheoid not developed. Metanotum: Hair 10-C

single, shorter than or about equal to 11-C; 11-C multiple, branched or forked. Hair

12-C single or 2-4 forked, longer than 10,11-C. Abdomen: Float hair (1-]) dendritic,

large. Hairs 2,3-I approximated, inserted cephalolaterad of the base of hair 1-I ; 2-I

multiple, moderately to strongly developed; 3-1 0.5 of to about equal to 2-I. Hair 4-I

multiple, short, inserted caudolaterad of 2,3-I. Hair 5-I single or 2-4 forked, long, in-

serted laterad of 4-I. Hairs 6,9-I single or double, moderately long to long. Hair 7-I

24 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

2-7 forked, strongly developed. Hair O-II-VII multiple, medium; 0-VIII 1-3 branched

(1-4). Hair 1-II,III multiple, moderately large to large; 1-IV-VII single, very long, 1.0-

_ 2.0 length of segment. Hair 2-II,III multiple, moderately large, inserted cephalolater-

ad of hair 1 of segment; 2-IV-VII usually 1-5 branched, inserted cephalolaterad or

cephalomesad of hair 1 of segment. Hair 3-II,III 1-3 branched, inserted well cephalad

of caudal margin of segment (except in benarrochi) and between hairs 1 and 2; 3-IV

multiple, moderately developed; 3-V single or 1-5 forked; 3-VI single or forked, in-

serted on caudal margin of segment mesad of hair 1-VI; 3-VII branched or forked.

Hair +I inserted cephalomesad of 5-II; 4-III-VII always inserted cephalad or cephal-

olaterad of hair 5 of segment; 4-II-IV branched, small to moderately long; 4-V-VII 1-

7 forked, moderately long; 4-VIII forked, inserted cephalomesad of hair 9-VIII. Hair

5-II multiple, small to moderately large; 5-III-VII inserted on caudal margin of seg-

ment between hairs | and 9; 5-III,I[V 1-13 branched, moderately large to large; 5-V-

VII single, occasionally double or triple, about equal to length of segment. Hair 6-II

1-3 branched, moderately long to long, inserted immediately cephalad of hair 7-II; 6-

II-VI 1-2 branched (1-6), moderately long; 6-VII single, moderately short. Hair 7-II

2-7 branched, equal to or shorter than 6-II; 7-III-VII inserted laterad of or arising

from oblique sublateral ridge, caudad of hair 8 of segment; 7-III-V 1-7 branched,

small to moderate; 7-VI,VII usually single (1-4 branched), moderately long to long.

Hair 8-IIJ-VII 1-7 branched, short, inserted mesad of oblique sublateral ridge. Hair

9-IT-VIII (caudolateral spines) highly variable, spiniform, inserted on caudolateral

margin of segment; 9-II minute, unpigmented; 9-III thin to bulletlike, minute to

short, pigmented or unpigmented; 9-IV-VIII darkly pigmented; 9-IV short to medi-

um, heavy, straight or slightly curved; 9-V-VIII moderately short to long, thin to

thick, straight or curved. Hair 10-III-V,VII inserted mesad of hair 7 and very near

caudal border of segment; 10-III single or 2-6 forked; 10-IV,V single, long; 10-VI ab-

sent; 10-VII single or double, moderately short to medium. Hair 11-III-VII single or

occasionally double, small. Hair 14-III-VIII minute and inconspicuous, inserted sub-

medially near anterior margin of segment. Terminal Segments: Hair 1-IX 1-3

branched, minute. Median caudal lobe short, dorsobasally covering genital and cer-

cal lobes. Genital lobe of female covering about basal 0.75 of cercal lobes. Cercal

lobe about 0.5 of segment VIII. Male genital lobe about as long as segment VIII and

usually mammillated distally. Paddle: Elliptical or obovate, often distally truncate

and emarginate, always longer than wide. Midrib moderately developed, not reach-

ing apex. Buttress strongly developed, serrated distally. Outer margin distad of but-

tress with short, fine spicules; inner margin with or without very short spicules. Hair

1-P single or double, strong, moderately short; 2-P forked, subequal to 1-P.

LARVAE. Head: Ovoid, usually slightly longer than wide, widest at ocular bulge;

light brown to black; with various patterns of pigmentation, from small patches to

extensive mottling. Collar darker than rest of head, highly sclerotized, narrow to

moderately wide. Mental plate with 3,4 teeth on each side of median tooth; median

tooth varied from narrow to broad and from sharply pointed to blunt. Labial plate

subtrapezoidal; maxillary suture not contiguous with posterior tentorial pit. Mouth

brushes with numerous, simple filaments. Median labral plate bilobed and small.

Mandible and maxilla normal. Hair 1-C strong, moderately long, curved ventromedi-

ally, arising from lateral lobes of median labral plate. Inner clypeal hairs (2-C) close-

ly approximated or widely spaced, single and simple, barbed or plumose with branch-

es short to long, occasionally dendritic. Outer clypeal hair (3-C) shorter than or

about equal to 2-C. Posterior clypeal hair (4-C) 1-5 branched; 4-C simple, forked,

branched or dendritic, inserted caudomesad of outer clypeal hairs. Hairs 5-7-C plu-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 25

mose, long, with long branches; 5-C longest; 5,6-C usually extending beyond anterior

margin of head; distance between hairs 5-C equal to or greater than distance between

5-C and 6-C. Hair 8-C branched, forked or dendritic, with 2-10 branches. Hair 9-C

usually slightly longer than 8-C and occasionally with a few more branches. Hair 10-

C 3 branched (2-4), small. Hair 11-C plumose, long, inserted basolaterad of antenna.

Hair 12-C 2-6 branched, small to large. Hair 13-C 4-6 branched, inserted immediately

caudad of 11-C, moderately long. Hair 14-C 2-5 branched, short, inserted caudoven-

trad of cibarial bar. Hair 15-C varied in length, inserted immediately mesad of maxil-

lary suture. Bmh branched or dendritic, moderately short to moderately long. An-

tenna: Same color as or darker than head with a darker basal band, slightly tapering

toward apex. About 0.5 length of head capsule, with spicules on mesal or ventral sur-

faces, spicules longer on mesal margin; dorsal surface occasionally with very short

spicules. Hair 1-A 2-7 branched (2-10), small to large, inserted on dorsolateral sur-

face about 0.25-0.33 from base. Hairs 2,3-A long, saberlike, usually serrated on baso-

mesal margin; 2-A inserted dorsomesad of 3-A at apex of antenna. Hair 5-A short,

spiniform, inserted at apex of antenna. Hair 6-A short, thick, blunt, peglike. Thorax:

Uniformly pigmented and with a few small sclerotized plates dorsomedially on meso-

and metathorax. Integument without spicules. Pro-, meso- and metathoracic pleural

group hairs all arising from large common tubercles; tubercles laterally modified into

a large spine. Submedian prothoracic group (1-3-P) with hair 1-P palmate with unser-

rated, narrow to broad, lanceolate leaflets or, in albimanus, plumose with filiform

branches; 2-P plumose, moderately long, with an elongate dorsoventrally flattened

shaft; 1,2-P with or without a common tubercle; 3-P single and simple, short. Hairs

4,5-P plumose, strongly developed, each arising from a separate tubercle; 4-P shorter

than 5-P. Hair 6-P single and simple, long, arising from same large tubercle as 5-C.

Hairs 7,8-P plumose, very long; 7-P inserted immediately ventrad of 6-P; 8-P arising

from strong tubercle mesad of prothoracic pleural group (9-12-P). Hairs 9,10,12-P

single and simple (except 9-P pectinate in albimanus), long; 9-P shorter than 10-P; 11-

P forked at various distances from base, shortest of pleural group, about 0.4-0.6

length of 10-P; 12-P longest of pleural group. Hair 13-P with a highly variable num-

ber of branches, moderately large. Hair 14-P moderately large to large, 5-15 branched

from short to long, dorsoventrally flattened stalk. Hair 1-M moderately long, strong-

ly plumose, with an elongate, dorsoventrally flattened shaft, arising from fairly large

tubercle. Hair 2-M single or forked, moderately short. Hairs 3,5-M single and simple,

long. Hair 4M branched or dendritic, short. Hair 6-M 2-4 forked near base, long.

Hair 7-M 3-5 branched, moderately large. Hair 8-M moderately plumose with 25-30

branches, long, inserted in small tubercle. Hairs 9,10-M single, long; 10-M about 1.1-

1.4 length of 9-M. Hair 12-M single, moderately short, about 0.33 length of 10-M.

Hair 13-M multiple, small. Hair 14-M numerously branched or weakly pectinate,

small. Hairs 1,2-T single; 1-T short; 2-T moderately short to moderately long; 1-T us-

ually about 0.5 length of 2-T. Hair 3-T always palmate and either brushlike or with

spreading leaflets, usually unpigmented. Hair 4-T 3,4 branched, very small. Hairs 5,

7,8-T strongly plumose, very long, arising from moderately developed tubercles. Hair

6-T 2,3 forked, moderately long. Hair 9-T single (except pectinate with 3-5 branches

in albimanus), long; 10-T single, longer than 9-T. Hair 12-T 2,3 forked, moderately

long. Hair 13-T 2,3 branched, large. Abdomen: Integument without spicules except

on anal segment. Large, strongly sclerotized, median tergal plates present on anterior

margins of segments I-VII. Usually small, median and submedian, sclerotized spots

present immediately caudad of tergal plates; median spot larger than submedian spot,

occasionally connected to plates. Chaetotaxy, in general, as in genus. Hair O-II-VII

26 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

2-9 branched (1-12), moderately small to moderately large (O-II very small in trinkae).

Hair 1-I-VII palmate, with unserrated, narrow to broad, lanceolate, blunt or pointed

leaflets; 1-I 8-18 branched, unsclerotized or weakly sclerotized; 1-II-VII about 20-30

branched. Hair 5-II,III 3-12 branched, small to moderately large. Hairs 6-I-III and 7-

[,II strongly plumose, very long. Hair 6-IV-VII single, very long. Hair 7-III-V 2-4

branched, moderately long. Hair 8-II-VI 2-4 branched, moderately small. Hair 9-I

usually 3-7 branched, rarely 6-9 branched (triannulatus); 9-II-VII 6-13 branched,

moderately large. Hair 11-I 2-5 branched (5-7 in triannulatus), large.. Hair 13-I 4-10

branched (2-4 in triannulatus, benarrochi and albimanus),; 13-II,1II small to moderate

(moderately large to large in triannulatus and albimanus); 13-IV 3-7 (2-8) branched

(rarely 10-13 in benarrochi). Segment VIII: Hair O-VIII 2-5 branched (rarely single

in some benarrochi), very small. Hairs 1,4-VIII moderately long; 4-VIII always long-

er than 1-VIII. Hairs 2,3-VIII branched, large, highly variable. Spiracular Lobe:

Pecten with 11-22 teeth, highly variable in length; teeth usually as follows beginning

ventrally: (1) 1 long, (2) numerous, mixed short to moderate, (3) 1 long, (4) 4-6

short to moderate and (5) some combination of long and short, or with 2,3 long;

teeth sabrelike, dorsally curved, serrated in basal half. Lateral arms of spiracular ap-

paratus varied in length, extending or not extending to spiracular openings. Hair 2-S

5-9 branched, moderately small. Hair 6-S single or forked. Hairs 8,9-S small to mod-

erately large, 3-7 branched (3-8), rarely 6-9 branched (ininii). Anal Segment: Saddle

strongly sclerotized, extending about 0.5 around segment; light brown to brown.

About caudal 0.6 of saddle and most of anal segment finely spiculose; caudal margin

with thicker, stronger spicules. Hair 1-X single, strongly developed, as long as or

longer than saddle; inserted on or not inserted on saddle. Hair 2-X strongly plumose,

15-20 branched, long. Hair 3-X 7-10 branched, longer than 2-X. Ventral brush with

8 pairs of long, strongly plumose hairs. Anal gills varying from less than length of

saddle to greater than 2.0 length of saddle.

2( 1),

3(2).

4(3).

5(4).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) pay |

KEYS TO GROUPS AND SPECIES

FEMALES

(3. galvaoi and 7. anomalophyllus not included)

Dark caudolateral scale tufts absent from abdominal segment II; palpal seg-

ment 4 all dark or yellow to golden brown on mediolateral surface, nev-

er white or cream; foretarsal eo 5 usually all dark (fig. 5) (Albima-

nus Group) . . . . » » «- dt, albimanus

Dark caudolateral scale Tats present on abecmindt segment II; palpal seg-

ment 4 with at least some white or cream on mediolateral surface; fore-

tarsal segment 5 variable (OswaldoiGroup). ..........2

Oswaldoi Group

Anterior mesepimeron (mep) with a conspicuous patch of light scales; fore-

tarsal segment 4 with a light band in apical 0.40-0.65; foretarsal segment

5 predominantly dark; hindtarsal segment 2 dark in basal 0.4-0.7; hu-

meral light spot of vein C small, 0.5-1.3 length of basal dark spot; small

species (fig. 7) (Triannulatus Subgroup) . . . . . 14. triannulatus

Anterior mesepimeron (mep) without a patch of light scales; foretarsal seg-

ment 4 predominantly dark, or if light in more than apical 0.3 then fore-

tarsal segment 5 about 0.5 apically light and/or hindtarsal segment 2 less

than 0.4 basally dark; humeral light spot of vein C small to large; mod-

erately large to large species (Oswaldoi Subgroup) . .... . Fa

Oswaldoi Subgroup

Hindtarsal segment 3 dark in basal 0.20-0.35; vein C predominantly dark,

subbasal, presectoral and sectoral dark spots fused into one large spot;

preapical dark spot of vein M extending uninterrupted onto vein M,..

A Ve, cui, 5 . . .12. rondoni

Hindtarsal segment 3 completely white: subbasal, presectoral and sectoral

dark spots of vein C not fused into large spot; preapical dark spot of |

vein Moot extending onto vein Mie. we ee oo SO. A

Hindtarsal segment 2 with basal dark band usually less than 0.25 length of

segment; vein C humeral light spot greater than 1.5 length of basal dark

SPOb i ee, ee

Hindtarsal segment 2 with basal dark baad cual al io: or ‘peeater than

0.25 length of segment, if less than 0.25 then vein C humeral Kevan spot

less than 1.5 length of C basaldark spot. . ... . eG

Foretarsal segment 4 all light to rarely more than 0.3 basally dark; midtar-

sal segment 4 with a light band in apical 0.15-0.25; foretarsal segments

3-5 predominantly cream to white, dark scales often present only on

dorsobasal surface of segment; foretarsal segment 2 light in apical 0.35-

0.55; foretarsal segment 3 light in apical 0.70-0.86. . . . . 6. ininii

6(4).

7(6).

8(7).

9(8).

10(8).

PC):

Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Foretarsal segment 4 all dark to at least 0.3 basally dark; midtarsal seg-

ment 4 all dark; dark basal bands on foretarsal segments 3-5 almost

completely encircling each segment, dark scales occasionally absent

from ventral surface; foretarsal segment 2 light in apical 0.20-0.45; fore-

tarsal segment 3 light in apicalO.50-0.85. ... . . . . 2. oswaldoi

Subcostal light spot of vein C usually greater than 0.5 (0.45-1.00) length

of subcostal dark spot; upper mesepimeron (mep) often with 1-4 light,

obovate scales; hindtarsal segment 2 usually dark in basal 0.25-0.35; hu-

meral light spot of vein C usually large, 1.8-3.5 (1.0-3.7) length of basal

dark spot (ig. Gy oi) oe% . . .8& rangeli

Subcostal light spot of vein C AOE aye 168s than 0. 5 length of subcos-

tal dark spot; upper mesepimeron (mep) usually without light scales;

hindtarsal segment 2 and humeral light spot of vein C variable . . .7

Hindtarsal segment 2 dark in about basal half, 0.40-0.55 (0.3-0.6); light

wing spots at least on veins C and R light cream to yellowish, not white

. 5. aquasalis

13. benarrochi

Hindtarsal segment 2 dark in less than basal 0.40, or if greater than 0.40

then light wing spots white, not light cream to yellowish .... .8

Vein C humeral light spot less than 2.0 length of basal dark spot . . .9

Vein C humeral light i ie to or greater than 2.0 length of basal dark

SOT he OR anv ea ee rower Pu he ee LO

Light spots on wing usually very light, white or very light cream; vein C

with humeral light spot 1.3-2.0 of basal dark spot (fig. 7) . 9. trinkae

Light spots on wing usually cream, at least on anterior veins; vein C with

humeral ee ee Q0.7-1.3 sy 7- 1. a of basal dark oe. (fig. 7)

. 10. nuneztovari

Light scales on wing (at least anterior veins) and coxae (usually) gray to

cream to yellow, not white; foretarsal segment 5 cream, gray or golden

iapica OPSO0.5 0" Af ae . . 4. noroestensis

Light scales on wing and coxae asitally: white or very light cream; foretar-

Sale ete OU banGcd ty kk Guan ela he sa a kw A

Midtarsal segment 5 usually cream in less than apical 0.3; foretarsal seg-

ment 2 with a cream to white band in apical 0.25 (0.18-0.35), segment

5 usually golden to brown, occasionally differentiated into 0.5 dark, 0.5

light; vein C humeral ak spot usualiy 2.0-4.0 (1.2-4.1) of basal dark

SODE Chigi yy yee alr eg. Ld, strodei

Midtarsal segment 5 cream in 1 apical 0. 2 0, fe foretarsal segment 2 witha

cream to white band in apical 0.31-0.46, segment 5 cream to white in

about apical 0.5; vein C humeral wee ats 2.0-2.5 (1.3-4.0) of basal

dark spot (fig. aye hay ae . . 9. trinkae

ZA.

3(2).

4(3).

5(4).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 29

MALE GENITALIA

Ventral claspette without setae, apex rounded and not expanded laterally

into large auriculate lobe; preapical plate of ventral claspette very small,

weakly sclerotized; apex of aedeagus slightly broader than long; sternite

IX short, subrectangular (fig. 8) (Albimanus Group) . . J. albimanus

Ventral claspette with setae, or if without setae then apex expanded later-

ally into large auriculate lobe; preapical plate of ventral claspette and

aedeagus variously developed; sternite IX moderately long to long, sub-

trapezoidal to subtriangular (Oswaldoi Group)

Oswaldoi Group

Ventral claspette without setae, apex expanded laterally into large, auricu-

late lobe, preapical plate small, oval and heavily sclerotized; apex of

aedeagus about 1.5 as long as wide (fig. 31) (Triannulatus Subgroup) .

. . 14. triannulatus

Ventral Claspene ante co at least on ‘Basil builds, apex with or without

lateral expansion, preapical plate various; apex of aedeagus less than 1.5

as long-as wide (Oswaldoi-Suberoup) woe ie ek ee ee 8

Oswaldoi Subgroup

Ventral claspette with apex rugose or deeply striated, and moderately to

strongly expanded laterally into rounded or pointed lobe, setae on later-

al margins not extending toward apex as far as level of apical margin of

preapical plate; apex of aedeagus without leaflets (Strodei Complex) . 4

Ventral claspette with apex neither rugose nor deeply striated nor laterally

expanded, setae extending to apex or at least as far as level of apical

margin of preapical plate; apex of aedeagus with or without leaflets

ACW! SON) Oe ee ee ee MO a G

Strodei Complex

Ventral claspette small, about 0.33 length of sidepiece, apex moderately

expanded laterally, apicolateral margins sharply angled and moderately

pointed; basal lobule of ventral claspette small, narrow, curving mesad,

with setae along basal margin short, about equal to or slightly longer

than width of aedeagus; preapical plate of ventral claspette heavily scler-

OMe (11g ZO) ee 6 el ONO ES Benarroct

Ventral claspette large, about 0. = length of sidepiece, apex strongly ex-

panded laterally, apicolateral margin produced into large, rounded lobe

with basal and lateral margins convex and apical margin weakly concave;

basal lobule of ventral claspette large, with setae along basal margin

long, about 2.0-3.5 width of aedeagus; preapical plate of ventral cla-

spette very weakly to moderately sclerotized’. 0. oo 6". Ge eee

Ventral claspette with setae on lateral margins extending toward apex to

base of apicolateral lobe, preapical plate weakly to moderately sclero-

tized and moderately well-defined (fig.27). . . . . . . IJ. strodei

Ventral claspette with setae usually on basal lobule only, not extending to

base of apicolateral lobe, preapical plate very weakly sclerotized and ill-

defined: (hig. Vb wi ai A a eo ieee in a ee ee

6(3).

7(6).

8(7).

9(8).

10(9).

LEC9).

Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Oswaldoi Complex

Apex of aedeagus with a pair of long, sclerotized, serrated leaflets about

0.4 length of aedeagus; ventral claspette with large basal lobule with se-

tae along basal margin 1.5-2.0 width of aedeagus, preapical plate oval to

semicircular and moderately sclerotized (fig. 19). . 7. anomalophyllus

Apex of aedeagus without leaflets or occasionally with very small, mem-

branous leaflets; ventral claspette with large or small basal lobule, pre-

apleal plate vVariousivdeveloped. = 6.6.44) Bee eT

Ventral claspette strongly conoid, with a very small median sulcus at apex,

setae on lateral margins (exclusive of basal lobule) short and extending

toward apex only as far as level of apical margin of preapical plate, setae

along basal margin of basal lobule long about 2.0-3.0 width of aedeagus,

preapical plate very large, crescent shaped and ae sclerotized (fig.

Wa) ee are . . 6. ininii

Ventral claspette not strongly coneid. either truncate ¢ or with a moderately

small to large median sulcus at apex, setae on lateral margins extending

to or nearly to apex, setae along basal margin of basal lobule short to

lone. preapical Pilate spice 116 tite 2. dw kk i ae eB

Ventral claspette with a concentration of long setae about 1.5 width of

aedeagus on basomesal margin of basal lobule and directed caudally into

mesal cleft, wide at apex (width 0.4-0.5 length of claspette) with abrupt-

ly angled, rounded lateral margins, aha plate small, oval and heavi-

ly sclerotized (fie. 21). a . . 8 rangeli

Ventral claspette without setae concentrated on Gasomesal margin of basal

lobule, shape and preapical plate varied: 232. 5 2. oe

Ventral claspette with apex moderately broad to broad, width at apex 0.4-

0.6 length of claspette, lateral margins of claspette not tapering appreci-

ably medially toward apex, lateral margins of apex abruptly angled, pre-

apical plate moderately small and semicircular to oval. . . . . . 10

Ventral claspette with apex moderately narrow to narrow, width at apex

about 0.3 length of claspette, lateral margins of claspette tapering

toward apex, lateral margins of apex evenly curved, preapical plate vari-

Dis deve ORO i) a ee ee ww LI

Length of aedeagus 1.33-1.60 (1.31-1.89) length of ventral claspette; ven-

tral claspette moderately short, 0.25-0.40 length of sidepiece, width of

apex 0.50-0.60 (0.50-0.64) length of claspette (fig. 25) . :

mee 10. nee iovari

Leneth of pedeaeur 1 00- 1 20 (1 00- i a3) ieneth of ventral claspette; ven-

tral claspette moderately long, 0.40-0.50 length of sidepiece, width of

apex 0.43-0.50 (0.38-0.54) length of claspette (fig. 23) . . 9. trinkae

Aedeagus truncate at apex; preapical plate of ventral claspette large, semi-

circular to oval, heavily sclerotized (fig. 13). . . . . 4. noroestensis

Aedeagus rounded at apex; preapical plate of ventral claspette variously

EVO ee eg ew ke he

12(11).

1Stl2).

32),

4(3).

a2);

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 31

Ventral claspette with setae along basal margin of basal lobule moderately

short about equal to or slightly longer than width of aedeagus, setae

usually not reflexed, preapical plate moderately small, circular to oval

and weakly to moderately sclerotized (fig. 15) . . . . . 5. aquasalis

Ventral claspette with setae along basal margin of basal lobule long about

2.0 or more width of aedeagus, setae often reflexed caudally, preapical

plate large and moderately to strongly sclerotized . . . .... 13

Ventral claspette with setae along basal margin of basal lobules about 2.0

width of aedeagus, preapical plate large, usually crescent shaped and

moderately to strongly sclerotized (fig. 10). . . . . . .2. oswaldoi

Ventral claspette with setae along basal margin of basal lobules very long

about 3.0 width of aedeagus, preapical plate strongly sclerotized, large

and circular to semicircular with small basolateral projections (fig. 12)

3. galvaoi

PUPAE

(3. galvaoi, 7. anomalophyllus and 12. rondoni not included)

Hair 9-VII long, usually more than 0.33 length of segment. . . .. .2

Hair 9-VII short, less than or equal to 0.33 length of segment. . . . .9

Hair 9-V less than 2.3 length of 9-IV; 9-VI relatively short, 9-VII 1.4-2.0

leneth of 9-VI1... ..... oo

Hair 9-V 2.3 or more length of so IV, rarely 7 0: 9-VI Itong, 9-VII 7 0- : 5

leneth of 9-VI ...<.. : : Se

Hair 9-VII usually less than 0.5 length of segment ve 13). ae

4. noroestensis

Hair 9 VII about 0. Sh or more ener of S eaeat LAE is i re

Sum of branches of hairs 1-III, 5-III and 0-VI less than 18 (fig. 27)

. . Ll. strodei

Sin of Bega ctics ol oe 1 IIL, oa I ana 0-VI greater than 20 (fig. 17)

Re let a aaa ae tee . 6. ininii

Pinna of trumpet very long, about 4.0-6.5 (3.4-8.0) length of meatus and

narrow, not appearing to taper toward apex; hair 9-V-VIII very long,

slender, acuminate; 9-III small, unpigmented, slightly longer than 9-I];

7-C with one branch very long, about 1.5-2.0 length of other branch(es);

6-I,II very long and 9-I subequal to 6-I; 3-I subequal to 2-I; 2-I 3-6

branched: size small (fic.31) . ... piace e 4. triannulatus

Pinna of trumpet usually equal to or less than 4. 0 length of meatus and

moderately wide, if longer than 4.0 then tapered or hair 9-V less than

2.3 length of 9-IV or 9-III about 2.0 length of 9-II; hair 9-V-VIII moder-

ately slender to heavy; 9-III small to moderate, variously pigmented; 7-C

variously developed; 6-I,II and 9-I short to nes 2,3-1 nae devel-

oped; size moderately large to large. . . . ER 3

6(5).

7(6).

8(7).

9(1).

10(9).

2(1).

Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Hair 9-III strongly pigmented, heavy and usually thick, at least 2.0 length

of 9-II; 6-II 2.0 or more length of 7-II; 2-I moderately developed, 3-5

branched (2-6), forked about 0.5 distance from base; 3-I subequal to 2-I

(Mis. Oy . . IL. albimanus

Hair 9-III Caualis jess than On 0 ieneth of 9 Il, y feaeer then weakly to

moderately pigmented and not thick and heavy- 2,3-I and 6-II varied

Hair 2-1 moderately short, usually with 4-8 (2-10) branches arising near or

atvbase Gigs Vo) uve . . . J. aquasalis

Hair 2-I moderately long to cae. foe. pases oie at least 0.20 dis-

Cane de ett he Mehndi A tise Pari eg Lg &

Sum of branches of hairs O-II and O-III usually greater than 8 (8-14); O-II

moderate, usually 5-7 branched (4-7); 7-C with branches subequal or

one branch slightly longer than other(s) (fig. 21). . . . .8& rangeli

Sum of branches of hairs O-II and O-III usually less than 8 ee 9); O-IT small,

single to triple (1-4); 7-C with one branch about 1.5 length of other

Dic Beles) pie Se Oe Nee ain Se 8 erinkae

Pinna of trumpet short, about 1.6-2.1 length of meatus, appearing apically

flared in lateral aspect, meatal cleft appearing narrow and basally point-

ed; hair 6-II equal to or slightly longer than 7-II (fig. 10). . 2. oswaldoi

Pinna of trumpet moderately long to long, 3.0-5.5 length of meatus, not

appearing apically flared in lateral aspect, meatal cleft moderately point-

ed to rounded at base; hair 6-II conspicuously longer than 7-II . . 10

Pinna of trumpet moderately long, 3.5-4.5 (3.0-4.8) length of meatus,

appearing broad medially and tapered toward apex in lateral aspect; hair

10-C less than 0.5 length of 12-C; 6-I usually more than 2.0 length of

The 25 eo 2 e . . 10. nuneztovari

Pinna of trumpet long, 4. 5: : l (4. 45. 5) nei of estas. not appearing to

taper toward apex in lateral aspect; hair 10-C more than 0.5 length of

12-C; 6-I about 1.5-2.0 length of 7-I (fig. 29) . . . . .13. benarrochi

LARVAE

(3. galvaoi and 12. rondoni not included)

Hair 1-P plumose, with filiform branches; 9-P,T pectinate, usually 4-6

branched (4-7); 13-I-IV moderately large to large; 4-C moderately long

to long, usually extending to anterior margin of head capsule (fig. 9)

(Albimanus Group) . . . . . I, albimanus

Hair 1-P palmate; 9-P,T single: 13 Fe IV etna to large: 4-C short or we

(OswaldoiGroup) ...... st Wager ~~)

Oswaldoi Group

Hairs 2-C widely spaced, clypeal index about 1.0-2.0, single to plumose . 3

Hairs 2-C closely op unite, ue oa index about oh 5-4.0, tis and

simple or barbed . . . oie ei Navies |

3(2).

4(3).

5(4).

6(5).

7(4).

8(7).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 33

Hair 11-I large, usually 5-7 branched (3-7); 13-I very large, usually 3

branched (2-4); lateral arm of spiracular apparatus long, directed lateral-

ly; 1-P with 15-20 (13-20) very narrow to narrow lanceolate leaflets

(rarely filiform); 2-C single, and simple or barbed (fig. 32) (Triannulatus

Subgroup) . . . . 14. triannulatus

Hair 1 1- I moderately jibes. 2-4 branched: 18 -[ ‘Sia to moderately large,

usually more than 3 branched; lateral arm of spiracular apparatus very

short to moderately long (except long in ininii); 1-P with usually 9-16

(8-18) narrow to broad lanceolate leaflets; 2-C single to plumose (Os-

WAlGOt SUCT OUD) ce cl ie a, aig et ana Mucha wees ea bape y hey

Oswaldoi Subgroup

Hair 3-C, and usually 2-C, plumose in about apical half, with distinct, mod-

erately 1OMe tO 16s WianOhes 3a ee ee ae ate Ma? aS

Fairs. 2. 5-Csinele, and sinmmle'os barbed fila hp eee ai a abt ee ha

Hair 1-A long, at least 2.0 width of antenna at point of insertion; 13-IV

small, usually 10-13 branched (6-13); 1-X moderately short, as long as

or slightly longer than saddle; 3-C with moderately long branches and

2 barbed (hig: 30). fo . . « L3. benarrochi

Hair 1-A short to moderately short, always less than 2.0 width of antenna

at point of insertion; 13-I[V moderately large, usually 5-7 branched

(3-8); 1-X longer than saddle; 2,3-C with branches subequal in aya or

branches of 3-C slightly longer than those of 2-C. . . . . iO

Hair 1-X inserted on saddle, on or near ventral margin; anal gills usually

short, about 0.5 length of anal segment; 2,3-C with simple branches,

rarely dendritic; lateral arm of spiracular apparatus very short; pecten

with median teeth mixed medium and short (fig. 16) . . . 5. aquasalis

Hair 1-X not inserted on saddle; anal gills long, as long as or longer than

anal segment; 2,3-C with usually dendritic branches; lateral arm of spi-

racular apparatus moderately long; pecten with median teeth mostly

SRSA ED Nee eee Oi Ie RE oe ee oe RIO OE

Hair 4-C single or forked, moderately long to long, usually extending to

near or beyond base of 2-C, if moderately long (0.30-0.60 length of 3-C)

then 13-V 46 branched (4- Dy aes : .8

Hair 4-C variously branched, short to maderatel: lore, usaally not ie

ing to base of 2-C, if moderately long (0.30-0.45 length of 3-C) then

13-V almost always 3 branched (3-5) at least on one side of larva . .9

Hair O-II very short, about 0.5 or less length of leaflets of 1-II, inconspicu-

ous, single to triple; 3-C 0.5-0.8 length of 2-C; 4-C long, 0.7-1.0 length

of 3-C; 13-IV moderately large, 3,4 branched, 1.5-2.0 length of leaflets

of 1 IV Ce ae Cie . . 9. trinkae

Hair O-II moderately long, subequal to or longer than length of leaflets of

1-II, conspicuous, usually 5-8 branched (4-10); 3-C 0.76-0.90 length of

2-C; 4-C moderately long, 0.3-0.6 length of 3-C; 13-IV moderate, usually

4-6 branched or _ aa to or bids nae than leaflets of 1-IV (fig.

ZO ik ieee . . . . LO. nuneztovari

34 Contrib. Amer. Ent. Inst., vol. 15, no. 7,1980

9(7). Hair 1-A long, about 2.0 width of antenna at point of insertion; lateral arm

: of spiracular apparatus long, directed laterally (fig. 18) . . . 6. ininii

Hair 1-A short, as long as or slightly longer than width of antenna at point

of insertion; lateral arm of spiracular apparatus short to moderately

ess 10

short, directed caudolaterally

Hair 5-I short, inserted 0.75-1.00 its length from lateral margin of abdo-

men; 2-III felatively short, 132.0 length of leaflets of 1-III; 2,13-IV

. 8 rangeli

10(9).

moderately short, slightly longer than leaflets of 1-IV (fig. 22)

Hair = I ‘moderately short, inserted less than 0. 75 its length from lateral

margin of abdomen; 2- il longer than 2.0 length of leaflets of 1-III;

2,13-IV PORTE. = about 1.5 Pee of leaflets of 1-IV (fig. 14)

. .4. noroestensis

Li). Hair 1-P with 13-17 narrow leaflets; 2-P 16-24 branched; 1-M 31-35

branched; 1,2-P rarely sharing common tubercle; 1-X inserted on saddle

near ventral margin, or rarely on ventral margin at base of indentation

. . Ll. strodei

(Mie.25).

Hair 1-P with 9- 12 od eth, fieag to Were eis: 2- Pp usually 12-14

branched (12-17); 1-M usually 20-30 branched (20-32); 1,2-P arising

from a common tubercle; 1-X inserted usually on ventral margin of sad-

dle at base of indentation, or less often on saddle near ventral margin

. 7, anomalophyllus

(fig. 20)

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 35

ALBIMANUS GROUP

1, Anopheles (Nys.) albimanus Wiedemann

Figs. 1, 4, 5, 8,9

1820. Anopheles albimanus Wiedemann 1820:10. TYPE: Holotype female, Santo Domingo

(Dominican Republic) [NMW]. Specimen in poor condition (abdomen and antennae mis-

sing, legs missing or incomplete except for complete left hindleg), with following labels,

//{red square] //albimanus/det Wiedem//albimanus/Wied/St. Domingo// (Belkin 1968b:8).

1900. Anopheles cubensis Agramonte 1900:460-464. TYPE: Syntypes males and females, sever-

al localities in Cuba [LU]. Synonymy with albimanus by Theobald (1907: 106-109).

1901. Anopheles argyrotarsis (!) albipes Theobald 1901:125-128. TYPE: Lectotype male with

the following labels, //16.XII.99//8.2.00 Jamaica Dr. Grabham//Type//Anopheles argyri-

tarsis var. albipes Theobald (Type)// [BM; designation of Belkin 1968b:8-9]. Originally

described from unspecified number of males and females from Jamaica, Guyana, Rio de

Janeiro, Antigua and India. Synonymy with albimanus by Coquillett (1906:8, 13).

1905. Cellia tarsimaculata Goeldi 1905:133. TYPE: Same as for albipes [BM; designation by

Belkin 1968b:8-9]. Unjustified emendation for albipes Theobald 1901.

1938. Anopheles albimanus bisignatus Hoffmann 1938:176-177. TYPE: Syntypes adults, Gon-

zalez (Tamaulipas), Nov; Tampico (Tamaulipas), June, Oct, Nov, Dec, Jan; Altamira (Ta-

maulipas), June; El Mante (Tamaulipas), July; San Jeronimo, Laguna de Tamiahua (Vera-

cruz), June, July, Mexico [UM], (Belkin, Schick and Heinemann 1965:34).

1938. Anopheles albimanus trisignatus Hoffmann 1938:177-178. TYPE: Syntypes adults, region

of Tampico (Tamaulipas), Mexico, Dec, Jan [UM], (Belkin, Schick and Heinemann 1965:

34).

Anopheles (Nyssorhynchus) albimanus of Dyar (1922:103; 1925:187, 188, 194; 1928:103, 434-

435); Root (1922a:322; 1922b:384-389, 390-391; 1923:267, 269, 276; 1924b:460-462; 1926a:

51; 1926b:709; 1932:779-782); Christophers (1924:38-39, 89); Barraud and Covell (1928:674);

-Lima (1928:94-95); Boyd and Aris (1929:309-399); Kumm (1929:6-8; 1941a:359; 1941b:93-

97, 99-100); Hill (1930:712); Carley (1931 :293-296); Edwards (1932:45); Senevet (1932:252;

1938:181-182; 1948c:437, 439); Townsend (1933a:101-102); Rozeboom (1936b:480-489;

1938a:95, 99-100; 1938b:289-301; 1942:237; 1963:110-114); Kumm and Ruiz (1939:437-438,

441); Lane (1939:19-21; 1944:263-265; 1949:401-402; 1953:256-259); Simmons (1939: 124-

125); Gabaldon (1940:3-7); Gabaldon, Lopez and Ochoa Palacios (1940:33); Gabaldon, Aqui-

lera and Arevalo (1941:59); Komp (1941a:88, 92-97; 1942:5, 40, 43, 67-69, 80, 115-117, 132,

154-156); Rozeboom and Gabaldon (1941 :95-97); Simmons and Aitken (1942:38, 45, 53, 60,

80-84); Cova Garcia (1943:467-472); Ross and Roberts (1943:35-36); Matheson (1944:115-

116); Floch and Abonnenc (1945:1-3; 1946b:3-5); Levi-Castillo (1945:2, 7, 92-104; 1949:9, 10,

15, 27, 32, 57, 58, 67, 72, 76, 81, 85); Carpenter, Middlekauff and Chamberlain (1946:80-83);

Gabaldon and Cova Garcia (1946a:19-32; 1952:181); Arnett (1947:194-196; 1950:106, 110,

112, 114); Vargas (1948:153-163; 1959:376, 382); Penn (1949:52, 68-69); Thompson (1950:

692-695); Vargas and Martinez Palacios (1950:119-123; 1953:326-327; 1956:119-122); Belkin

(1952:121, 123, 130; 1965:14; 1968b:8-9); van der Kuyp (1954:49-50); Carpenter and La Casse

(1955:55-57); Frizzi and Ricciardi (1955:403, 404, 406); Horsfall (1955:172-177); Perez Vigue-

ras (1956: 180-186); Vargas V. (1956:27, 29; 1957; 1958a; 1958b; 1961a:97-105; 1961b:153-

170); Bejarano (1957:326-332); Rozeboom and Kitzmiller (1958: 244); Schreiber and Guedes

(1959b:128-129; 1960:356-357); Stone, Knight and Starcke (1959:30); Forattini (1962:371-

376); Montchadsky and Garcia Avila (1966:32); Porter (1967:36); Elliott (1968:244); Keppler

and Kitzmiller (1969:31); Belkin, Heinemann and Page (1970:24-27); Keppler, Kitzmiller and

Rabbani (1973:42-49); Cova Garcia and Sutil O. (1976:30; 1977:14, 45, 63, 87); Cova Garcia,

Pulido F. and Amarista M. (1977:157); Knight and Stone (1977:60).

Anopheles albimanus of Robineau-Desvoidy (1827:411); Wiedemann (1828:13); Dyar and Knab

(1906a:175-176); Busck (1908:57); Darling (1909: 2051-2053); Knab (1913:34-37, 40-42);

Christophers (1915:390-391); Zetek (1915:221-271); LePrince and Orenstein (1916:43-114);

Howard, Dyar and Knab (1917:979-984); Johnson (1919:424); Evans (1921:446-451; 1922:

36 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

213); Root (1924a:207); Gowdey (1926:74); Hoffman (1926:377; 1930:358); Senevet (1931:

76-79); Earle (1932:381-384); Hoffmann (1932:523-529; 1938:167-180); Clark (1934:643);

Curry (1934:644-651); Hill (1934:425-428); Martini (1935:14, 15); Rozeboom (1935:521-528;

1936a:471-478; 1941:98-104; 1953:1116; 1962:668); Clark, Komp and Jobbins (1940:62-67);

Kumm, Komp and Ruiz (1940:388-391, 419); Vargas (1940a: 199-200; 1940b:67; 1941:112,

114, 116, 118; 1976:88, 89); Carr, Melendez and Melendez (1941:749-750); Kumm and Ram

(1941:559); Carr and Hill (1942:600-601, 604, 606); Carr and Melendez (1942:59-60); Carr,

Melendez, Ros and Melendez (1942:70-71); Kumm and Zuniga (1942:401, 404; 1944:8-15);

Kumm, Bustamante and Herrera (1943:374); Russell, Rozeboom and Stone (1943:20, 26, 29,

30, 41); Eyles (1944:2, 10-11); King, Bradley and McNeal (1944:39); Weathersbee (1944: 25-

28); Carpenter, Chamberlain and Wanamaker (1945:402); Hill and Hill (1945:3; 1948:36-37);

Paul and Bellerive (1947:63-66); Thompson (1947:78); Castellanos, Murrieta, Lassmann and Or-

tiz (1949:34-35); Burgess (1950:108); Senior-White (1950:4-5); Muirhead-Thomson (1951:

1114-1117); Charles and Senevet (1953:1109-1115); Rozeboom (1953:1116; 1962:668); van

der Kuyp (1953:144, 145); Floch (1954:4); Vargas and Martinez Palacios (1955:82-121, 123);

Jensen and Jones (1957:464-469); Vargas V. (1958c:1, 3); Foote and Cook (1959:112-113);

Cova Garcia (1961 :33-34, 68, 85-86, 113, 122-123, 154, 178); Fauran (1962:73-80; 1964:52-

53); Hobbs (1962:245-251; 1973 :420-423); Vincke and Pant (1962:2-11); Davidson (1963:25-

33); Rachou, Lyons, Mourra Lima and Kerr (1965:32-61); Fauran and Courmes (1966: 105);

Stojanovich, Gorham and Scott (1966a:11, 19, 28; 1966b:19, 31, 38); Taylor (1966:393-397);

Gorham, Stojanovich and Scott (1967:14, 45, 49; 1973:110, 136, 140); Scherer, Campillo-

Sainz, Dickerman, Diaz Najera and Madalengoitia (1967:79-91); Central America Malaria Re-

search Station (1968:8-37; 1969:5-53; 1970:5-41; 1971:9, 12-18, 24-41, 48-50, 55-56, 70-71;

1972:5-13, 17, 21-27, 31-32, 37-41, 57); Wilton and Fay (1970:628-632); Morales-Ayala (1971:

139); Ali and Rozeboom (1972:574-579); Breeland (1972a: 751-754; 1972b:62-72; 1972c:99-

106); Elliott (1972:757); Rabbani and Kitzmiller (1972:421-432); Wilton and Fetzer (1972:

459-460); Wilton, Fetzer and Fay (1972:23-27); Lofgren, Boston and Borkovec (1973:187-189);

Shelton (1973:3-4, 8, 10); Hobbs, Lowe and Schreck (1974:389-393); Cova Garcia and Sutil O.

(1975a:21; 1975b:212); Lowe, Schreck, Hobbs, Dame and Lofgren (1975:160-168); Warren,

Richardson and Collins (1975:549-551); Garcia-Aldrete and Pletsch (1976: 76, 79); Pletsch and

Garcia-Aldrete (1976:87-92); Bautista-Garfias, Mercado-Sanchez and Morilla-Gonzalez (1977:

15-18); Warren, Collins, Richardson and Skinner (1977:607-61 1).

Anopheles (Anopheles) albimanus of Bonne and Bonne Wepster (1925:516).

Anopheles (Cellia) albimanus of Dyar (1918:151).

Cellia albimana of Coquillett (1906: 13); Theobald (1907: 106-109 in part; 1910:69-70 in part);

Surcouf and Gonzalez-Rincones (1912:274); Lutz, Souza Araujo and Fonseca Filho (1919:85);

Neiva and Pinto (1922a:321; 1922b:356-357); Boyd (1926:31, 48).

Nyssorhynchus albimanus of Blanchard (1905:202-204); Autran (1907:10-11); Surcouf and Gon-

zalez-Rincones (1912:271-272); Townsend (1934:493, ?).

Cellia albipes of Theobald (1903:110-113; 1905a:15-17; 1905b:11); Prout (1909:487).

Nyssorhynchus cubensis of Blanchard (1905:204-205); Surcouf and Gonzalez-Rincones (1912:272-

273).

Cellia argyrotarsis (!) of Theobald (1905a:4; 1905b:11; 1907:105; 1910:68); Prout (1909:487).

Anopheles argyrotarsis (!) of Theobald (1901:123-125); Johnson (1919:425).

Laverania argyrotarsis (!) of Theobald (1902:183).

Anopheles (Nyssorhynchus) rondoni of Martini (1935:14, 24).

FEMALE (fig. 5). Wing: 3.16 mm. Proboscis: 2.05 mm. Palpus: 1.95 mm. Fore-

femur: 1.45 mm. Abdomen: about 2.5 mm. Head: Integument brown to dark

brown. Interocular width 3, 4 ommatidial diameters. Proboscis about 1.4 length of

forefemur. Posterior vertex and occiput with large, erect, white, truncate, weakly

forked scales becoming darker posteriorly; anteriorly, vertex with a few, small, obo-

vate, decumbent scales along upper orbitals and longer, narrow, white, decumbent

scales along interorbital line. Frontal tuft with 14-20 long, white, setiform scales.

Proboscis brown to dark brown, with decumbent scales and short setae; appearing

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 37

metallic dark copper when viewed at angle; labella lighter than labium. Palpal seg-

ments 1-3 predominantly same color as labium; segments 2, 3 with a small cream to

white band at apex; segment 4 either all dark or yellow to golden brown on medio-

lateral surface, never white or cream; segment 5 white. Thorax: Pruinose areas of

scutum silver gray to reddish; nonpruinose regions light brown to dark brown. Med-

ian anterior promontory area with 10-15 long, erect, white scales and a few, long,

light setae. Upper humeral tuft scales similar to those on median anterior promon-

tory, lower tuft scales shorter, darker and obovate or cuneate. Prescutellar region

moderately large, dark, horseshoe shaped to triangular; bare space extending onto

median portion of scutellum. Ppl with 1, 2 long, dark setae. Sp with about 2-8 light

setae and a few, lanceolate, light scales. Pra with 5-10 long, dark setae and a patch of

long white scales. Upper stp with 2-4 long, dark setae and approximately 10 cream

to yellow, truncate, spatulate scales in a horizontal arc. Lower stp with 1, 2 long,

dark setae and a small patch of scales similar in shape and color to those on upper stp.

Upper mep with 5-10 long, dark setae oriented horizontally or diagonally; scales ab-

sent. Legs: Light scales on coxae generally white. Femora and tibiae as in section

description; midfemur with anteroapical spot and knee spot moderate to large. Fore-

tarsal segment 1 with less than apical 0.15 cream to white; segment 2 cream to white

in apical 0.15-0.35; segment 3 cream to white in apical 0.1-0.6; segments 4, 5 normal-

ly all brown, occasionally apex of 5 with cream to light brown scales; specimens from

Caribbean islands with smaller apical light bands on foretarsus. Midtarsus usually

with fewer extensive light areas than in Oswaldoi Subgroup, appearing very dark; seg-

ment | and occasionally 2 with a very small, golden, apical band; segments 3, 4 with-

out an apical light band; segment 5 with or without a golden apical band. Hindtarsal

segment | with a yellow to white apical band, often indistinct; segment 2 dark in bas-

al 0.4-0.6 (0.4-0.8); apical portion of segment 2 and all of segments 3, 4 white; very

rarely base of segments 3,4 with a brown ring; segment 5 brown in about basal 0.5,

remainder cream to white. Wing: Apex moderately rounded, not strongly tapered.

Light spots normally cream, particularly on veins C and R; dark spots varying from

light brown to almost black; considerable variation in lengths of light and dark spots.

Vein C usually with large dark spots; presectoral and/or sectoral light spots often ab-

sent; C humeral light spot about 1.0-2.0 (0.95-2.50) of basal dark spot (basal dark

spot usually greater than or equal to 0.5 of humeral light spot); subcostal light spot

about 0.25-0.40 of subcostal dark spot; preapical light spot 0.25-0.45 of preapical

dark spot. R with 2 dark spots, presectoral and sectoral, moderately long and sub-

equal in length. R,,3 about 0.5 of R,. Rs-R,,3 predominantly dark, with 2 or 3 us-

ually small, light spots. Base of M usually light followed by a moderately long to long

sectoral dark spot; preapical dark spot of variable length. Cu sectoral dark spot not

reaching furcation. Vein A subcostal dark spot moderately large, about 0.2 length of

vein. Apical light fringe spot small to moderate; small, inconspicuous, light fringe

spots at apices of Rais, Mi42, M314, Cu,, Cu, and A; a moderate size, light fringe

spot at level of 0.5 distance from base of A. Abdomen: Integument of tergites

brown to dark brown; occasionally light mottling present on segments II-VII. Ter-

gite I with light to dark, long setae laterally and along apical margin; without scales.

Long setae in posterior 0.5-0.7 of tergites I-VII, and on mediolateral, lateral and api-

cal margins; shorter setae more medially. Tergites II-VII with cream to golden, small,

obovate scales in subtriangular pattern in proximal segments, and more extensive sub-

rectangular pattern in apical segments; occasionally with darker median scales on ter-

gites II-VII; caudolateral margin with large, conspicuous, outstanding, dark scale tufts

on segments III-VII. Sternites II-VII with submedian and apical setae and submedian

38 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

white scales; caudomedian and caudolateral dark scale tufts on sternites ITI-VII simi-

lar to those on tergites, with a few light scales occasionally interspersed. Sternite VIII

densely covered with small, cream to golden scales, medially with longitudinal stripe

of dark scales.

MALE (fig. 5). Wing: 3.05 mm. Proboscis: 2.35 mm. Forefemur: 1.5 mm. Ab-

domen: about 2.75 mm. Essentially as in female except for sexual characters. Head:

Palpal segment 4 with a basal and an apical dark band; mediolaterally with mixed

light and dark scales; extreme base and apex with a few light scales; ventral surface

usually dark scaled. Antenna: About 0.7 length of proboscis. Legs: Forefemur 0.6

length of proboscis. Basal plantar surface of foretarsal segment 5 with about 5-8

moderately long, spiniform setae; setae longest toward apex. Claw on foreleg long,

slightly curved; submedian tooth about 0.25 length of claw, acuminate and recurved

at apex; external basal tooth subequal to submedian tooth and decurved.

MALE GENITALIA (fig. 8). Segment VIII: Tergite and sternite medially with

numerous broad scales; scales more densely distributed on tergite. Posterior border

of sternite with row of long setae. Segment IX: Sternite short, curved and subrec-

tangular, anterior border almost parallel with posterior border, tapering at lateral

margins. Slight, inconspicuous, ribbonlike, nonspiculose thickening (anterior apo-

deme) along anterior border of sternite. Sidepiece: Moderately narrow. Tergal sur-

face usually with 4, 5 long, submedian tergomedial bristles and one long apicolateral

bristle at base of clasper. Bristles laterad and mesad of tergomedial bristles 0.5 length

of tergomedial bristles. Parabasal spine usually at least 2.0 length of its tubercle. Ac-

cessory spines long; longer more dorsal spine about 0.5 length of sidepiece; more ven-

tral spine 0.58-0.70 of dorsal spine. Internal spine subequal to shorter accessory

spine, apically retrorse. Clasper: Ventromesal margin with row of short spicules.

Spiniform narrow and moderately short, subequal to seta b. Dorsal Claspette: About

0.5 length of sidepiece. Pedicel moderately long and narrow; base moderately curved

mesad. Leaflets broad and lanceolate; longest equal to or slightly less than 0.5 length

of claspette; basomesal projection of dorsal leaflet small. Ventral Claspette: About

0.32-0.40 of sidepiece; without setae; apex of ventral lobe produced into a pair of

rounded, striated lobes, separated by a moderately shallow median sulcus. Immedi-

ately basad of sulcus a very small, circular to oval, weakly sclerotized preapical plate.

Ventral surface of basal lobule produced into a large, inflated, ovoid, bulbous lobe,

often deflated and not apparent on slide. Phallosome: Aedeagus about 1.4 length of

ventral claspette; moderately wide; apex rounded, slightly broader than long, with

distinct membranous border; without leaflets; subapical, ventromesal, triangular pro-

jections nearly meeting or meeting on midline of ventral surface.

PUPA (fig. 8). Abdomen: 2.7 mm. Trumpet: 0.5 mm. Paddle: 0.75 X 0.55 mm.

Cephalothorax: Strongly pigmented, light brown to dark brown. Wing cases often

with longitudinal dark stripes. Hairs 1-3-C 1-3 branched; 2-C shorter and weaker

than 1,3-C. Hairs 4,5-C of subequal length; 4-C 2,3 branched (1-3); hair 5-C 2,3

branched (1-4). Hair 6-C single or forked. Hair 7-C 2-4 branched (2-5) near base,

often one branch longer than other(s). Hairs 8,9-C subequal in length; 8-C single (1-

3 branched), heavy; 9-C 1-3 forked about 0.3-0.5 from base. Trumpet: Weakly pig-

mented, light yellow to brown. Pinna moderately sclerotized and open; long, about

3.3-4.0 (2.9-4.5) length of meatus; in lateral aspect, appearing broad medially and

tapered toward apex. Meatal cleft broad, and pointed or rounded at base. Metano-

tum: Hair 10-C single, subequal to 11-C. Hair 11-C 2-4 branched (2-6). Hair 12-C

1,2 forked 0.25-0.50 from base, 1.5-2.0 length of 10-C. Abdomen: Hairs with fewer

branches in general than remaining species in section. Strongly pigmented, light

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 39

brown to dark brown, often with darker, median, longitudinal stripe. Float hair 1-I

with 8-13 major branches, extensively dendritic apically. Hair 2-1 3-5 forked (2-6)

about 0.5 from base, moderately developed; 3-I single, subequal to 2-I. Hair 4-I 3-6

forked (3-7) about 0.3-0.5 from base. Hair 5-I single to triple, long. Hair 6-I single,

very long, usually 2.0 length of 7-I. Hair 7-I 2-4 branched. Hair 9-I single, very long,

slightly shorter than or equal to 6-I. Hair O-II-IV 2-5 branched (2-6); 0-VII 1-3

branched (1-5). Hair 1-II 5-9 branched (5-11); 1-III 3-6 branched (2-7); 1-I'V-VII sin-

gle or rarely double, very long, about 1.5-2.0 length of segment; 1-IV slightly shorter

than 1-V-VII. Hair 2-III 3-5 branched (2-6). Hair 3-IV 3-6 branched (3-7), inserted

very close to hair 4-IV; 3-V 1-3 branched. Hair 5-III 4-8 branched (410), large. Hair

6-II single, very long, 2.0 or more length of 7-II; 7-II 2-5 branched (2-6). Hair 6-III-

VI single, long. Hair 7-III-V 2-5 branched (2-6), small to moderately small. Hair 8-

III-VII 2-5 branched (2-6), small. Hair 9-II very small, unpigmented; 9-III pigmented,

small, about 2.0 length of 9-II; 9-IV thick, about 2.0 of 9-III and 0.3 of 9-V,VI; 9-V-

VII slightly curved, acuminate; 9-VIII almost straight; 9-VII, VIII long, about 0.5 of

segment and about 1.2 of 9-V,VI. Hair 10-III 2,3 branched (1-4), moderately long.

Hair 4+VIII 1-3 forked. Terminal Segments: Apex of male genital lobe with small,

short, moderately distinct, mammilliform protuberance; lobe broad and heavy. Pad-

dle: Large, obovate, broad and emarginate at insertion of hair 1-P. Midrib with later-

al margin more sclerotized than mesal margin, distally becoming indistinct. External

buttress 0.65 length of paddle, distally very finely serrated; moderately distinct, short

(0.02 mm), fine, filamentous spicules distad of buttress, extending around apex a

short distance and becoming shorter along inner margin. Hair 1-P single, long (0.1

mm); 2-P 1,2 forked, subequal to 1-P.

LARVA (fig. 9). Head: 0.6 mm. Antenna: 0.3 mm. Anal Saddle: 0.45 mm.

Head: Integument usually extensively mottled, varying from greenish brown to dark

reddish brown. Collar moderately wide (about 0.06 mm) and dark. Mental plate

strongly sclerotized, almost black; median tooth moderately broad, about 1.5 as wide

as adjacent tooth, tapered to a point. Inner clypeal hair (2-C) with about 8-10 barbs;

outer clypeal (3-C) plumose with 8-10 short to moderately long branches; hairs 2-C

widely spaced, clypeal index about 1.25. Hair 4-C single or forked apically, moder-

ately long to long, usually extending to anterior margin of head capsule. Hairs 8,9-C

2,3 branched (2-4), large. Hairs 10,12-C 2-4 and 3-5 (3-6) branched respectively,

moderately large. Hairs 13,15-C dendritic, strong and fairly long. Antenna: Mesal

margin with numerous, moderately stout, acuminate spicules; ventral surface with

fewer, shorter, stouter spicules; few or no spicules on dorsal surface. Hair 1-A 2-4

branched (2-5), small, slightly longer than width of antenna, inserted on basal third

of antenna. Hair 4-A 2,3 forked (2-6), long. Thorax: Integument moderately to ex-

tensively pigmented, tawny to dark brown. Hair 1-P plumose, with a short to elon-

gate flattened shaft and 21-24 (12-27) filiform branches; 2-P like 1-P but longer, 19-

21 branched (19-29); hairs 1,2-P sharing common tubercle. Hair 9-P pectinate with

4-6 (4-7) long branches, long, subequal to 10-P. Hairs 10,12-P single and long. Hair

11-P 2,3 forked (2-4). Hair 14-P 5-10 branched (5-12), with a short flattened stalk.

Hair 1-M 31-36 branched (21-46), with a broad, elongate, flattened shaft. Hair 2-M

usually forked, 1-3 branched. Palmate hair 3-T with 10-14 (9-16) unsclerotized,

moderately long, lanceolate leaflets. Hair 9-T pectinate with 3-5 branches (1-7) in

apical half, very long. Abdomen: In general, the hairs with fewer and longer branch-

es than in remaining species. Hair O-II-VI 2-4 branched (2-6); 0-VI smallest; 0-VII 2

branched (1-4), small. Palmate hair 1-I with 12-14 (11-17) semitransparent, lanceo-

late leaflets; 1-II-VII with long, broad, heavily pigmented, lanceolate leaflets; 1-II,VII

40 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

smaller than 1-III-VI. Hair 2-II, II 3,4 branched (2-4), large; 2-IV,V single or rarely

double, long; 2-V about 2.0 length of 2-IV. Hair 5-I 3 branched (3,4), moderate in

size, located close to lateral margin; 5-III 5-9 branched (4-10), large. Hair 11-I 3,4

branched (2-5), large. Hair 13-I 3,4 branched (2-4), moderately large; 13-II 5,6

branched (3-8), moderately large; 13-III,IV 3,4 branched (2-4), large; 13-V 3-5

branched, very large; 13-VI 6-9 branched (3-10), moderately small. Tergal plate VII

usually less than 2.0 size of plate VI. Spiracular Lobe: Pecten with 14-18 (13-21)

acuminate teeth and with a conspicuous row of thin spinules on basal half of dorsal

surface; teeth usually as follows beginning ventrally: (1) 1 long, (2) 5-12 mixed short

to moderate, occasionally with a long interspersed, (3) 1 long, (4) 3-5 mixed short to

moderate and (5) 2,3 long or combination of short and long. Lateral arm of spiracu-

lar apparatus very short to short, extending a short distance below the spiracular

openings. Hairs 8,9-S 3,4 forked (2-5), with moderately long branches. Anal Seg-

ment: Hair 1-X single, very long, about 2.0 length of saddle, strongly developed; al-

most always inserted on saddle. Fine spicules over surface of anal segment except at

base; spicules along apical border stronger and longer. Ventral brush (4-X) with 8

pairs of strongly developed, plumose hairs. Anal gills usually shorter than or equal to

length of saddle, occasionally very short.

DISCUSSION. The monotypic Albimanus Group can be distinguished from the

Oswaldoi Group in the female by the combination of (1) palpal segment 4 all dark or

yellow to golden brown on mediolateral surface, never white or cream, (2) dark cau-

dolateral scale tuft absent from abdominal segment II and (3) foretarsal segment 5

usually all dark; in the male genitalia by (1) sternite IX short, subrectangular, (2) ven-

tral claspette completely without setae and with ventral surface produced into a pair

of large, inflated, ovoid, bulbous lobes, (3) preapical plate very small, circular to oval,

weakly sclerotized and (4) apex of ventral claspette rounded with a shallow median

sulcus; in the pupa by the combination of (1) pinna of trumpet long, broad medially

and appearing tapered toward apex, (2) hair 2-I 3-5 forked (2-6) about half distance

from base, moderately developed, (3) hair 9-I very long, slightly shorter than or equal

to 6-I, (4) hair 6-II single, very long, 2.0 or more length of 7-II, (5) hair 9-[V about

2.0 length of 9-III and (6) hair 9-VII, VIII long, about 0.5 length of segment; and in

the larva by the combination of (1) inner clypeal hair (2-C) barbed and clypeal index

about 1.25, (2) hair 4-C moderately long to long, (3) hair 1-P plumose with filiform

branches, (4) hair 9-P,T pectinate, (5) hair 13-I,III,[V few branched, moderately large

to large and (6) hair 1-X almost always inserted on saddle.

The external morphology of albimanus is very constant throughout its range. Usu-

ally the amount of intrapopulational variation is as great as or greater than the varia-

tion which occurs among populations, with the following exceptions. Many adults

from southern Texas exhibit a greater proportion of dark basal banding on hindtarsal

segment 2, often as much as 0.8 the length of the segment. Adults from the Antilles

usually have proportionally smaller light areas on the foretarsus than those from the

mainland. The extreme condition was observed in 4 specimens from Barbados that

completely lack light apical rings on foretarsal segments 2 and 3. Similarly, hindtar-

sal segment 5 is predominately dark in several adults from Puerto Rico. The pupae

from the Antilles often exhibit a greater number of branches than mainland speci-

mens on many of the abdominal hairs. Noticeable characters in the larvae which vary

among populations from different islands in the Antilles and also among different pop-

ulations on the mainland are, (1) length of the median shaft of hair 1-P, which is usu-

ally moderately long but occasionally very short, (2) number and arrangement of pec-

ten teeth and (3) size of the anal gills. Another variation, although not showing any

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 41

geographical pattern, occurs in several adults from Guayaquil (Ecuador), Acapulco

(Mexico) and Barcelona (Venezuela); palpal segment 4 may be predominately light

creamish brown, similar to that of triannulatus. Several of the specimens from Barce-

lona have a light apical band on foretarsal segment 4, also similar to that of triannula-

fus.

The small amount of morphological variation in albimanus is correlated with its

genetic compatability and the stable banding pattern of the salivary chromosomes

found among populations from several different localities. Hobbs (1962) examined

5 laboratory strains of albimanus from El Salvador, Guatemala, Mexico, Jamaica and

the Gorgas Memorial Laboratory colony in Panama. He found that there were no

heterozygous inversions and that the Mexico, El Salvador and Panama strains were

interfertile. He concluded that there was no evidence for the existence of isolated

strains and that albimanus is not polymorphic to any great degree. Similarly, Kep-

pler and Kitzmiller (1969) showed complete reciprocal fertility between 4 field pop-

ulations of albimanus from Mexico, Guatemala, Nicaragua, Costa Rica and one labor-

atory colony from El Salvador. Keppler, Kitzmiller and Rabbani (1973) examined

the salivary chromosomes from the above 5 strains and found them to be “‘remarkab-

ly uniform,” and after examining over 1000 slides discovered no individuals heterozy-

gous for an inversion.

I believe that albimanus diverged from the main phyletic line very early in the evo-

lution of the Albimanus Section (fig. 4). An. albimanus is the least derived species in

the section and retains several ancestral characters which it shares with several species

in the Argyritarsis Section of Nyssorhynchus. Almost none of these ancestral charac-

ters are found in the Oswaldoi Group except for a few in triannulatus. These charac-

ters are essentially those that distinguish the adult, male genitalia and larva of albima-

nus from the Oswaldoi Group, with the exception of the development of hair 9-P,T

in the larva and the short ninth sternite in the male genitalia; the latter 2 characters

seem to be derived since they occur nowhere else in the section.

Cytogenetically, albimanus is also quite distinct. In comparing the salivary gland

chromosomes of nuneztovari with albimanus, aquasalis and An. (Nys.) darlingi, Kitz-

miller, Kreutzer and Tallaferro (1973:443) stated that the similarities between nunez-

tovari and albimanus are not as “striking” as between darlingi and aquasalis. With re-

spect to the banding of salivary chromosomes, aquasalis seems to be more like dar-

lingi (of the Argyritarsis Section) than like albimanus. In comparing 3 species (aqua-

salis, oswaldoi, albimanus), Kitzmiller and Chow (1971:80-81) saw more homology

between the salivary chromosomes of aquasalis and oswaldoi than they did between

aquasalis and albimanus. Further they state, ‘“‘If the albimanus salivary gland map is

used as the standard map, it seems that many bands may probably have been added

to aquasalis and oswaldoi by some process, perhaps duplication or translocation dur-

ing evolution.”’ The combination of these findings clearly indicate that albimanus is

relatively remote from the remaining species in the Albimanus Section.

An. albimanus most likely evolved somewhere in Central America, radiating from

its center of origin north and south along the Pacific and Atlantic seaboards. The

species probably invaded the Greater Antilles via or south of the Yucatan Peninsula,

becoming distributed throughout the Greater Antilles and the larger, more northern

islands of the Lesser Antilles. Why albimanus has not invaded the southern islands of

the Lesser Antilles is not known. The most likely explanation is that albimanus is

still radiating from its center of origin and therefore has not yet been able to colonize

the more southern islands in the Lesser Antilles. The rate at which colonization oc-

curs is dependent on many factors and is different for each individual species. There

42 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

is considerable evidence that the Lesser Antilles are geologically more recent than the

Greater Antilles (Martin-Kaye 1969; LePichon and Fox 1971), which might partially

explain why albimanus has not yet extended its distribution into the Lesser Antilles.

Hoffmann (1938:176-178) described 2 interesting variations or mutants of albi-

manus (bisignatus and trisignatus) from the states of Tamaulipas and Veracruz, Mex-

ico. These 2 “‘varieties’ are distinguished by bisignatus having an additional black

ring at the base of hindtarsal segment 3, and ¢trisignatus having extra black rings at the

bases of hindtarsal segments 3 and 4. Rozeboom (1963) established a colony of bi-

signatus and found that trisignatus individuals were fairly common in the F, genera-

tion. He stated that trisignatus apparently represents a more marked expression of

the bisignatus gene or genes and that the trait was recessive. The occasional occur-

rence of these mutants probably led Martini (1935:24) to report rondoni from Mexi-

co, most likely based on a female of bisignatus. I have examined specimens exhibit-

ing either of these characters from Costa Rica, Guatemala and Texas.

BIONOMICS. Because of its medical importance, the literature on the bionomics

of albimanus is voluminous. Several good reviews and/or studies concerning the nat-

ural history of albimanus include Breeland 1972a, c; Central America Malaria Re-

search Station 1969-1972; Horsfall 1955:172-177; and Foote and Cook 1959:112-

113.

The immatures have been found in the following aquatic habitats by the project

“Mosquitoes of Middle America’: potholes, drainage areas, spring seepages, borrow

pits and ground pools (33%); ponds, lakes, dam spills and reservoirs (16%); streams

and canals (21%); ditches (6%); swamps (6%); marshes (5%); crab holes (4%); animal

and vehicle tracks (4%); rock holes (1%); and miscellaneous (4%). The immatures

are tolerant of brackish water and were found in brackish habitats in about 3% of the

collections. Previous studies indicate that albimanus can tolerate water up to and

possibly greater than 0.67 seawater (Taylor 1966:394; van der Kuyp 1953; Horsfall

1955:172-177; Central America Malaria Reaearch Station 1970). The immatures are

almost always collected in full sunlight or partial shade, although occasionally they

are collected in deep shade. The breeding sites are usually characterized by abundant

vegetation with scum and algae, and often are turbid with a muddy bottom. The im-

matures are often collected in polluted waters. The breeding sites are usually found

in areas of secondary growth such as plantations, open woodlands and meadows.

Aquatic plants often associated with the immatures are Pistia sp, Elodea sp, Naias sp,

Chara sp and Utricularia sp.

At least 51 species in the family Culicidae have been found associated with albima-

nus including 2 species of the Albimanus Section, triannulatus and aquasalis. An. al-

bimanus was most often collected with the following species in the majority of the

habitats in which this species occurs: Culex (Mel.) atratus, Cx. (Mel.) erraticus, Cx.

(Cux.) nigripalpus, Cx. (Cux.) coronator group, Anopheles (Ano.) grabhamii, Urano-

taenia ( Ura.) socialis; and to a lesser extent, Psorophora (Gra.) confinnis group and

Cx. (Mel.) pilosus. Species commonly found in ground pool type habitats with albi-

manus are An. (Ano. ) neomaculipalpus, An. (Ano.) pseudopunctipennis, Aedes

(Och.) taeniorhynchus, Ae. (Och.) tortilis, Ae. (Och.) scapularis, Cx. (Mel.) bastagar-

ius, Cx. (Mel.) chrysonotum, Cx. (Mel.) dunni, Cx. (Mel.) edueator, Cx. (Mel.) io-

lambdis, Cx. (Mel.) spp, Ur. (Ura.) geometrica and Ur. (Ura.) lowii; and occasionally

An. (Ano.) apicimacula, An. (Ano.) punctimacula, Ae. (Och.) angustivittatus, Ae.

(Och. ) sollicitans, Cx. (Mel.) conspirator, Cx. (Cux.) declarator group, Ur. (Ura.) pul-

cherrima, Ps. (Gra.) cingulata group, Ps. (Gra.) jamaicensis and Chagasia sp. Species

associated with albimanus in ponds, lakes and marshes are An. (Ano.) crucians, Aede-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 43

omyia (Ady.) squamipennis, Cx. (Mel.) madininensis, Cx. (Mel.) ocossa, Cx. (Cux.)

maracayensis (?), Mansonia (Man. ) dyari, Ma. (Man.) titillans, Ps. (Gra.) pygmaea, Ur.

(Ura.) apicalis, Dixella sp, Corethrella sp 23 and Co. sp 36. In streams and ditches

the following have been collected with albimanus: An. (Nys.) argyritarsis, Cx. (Mel.)

inhibitator and Ps. (Pso.) howardii. Found rarely with albimanus in crab holes have

been Cx. (Cux.) habilitator, Cx. (Cux.) janitor, Deinocerites pseudes and De. cancer.

An. albimanus is primarily restricted to the humid lowlands of Middle America,

being most numerous up to an altitude of about 400 m, but occasionally it is collec-

ted at higher elevations. Molley (in Hoffmann 1932:528) collected albimanus near

Guatemala City, Guatemala at 1372 m, and Clark in the same paper claims to have

collected albimanus at 915 m in Costa Rica and 457 m in Haiti. The highest record-

ed altitude I have found was in Morelia, Mexico at 1941 m (Vargas and Martinez Pa-

lacios 1950:5).

The seasonal distribution of albimanus seems to vary from one locality to the next.

Rachou, Lyons et al. (1965:37) stated that in El Salvador, in 6 of the 8 localities

studied, albimanus was most abundant during the wet season and, in 2 of the 8, was

most numerous during the dry season. The Central America Malaria Research Station

(1969, 1970) and Breeland (1972b) reported that in some areas along the Pacific

Coast in El Salvador, albimanus is predominant during the dry season because of the

abundance of estuaries, rivers and permanent bodies of water along streams that get

flushed out during the rainy season. However, in other areas albimanus is most abun-

dant during the rainy season, in habitats such as ditches, flooded pastures and marsh-

es (Breeland 1972a; Kumm and Zuniga 1944; Vargas and Martinez Palacios 1953;

Paul and Bellerive 1947:63-66; Elliott 1968:242).

Most reports indicate that the adults of albimanus are most active from dusk to

midnight (Breeland 1972a), sometimes beginning a little later, as in Colombia at

2200-2400 hours (Elliott 1972:757) and occasionally earlier, 1730-2100 hours in

Haiti (Taylor 1966). Daytime adult resting sites include almost any protected area

such as large rocks in wooded areas (Breeland 1972c; Central America Malaria Re-

search Station 1970). Wilton and Fetzer (1972) showed human breath stimulated

mating, which was at a maximum one hour after sunset in the laboratory, agreeing

with the time Rozeboom (1941) observed in nature.

Taylor (1966) reports that in Haiti 75-92% of the biting activity was outdoors. El-

hott (1968) also found that biting is greater outdoors than indoors. However, several

workers report that in some areas, albimanus is the predominant mosquito biting in-

doors. Kumm, Komp and Ruiz (1940:388-389) stated that 96.4% of the anophelines

captured in houses in Costa Rica were albimanus and 93.3% of those collected in sta-

bles. Kumm and Zuniga (1942:400) reported that in El Salvador, 87.6% of the house

captures were albimanus. The conclusion of the 1970 report of the Central America

Malaria Research Station was that albimanus may be more zoophilic than anthropo-

philic and more exophilic than endophilic but because of the ““overwhelming’’ num-

bers becomes a serious problem. Host preference tests generally indicate that albima-

nus feeds principally on man and domestic animals, such as horses, cows, pigs, dogs,

goats and chickens, and on nondomestic animals such as raccoons. It also prefers

larger hosts to smaller (Breeland 1972a; Central America Malaria Reaearch Station

1970; Taylor 1966; Hill 1934; Thompson 1950; Weathersbee 1944).

An. albimanus is capable of flying considerable distances. LePrince and Orenstein

(1916:114) released adults marked with an aqueous solution of an aniline dye, and

recaptured individuals 366 to 1905 m from the release site. Hobbs, Lowe and

Schreck (1974) reported release-recapture studies using fluorescent powders during

44 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

the dry season in El Salvador; 85,750 adults were released of which 924 were recap-

tured up to 11 days after release. The maximum distance the mosquitoes traveled

was 3 km from the release site. The females were recaptured an average of 548 m

from the release site and the males an average of 460 m. Lowe, Schreck et al. (1975)

performed the same experiment at the same locality during the wet season, finding

that the adults survived up to 14 days and were captured an average of 1000 m from

_ their release sites.

MEDICAL IMPORTANCE. Kumm (1941a) stated that in 17 of 20 countries in

the Caribbean region albimanus is the principal vector of malaria. Throughout its

range albimanus is a very important vector of malaria except on the islands of Guade-

loupe and Marie Galante in the Lesser Antilles. In some areas such as El Salvador it

may be the only vector (Breeland 1972a). As early as 1909 Darling experimentally

infected albimanus. He fed 100 mosquitoes on patients infected with Plasmodium

vivax or P. falciparum and found 70.8% of the females became infected. Rozeboom

(1938b: 291-292) in one experiment infected 87.5% of the females of albimanus

feeding on a patient with P. falciparum. For all Rozeboom’s experiments in which a

total of 113 albimanus fed on infected patients, 32.7% became experimentally in-

fected.

Hill (1928:355) was the first to report finding albimanus naturally infected. Since

then albimanus. has been reported to be naturally infected in virtually every country

in which it occurs. In general, natural infection rates have been reported to be ex-

ceptionally low. At the Central America Malaria Research Station (1970) a total of

434 salivary glands and 241 midguts of different females were dissected and none

was infected. Likewise, Rozeboom (1938b:297) reported a natural infection rate of

1.1% in Santa Rosa, Panama. Because of the low infection rate, albimanus is consid-

ered by many to have the attributes of a relatively poor malaria vector. Nevertheless,

Hobbs, Lowe and Schreck (1974) stated that even during the dry season albimanus is

capable of surviving 11 days, a time long enough to become infective. The reason

that albimanus is such an effective and successful vector is probably partly dependent

upon its longevity and most importantly “because of its close contact with human

populations and its relatively high population level...” (Belkin, Heinemann and

Page(l9/0: 26). ) |

Warren, Richardson and Collins (1975) described 4 inbred morphological pupal

phenotypes (pupal pleiomorphism) of albimanus that had been selected from a strain

“established in September-October 1971 from female mosquitoes collected in stables

near Lake Apastepeque, El Salvador.” Interestingly, Warren, Collins et al. (1977) re-

ported that the females of 3 of these phenotypes show considerable variation in their

susceptibility to infection by Plasmodium sp. Their results indicate that all the phen-

otypes are much less susceptible to infection with P. falciparum than with P. vivax.

The relative susceptibility of the different variants to infection with P. vivax was as

follows: the green nonstriped phenotype was the most refractory; the brown, non-

striped phenotype intermediate; and the brown striped the most receptive.

A possible arbovirus in the Bunyamwera Group has been isolated from albimanus

in Tlacotalpan, Veracruz, Mexico (Scherer, Campillo-Sainz et al. 1967). The Tlaco-

talpan virus was recovered in 1961 from 160 specimens collected at a pig baited,

modified Magoon trap. The native population, cattle and pigs at several localities

along the southeastern coast in the states of Veracruz and Tabasco, and one cow

from the western central coast in the state of Nayarit showed antibodies to the virus.

Recently, Bautista-Garfias, Mercado-Sanchez and Morilla-Gonzalez (1977) experi-

mentally infected females of albimanus with Venezuelan Encephalitis virus TC-83

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 45

by permitting them to feed on infected suckling mice. In a second experiment, 10,

11, 12 and 13 days after exposure to the virus, the females were again allowed to

feed on mice. On their second feeding, the mosquitoes that had had an infected

blood meal 10 days earlier transmitted the virus to one of the 6 mice to which they

were exposed. The infected mouse died on the fifth day after exposure, and the vi-

rus was then “isolated and neutralized with a specific VEE antiserum.”

DISTRIBUTION (fig. 1). An. albimanus is predominantly a tropical lowland spe-

cies most abundant at elevations less than 400 m and occuring in greatest abundance

in coastal plains or along waterways running to the coast. An. albimanus occurs as

far north as the Brownsville area in Texas on the Gulf of Mexico. It extends south

through Mexico and Central America and east along the Caribbean coast of Colom-

bia to the Paria Peninsula in Venezuela. On the Pacific side it extends from Baja

California Sur, Mexico to northern Peru (Piura). An. albimanus occurs commonly

throughout the Greater Antilles, the Bahamas, the Florida Keys, the Cayman Islands

and the Leeward Islands in the Lesser Antilles as far south as Montserrat and Antigua;

it is found only occasionally on Guadeloupe (Basse Terre) and Marie Galante. An.

albimanus does not occur in the Windward Islands of the Lesser Antilles, Trinidad,

Tobago or farther southeast than the Paria Peninsula in Venezuela. It was reported

to have occured in the past on Barbados (Charles and Senevet 1953; Rozeboom

1953), but it has not been found there recently. :

Material Examined: 5467 specimens: 1026 males, 142 male genitalia, 2566 fe-

males, 624 pupae, 1109 larvae; 516 individual rearings: 228 larval, 214 pupal, 74 in-

complete.

ANTIGUA (258 specimens: 32M, 10Mgen, 33F, 57P&p, 126L&1; 44 ind rear:22 1, 19p, 3inc).

St. George: Emersons Pond, ANT 34. Fitches Creek, ANT 66, 67. St. John: Golden Grove Mill,

ANT 32, 57. Potters, ANT 52,53, 79. St. Mary: Bolands, ANT 61. St. Paul: Liberta, 8 Sep 1938,

H. Box, 1M [BM1939-219] ; same locality, ANT 89, 91. St. Peter: All Saints, ANT 36. St. Philip:

Freetown, ANT 69, 70, 72, 73, 75, 76. Parish and locality not specified: 11 Apr 1900, W. Forrest,

1F on slide [BM] ; 27 Nov 1900, same collector, 1M; 2 Dec 1900, same collector, 2F.

BAHAMA ISLANDS. Eleuthera (Island) (14F): Governor’s Harbour, 31 Mar 1953, L. Giovan-

noli, E. Hayden, 3F. James Cistern, 1 Apr 1953, L. Giovannoli, E. Hayden, 11F. New Providence

(Island) (55 specimens: 2M, 3F, 12P&p, 38L&l; 8 ind rear: 5p, 3inc): Harold Pond, BAH 46. Lake

Cunningham, BAH 3. Yamacraw Beach, BAH 18. San Salvador (Island) (3F): Cockburn Town, 18

Mar 1953, L. Giovannoli, 2F. Locality not specified, 4 Sep 1923, P. Bartson, 1F.

BARBADOS (1M, 3F). Locality not specified, received 24 Apr 1928, 1M, 3F [JH].

BARBUDA (2L). Low Pond, 5 Jul 1955, Hummelinck, 2L.

BELIZE (170 specimens: 17M, 2Mgen, 150F, 11). Belize: Belize, BH A164; same locality, KO

115A-13, 14, 1M, 20F. Freetown, BH A186. Ladyville, BH 362. Cayo: Cayo, BH L37. Central

Farm, BH L20. Corozal: Locality not specified, KO 115A-19, 5M, 30F. Stann Creek: Stann

Creek, 30 Jul 1941, W. Kumm, 11F; same locality, KO 115A-17, 6M, 45F; same locality, KO 115A-

23, 1M, 12F; same locality, KO 115A-24, 1M, 16F; same locality, 1F; same locality, 1M. Toledo:

Punta Gorda, 9F; same locality, KO 115A-12, 1F. District and locality not specified: 1F.

CAYMAN ISLANDS. Grand Cayman (Island) (47 specimens: 3M, 2Mgen, 37F, 1P, 4L): North

Sound Estate, CAY 119. West Bay, CAY 2, 10B. Locality not specified, CAY 11A.

COLOMBIA (30 specimens: 24M, 5Megen, 1F). Atlantico: Calamar, 1966, J. Bequaert, 1F

[MCZ]. Cordoba: Monteria, COL 421 , 422.

COSTA RICA (179 specimens: 33M, 14Mgen, 100F, 19p, 13L&l; 16 ind rear: 71, 9p). Cartago:

Atirro, CR 590. Concepcion, 26 Oct 1920[?], A. Alfaro, SF. Turrialba, CR 394; same locality, 6

7, 19 Sep 1920, A. Alfaro, 7F. Guanacaste: Liberia, 27 Jul 1962, F. Truxal, 1F; same locality, H.

Kumm, 1Mgen [CU]. Palo Verde, CR 438. Heredia: Bolson, Jan 1921, A. Alfaro, 7F. Limon:

Bananito, CRR 53, 1F. Cahuita, H. Kumm, no. 183, KO H-18-9, 1M. Finca La Lola, CR 160. Li-

mon, F. Knab, no. 3151a, 1F; same locality, 1F (A-20) [CU] ; same locality, 1F. Matina, 16 Jul

1925, CRR 20, 3M, 8F; same data, CRR 21, 4F; same data, CRR 22, 6F; same data, CRR 25, 3F;

46 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

same data, CRR 26, 4F; same data, CRR, 27, 3F; same data, CRR 28, 2F; same locality, 3 Sep

1925, CRR 37, 1F. Pueblo Nuevo, CR 546. Puerto Viejo, 12, 15,16 Jan 1921, A. Alfaro, 4F.

Westfalia, CR 473. Zent, CR 543; same locality, 15 Jul 1925, CRR 16, 1F; same data, CRR 17,

2F; same locality, 2 Sep 1925, CRR 36, 1F. Puntarenas: Boca del Rio Barranca, CR 1, 29. Do-

minical, 28 May 1943, T. Aitken, CRM 19, 1F; same locality and collector, 1943, CRM 25, 4F;

same data, CRM 26, 2M, 5F. Nicoya, H. Kumm, KO H-18-5,5M. Puntarenas, CR 117. Tarcoles,

CR 569. Province not specified: Bamos, 1F. Barranca, H. Kumm, no. 82, 1M [BM1938-696] ;

same locality, 1F (82A) [CU] ; same locality, 1F (A-17) [CU]. Las Canas, 2F (A-5). Margarita, 21

Jan 1930, W. Komp, 1M (var. trisignatus). San Antonio, 20 Jul 1925, CRR 31, 1F; same locality,

27 Aug 1925, CRR 35, 2F. Santa Cruz, 10 Dec 1937, 1F. Locality not specified, 1938, H. Kumm,

CRR 132, 2M, 2Mgen.

CUBA (81 specimens: 37M, 3Mgen, 41F). La Habana: Habana, Oct 1900, Carroll, 1F; same

locality, 14 May 1902, Taylor, 1M; same locality, Cornell U. Expedition, 1M (1027-693) [CU].

San Antonio de los Banos, J. Pazos, 1F (201). Oriente: Banes, 1F (22) [MCZ] ; same locality, 1M,

3F (24) [MCZ] ; same locality, 1M (25) [MCZ] ; same locality, 1M, 1F (26) [MCZ]; same locality,

2F (28) [MCZ] ; same locality, 2M, 4F (32) [MCZ] ; same locality, 1F (84) [MCZ] ; same locality,

2F (36) [MCZ]. Guantanamo, U. S. Naval Station, 27 Oct 1953, K. Knight, coll. 449, 6M, 5F;

same data except 28 Oct, coll. 454, 1M. Guaro, 3M, 1F (6) [MCZ]. Levisa, near Nicaro, 9 Jul

1953, K. Knight, coll. 381, 1M. Preston, 2M (13) [MCZ]. Province not specified: Soledad, 12

Apr 1926,.2M, 4F (45) [MCZ] ; same locality, 12 Dec 1926, 1F (45) [MCZ]; same locality, 1926,

1M (43), 2F (45) [MCZ]. Locality not specified, Sep 1934, 8M; J. Pazos, 12F; K. Knight, coll.

447, 1M; CUB 32. 3

DOMINICAN REPUBLIC (262 specimens: 29M, 7Mgen, 131F, 41P&p, 54L&l; 37 ind rear: 171,

18p, 2inc). Barahona: La Haya, Jan 1928, 2F. Distrito Nacional: Arroyo Hondo, RDO 4, 6.

Finca Engombe, RDO 28, 39, 69, 148, 159, 200, 256, 257, 267. Rio Haina, RDO 150, 204. Nai-

boa, RDO 162. El Seibo: Rio Magua, RDO 189. La Vega: Jima Abajo, RDO 5. Rincon, RDO

11. Samana: Santa Barbara de Samana, RDO 44. Sanchez Ramirez: La Mata, RDO 16. San Cris-

tobal: Boruga, Canada, RDO 174. La Toma, RDO 23. San Cristobal, RDO 144. San Francisco

mines, RDO 181; same locality, A. Busck, Sep 1905, 1F. Villa Altagracia, 10 Jun 1941, 6M, OF.

San Pedro de Macoris: San Pedro [de] Macoris, 31 Mar 1928, P. Ricart, 1M. Valverde: Arroyo de

Aqua, RDO 232.

ECUADOR (105 specimens: 12M, 1 Mgen, 62F, 2p, 28L&1; 2 ind rear: 11, 1p). El Oro: Locality

not specified, 1943, E. Hopkins, ECUK 43, 1F. Guayas: Chongon, ECU 132. Guayaquil, Jan

1939?, H. Hanson, KO 155-28, 1F; same locality, Apr 1940, J. Murdock, 5M, 10F; same locality

and collector, KO 115-23, 2M, 2F; same locality, Aug 1943, KO 111-4, 1F; same locality, 4 Mar

1964, R. Schuster, GAL 16, 14F; same locality, F. Campos R., 2M, 13F; same locality, ECU 102,

177. El Triunfo, ECU 2. Los Rios: Pichilingue, 16 Mar 1946, E. Hambleton, 8F. Province not

specified: Rio Milagro, Oct 1943, R. Levi-Castillo, KO 115A-4, 1M, 1F. Locality not specified, F.

Campos R., 1F; R. Levi-Castillo, 6F.

EL SALVADOR (39 specimens: 11M, 7F, 9p, 12L&1; 91 ind rear). Cuscatlan: Locality not spe-

cified, Aug-Nov 1940, F. Figueroa, 1M, 4F. La Libertad: Congrejera, SAL 56. Hacienda Melara,

SAL 55. San Salvador: [lopango, 1M, 1F. San Vicente: Laguna de Apastepegue, SAL 54. Usulu-

tan: Espiritu Santo [Island], 602, 2M.

GUADELOUPE. Basse Terre (1F,3L): Jarry, FWI 1030. Viard, FWI 943. Marie Galante (Is-

land) (1M, 1Mgen, 7L): Marie Galante Airport, FWI 921. St. Louis, FWI 264. Locality not speci-

fied, LAR 16.

GUATEMALA (107 specimens: 12M, 6Mgen, 25F, 25P&p, 39L&l; 23 ind rear: 9 1, 11p, 3inc).

Escuintla: Amatitlan, GUA 45, 46, 49. Puerto de San Jose, GUA 35. Izabal: El Cedro, GUA 95.

Entre Rios, GUA 104. Finca Yuma, GUA 92. Morales, GUA 75. Pt. Barrios, 18 Aug 1903, W.

Stone, 1F. Peten: Pueblo Nuevo, Nov 1941, 8F. [Laguna] San Juan Acul, Apr 1942, 2F. Retal-

huleu: Champerico, GUA 21, 126. Zacapa: Teculutan, GUA 14. Department not specified: Es-

quipulas, 1939, 1M, 3F (var. bisignatus).

HAITI (252 specimens: 57M, 2Mgen, 117F, 37P&p, 39L&l; 19 ind rear: 81, 9p, 2inc). Artibo-

nite: Gonaives, HAR 12. Nord: Limbe highway, HAR 42. Nordouest: La Pointe, HAT 118.

Ouest: Arcahaie, HAT 9, 11. Carrefour, HAT 3, 7, 8. Petit Goave, HAR 33, 35. Port au Prince,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 47

HAT 2, 109, 110, 132, 134; same locality, 30 Mar, E. Petersen, 3M, 1F; same locality and collector,

Feb 1924, 1M, 1F. Thor, HAT 105. Sud: Godet, HAT 12. Department not specified: Mar Franc,

9 Jul 1941, Jobbins, 3F. P. Margot Rd., 25 Jun 1941, Jobbins, 7F. Locality not specified, Nov

1928, S. Cook, 1M, 3F; 29 Oct 1929, ?S. Cook, HAR 8; Oct 1929, S. Cook, HAC 2; 1 Jan 1930,

2S. Cook, HAR 11; 27 Jun 1941, Jobbins, 4M; no data, HAR 43.

HONDURAS (111 specimens: 11M, 1Mgen, 74F, 13p, 12L&l; 13 ind rear: 81, 4p, linc). Atlan-

tida: Humazones, HON 59. Ceiba, 28 Jul 1945, 2M, 1F. Lancetilla, HON 29, 36. Las Brisas,

HON 60. Colon: Puerto Castilla, HON 97, 99, 100; same locality, K. Maxwell, 3M, 3F. Truxillo

[Trujillo] , 28 Nov 1942, J. Duncan, KO 115A-11, 48F; same data, 5F. Cortes: Finca Chasnigua,

HON 6. Puerto Cortes, HON 67. Tegucigalpa: Locality not specified, Jul 1945, KO 115-3, 1M.

Yoro: Progreso, 26 Oct 1940, W. Komp, IF.

JAMAICA (565 specimens: 71M, 10Mgen, 117F, 167P&p, 200L&1; 143 ind rear: 601, 65p, 18

inc). Clarendon: Milk River Bath, JA 868. Kingston and St. Andrew: Ferry, JA 350, 781, 896.

Kingston, various dates 1902-1906, M. Grabham, 9M, 5F. Newstead, JA 962. Temple Hall, JA 46,

47,48. St. Ann: Claremont, JA 742. Delight, JA 757. Runaway Bay, JA 766. St. Catherine:

Caymanas, JA 1, 4, 29, 30, 32, 210, 743, 744, 746, 747. Central Village, JA 759. Congrieve Park,

JA 22. Grange Farm, JA 20. Great Salt Pond Farm, JA 26. Gregory Park, JA 16, 17, 18. Naggo

Head, JA 8, 10, 11, 23. Passage Fort, JA 12. Port Henderson, JA 9. Rio Cobre Dam, JA 819.

Spanish Town, JA 24, 27, 35, 36. St. Elizabeth: Luana, JA 357. Black River, 2F. St. Mary: Cas-

tleton Botanical Gardens, JA 807. Fort Stewart, JA 808. St. Thomas: Albion Estate, JA 803,

873. Amity Hall, JA 58; same locality, 5 Dec 1962, T. Aitken, no. 53, 1F, 1L; same locality, 11

Dec 1962, T. Aitken, no. 136, 1P, 2L, 11. Chiswick, JA 65. Dalvey, JA 91, 148. Duckenfield

Hall, JA 54, 55. Golden Grove, JA 56, 63. Grants Pen, JA 75, 77, 413, 800, 813, 814, 871, 872.

Hampton Court, JA 149. Holland Bay, JA 57. White Bay, JA 121, 169. Winchester, 4 Dec 1962,

T. Aitken, no. 5, 1F; same locality and collector, 5 Dec 1962, no. 25, 3F; same locality and collec-

tor, 6 Dec 1962, no. 44, 1F; same locality and collector, 7 Dec 1962, no. 38, 1F. Winchester

House, JA 67, 205. Locality not specified, Mar 1928, M. Boyd, 2M, 9F. Westmoreland: Crab

Pond Bay, JA 782, 785. Negril, JA 231,935. Parish not specified: Cave Stream, R. Hill, no. 28,

1M, 1F. Locality not specified, 7 Dec 1899, F. Cundall, M. Grabham, 1F [BM]; 8 Feb 1900, M.

Grabham, 1M, 1F [BM]; 19 May 1905, N. Y.S., 1M, 1F; 1910, F. Theobald, 1L (396) [BM] ;

1932, Washburn, 1F [BM1933-6].

MEXICO (212 specimens: 10M, 7Mgen, 160F, 11p, 24L&1; 10 ind rear: 51, 5p). Baja California

Sur: San Jose del Cabo, 23 Jan 1979, A. Bourge, D. Pletsch, 4F. Guerrero: Acapulco Airport, 4

Feb 1975, D. Pletsch, 1F. Jalisco: Puerto Vallarta, MEX 457. Nayarit: San Blas, 25-29 Jun 1956,

W. McDonald, 14F, 2L. Oaxaca: Tehuantepec, MEX 106. Quintana Roo: Cancun, 22 Aug 1975,

D. Pletsch, 1F. Tabasco: Tenosique, MEX 575. Villahermosa, 20 Jan 1942, 3F. Tamaulipas: Gon-

zalez, MEX 203, 204. Padilla, MX 13. Tampico, MEX 205, 206, 207, 208. Veracruz: El Naranjo,

MEX 101. St. Lucrecia, Oct 1911, F. Urich, 1F. Tampico, 3 Sep 1902, Goldberger, 1M, 6F; same

locality, 22 Jan 1921, J. Prince, 36F; same locality and collector, 10 Mar 1921, 1F; same locality

and collector, 17 Dec 1921, 1F. Veracruz,3F. Yucatan: [Isla de] Cozumel, 9 Dec 1942, D. Hall,

4OF. Progreso, 25 Oct 1927, C. Hoffmann, 1F [AMNH]. State not specified: ““Boleta 408,” KO

115A-1, 1M. Intapa, 31 Oct 1973, D. Pletsch, 1F. Locality not specified, MEX 742.

MONTSERRAT (136 specimens: 11M, 6Mgen, 20F, 36P&p, 63L&1; 26 ind rear: 22 1, 1p, 3inc).

St. Anthony: Delvins Village, MNT 81. Elberton Estate, MNT 2,3. Foxs Bay, MNT 9, 10, 11, 39,

41, 43, 84, 88, 93. St. Peter: Happy Hill Village, MNT 15. Old Road Estate, MNT 50. Olveston

Estate, MNT 24, 55. Parish and locality not specified: LAR 14.

NEVIS (145 specimens: 14M, 18F, 31P&p, 82L&1; 23 ind rear: 141, 4p, 5inc). St. Paul Charles-

town: Charlestown, LAR 11. St. Thomas Lowland: Cades Bay, NVS 13. Cotton Ground, NVS

14. Paris Pond, NVS 1, 2,46. Pinneys Estate, NVS 17, 47, 18, Vaughans, NVS 16.

NICARAGUA (215 specimens: 35M, 7Mgen, 115F, 15P&p, 43L&l; 14 ind rear: 81, 6p). Chin-

andega: Corinto, 13 Sep 1943, H. Crowell, KO 115-9, 16F; same locality and collector, 15 May

1945, KO 115-4, 1F; same locality and collector, 13 Sep 1945, KO 115-11, 11F. Granada: Grana-

da, NIR 2, 2F; same locality, 26 Nov 1924, NIR 3, 1F; same locality, NIR 6, 3M, 1Mgen, 6F.

Leon: Las Penconas, NI 22. Pantano El Esparto, NI 19, 29. Puerto Somoza, NI 16, 20, 25, 28.

Simonillo, NI 30. Managua: Lago de Managua, NIR 5, 2M, 1Mgen, 4F. Managua, NIR 17, 3M,

48 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

10F. Tipitapa, 3 Jan 1925, NIR 16, 1M, 1Mgen, 4F. Rivas: Rivas, 21 Nov 1924, NIR 22, 1M, IF.

Zelaya: Bluefields, 13 Aug 1943, P. Woke, KO H-18-14, 1M, 2F; same locality, NI 39; same local-

ity, 9F. Rama, 17 Aug 1943, P. Woke, 2M, 5F. Department not specified: El Recreo, 20 Aug

1943, P. Woke, KO H-18-13, 1M. “Pila Office,’ NIR 14, 5M, 1Mgen, 6F; same locality, NIR 15, 7F.

Locality not specified, NIR 4, 2M, 1Mgen, 1F; NIR 9, 10M, 1Mgen, 4F; NIR 10, 1M, 2F; NIR 13,

3F; NIR 19, 1F; NIR 20, 1M; 22 Nov 1942, 4M, 1F.

PANAMA AND CANAL ZONE (1770 specimens: 394M, 47Mgen, 1012F, 108P&p, 209L&1; 100

ind rear: 26 1, 46p, 28inc). Bocas del Toro: Almirante, PA 669; same locality, Jan 1930, PAX 6,

16M, 19F. Big Creek, PA 652. Milla 2, PA 337. Punta de Pena, PA 238. Canal Zone: Ancon, Jul

1941, PAX 181, 2M, 6F; same locality, 7M,44F [CU]. Chulupi, Rio Chagres, 31 Mar 1944, Pierce,

KO 115A-32, 7M, 27F. Corozal, 3 Jul 1922, J. Shropshire, 1F [MCZ]. Cristobal, 13 Jul 1944,

Sterns, et al, ASM 68-1, 3M, 2F, 10L [CU] ; same data, 49-68-1 [ASM 68-1], 2M, 1Mgen. Culebra,

3 May 1925, D. Baker, 3F [MCZ] ; same locality and collector, 1925, 1M. Empire, 27 Oct 1924, J.

Zetek, 1F [MCZ]. Escobal, PA 1158, 1159. Fort Clayton, 21 Mar 1925, D. Baker, 1F [MCZ];

same locality, Nov 1939, W. Komp, 1M, 3F; same locality, 1944, ASM 66-1, colony, 4M, 1 Fgen,

3L [CU], 1F; same locality, Nov 1939, W. Komp, KO H-18-8, 1M. Fort Kobbe, 27 Oct 1939, PAX

149, 5M; same locality, 6 Feb 1963, F. Padilla, CZ 23, 1L; same locality, 10 Feb 1964, J. Rodri-

quez, CZ 118, 1 lpF, 1L. Fort Thomas, 3F [MCZ]. France Field, 19 Nov 1924, J. Zetek, 1F

[MCZ] ; same locality, 1925, Baker, 1M, 2F. Gamboa, Rio Chagres, 4 Oct 1944, Wood, Middle-

kauff, ASM 195-1, 2L [CU]. Gatun, Jan-Feb 1913, J. Zetek, 4F [AMNH], 92F. Juan Mina, Rio

Chagres, 23 May 1939, PAX 112, 3M, 16F; same locality, PA 1122, 1123, 1125, 1160. Madden

Dam, PA 1137. Mindi, May 1941, KO 115-10, 1M, 2F; same locality, KO 115-17, 5F. Mojo Pollo,

Rio Chagres, 31 Mar 1944, KO 115A-15, 9M, 3F. Pacific side, Feb 1943, KO 115A-25, 1F. Para-

iso, A. Jennings, 1M [CU]. Locality not specified, Feb 1933, D. Curry, 1M, 3F [BM1933-504] ;

Jun 1941, KO 115A-9, 2M, 1F; Jun 1941, KO 115A-30, 3M, 4F; 1941, KO 115-12, 2F; Jan 1943,

KO 115-44, 8M, 9F; Jan 1944, KO 112-10, 1F; 6 Jul 1950, 4F; H. Crowell, KO 115-8, 7M, 3F; KO

115A-29, 1F; KO 115-45, 7M, 19F. Colon: Portobelo, PA 604. Santa Rosa, 28 Jul 1943, G. Fair-

child, KO 115A-26, 3F; same locality, PA 43. Darien: Pucro, PA 631. Los Santos: Cape [Punta]

Mala, 3 Feb 1944, Marucci, Wood, KO 115A-2, 3M, 3F. Panama: Chepo, PA 1064. Isla de San

Jose, 15 Mar 1944, KO 115-27, 2F; same locality, 25 Jul 1944, KO 115A-7, 2F. Juan Diaz, PA

749, 801, 836. Juan Franco, 6 Jul 1941, PAX 177, 1M, 4F. Juan Mina, 6 Aug 1943, G. Fairchild,

KO 115A-28, 23F; same locality and collector, 31 Aug 1943, KO 115A-27, 14F; same locality, date

and collector, 5F; same locality and collector, 3 Sep 1943, KO 115A-18, 25F; same locality and

collector, 5 Oct 1943, KO 115A-10, 18F; same locality, 17 Jul 1949, 11 [JH] ; same locality, 25 Jul

1949, 121 [JH] ; same locality, G. Fairchild, KO 115A-16, 6F. La Chorrera, 4 Jul 1944, 49-36-2

[ASM 36-2], 1M, 1Mgen, 2F; same locality, 6 Nov 1944, ASM 270-1, 6L [CU] ; same locality, 2

Dec 1944, Wood, Griffing, ASM 328-1, 3L [CU] ; same locality and date, ASM 329-1, SL [CU];

same locality, 7 Dec 1944, ASM 334-1, 1Fgen, 11L [CU]; same locality, 7 Dec 1944, 1F; same lo-

cality, 13 Jan 1945, ASM 378-1, 1M [CU] ; same locality, 23 Jan 1945, ASM 382-2, 2M [CU];

same locality, 16 Jul 1945, ASM 655, 1F; same locality and date, 5F; same locality, 27 Nov 1944,

Wood, Griffing, ASM 299-2, 1L [CU]. Las Guacas, PA 13. Pacora, PA 1063. Paitilla, PA 755,

780. Panama City, 15 Sep 1935, PAR 25, 4M, 3Mgen. Panama Viejo, 27 Jul 1935, PAR 18, 1M,

1Mgen, 44F; same locality, 17 Nov 1944, R. Arnett, K. Frick, ASM 283-1, 1L [CU]. Tocumen, PA

547, 1147. Province not specified: Aguadiente, 31 Jan 1935, PAR 75, 74M, 3Mgen. Lagarto, 26

Jul 1945, Van Doran, ASM 680-2, 2F. Rio Chagres, 12 Feb 1943, KO 115A-8, 17M, 18F; same

locality, 25 Jul 1945, Van Doran, ASM 679-1, 1F. Rio Pescado, 7 May 1937, 1M, 2F. Sabanas Rd.,

30 Oct 1934, 1M, 1F. Laboratory colonies, PA 1178, PAR 118A, 120, 123, PAX 60, 61, 62, 63, —

64, 65; colony, 17 Aug 1937, 3M, 2Mgen, 9F; colony, 31 Jul 1937, 1M, 1Mgen, 20F. Locality not

specified, 28 Mar 1935, PAR 78, 1Mgen; 2 Oct 1935, PAR 92, 12M, 3F; 31 Jul 1937, 1M, 14F; no

data, 3F.

PERU (10 specimens: 1M, 1Mgen, 2F, 3p, 3L&1; 3 ind rear: 21, 1p). Piura: Mallares, PER 18.

PUERTO RICO. Culebra Island (2M): Locality not specified, 20 Jan 1954, H. Hurt, 1M (567);

same collector, 21 Jan 1954, 1M (569). Puerto Rico (Island) (416 specimens: 185M, 4Mgen, 151F,

27P&p, 49L&l; 16 ind rear: 51, 9p, 2 inc): Aguirre, 10 May 1920, H. Green, 2M, 2F. Arecibo, Mar

1942, H. Pratt, 2L. Carolina, Mar 1942, W. Hoffman, H. Pratt, 7p. Catano, PRA 31; same locality,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 49

10 Nov 1943, H. Pratt, 1L. Cayey, 2 Dec 1943, J. Maldonado-Capriles, H. Pratt, 5L. Dorado, PR

151. Fort Buchanan, 14 Dec 1942, T. Aitken, 2M, 2F; same locality, 13 Jan 1943, H. Pratt, 1F;

same data except 19 Jan, 2M; same data except 26 Jul, 1Mgen; same locality, PRA 69. Gurabo, PR

123; same locality, H. Pratt, 2L. Juan Diaz, Jul 1944, KO 106-17, 2M. Lares, 28 Apr 1944, H.

Pratt, 1L. Levittown, PR 160, 166. Loiza, KO 115-1, 53F; same locality, KO 115-7, 6F. Loiza

Aldea, 12 Sep 1935, R. Watson, 2F. Losey Field, 18 Nov 1942, A. Pritchard, 7L; same locality,

1944, H. Pratt, KO 115-5, 30M; same locality, KO 115-13, 42M; same locality, KO 115-16, 32M.

Mayaguez, 25 Feb 1936, G. Tulloch, 9M, 14F; same locality, PR 116. Puerto Arturo, 30 Oct 1940,

KO H-18-10, 3M, 2Mgen, 4F. Punta Salinas, 28 Mar 1942, W. Hoffman, H. Pratt, 3P. San Juan,

Jan 1914, 1M, 2F; same locality, 3L. Tortuguero, Camp, 6 Feb 1943, G. Bradley, 3F. Tortuguero,

Laguna, 25 Aug 1942, E. Charneco, 2L; same locality, 15 Dec 1944, H. Pratt, 2L. Yauco, 26 Dec

1942, J. Maldonado-Capriles, 5L. Locality not specified, Feb 1914, W. Wippitt, 2M, 6F; KO 115-2,

40M; H. Pratt, PRX 13, 2M, 1F; no data, 1M, 3F. Vieques Island (21 specimens: 5M, 1Mgen, 7F,

2p, 6L&l; 2 inc ind rear): Locality not specified, 27 Feb 1942, H. Hurlbut, 1 lp (AA 2-27-42-1); 3

Mar 1942, H. Hurlbut, 1 lp (AA 3-3-42-3); 1942, H. Hurlbut, 5M, 1Mgen, 6F, 4L; 10 Jan 1943, H.

Pratt, TF,

SAINT MARTIN (1L). Old Battery Cistern, 3 Jun 1955, Hummelinck, 1L (529b).

UNITED STATES. FLORIDA (24 specimens: 11F, 1p, 12L&l; 1 inc ind rear). Monroe Co.:

Big Pine Key, 26 Mar 1948, J. Haeger, 1 IP (274-101) [CU]. Key West, 5 Sep 1946, Fernandez,

1F; same data except 25 Sep, 1F; same data except 26 Sep, 1F; same data except 17 Oct, 1F; same

locality, C. Gardner, 2F. Marathon Key, 7 Aug 1946, USPHS, 1F. Stock Island, 31 Oct 1946, Fer-

nandez, 1F; same locality, 15 Oct 1959, W. Warner, 1F; same data except 20 Oct, 1F. Vaca Key,

27 Aug 1947, W. Buren, 1F. Locality not specified, “Sta 338,” 7 Dec 1949, J. Haeger, 11L. TEX-

AS (97 specimens: 8M, 4Mgen, 65F, 20L). Cameron Co.: Brownsville, 13 Oct 1923, R. Turner,

13F; same data except 24 Oct, 2F; same locality, 29 Nov 1939, 2F; same locality, reared 14 Dec

1939, 2M; same locality, 6 Aug 1942, W. Gordon, 1M, 1Mgen, 13F [CU]; same data except 10

Aug, 1M, 1Mgen, 5F [CU]; same locality, 13 Sep 1942, E. Garbera, 1F; same locality, 26 Sep 1942,

E. Ross, 1L [ANS]; same locality, 28 Sep 1942, E. Ross, H. Roberts, 3F; same locality, 12 Oct

1942, T. Burns, 5F; same locality, 19 Nov 1942, E. Gerberg, IMgen, 3F [CU]; same locality, 1942,

1M, 1Mgen [CU]; same locality, 5 Jun 1943, W. Gordon, 5F [CU] ; same locality, Jan 1944, C.

Joyce, 1M; same locality, Oct 1944, R. Usinger, 5L; probably same locality, KO 115-25, 12F.

Combes, Oct 1944, R. Usinger, 7L. Harlingen, Jan 1944, Col. Pfrejer, 1L. Locality not specified,

11 Jun 1937, T. McGregor, 1M. Hidalgo Co.: Edinburg, Oct 1944, R. Usinger, 3L. McAllen, 4 Nov

1923, R. Turner, 1F. County not specified: Rancho Viejes, Resaca, Oct 1944, R. Usinger, 2L.

Yturria, Oct 1944, R. Usinger, 1L.

VENEZUELA (111 specimens: 6M, 1Mgen, 85F, 6P&p, 13L&1; 6 ind rear: 41, lp, linc). Anzo-

ategui: Barcelona, 8-15 Aug 1940, VZK 45, 1M; same locality, 25 Sep 1944, VZK 17, 1M, 2F;

same locality, 26 Sep 1944, W. Komp, KO 115A-33, 13F; same data, KO 115-37, 11F; same data,

KO 115-40, 10F; same data, KO 115-43, 18F. Puerto La Cruz, 27 Sep 1944, VZK 24A, 3M, 5F;

same locality, 26 Sep 1944, KO 111-21, 18F; same locality, 25 Sep 1944, VZK 19, 1F. Aragua:

Turiamo, 13 Aug 1944, E. Winton, KO 111-12, 2F. Carabobo: Boca de Yaracuy, VZ 259. La Ca-

brera Peninsula, VZ 93, 94, 95. Puerto Cabello, 12 Jul 1927, F. Root, 1M [BM1929-194]. Zulia:

Independencia, Nov 1949, A. Gabaldon, 3L (1375. 210, Zonet, Lot 98). State and locality not

specified: 14 May 1940, VZK 44, LF.

VIRGIN ISLANDS. St. Croix (Island) (SL): Bethlehem, Nov 1939, H. Beatty, 2L. Caledonia,

VI 12. St. John (Island) (1 lpM): Leinster Bay, VI 6. St. Thomas (Island) (1M, 1F): Locality not

specified, LAR 1A.

50 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

OSWALDOI GROUP

FEMALES. Head: Light to dark. Palpal segments 2 and 3 predominantly dark

with a small to medium white band at apices; erect outstanding scales on segments 1,

2 and basal 0.3-0.5 of 3, becoming less outstanding toward apex of 3; segment 3 with

or without dorsolateral, golden to white speckled, longitudinal stripe; segment 4

with at least some white or cream on mediolateral surface, base and apex usually

with brown band; segment 5 completely white or with small dark basal band. Thor-

ax: Pruinose and scale patterns and color same as in section description. Occasional-

ly upper mep with few light scales. Anterior mep with light scale patch in triannula-

tus only. Legs: Foretarsal segments 2 and 3 basally dark at least on dorsal surface

and light in about apical 0.2-0.9 of segment; segment 4 usually all dark at least on

dorsal surface, occasionally with a few golden to cream, apical scales or with a broad-

er apical light area (triannulatus), rarely predominantly light (some ininii); segment 5

usually dark in about basal 0.5 and light in apical 0.5, but varying from predominant-

ly dark (triannulatus) to predominantly light (some strodei and ininii). Midtarsal seg-

ments 1, 2 and 3 with a small to moderate, apical, light area, basally predominantly

dark at least on dorsal surface; segment 4 all dark at least on dorsal surface, except in

ininii; segment 5 with apical 0.3-0.5 golden to white, rarely completely cream (some

ininii), usually dark at least on dorsal surface in basal 0.5-0.7. Hindtarsal segment 2

with basal 0.10-0.85 dark, apical remainder white; segment 3 usually all white, rarely

dark at least on dorsal surface in basal 0.3 (0.20-0.35) of segment (rondoni and mu-

tants); segment 4 light, very rarely basally dark (mutants). Dark basal areas and light

apical areas of foretarsal segments 2-5, light apical areas of midtarsal segments 1-5

and dark basal area of midtarsal segment 5 appearing as complete or incomplete, light

and dark bands or rings; light bands almost always complete (entirely encircling seg-

ment); dark basal bands usually appearing complete or almost complete with light

scales usually on ventral surface. Hindtarsal segment 2 with dark basal area appear-

ing as a dark basal band; segment 5 with basal dark and apical light areas appearing as

complete bands. Wing: Wing spots as in section description, highly variable and im-

portant in species diagnoses. Abdomen: Caudolateral scale tufts present on tergites

and sternites II-VII. Sternite I usually with a few, moderately long to long setae and

occasionally some inconspicuous obovate to lanceolate, light scales in basal half.

MALES. Essentially as in females except for sexual characters. Head: Palpal seg-

ment 4 with a subbasal and a subapical dark band, ventral surface usually with some

dark scales, rest of median surface predominantly light. Antenna: 0.75-0.85 length

of proboscis. Legs: Forefemur 0.55-0.70 length of proboscis. Basal plantar surface

of foretarsal segment 5 with short to long, spiniform setae.

MALE GENITALIA. Segment IX. Sternite moderately long to long, with a short

to long, variously shaped, nonspiculose apodeme along anterior border. Sidepiece,

Clasper: As in section description. Claspettes: Ventral claspette with setae at least

on basal lobules in Oswaldoi Subgroup, or without setae and apicolaterally produced

into auriculate lobes in the Triannulatus Subgroup. Preapical plate and apex of ven-

tral claspette variously developed. Phallosome: Triannulatus Subgroup without leaf-

lets; Oswaldoi Subgroup with or without leaflets.

LARVAE. Moderate to large, rarely small (triannulatus). Head: Moderately to

heavily sclerotized. Median tooth of mental plate narrow to broad. Hairs 2-C widely

spaced except in anomalophyllus and strodei. Hair 3-C equal to or shorter than 2-C.

Hair 4-C usually short and branched, except in nuneztovari and trinkae. Hairs 8,9-C

dendritic or weakly plumose, moderately small to large. Hairs 10,12,13-C large in

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 51

triannulatus and ininii, small to moderate in remaining species. Hair 15-C moderately

large to large, single to numerously branched. Collar narrow to wide, heavily sclero-

tized. Antenna: Weakly to heavily pigmented. Spicules on mesal margin short and

thin to long and stout. Hair 1-A small to moderately large or occasionally large (in-

inii, benarrochi). Thorax: Moderately to heavily pigmented. Submedian prothorac-

ic group (1-3-P) with or without 1,2-P sharing a common tubercle; 1-P palmate. Hairs

9,10,12-P single and long; 12-P longer than 9,10-P; 9-P subequal to 10-P; 11-P double

or rarely triple, branched about 0.25 from base, about 0.5 or less length of 12-P.

Hair 14-P with a moderately short to long, flattened shaft. Mesothoracic hair 1-M us-

ually ovoid to elliptical in outline, with elongate flattened shaft. Metathoracic hair 2-

T moderately long to long. Hair 3-T palmate, transparent to weakly pigmented,

brushlike or with spreading lanceolate leaflets. Hairs 9,10-T single and long, 9-T

shorter than 10-T. Abdomen: Hair 0-II-VII with few to numerous branches, usually

more than 3 except in some ¢trinkae. Palmate hair 1-I unpigmented to weakly pig-

mented, leaflets narrow to moderately wide; 1-II-VII usually strongly sclerotized,

with long, narrow to broad, lanceolate leaflets. Hair 2-II highly variable, 3-10

branched. Hair 5-I small to large, 2-12 branched. Hair 11-I moderately large to large.

Hair 13-I more than 3 branched (4-9), except in triannulatus and benarrochi. Hair

13-II, III small to moderately large, numerously branched (except 13-II in triannula-

tus); 13-IV small to moderate, never as large as in albimanus; 13-V moderately large

to very large. Spiracular Lobe: Pecten highly variable. Lateral arms of spiracular ap-

paratus varying from very short and directed caudolaterally to long and directed later-

ally. Anal Segment: Covered with spicules. Saddle reddish brown. Hair 1-X short

to long, inserted on or not inserted on saddle. Anal gills short to long.

DISCUSSION. The species comprising the Oswaldoi Group are distinguished from

that in the Albimanus Group in the females by (1) palpal segment 4 with at least

some white or cream on mediolateral surface, (2) presence of dark caudolateral scale

tufts on abdominal segment II and (3) foretarsal segment 5 usually with at least an

apical light band to occasionally completely light, except in some strodei and triannu-

latus where it is all dark; in the male genitalia by (1) sternite [X moderately long to

long, subtrapezoidal or subtriangular, (2) ventral claspette with setae, or if without

setae then apex of claspette expanded laterally into large auriculate lobes and (3)

ventral surface of ventral claspette without bare, inflated, ovoid, bulbous lobes; in the

larvae by (1) hair 4-C usually short except in trinkae and nuneztovari where it is long-

er, (2) hair 1-P palmate, (3) hair 9-P,T single and (4) hair 13-I,IJI,1V small to moder-

ately large, or if large then hair 11-I large and 5-7 branched.

I am separating the Oswaldoi Group into 2 distinct subgroups, the monotypic Tri-

annulatus Subgroup and the Oswaldoi Subgroup composed of 12 species (fig. 4). Al-

most all the characters separating the Oswaldoi Group from the Albimanus Group are

derived, with the exception of those mentioned on page 14. An. triannulatus is the

least derived species in the group, retaining many characters that are intermediate be-

tween the Albimanus Group and the Oswaldoi Subgroup. For example, in the fe-

male of triannulatus, (1) palpal segment 4 is usually darker than in the Oswaldoi Sub-

group, but not as dark as in albimanus and (2) foretarsal segment 5 is predominantly

dark; and in the larva, (1) hair 1-P is not plumose as in albimanus, but has numerous

and very slender, almost filiform branches and (2) hair 13-I,III,IV is relatively large

and few branched, as in albimanus.

The Oswaldoi Subgroup is a collection of very closely allied species. Their external

morphology is interspecifically very similar and usually highly variable intraspecifical-

ly. For these reasons the keys are not always reliable, particularly in some cases for

52 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

the adult females and pupae. In most instances, the adult females of aquasalis, rang-

eli, rondoni, oswaldoi, ininii and nuneztovari are easily identified. An. aquasalis can

be confused with benarrochi; however, since the distributions of the 2 species are so

different, this usually is not a problem. An. trinkae is difficult to identify because

the humeral light spot on the costal vein is not always less than twice the length of

the basal dark spot. An. oswaldoi may be confused with strodei, ininii and occasion-

ally noroestensis, so it is necessary to refer to the secondary characters listed in the

key. An. anomalophyllus is very similar to strodei, but it is rare and encountered on-

ly in Panama and Costa Rica. Again it is reemphasized that reference should be made

to the individual species description, discussion, bionomics and distribution sections

when attempting to identify any specimens in this group.

The Oswaldoi Group is more or less restricted to tropical Central and South Amer-

ica, and is predominantly of South American origin. Only the relict species, anomalo-

phyllus, appears to have evolved in Central America. The other species occurring in

Central America, aquasalis, oswaldoi, triannulatus and strodei, probably have invaded

this area in recent times. An. strodei extends the farthest north, having been collec-

ted in the state of Veracruz, Mexico (Vargas and Martinez Palacios 1956:142). On

the Pacific side of the Andes, northern Peru is the southernmost limit of the group.

Only one species, aquasalis, has invaded the Lesser Antilles, extending as far north as

the island of Antigua. An. aquasalis is capable of breeding in brackish water which

may explain its success in colonizing these Caribbean islands. East of the Andes the

group extends as far south as the northern provinces of Argentina.

OSWALDOI SUBGROUP

FEMALES. Moderately large to large. Head: Palpal segment 4 predominantly

white, usually with a basal and an apical dark band, and often with dark scales of

varying amounts on ventral surface. Thorax: Integument light brown to almost

black. Median anterior promontory area with long white scales often extending dor-

sad onto anterior of acrostichal line. Anterior mep without scales. Legs: Foretarsal

segments 1-4 with varying amounts of white; segment 5 usually with apical 0.5 or

more golden to white, except in some strodei. Midtarsal segment 3 with a small, api-

cal, cream band. Wing: Variable. Vein C humeral light spot usually large except in

most specimens of nuneztovari and trinkae.

MALES. Essentially the same as in female except for sexual characters. Head:

Palpal segment 4 with a basal and an apical light band and with or without a subbasal

and a subapical dark band, mediolaterally extensively cream to white, ventral surface

dark. Thorax: Anterior mep without a scale patch.

MALE GENITALIA. Segment IX: Sternite moderately long to long, subtriangu-

lar or subtrapezoidal, with short to long, variously shaped, nonspiculose apodeme

along anterior border. Sidepiece, Clasper: As in group description. Claspettes: Dor-

sal claspette as in species descriptions. Ventral claspette never bare, with short to

long setae on basal lobules, or on basal lobules and on lateral and ventral surface of

fused portion of lobe, setae extending to or near to apex. Preapical plate variable in

shape and pigmentation. Phallosome: Aedeagus with or without leaflets; apex less

than 1.5 as long as wide.

LARVAE. Moderate to large. Head: Hairs 2,3-C single and simple, barbed or plu-

mose with short to long branches in about apical half; hairs 2-C narrowly or widely

spaced. Antenna: Hair 1-A short to moderately long, except in benarrochi and ininii.

Thorax: Submedian prothoracic group (1-3-P) with hair 1-P with fewer than 16 (ex-

cept in some strodei and ininii) narrow to broad, lanceolate leaflets; 1,2-P with or

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 53

without common tubercle. Hair 14-P with a short to moderately long, flattened

shaft. Metathoracic hair 3-T with semitransparent or weakly pigmented leaflets. Ab-

domen: Palmate hairs narrow to broad. Hair 11-I moderately large, 2-4 branched.

Hair 13-I small to moderately large, usually more than 3 branched; 13-III usually

small and numerously (5-14) branched; 13-IV small to moderate. Sclerotized tergal

plate VII equal to or less than about 1.7 size of tergal plate VI. Spiracular Lobe:

Pecten variable. Lateral arms of spiracular apparatus very short to moderately long,

except long in ininii. Anal Segment: Covered with spicules. Hair 1-X moderately

long to long, inserted on or not inserted on saddle. Anal gills short to long.

DISCUSSION. Members of the Oswaldoi Subgroup can be distinguished from the

Triannulatus Subgroup in the female by (1) palpal segment 4 more predominantly

white, (2) absence of anterior mesepimeral scales, (3) absence of a large apical light

band on foretarsal segment 4, except occasionally in nuneztovari and ininii and (4)

foretarsal segment 5 half to completely light, except in some strodei; in the male gen-

italia by the presence of setae at least on the basal lobules of the ventral claspette;

and in the larvae by (1) palmate hair 1-P usually with fewer than 16 narrow to broad

leaflets, (2) hair 14-P with a short to moderate shaft, (3) hair 1 1-I moderately large,

2-4 branched, (4) hair 13-I small to moderately large, usually more than 3 branched,

(5) hair 13-III usually small and numerously branched and (6) lateral arms of spiracu-

lar apparatus short to moderately long, except in ininii.

The relationships of the species within the Oswaldoi Subgroup is very difficult to

determine because of the tremendous number of overlapping characteristics of the

species in each of the different stages. The derived features of the male genitalia serve

to link together the taxonomically important characters in the other life stages. Up-

on examination of the ventral claspette of the male genitalia, 2 distinct phyletic lines

become apparent, which I am calling the Oswaldoi Complex and the Strodei Complex

(fig. 4). Of these 2 complexes, the Strodei Complex is more closely akin to the Tri-

annulatus Subgroup than is the Oswaldoi Complex. This was also evident to Gabal-

don (1940:4), when he placed strodei and later (Gabaldon and Cova Garcia 1952:

178) rondoni and benarrochi in the same group as triannulatus in the Subseries Tri-

annulatus, based on the morphology of the male genitalia. Other characters linking

triannulatus to the Strodei Complex are the slender collar and numerous slender

branches of hair 1-P in the larva, and the shape of the trumpet and the caudolateral

spines in the pupa. The relationship of benarrochi to triannulatus, and to rondoni

and strodei is not clear. However, the male genitalia of benarrochi are more like stro-

dei than any other species in the Oswaldoi Subgroup. The Oswaldoi Complex is the

more derived complex in the Oswaldoi Subgroup.

OSWALDOI COMPLEX

FEMALES. No diagnostic characters separating this complex from the Strodei

Complex in the adult females are apparent.

MALES. Essentially as in females except for sexual characters.

MALE GENITALIA. Segment IX: Sternite long, about 0.5 length of phallosome,

approximating shape of an isosceles trapezoid or a truncated triangle. Apodeme

across anterior border short to long. Sidepiece: Parabasal spine moderately short to

moderately long. Ventral Claspette: Nonrugose along apical margin, with setae on

basal lobules and on lateral and ventral surface extending to or near to apex, apex

not laterally expanded. Preapical plate moderately small to large, usually distinct.

Phallosome: Aedeagus with or without leaflets, apically rounded or truncate (noro-

estensis).

134 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

LARVAE. Moderately large to large. There are no good diagnostic characters in

the larvae to separate this complexfrom Strodei Complex. Head: Variable branch-

ing of inner and outer clypeal hairs (2,3-C); inner clypeals (2-C) widely spaced, ex-

cept in anomalophyllus. Hair 4-C usually short, except in nuneztovari and

trinkae. Hairs 10,12,13-C large in ininii, small to moderately large in remaining spe-

cies. Collar always moderately wide to wide, heavily sclerotized. Antenna: Hair 1-A

small to moderately large, except in ininii. Hairs 2,3-A relatively large, particularly

in ininii. Thorax: In many but not all species, a large proportion of specimens with

hairs 1,2-P of submedian prothoracic group (1-3-P) sharing a common tubercle, or

base of 1-P not sclerotized and 1-P apparently arising separately from 2-P. Hair 14-P

with short to moderately long, flattened shaft. Abdomen: Hair O-II small to moder-

ately large. Hair 5-I moderately large (except in rangeli) and close to lateral margin

of abdomen. Hair 13-I-III small, numerously branched; 13-IV larger than 13-I-III,

with fewer branches than 13-VI. Spiracular Lobe: Lateral arms of spiracular appara-

tus short, except moderately long in noroestensis and oswaldoi, and long in ininit.

Anal Segment: Hair 1-X inserted on or not inserted on saddle. Anal gills short to

long.

DISCUSSION. In general, the male genitalia and the larvae offer the best charac-

ters for separation of the species in the Oswaldoi Complex. The Oswaldoi Complex

is distinguished from the Strodei Complex in the male genitalia by (1) presence of se-

tae extending to the apex of ventral claspette or to at least as far as level of apical

margin of preapical plate, (2) apex of ventral claspette not laterally expanded and (3)

parabasal spine slightly shorter, usually extending neither to far margin of aedeagus

nor to tubercle of other parabasal spine.

The species in this complex are very closely related. On the basis of the male geni-

talia, there seem to be 2 primary phyletic lines. An. oswaldoi, galvaoi, noroestensis,

aquasalis, ininii and possibly anomalophyllus form one line, and trinkae, nuneztovari

and rangeli compose the other (fig. 4). I have not designated these phyletic lines as

separate taxa due to their close phylogenetic relationship and because it would tend

to excessively complicate the presently proposed hierarchy.

The species of the first phyletic line, aquasalis, noroestensis, galvaoi, oswaldoi, in-

inii and anomalophyllus, are characterized by the tapering, rounded apex of the ven-

tral claspette, with a distinct crescent shaped or oval preapical plate. An. aquasalis

and oswaldoi are fairly widely distributed in Central and South America, aquasalis

primarily along the Atlantic Coast and oswaldoi in the interior. An. noroestensis is

widely distributed from the northeast of Brazil south to the northern states of Argen-

tina, but is not found in northwestern South America or Central America. The latter

3 species are often broadly sympatric, but they usually occupy distinct niches. An.

aquasalis is primarily associated with brackish water, oswaldoi with the forest and

noroestensis with open drier areas. An. galvaoi seems to be primarily restricted to the

high Amazon basin extending south into Mato Grosso and Sao Paulo and possibly

east into Goias and Bahia. It would seem to be sympatric with oswaldoi throughout

most of its range, although it is not clear if the range of galvaoi overlaps with that of

noroestensis, its closest ally. An. anomalophyllus and, to a lesser extent, ininii retain

many primitive characters which clearly distinguish them from the remaining species.

Both species have restricted known distributions. An. anomalophyllus is a rare spe-

cies, to date found only in Costa Rica and northern Panama. An. ininii has been col-

lected only in French Guiana and Para, Brazil. The ventral claspettes of ininii and an-

omalophyllus look very much alike in outline and are, in general, similar to those of

oswaldoi, aquasalis, galvaoi and noroestensis. The aedeagus of both species has leaf-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 55

lets, very strongly developed in anomalophyllus and weakly developed in ininii. On

the basis of the male genitalia, adult and larva, ininii seems to be clearly affiliated

with the first phyletic line. In the case of anomalophyllus, the male genitalia corre-

spond to the type in the Oswaldoi Complex, but the larva and adult look very much

like the strodei type. For this reason I am unsure of the phylogenetic relationship

between anomalophyllus and the rest of the species in the Oswaldoi Subgroup. One

possibility is that anomalophyllus is an annectent relict form, intermediate between

the Oswaldoi Complex and the Strodei Complex.

The other phyletic line in this complex, composed of rangeli, trinkae and nunezto-

vari, is again based primarily on the structure of the ventral claspette of the male

genitalia, correlated with characters in the other stages. The ventral claspette of these

species is broad at the apex and has setae on the ventral and lateral surfaces of the

fused portion of the lobe. The adults of rangeli, trinkae and nuneztovari are exceed-

ingly alike, making species diagnosis difficult. It is possible that the 2 latter species

have not differentiated enough to become reproductively isolated, in which case they

would be considered subspecies rather than species. As will be discussed later, this is

really not so important, since they appear to be allopatric, and differ significantly in

the immature stages and male genitalia. For these reasons, I feel justified in treating

these 2 as separate species. The species in this second phyletic line are restricted to

the Caribbean, Orinoco and Amazon drainage systems, with extensions into Bolivia

and Mato Grosso.

2. Anopheles (Nys.) oswaldoi (Peryassu)

Figs. 2, 4, 6, 10, 11

1922. Cellia oswaldoi Peryassu 1922:179. TYPE: Syntypes males and females, Vale do Rio

Doce (Espirito Santo) and Baixada Fluminense (Rio de Janeiro), Brazil, Mar and Apr

[Museu Nac Rio de Janeiro] (Belkin, Schick and Heinemann 1971:6).

1932. Anopheles (Nyssorhynchus) tarsimaculatus var. aquacaelestis Curry 1932:566-571.

TYPE: Syntypes male with genitalia slide, female and larva, Atlantic side of Canal Zone,

Panama, no type locality cited, although Colon Hospital, 20 Jun 1929, C. H. Bath and

lower Chagres River mentioned [LU]. Synonymy with oswaldoi by Senevet and Abon-

nenc (1938:487-493).

1942. Anopheles (Nyssorhynchus) konderi Galvao and Damasceno 1942:115-118. TYPE: Syn-

types male(s), larva(e), pupa(e), right margin of Solimoes [Amazon] , 300 km from Man-

aus, Coari (Amazonas), Brazil [LU, no material in FMSP, 651; apparently lost] (Belkin,

Schick and Heinemann 1971:6). Synonymy with oswaldoi by Lane (1953:262).

Anopheles (Nyssorhynchus) oswaldoi of Senevet (1934:49-52; 1948a:277-279; 1948c:437); Gal-

vao and Lane (1938:175-177; 1941:10); Rozeboom (1938a:101; 1938b:289-290; 1942:239-

240); Senevet and Abonnenc (1938:493); Lane (1939:25-26; 1944:262-268; 1949:403; 1953:

261-263); Gabaldon (1940:5); Gabaldon and Aguilera (1940:65-66); Gabaldon, Cova Garcia and

Lopez (1940:10, 11); Galvao (1940:426-432; 1943:141, 143, 149, 151); Komp (1941a:93, 95,

97; 1942:26, 39, 40, 43, 74-75, 80, 124-125, 132, 160-161); Rozeboom and Gabaldon (1941:

92-98); Floch and Abonnenc (1942a:2; 1942b:2; 1946b:3-5; 1947:6; 1951:49-50); Fonseca and

Fonseca (1942:93, 94); Galvao and Damasceno (1942:121); Oliveira and Verano (1942:353-

358); Simmons and Aitken (1942:39, 46, 54, 62, 95-96); Cerqueira (1943:19); Galvis (1943:93);

Ramos (1943:51-52, 56-60); Causey, Deane and Deane (1944:2, 4, 5; 1946:27); Correa and Ra-

mos (1944b:130; 1944c:11-12); Deane, L. M., Causey and Deane (1946:7-9; 1948:874-876);

Deane, M. P., Causey and Deane (1946:38, 45); Gabaldon and Cova Garcia (1946b:99-103);

Amaral and Penido (1947:164, 168, 173-178); Arnett (1947:197); Vargas (1948: 158; 1959:377,

383); van der Kuyp (1949a:67-68); Levi-Castillo (1949:9-11, 13, 15, 28, 67, 72, 76, 82, 86);

Bejarano and Duret (1950:150, 153); Downs (1950:29-30); Duret (1950a:371; 1950b:302);

Rey and Renjifo (1950:534, 537); Carvalho and Rachou (1951:475, 480); Rachou and Ferraz

56 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

(1951:542, 547-553); Rachou and Ricciardi (1951:424-426, 432-437); Senior-White (195 1a:

330); Horsfall (1955: 184); Senevet and Andarelli (1955:339); Vargas V. (1956:29; 1957; 1958a;

1958b); Bejarano (1957:326-333, 335-336; 1959:305, 315, 325); Castro (1959a:174); Stone,

Knight and Starcke (1959:33-34); Cerqueira (1961:126); Fauran (1961:11); Forattini (1962:

421); Garcia and Ronderos (1962:137-139, 158-159); Belkin, Schick and Heinemann (1965:44;

1971:61); Morales-Ayala (1971:139); Consolim and Galvao (1973:177); Cova Garcia and Sutil

O. (1976:31; 1977:23, 47, 62, 88); Knight and Stone (1977:63-64).

Anopheles oswaldoi of Kumm (1932:1); Gabaldon, Cova Garcia and Arevalo (1940: 26, 28-32); Ga-

baldon, Ochoa Palacios and Perez Vivas (1940:42-55); Rozeboom (1941:102-103); Vargas

(1941:118); Downs, Gillette and Shannon (1943:29, 30, 33); Russell, Rozeboom and Stone

(1943:26, 30, 37, 42, 48); Floch and Abonnenc (1946a:2); Vargas V. (1958a; 1958c:1, 3);

Foote and Cook (1959:129-130); Cova Garcia (1961:34-35, 69, 86-87, 113, 123-124, 154); Ma-

ciel (1962:477); Vincke and Pant (1962:2-7, 10, 12, 14); Mattos and Xavier (1965:273); Stojan-

ovich, Gorham and Scott (1966a:12, 18, 34; 1966b:21, 29, 43); Gorham, Stojanovich and Scott

(1967:15, 47, 58; 1973:112, 139, 149); Cova Garcia and Sutil O. (1975a:23; 1975b:21 1); Pan-

day (1977:732, 734); Cova Garcia, Pulido F. and Amarista M. (1978:157).

Anopheles (Nyssorhynchus) oswaldoi var. oswaldoi of Galvao and Lane (1937e:213-216; 1938:

175); Galvao (1938a:51, 52, 53-54, 58; 1938b:101-103); Galvao and Amaral (1938: 13); Fonse-

ca and Fonseca (1942:118).

Anopheles oswaldoi oswaldoi of Vargas (1943:59).

Anopheles (Nyssorhynchus) tarsimaculatus var. oswaldoi of Christophers (1924:40, 90); Galvao

and Lane (1937a:77, 78; 1938:173-174).

Anopheles tarsimaculata var. oswaldoi of Root (1926a:51, 89, 107-108).

Nyssorhynchus (Nyssorhynchus) tarsimaculatus var. oswaldoi of Lima (1928:96-98).

Nyssorhynchus aff. oswaldoi (?) of Townsend (1934:495-497).

Cellia oswaldoi of Boyd (1926:31, 37).

Anopheles (Nyssorhynchus) tarsimaculatus in part of Root (1922a:322; 1926b:684-700, 709-711;

1932:781); Christophers (1924:41, 90); Dyar (1925:187, 188, 195; 1928:439); Davis (1928:

549-553; 1933:278, 280); Shannon and Davis (1930:488); Senevet (1932:251; 1937:358-360);

Shannon (1933:124-133, in part?); Komp (1936a:161-162); Galvao, Lane and Correa (1937:40,

41); Lane (1939:29); Pinto (1939:390-398); Leeson and Buxton (1949:251, ?).

Anopheles tarsimaculatus in part (?) of Pinto (1930:154, 156).

Anopheles (Nyssorhynchus) tarsimaculata in part of Root (1924b:46 1-463).

Anopheles tarsimaculata in part of Dyar (1923:185); Godoy and Pinto (1923:29-33); Senevet

(1934: 29).

Anopheles (Anopheles) tarsimaculata in part of Bonne (1923:127-128); Bonne-Wepster and Bonne

(1923:127); Root (1923:276; 1924b:460-463); Bonne and Bonne-Wepster (1925:511-515).

Cellia tarsimaculata in part of Neiva and Penna (1916:94); Peryassu (1921b:183); Bonne (1924:

133-137).

Anopheles (Nyssorhynchus) aquacoelestis (!) of Senevet and Abonnenc (1938:487-493).

Anopheles (Nyssorhynchus) konderi of Galvao (1943:143, 148, 149, 151); Causey, Deane and

Deane (1944:2, 4, 5; 1946:27); Correa and Ramos (1944b:130); Deane, L. M., Causey and

Deane (1946:8; 1948:876-877); Deane, M. P., Causey and Deane (1946:38, 42, 45); Vargas

(1948:158); Lane (1949:403; 1953:262); Levi-Castillo (1949:11, 15, 28, 67, 72, 76, 81, 86);

Castro, Garcia and Bressanello (1959:549).

Anopheles konderi of Cova Garcia (1964:200).

Cellia albimana in part (?) of Neiva (1909:69-77).

FEMALE (fig. 6). Wing: 3.4 mm. Proboscis: 2.3 mm. Palpus: 2.25 mm. Forefe-

mur: 1.5mm. Abdomen: about 2.8 mm. Head: Integument brown to dark brown.

Proboscis about 1.5 length of forefemur. Vertex with many long, narrow, erect, cu-

neate scales, each with a narrow threadlike base. Palpal segment 2 occasionally with

dorsal light scales; segment 3 with a fairly extensive, conspicuous, dorsal or dorsola-

teral, longitudinal light stripe; apex of segment 2 with a small to moderate band of

whitish scales; apex of segment 3 with a moderately large band of white scales, 0.18

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 57

(0.15-0.25) of segment; segment 4 predominantly white with a moderately large, ba-

sal, dark band and a small, apical, dark band, with or without a usually inconspicu-

ous, dark, ventral stripe; segment 5 completely white. Antenna: Dorsomesal and

mesal surfaces of flagellar segment 1 distally with a large, very prominent patch of

long white scales. Flagellar segments 2-13 with 7-9 long, silver setae in basal whorl.

Thorax: Integument brown to very dark brown. Median anterior promontory with

numerous long, white, setiform scales extending dorsad onto anterior of acrostichal

line. Silver to cream scales on scutum slightly longer than those of aquasalis. Prescu-

tellar space usually horseshoe shaped. Sp with 4-9 moderately long, silver setae. Pra

with about 7-10 long, silver setae and a patch of gray to silver scales. Upper stp with

4 (2-6) long, light to dark setae in horizontal to vertical arc and a patch of gray to sil-

ver scales in horizontal row. Lower stp with 1, 2 long setae and a small patch of gray

to silver scales. Legs: Coxal light scales usually very light gray to white, usually not

cream. Midfemur with anteroapical spot and knee spot conspicuous and moderate in

size. Foretarsal segment 2 with apical 0.35 (0.20-0.45) cream to white; segment 3

with a large, 0.65-0.85 (0.50-0.85), apical, cream to white band, usually more exten-

sive than in aquasalis; segment 4 all dark to at least 0.3 basally dark; segment 5 gold-

en to light cream in about apical 0.5. Midtarsal segment 2 with a golden to cream

band in apical 0.10 (0.05-0.30); segment 3 with a small, golden to cream, apical band;

segment 4 all dark; segment 5 golden to white in apical 0.3-0.5. Hindtarsal segment 2

with a small dark band in basal 0.18 (0.12-0.25), remainder white. Wing: Light spots

on vein C and R cream to very light cream, not white; more posterior veins may be

white. Vein C humeral light spot 1.6-3.0 (1.1-3.8) of basal dark spot; subcostal light

spot about 0.11-0.35 (0.1-0.5) of subcostal dark spot, almost never greater than 0.5;

preapical light spot about 0.3 (0.25-0.40) of preapical dark spot; apical dark spot us-

ually very small and inconspicuous. R sectoral dark spot usually conspicuously smal-

ler than presectoral dark spot. Rs-R,,3 variable, more or less predominantly dark,

with 3, 4 dark spots and 3, 4 light spots. R3; with or without small apical dark spot.

Ry.s5 subcostal.and preapical dark spots usually moderately small. M highly variable,

from almost all light, with few dark scales, to predominantly dark; preapical dark

spot not extending onto M,,,. Apical light fringe spot usually moderately long, con-

spicuous, occasionally divided; light fringe spot at apex of Ry,; moderately large;

additional, inconspicuous, light fringe spots at apices of M,,., M34, Cu,, Cu, and A;

a moderate size, light fringe spot at level of 0.5 distance from base of A. Abdomen:

Sternite I with a few setae and a few inconspicuous, lanceolate scales. Scales of cau-

dolateral tufts on tergites and sternites long, very broad, outstanding, dark and obo-

vate.

MALE. Wing: 3.4 mm. Proboscis: 2.7 mm. Forefemur: 1.5 mm. Abdomen:

about 2.7 mm. Essentially as in female except for sexual characteristics. Head: Pal-

pal segments 2 and 3 with cream scales along dorsomesal border; segment 4 predomi-

nantly cream to white, usually with a basal or a subbasal, and occasionally an apical,

small, dark band; dark scales on ventral surface usually visible only when viewed ven-

trally. Antenna: About 0.75 length of proboscis. Flagellar segment 1 with long, con-

spicuous, oblanceolate scales on dorsomesal margin. Legs: Forefemur about 0.65

length of proboscis. Basal plantar surface of foretarsal segment 5 with 6-8 moderate-

ly long, thick, spiniform setae. Claw on foreleg long and curved; submedian tooth

0.20-0.25 length of claw, moderately acuminate and recurved at apex; basal tooth

moderately short and decurved, slightly less than length of submedian tooth.

MALE GENITALIA (fig. 10). Segment VIII: Tergite and sternite with moderate-

ly narrow, obovate scales; scales broader on sternite than on tergite. Segment IX:

6 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Sternite similar to aquasalis, curved, subtrapezoidal and moderately long, about 0.2

length of sidepiece. Anterior apodeme subtrapezoidal or subtriangular, and long

(longer than in aquasalis), about 0.25-0.33 length of sternite. Sidepiece: Moderately

narrow, conical. Tergal surface usually with 3-5 long, submedian tergomedial bristles

and 1, 2 subapicolateral bristles; with moderately long bristles mesad of tergomedial

bristles. Tubercle of parabasal spine large, appearing equal to or greater than 0.5

length of parabasal spine. Basal apodeme about 0.2 length of sidepiece. Longer,

more dorsal accessory spine about 0.5 length of sidepiece; more ventral spine 0.70-

0.85 of longer spine. Internal spine strongly retrorse apically. Clasper: Spiniform

attenuate, thin and long, subequal to seta b. Dorsal Claspette: Pedicel moderately

narrow (narrower than in aquasalis); base rounded and weakly curved mesad. Leaf-

lets moderately broad; basomesal projection of dorsal leaflet usually reduced. Ven-

tral Claspette: About 0.5 length of sidepiece. Lateral margins tapered toward nar-

row apex, width at apex about 0.33 or less length of claspette and about as wide as

or slightly wider than apex of aedeagus. Basal lobule very large, expanded laterally,

with long strong setae distributed along basal margin often reflexed and caudally di-

rected; setae along basal margin about 2.0 width of aedeagus; setae not concentrated

on basomesal margin. Lateral and ventral surfaces (exclusive of basal lobule) with se-

tae about equal to or slightly less than width of aedeagus; setae extending to or near-

ly to apex. Apex with rounded lateral margins and a moderately shallow median sul-

cus; sulcus with moderately sloping sides. Preapical plate large, usually crescent

shaped, moderately to heavily sclerotized, located about 0.3-0.4 length of claspette

basad of apex. Transparent membranous area immediately basad of preapical plate

bordering vertex of mesal cleft. Refringent structure moderately thick, in shape of

inverted horseshoe or V, usually without lateral arms and moderately pointed at ver-

tex. Phallosome: Aedeagus about 1.0-1.2 length of ventral claspette; apex rounded,

about as wide as long or slightly longer than wide; usually without, occasionally with

very small, membranous leaflets. |

PUPA (fig. 10). Abdomen: 2.5 mm. Trumpet: 0.45 mm. Paddle: 0.75 X 0.55

mm. Cephalothorax: Brownish yellow to brownish black. Wing cases occasionally

with dark, heavily sclerotized, longitudinal stripes. Hair 2-C slightly shorter than 1,

3-C; 1-C 1,2 branched; 2-C 1-3 branched; 3-C 2,3 branched (2-4). Hairs 4,5-C 1-3

forked. Hair 6-C 1,2 forked (1-3). Hair 7-C with 2-4 subequal branches, moderately

long. Trumpet: Pinna moderately heavily pigmented, dark amber; short, about 1.6-

2.1 (1.5-2.2) length of meatus; in lateral aspect, appearing truncate and flared apical-

ly, usually not appearing to taper toward apex. Metanotum: Hair 10-C single and

moderately short (0.1 mm), slightly shorter than 11-C. Hair 11-C 2-4 branched (1-4)

near base. Hair 12-C single, more than 2.0 length of 10-C. Abdomen: Hairs in gen-

eral shorter than those of aquasalis. Hair 2-1 3-7 branched, dendritic, moderately

long; 3-I about 0.50-0.75 length of 2-I. Hair 4-I variable, 3-5 branched (2-5). Hair 5-

I 1-3 forked (1-4), moderately long. Hair 6-I single or double, moderately long. Hair

7-I 2,3 branched (2-4), 0.60-0.75 length of 6-I. Hair 9-I 1,2 branched, longer than 7-

I and shorter than 6-I. Hair O-II-VI 3-6 branched (2-6); 0-II,III usually with 1,2 more

branches than O-IV-VII; 0-VII 3,4 branched (3-5). Hair 1-II 6-9 branched (49) and

1-III 7-13 branched (3-13), both strongly developed and with median branches long-

er than lateral; 1-I[V-VII single and long, about 1.5-1.7 length of segment. Hair 3-IV

3-5 branched (2-6); 3-V 2,3 forked (1-4), moderately long. Hair 5-III 5-8 branched

(4-8), moderate; 5-IV 3-5 branched (2-5), larger and stronger than 5-III. Hair 6-II

single, equal to or slightly longer than 7-II; 6-III single or double, moderately long.

Hair 7-II 2,3 branched (2-4); hairs 7-III-V and 8-III-VII short, with 4 or fewer

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 59

branches, usually 2,3 branched; 7-VI single or double, moderately long; 7-VII single

and long. Hair 9-II small and unpigmented; 9-III about 2.0 length of 9-II, weakly pig-

mented; 9-IV at least 2.0 of 9-III; 9-[V-VIII heavily pigmented; 9-IV,V shorter than

9-VI, VII; 9-V-VIII weakly curved, stout, short, usually 0.33 or less length of segment.

Hair 10-III 1-3 forked (1-4), moderate. Hair 4-VIII 2-4 forked (1-4), moderately

long. Terminal Segments: Male genital lobe with distinct, apical, mammilliform pro-

tuberance. Paddle: Very similar to that of aquasalis in size and shape. Subapex of

paddle weakly to moderately emarginate at insertion of hair 1-P. Midrib distinct ba-

sally, indistinct distally. External buttress 0.65 length of paddle. External margin

distad of buttress with fine short filamentous spicules extending around apex and ter-

minating 0.5 from base on inner margin. Hair 1-P single, moderately long. Hair 2-P

single or forked, subequal to 1-P.

LARVA (fig. 11). Head: 0.55 mm. Antenna: 0.35 mm. Anal Saddle: about 0.30

mm. Head: Moderately pigmented, light brown, with brown to dark brown mot-

tling. Median tooth of mental plate moderately broad and pointed. Inner and outer

clypeal hairs (2,3-C) plumose, with moderately long to long, usually dendritic branch-

es in apical half; 2,3-C subequal in length; hairs 2-C widely spaced, clypeal index

about 1.65. Hair 4-C 2-4 branched, moderately short, inserted moderately close to

anterior margin, distance between insertion of 4-C and 2-C subequal to distance be-

tween insertion of 2-C and 3-C. Hairs 8,9-C highly variable, 4-7 (3-8) and 4-7 (4-10)

branched respectively, moderately large, length about 1.5 distance separating hairs 5-

C. Hair 12-C 2-4 branched, subequal to hairs 8,9-C. Hair 15-C 3-5 branched (2-5),

moderately long. Collar dark brown, heavily pigmented and broad dorsomedially

(about 0.06 mm). Antenna: Pigmentation similar to head capsule. Mesal margin

with numerous, moderately long spicules becoming shorter and stouter apically; a

few short spicules on ventral surface. Hair 1-A 5-8 branched (3-8), short, about as

long as width of antenna at point of insertion. Thorax: Integument gray to dark yel-

lowish brown or grayish black. Inner submedian prothoracic group (1-3-P) with or

without hairs 1,2-P sharing common tubercle; palmate hair 1-P with 10-12 moderate-

ly narrow leaflets; 2-P 13-18 branched (12-18), 3.0-4.0 length of 1-P. Hair 14-P 6-8

branched from a short to moderately short, flattened stalk, lateral branches shorter

than median. Mesothoracic hair 1-M strongly plumose, 23-29 branched. Metathorac-

ic hair 2-T long, extending beyond caudal margin of thorax. Palmate hair 3-T moder-

ately large, with 9-13 semitransparent, narrow leaflets. Abdomen: Integument as in

thorax; more pigmented medially. Hair O-II-VII 5-8 branched (4-8), moderately large

to large. Palmate hair 1-I with 10-12 (8-16) moderately narrow, semitransparent leaf-

lets; 1-II-VII with 18-25 moderately broad, spreading leaflets; 1-III-VI larger than 1-

IT, VII. Hair 2-1 5-8 branched, large; 2-III 4-7 branched, large; 2-IV single, moderate-

ly long; 2-V 2,3 branched, large. Hair 5-I 2,3 branched, moderate; 5-II 7-12 branched,

moderate. Hair 9-I 3,4 branched (3-6), long. Hair 13-I,II,III 7-10, 6-11 and 8-11 (7-

11) branched respectively, small; 13-IV 5-7 branched, moderately large; 13-V 3-5

branched (3-6), very large, with long branches extending beyond caudal margin of

segment V; 13-VI 8-10 branched, about equal to 13-IV. Spiracular Lobe: Pecten

with 13-16 teeth; most of median teeth subequal and of moderate length; serrations

on dorsobasal margins of teeth moderately short. Lateral arm of spiracular apparatus

moderately long, projecting caudolaterad or laterad, rounded at lateral margins, usu-

ally reaching spiracular openings. Hairs 8,9-S 3-5 branched. Anal Segment: Almost

entire surface except for base covered with fine spicules, more strongly developed at

apex. Hair 1-X longer than saddle; not inserted on saddle; inserted immediately ven-

trad of ventral margin of saddle and usually slightly more than 0.5 distance from base

60 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

of segment. Anal gills long (0.5 mm), as long as or longer than anal segment.

DISCUSSION. An. oswaldoi can be distinguished from other species in the Oswal-

doi Subgroup in the female by the combination of (1) foretarsal segments 2 and 3

cream to white in apical 0.20-0.45 and 0.50-0.85 respectively, (2) foretarsal segment

4 all dark to more than 0.3 dark basally, (3) midtarsal segment 4 all dark, (4) hind-

tarsal segment 2 less than 0.25 dark basally and (5) vein C with a small, basal, dark

spot, and humeral light spot 1.6-3.0 (1.1-3.8) length of basal dark spot; in the male

genitalia by the combination of (1) basal lobules of ventral claspette very large, ex-

panded laterally, with long setae along basal margin which are about 2.0 width of ae-

deagus, (2) lateral and ventral surfaces of ventral claspette (exclusive of basal lobules)

with setae equal to or slightly less than width of aedeagus, (3) preapical plate large,

usually crescent shaped, moderately to heavily sclerotized and (4) aedeagus usually

without leaflets; in the pupa by (1) pinna of trumpet short, about 1.6-2.1 length of

meatus, appearing truncate and flared apically, (2) meatal cleft pointed at base, (3)

hair 9-V-VIII short, usually 0.33 or less length of segment, (4) hair 6-I] equal to or

slightly longer than 7-II and (5) hair 2-I 3-7 branched, moderately long; and in the

larva by the combination of (1) hairs 2,3-C plumose, with usually dendritic branches,

hairs 2-C widely spaced, clypeal index about 1.65, (2) hair 4-C moderately short, dis-

tance between insertion of 4-C:and 2-C subequal to distance between insertion of 2-

C and 3-C, (3) hairs 8,9-C moderately large, length about 1.5 distance separting hairs

5-C, (4) lateral arm of spiracular apparatus moderately long, (5) median teeth of pec-

ten mostly subequal, (6) hair 1-X not inserted on saddle and (7) anal gills long, as

long as or longer than anal segment.

The populations of oswaldoi examined in this study do not exhibit much interpop-

ulational variation. In the larvae, the clypeals (2,3-C) of the Panamanian populations

do not show as extensive branching as the South American populations. Frequently

the branching of the clypeals is simple in Panamanian specimens rather than dendri-

tic. In the male genitalia, the Panamanian populations sometimes have very small,

membranous, basolaterally directed leaflets similar to those in ininii, rangeli and nu-

neztovari. It is possible that these leaflets are occasionally present in other popula-

tions, but I have not been able to observe them. However, the diagnostic key charac-

ters are constant, permitting identification of all stages.

An. oswaldoi is most closely related to noroestensis, galvaoi and aquasalis. The

male genitalia of these species are very similar in general appearance; this is particular-

ly evident when comparing oswaldoi with galvaoi and noroestensis. The shape of the

preapical plate and the length of the setae on the ventral claspette of galvaoi and nor-

oestensis, and the truncate aedeagus of noroestensis are the only characters which

differentiate the male genitalia of these species from oswaldoi. There is also consid-

erable overlap in the adults of oswaldoi and noroestensis, which sometimes makes

species identification difficult. The larvae of aquasalis and noroestensis are similar to

those of oswaldoi, differing mainly in the characters given above. The pupa of noro-

estensis is distinguished from that of oswaldoi only by a longer pinna and a slightly

longer hair 9 on segments V-VIII.

Although oswaldoi may occur sympatrically, in the broad sense, with aquasalis and

noroestensis, it is ecologically isolated from these species, breeding in fresh, usually

well shaded water in the interior of forests. The center of origin for oswaldoi was

probably a forested region in tropical South America. An. oswaldoi has extended its

range through the isthmus of Panama as far north as Costa Rica.

Peryassu described oswaldoi as a species of Cellia in 1922. The following year

Dyar (1923:185) listed oswaldoi as a synonym of An. tarsimaculata Goeldi 1905.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 61

Bonne (1923:128; 1924:134, 135), and Bonne and Bonne-Wepster (1925:514)

recognized 2 “‘races” of tarsimaculatus existing in Surinam: one that was exo-

philic in the interior of the country, characterized by a very small dark ring on the

hindtarsal segment 2 (~oswaldoi), and the other a coastal species (=aquasalis). Root

(1924b:461, 462), after studying a series of mounts of the male genitalia of “oswal-

doi’’ and tarsimaculata, found no genitalic differences, and therefore synonymized

oswaldoi again with tarsimaculata. Lima (1928:96-98) described constant differen-

ces between tarsimaculatus and oswaldoi in the adult, pupa and larva, but maintained

that oswaldoi was only a variety of tarsimaculatus. In 1932, Curry described aquasa-

lis and aquacaelestis as varieties of tarsimaculatus. He stated that the variety aquacae-

lestis “is undoubtedly quite similar in appearance to A. oswaldoi from Brazil, which

Root (1926b:709-711) places in the synonymy of A. tarsimaculatus.” Senevet

(1934:49-52) revalidated oswaldoi, describing the pupa in some detail, and, in 1938,

Senevet and Abonnenc synonymized aquacoelestis [sic] with oswaldoi. Galvao and

Lane (1937e:218-221) stated that the species, oswaldoi, really consists of 3 types:

oswaldoi oswaldoi Peryassu 1922, oswaldoi var. metcalfi Galvao and Lane 1937 and

oswaldoi var. noroestensis Galvao and Lane 1937. In 1942, Galvao and Damasceno

recognized noroestensis as a species separate from oswaldoi and considered metcalfi

to be anomen nudum (sic, 7nomen dubium]. In this same paper they described

konderi as a species very closely related to oswaldoi, differing only in the shape of

the apex of the aedeagus. I have not had the opportunity to examine any material

from the type locality of konderi. However, since the proportions of the aedeagus

appear to change depending on how the genitalia are mounted, and some variation

does occur between different populations, I concur at this time with Lane (1953:

262) in synonymizing konderi with oswaldoi.

BIONOMICS. The immature stages of oswaldoi are found in the interior, usually

in or on the margins of tropical forests, frequently along roads, in cultivated fields or

grasslands adjacent to forested areas. The breeding sites are in permanent or tempor-

ary ground pools, margins of ponds or lakes, swamps and streamside pools; the sites

are found most often in deep to partial shade, less often in full sun. The immature

stages are commonly collected in the grassy margins of pools with muddy bottoms,

and algae and abundant flotage are often present. The water is always fresh and may

be turbid or clear. Galvao (1938b) reared oswaldoi in the laboratory at a mean tem-

perature of 25.7°C and found that the time necessary for development from egg to

adult was 12 days. The immatures are commonly encountered in association with

triannulatus and less often rangeli. Occasionally oswaldoi has been collected with in-

inii and nuneztovari in Para, Brazil. Other species found in association with oswaldoi

are Anopheles (Ano. ) neomaculipalpus, An. (Ano.) punctimacula, Culex (Cux.) coro-

nator group, Uranotaenia (Ura. ) geometrica and Ur. (Ura.) lowii.

The adults of oswaldoi are largely restricted to the forest. They are exophilic and

zoophilic. Of the 6470 adults of oswaldoi Deane, Causey and Deane (1948:876) cap-

tured in Para, Brazil, 83 or 1.3% were from inside houses. Rey and Renjifo (1950:

537) in 3 months, September through November, captured 2414 anophelines inside

houses in northern Colombia of which only 0.75% were oswaldoi. Correa and Ramos

(1944c:11) reported that from the Ilha de Santo Amaro, in the state of Sao Paulo,

Brazil, 5.2% of the anophelines collected in houses were oswaldoi. Although primar-

ily zoophilic, oswaldoi feeds freely on man inside the forest, such as in the Mojinga

Swamp in Panama or in the forest of French Guiana (Rozeboom 1941:102-103;

Floch and Abonnenc 1947:6; Curry 1932). Rozeboom (1942a:240), in the interior

of the cacao growing districts of Trinidad, collected twice as many females of oswal-

62 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

doi on animal bait as on human bait and found none inside houses. Deane, Causey _

and Deane (1948) reported that the peak of biting activity of oswaldoi was between

1800 and 1900 h.

MEDICAL IMPORTANCE. An. oswaldoi has been experimentally infected with

Plasmodium vivax and P. falciparum by Fonseca and Fonseca (1942:106-111) in the

state of Sao Paulo, Brazil and by Rozeboom (1942a:240) in Trinidad. However, it

has never been found naturally infected or otherwise implicated in transmitting ma-

laria. Deane, Causey and Deane (1948:876) dissected 540 females from the north-

east of Brazil and found none infected with Plasmodium. Lucena (1940b) and Cor-

rea and Ramos (1942b) reported finding oswaldoi var. metcalfi naturally infected in

Brazil; however, these investigators were probably dealing with noroestensis or aqua-

salis and not oswaldoi. An. oswaldoi, at present, does not seem to be a vector of ma-

laria.

DISTRIBUTION (fig. 2). An. oswaldoi is distributed throughout South America

east of the Andes as far south as the northern provinces (Formosa, Missiones, Salta,

Tucuman) of Argentina. It occurs throughout most of Colombia, Venezuela, the

Guianas, Brazil, Paraguay and Bolivia; it also occurs in Ecuador and Peru east of the

Andes. Northward, it extends into Panama and Costa Rica. It is also found in Trini-

dad, but not in Tobago or any other islands of the Antilles. It is not certain how far

south oswaldoi occurs west of the Andes.

Material Examined: 1376 specimens: 244 males, 117 male genitalia, 561 females,

273 pupae, 181 larvae; 259 individual rearings: 129 larval, 118 pupal, 12 incomplete;

9 progeny rearings.

BOLIVIA (7F). Beni: Guayaramerin, 18 Feb 1944, 1F [MCZ] ; same locality, 3 Jun 1947, SF

(47-12258). Pando: Ingavi, Abuna?, 29 Dec 1943, 3111, [KO] 119A-30, 1F.

BRAZIL. Amazonas (9F): Manaus, Jun 1931, R. Shannon, 4F; same locality, F. Urich, 1F.

Rio Amazonas above Manaus, 1914-18, R. Moffat, 1F [BM]. Rio Brancos, Aug 1930, 2F. Tonan-

tins, Mar 1931, 1F. Bahia (1M, 2Mgen, 2F): Bonfim, 26 Jan 1930, 2F. Salvador, 1936, R. Shan-

non, 1M, 1Mgen. Locality not specified, M. Boyd, 1Mgen. Mato Grosso (1M, 1Mgen, 43F): Mara-

caju, Jun 1937, 1M, 1Mgen. Westborder, May 1931, R. Shannon, 1F (151). Locality not specified,

May 1931, R. Shannon, 42F. Para (60 specimens: 8M, 9Mgen, 34F, 6p, 3 1; 6 ind rear: 21, 4p):

Altamira, 150 km W of, 5 Nov 1974, J. Reinert et al., coll. 77, 1 pF. Belem, 14 Apr 1941, W.

Komp, BRAK 2A, 3F; same locality, 16 Apr 1941, 1 1; same locality, 17 Apr 1941, BRAK 3, 4M,

3Mgen; same locality, Mar 1944, MacCreary, Bricken, 1Mgen; same locality, det. Deane, 1M [CU];

same locality, W. Komp, 3Mgen. Bacuri, 28 Oct 1974, J. Reinert, et al, coll. 62, 1Mgen, 2 IpM, 1

pM, 2 pF. Fordlandia, 31 Jun, R. Shannon, 1M, 1Mgen. Maraba area, 4 May 1976, 1F (M-6); same

locality, 2 Jun 1976, 1F (R-3); same locality, 5 Jul 1976, 11F (M-6); same locality, 7 Jul 1976, 1F

(M-8); same locality, 19 Oct 1976, 4F (1); same locality, 20 Oct 1976, 1F (1); same locality, 21 Oct

1976, 1F (1); same locality, 11 Nov 1976, 1F (1); same locality, 1976, D. Roberts, 3F. Locality

not specified, N. Davis, 4F. Rio de Janeiro (1M, 2Mgen, 11F): Bangu, 11 Jun 1924, F. Root, 1

Mgen. Porto das Caixas, 3 Mar 1925, 1F (37); same locality, 29 May 1925, F. Root, 1M and 1Mgen

(710210-1, lectotype of metcalfi). Rio de Janeiro, L. Saunders, 3F [BM1924-240] ; same locality,

M. Boyd, 1F [BM 1925-336] ; same locality and collector, 6F. Rondonia (9F): Porto Velho, May

1931, R. Shannon, 5F. Guajara Mirim, May 1931, R. Shannon, 3F. Sao Paulo (6M, 2Mgen, 5F, 1

Ip): Caraguatatuba, 1M, 1F. Juquia, Nov 1938, [?] J. Lane, 1F (1.116); same locality and collec-

tor, 1F [CU]. Santos, Guanuja, 1F. Sao Vincente, 1938, A. Galvao, 4M, 1F; same locality, 1M;

same locality, 1Mgen (97-16), 1 lp (15-12) [JH]. Locality not specified, 1938, A. Galvao, 1Mgen.

COLOMBIA (57 specimens: 13M, 9Mgen, 6F, 11p, 18L&l; 2 inc ind rear). Boyaca: Puerto Boy-

aca, COM 651. Caqueta: Tres Esquinas, COM 652. Meta: Restrepo, Rd. to Cumaral, Jul 1935,

KO 120A-4, 1M. Villavicencio, Jun 1947, CV 671, 2 lp, 9p, 15L, 11; same locality, 1947, L. Roze-

boom, CV-P 38, 1F, E (progeny); same locality, Jun 1947, [CV] 671.1, 1Mgen [JH]. Puerto Lo-

pez, COM 653. Puerto Porfia, COM 649. Locality not specified, 25 Nov 1936, COR 155, 2F.

Department not specified: Puerto Nino, 21 Feb 1922, F. Miller, 1F. Locality not specified, 1943,

[KO] 119A-21, 2F.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 63

COSTA RICA (1Mgen). Limon: Puerto Viejo, Apr 1938, H. Kumm, 1Mgen.

GUYANA (8 specimens: 3M, 3F, 2L). East Demerara-West Coast Berbice: Mackenzie, Sep

1945, 111A-310, 2F; same locality, 15 May 1945, G. Giglioli, [KO] 119A-41, 3M, 1F; same local-

ity, Sep 1949, 2L.

PANAMA AND CANAL ZONE (878 specimens: 144M, 65Mgen, 281F, 251P&p, 137L&l; 249

ind rear: 127 1, 113p, 9inc; 9 progeny rearings). Canal Zone: Atlantic side, May 1939, PAX 114,

3M, 5F. Escondido River, 19 May 1939, PAX 108, 2F. Fort Davis, 13 Mar 1923, D. Baker, 1F;

same locality and collector, 30 Mar 1925, 1M; same locality, 12 Feb 1940, PAX 168, 4M, 2F.

Fort Sherman, 8 Feb 1945, ASM 410-1, 1F; same locality, May 1945, 1M. France Field, 30 Aug

1924, J. Zetek, 1M, 1F; same locality and collector, 1 Nov 1924, 4M, 5F; same data, 1M, 2F

[MCZ] ; same locality, 11 Nov 1924, 1M, 8F; same locality, 23 Nov 1924, C. Mopp, 2M, 2F; same

locality, 7 Mar 1925, D. Baker, 1F; same locality and collector, 9 May 1925, 1M, 1F; same locality,

6 Feb 1940, 2M, 4F. Gatun, Jan 1913, J. Zetek, 1Mgen; same locality, 7 Jul 1931, D. Curry, PAX

25, 3F. Mojinga Swamp, Oct 1935, PAX 54, 9M, 11F; same locality, 28 Oct 1948, No. 113, 1L;

same locality, 2 Jul 1949, 16Mgen [JH] ; same locality, 20 Jul 1949, 14Mgen; same locality, PA

729, 1175, 1176 (6 progeny rearings), 1177 (3 progeny rearings); same locality, 12F. [Rio] Cha-

gres, 17, 19 Aug 1953, S. Carpenter, 20F; same locality and collector, 10 Sep 1953, 26F. Locality

not specified, D. Curry, 1F [BM1933-504];5 Aug 1909, A. Jennings, 1Mgen. Darien: Paya, GG

1-126, 1-129, 1-133, 1-137, 1-138, 1-141. Rio Tuira, GG 1-159. Panama: Juan Diaz, 21 Mar

1935, PAR 16, 3F. La Chorrera, 29 Jan 1945, ASM 397-1, 1F. Province not specified: Rio Trini-

dad, 9 Jun 1912, A. Busck, 2F; same locality and collector, 3 Jul 1912, 1F. Locality not specified,

5 May 1952, S. Carpenter, 1F (620926-6); 30 Jul 1936, PAR 104, 3F.

PERU (71 specimens: 2Mgen, 67F, 2L). Huanuco: Tingo Maria, May 1946, E. Hambleton, 2

Mgen; same locality, [21946], Emilio-Viale, 2L. Loreto: Iquitos, Mar-Apr 1931, R. Shannon, 67F.

SURINAME (23 specimens: 11M, 4Mgen, 8F). Marowijne: Moengo, 1944, Guicherit, 3F; same

locality and collector, Oct 1945, 1M (46-V-8d), 1M (46-V-8e), 1Mgen. Paramaribo: Paramaribo,

J. Bonne-Wepster, 4M, 1Mgen. District not specified: Billeton, 5 Jul 1943, 1F. “‘Leidino XIa,”

21 Jun 1946, E. van der Kuyp, 5M, 2Mgen. Locality not specified, J. Bonne-Wepster, BB 897, 4F.

TRINIDAD AND TOBAGO. TRINIDAD (158 specimens: 52M, 14Mgen, 71F, 4p, 17L&l; 1 p

ind rear). Caroni: Tabaquite, 22 Aug 1945, 2M, 5F; same data, [KO] 119A-31, 2F; same data, P.

Woke, 3M, 2F; same locality and date, [KO] 119A-40, 1F; same data, [KO] 119A-42, 3M, 3F;

same locality, 23 Aug 1945, 1 1;same data, [KO] 119A-33, 1F; same locality, 22 Aug 1945, [KO]

119A-39, 1M, 4F; same locality, 23 Aug 1945, [KO] 119A-29, 1F. Nariva: Charuma Forest, 27

Aug 1964, A. Guerra, TR 643, 1L. Nariva Swamp, 16 Nov 1961, T. Aitken, 1p; same locality, 18

Feb 1964, TR 77, 1L. Navet, 20 Aug 1964, A. Guerra, TR 610, 2L. St. Andrew: Cumuto, 2 Jun

1941, TRR 1, 2M, 2Mgen, 1F; same locality, 27 Jun 1941, TRR 15B, 1M, 1Mgen, 2F; same local-

ity, 7 Aug 1941, TRR 26, 2F; same locality, TRR 64, 2M, 1Mgen, 3F. Mount Harris, 10-16 Jul

1964, F. Powdhar, 1F. Sangre Grande, 1941, W. Downs, coll. E-43-44, 3Mgen. Valencia, 29 Aug

1945,11. St. George: Arima, 21 Aug 1938, E. de Verteuil, 1M [BM]; same locality, 8 Jul 1965,

A. Guerra, TR 1247, 1Mgen, 1 pM. Curepe, 21 Jul 1931, E. de Verteuil, 1M [BM1931-562]. Die-

go Martin River, 8-26 Feb 1941, W. Komp, TRK 35, 17M, 13F. Fort Read, Oct 1944, 1F. Four

Roads, 18 Mar 1941, W. Komp, [KO] 119A-6, 3M, 5F; same locality, 5 Jun 1941, TRR 3, 13M, 2

Mgen, 9F. Maraval, 1946, 5F. County not specified: St. Helena, 24 Oct 1936, TRK 33, 1M, 1M

gen, 8F. Locality not specified, W. Downs, 1Mgen (301-3), 1Mgen (409-1), 1Mgen (413-2) [JH] ;

Mar 1941, W. Komp, KO 111-7, 1M; 22 Aug 1945, 2p, 1L, 101; no data, 1F.

VENEZUELA (16 specimens: 4M, 6Mgen, 5F,1L). Aragua:. Maracay, 1F. Monagas: Caripito,

10 Jun [1935], 1F. Portuguesa: Acarigua, 1Mgen. State not specified: Rancho Cachipo, 5 Sep

1936, Quire, [KO] 119A-7,3M, 2F. Juasipati, May 1929, 2Mgen (52). San Joaquin, 16 Dec 1944,

E. Winton, KO 111-16, 1M. Locality not specified, VZ 421; Jul 1937, P. Anduze, 1L; 1938, A.

Gabaldon, 2Mgen; 1949, 1Mgen.

64 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

3. Anopheles (Nys.) galvaoi Causey, Deane & Deane

Figs. 3, 4, 6, 12

1943. Anopheles (Nyssorhynchus) galvaoi Causey, Deane and Deane 1943:293-296. TYPE:

Lectotype female (R426), marked capanemai in register, Rio Branco (Acre), Brazil, O. R.

Causey [FMSP, 694; designation by Belkin, Schick and Heinemann 1971:5].

Anopheles (Nyssorhynchus) galvaoi of Galvao (1943:142, 143, 145, 146, 148, 149, 151); Causey,

Deane and Deane (1944:1, 2, 5, 6; 1945:248; 1946:27, 31); Correa and Ramos (1944b: 131);

Galvao, Lane and Unti (1944:45, 46); Deane, L. M., Causey and Deane (1946:7, 8, 9, 10, 12;

1948:877); Deane, M. P., Causey and Deane (1946:43, 45); Coher (1948:88); Senevet (1948a:

279); Lane (1949:403, 410, 413, 414; 1953:269-270); Levi-Castillo (1949:11, 15, 28, 58, 67,

72, 76, 82, 86); Carvalho and Rachou (1951:475, 480); Rachou and Ricciardi (1951:424, 425-

426); Bejarano (1957:326, 330, 331); Vargas (1959:370); Cerqueira (1961:124); Forattini

(1962:325, 370, 415, 418-419, 476, 484, 491); Garcia and Ronderos (1962:137, 138, 139,

158); Consolim and Galvao (1973:177, ?).

Anopheles galvaoi of Foote and Cook (1959:29); Maciel (1962:477-478); Mattos and Xavier (1965:

272); Gorham, Stojanovich and Scott (1967:18, 47, 54).

FEMALE (fig. 6). Wing: about 3.5 mm. Proboscis: 2.3 mm. Palpus: 2.2 mm.

Forefemur: 1.5 mm. Abdomen: about 3.1 mm. Description based on only 4 dam-

aged females (2 paratypes and 2 topotypes). Head: Integument dark brown. Pro-

boscis about 1.5 length of forefemur. Vertex with numerous, long, erect, cuneate or

weakly forked, white scales. Palpal segments 2 and 3 with an indistinct, dorsolongi-

tudinal, light stripe; apex of segment 2 with a small, light, apical ring; segment 3

with a moderately large, light apical band; segment 4 with a basal and an apical, mod-

erately small, dark band, remainder light without distinct ventral stripe of dark scales;

segment 5 light. Antenna: Flagellar segment 1 with a patch of long, white, oblance-

olate scales distally on dorsomesal surface, shorter scales mesally; with small light

scales basally on lateral surface. Flagellar segments 2-13 each with about 6-9 long,

silver setae in a basal whorl. Thorax: Integument brown. Median anterior promon-

tory area with long, creamish to white, setiform scales extending dorsad onto most

anterior part of acrostichal line. Scutum with moderately short (0.7 mm), obovate,

creamish scales. Prescutellar space horseshoe shaped. Many of setae and scales mis-

sing or not apparent on pleuron. Upper stp with 2, 3 long, dark setae and many,

small, obovate, cream scales in horizontal arc. Lower stp with one long, dark seta

and a patch of small, obovate, cream scales. Upper mep with about 4 long, cream

setae. Legs: Light scales on coxae grayish white to white. Midfemur with antero-

apical spot and knee spot moderately large, distinct and light cream to white. Fore-

tarsal segment 2 cream to white in apical 0.13-0.24; segment 3 cream to white in api-

cal 0.16-0.40; segment 4 with a few, light, apical scales; segment 5 cream to white in

apical 0.30-0.45. Midtarsal segment 2 creamish white to white in apical 0.08-0.11;

segment 3 with a few, light, apical scales; segment 4 all dark or apically slightly light-

er; segment 5 creamish white to white in about apical 0.5. Hindtarsal segment 2

brown in basal 0.42-0.55; segment 5 cream in about apical 0.5. Wing: Light scales

on wing white except for creamish white scales on vein C. Vein C humeral light spot

large, 2.1-3.6 of basal dark spot; basal dark spot small; presectoral light spot absent

or consisting of only a single scale, so that presectoral and sectoral dark spots fuse in-

to a single, large, dark spot; subcostal light spot 0.24-0.45 of subcostal dark spot; pre-

apical light spot 0.35-0.48 of preapical dark spot; apical dark spot small. R presector-

al dark spot small; sectoral dark spot very small, in one specimen consisting of only 2

dark scales. R3 with 3 light spots and 3 occasionally indistinct, dark spots; apical

dark spot small. R4,; subcostal dark spot moderately small; preapical dark spot

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 65

moderately small to moderately large. M sectoral dark spot moderately small and in-

conspicuous, to moderately large with a few interspersed light scales. Cu sectoral

dark spot moderately small. Cu, subcostal dark spot small, in one specimen consis-

ting of only a single scale. Vein A subcostal dark spot small to moderate, about 0.15

of vein. Apical light fringe spot moderately small and divided; additional light fringe

spots at apices of Rais, Mis2., M34, Cu,, Cu, and A; a moderate size, light fringe spot

at level of 0.5 distance from base of A. Abdomen: Sternite I with a few, light, obo-

vate, submedian scales. Dark caudolateral scale tufts large, each scale about 0.1 mm

long. Tergites II-VII densely covered with cream to golden, narrow, lanceolate scales

in a subtriangular pattern on segments II, III, becoming more extensive and subrec-

tangular on distal segments.

MALE. Wing: about 3.3 mm. Forefemur: 1.5 mm. Description based only on a

single paratype male. Essentially as in female except for sexual characters. Head:

Palpal segments 2 and 3 with a dorsal stripe of cream to white scales, stripe longer on

2 than on 3; segment 4 witha light basal band, and a subbasal and a subapical, dark

band, mediolaterally predominantly light, ventral surface inconspicuously dark. An-

tenna: Flagellar segment 1 with long, oblanceolate, white scales on dorsomesal sur-

face. Legs: Claw on foreleg weakly curved; submedian tooth recurved at apex, about

0.25 length of claw; basal tooth subequal to submedian, decurved.

MALE GENITALIA (fig. 12). Description based on only 2 specimens (one para-

type and one from Sao Paulo). In general very similar to those of oswaldoi and noro-

estensis. Segment VIII: Tergite and sternite with narrow, long, obovate scales; scales

slightly broader on sternite than on tergite, darker medially on tergite. Segment IX:

Sternite as for those of oswaldoi and noroestensis; subtrapezoidal, about 0.25 length

of sidepiece. Anterior apodeme moderately long, about 0.2 length of sternite. Side-

piece: Moderately broad and subconical. Tergal surface with 5 alveoli for submedi-

an tergomedial bristles, 2 long, strong bristles remaining on specimens; 1, 2 long sub-

apicolateral bristles; 5, 6 moderately short to moderately long bristles mesad of ter-

gomedial bristles, subbasal bristles longest. Parabasal spine moderately long and nar-

row; tubercle of spine moderately long, 0.3-0.4 of spine; combined lengths of tuber-

cle and spine about 0.3 of sidepiece. Basal apodeme moderately short, about 0.2

length of sidepiece. Longer, more dorsal accessory spine slightly longer than 0.5 of

sidepiece, curving caudally at apex; more ventral spine about 0.8 of longer spine. In-

ternal spine subequal to shorter accessory spine, strongly retrorse apically. Clasper:

Spiniform long, slender and apically attenuate; seta b 1.5-2.0 length of spiniform.

Dorsal Claspette: About 0.5 of sidepiece. Pedicel moderately narrow; base curved

mesad. Leaflets about 0.40-0.47 length of claspette; dorsal leaflet with prominent

basomesal projection. Ventral Claspette: About 0.5 length of sidepiece. Lateral

margins tapered toward narrow apex, width at apex about 0.35 length of claspette

and 1.25-1.50 width of apex of aedeagus. Basal lobule very large, expanded laterally

and covered with very long, occasionally caudally reflexed setae distributed uniform-

ly over lobule; setae on basal margin about 3.0 width of aedeagus. Ventral and later-

al surfaces (exclusive of basal lobule) covered with short setae about 0.5 width of ae-

deagus; setae extending to or nearly to apex. Apex with rounded lateral margins sep-

arated by a moderately shallow V shaped median sulcus; lateral margins tapering api-

cally. Preapical plate distinct, large, circular to semicircular with small basolateral

projections directed basally, heavily sclerotized, located about 0.25 length of clasp-

ette basad of apex. Transparent membranous area small, not very distinct, with later-

al margins projecting basolaterally. Refringent structure very distinct, in shape of in-

verted V, lateral arms not visible. Phallosome: Aedeagus equal to or slightly longer

— 66 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

than ventral claspette; apex rounded, about as wide as long (possibly shorter than os-

waldoi); leaflets absent.

PUPA, LARVA. No specimens available.

DISCUSSION. An. galvaoi can be distinguished from other species in the Oswal-

doi Complex in the male genitalia by the combination of (1) ventral claspette with

basal lobule very large, covered with very long, uniformly distributed setae about 3.0 |

width of aedeagus, and ventral and lateral surfaces (exclusive of basal lobules) with

short setae about 0.5 width of aedeagus, (2) preapical plate heavily sclerotized, cir-

cular to semicircular with small basolateral projections and (3) apex of aedeagus

rounded, about as wide as long (possibly shorter than oswaldoi), without leaflets.

The female of galvaoi is apparently very similar to those of aquasalis and benarrochi.

Since there were only 4 damaged specimens available to me, I was not able to differ-

entiate the female of galvaoi from those of the latter 2 species. Consequently, I have

not included galvaoi in the key to females. Forattini (1962:418-419) states that Cor-

rea and Ramalho (1958) can distinguish galvaoi from aquasalis on the basis of the rel-

ative lengths of the dark basal spot and humeral light spot of vein C. Forattini re-

ports that in aquasalis the basal dark spot is about 0.5 the length of the humeral (B,)

spot; whereas, in galvaoi, the basal dark spot is clearly less than 0.5 the length of the

humeral light spot. For aquasalis this is not always true. Because aquasalis and gal-

vaoi are not sympatric, there should be no problem in recognizing aquasalis. A prob-

lem also arises in that the wing of benarrochi has the same characteristics as described

above for galvaoi, and it is possible that these 2 species occur sympatrically. The

male genitalia of galvaoi are very similar to those of oswaldoi and noroestensis, gal-

vaoi differing only by (1) longer setae on basal lobule of ventral claspette and shorter

setae on ventral and lateral surfaces of the fused portion of ventral claspette when

compared to oswaldoi, (2) circular or semicircular shape of preapical plate when com-

pared to oswaldoi and (3) rounded apex of the aedeagus in contrast to noroestensis.

Neither the larva nor the pupa of galvaoi was available for this revision. However,

the larva has been described by M. P. Deane, Causey and Deane (1946:36). From

their description and key (p 43) it seems that galvaoi most closely resembles goeldii

=nuneztovari), noroestensis and rangeli. With regard to the larva, these investigators

stated, “*.. .it [galvaoi] can usually be separated from A. goeldii in which the poster-

ior clypeal hairs are longer and simple or divided far from the base. In A. goeldii, A.

noroestensis and A. rangeli the anterior clypeal hairs have even less numerous and

shorter branches, and the leaflets of the abdominal palmate tufts of segments 5 to 7

are pointed at the tips.”’ In their key, galvaoi is distinguished from noroestensis and

rangeli by palmate hairs 1-V-VII possessing apically truncate leaflets. The difficulties

of correctly identifying galvaoi are best summarized by Rachou and Ricciardi (1951:

426): “Para se julgar dessa dificuldade, basta atentar-se para o fato de que esse mos-

quito, segundo os autores da especie, se assemelha ao A. tarsimaculatus (A. aquasalis)

nas marcacoes do alado, ao A. oswaldoi na terminalia do macho, ao A. noroestensis

na fase larvaria e possue ovos do tipo oswaldoi, donde se conclue que para se fazer

um diagnostico conscientodas dessa especie e indispensavel que se analise todo esse

conjunto, isto e todos as suas fases evolutivas.’”’

Based on the ventral claspette of the male genitalia, galvaoi shows the greatest af-

finity to oswaldoi and noroestensis. The larva seems to be very similar to that of nor-

oestensis. For these reasons I am tentatively placing galvaoi between oswaldoi and

noroestensis in this revision. The center of origin for galvaoi was probably within its

present distribution, possibly in some refugium in the upper Amazon basin or along

the eastern slope of the Andes.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 67

BIONOMICS. Very little has been written about the natural history of galvaoi.

Deane, Causey and Deane (1948:877) found the larvae in puddles containing grass

and algae exposed to the sun. Forattini (1962:415) reports that the larvae are found

in similar habitats to those of rondoni, such as puddles and marshes. The females are

said to be crepuscular and have been collected on animal bait but not inside houses

(Deane, Causey and Deane 1948).

MEDICAL IMPORTANCE. An. galvaoi is of no known medical importance.

DISTRIBUTION (fig. 3). An. galvaoi was described from the territory of Acre,

Brazil. This species has been reported also to occur in Brazil in the following territor-

ies or states: Amazonas, Rondonia, Mato Grosso, Para, Sao Paulo, Rio de Janeiro (?)

(Carvalho and Rachou 1951:475, 480); Goias (?) (Mattos and Xavier 1965:272); and

Bahia (?) and Parana (?) (Rachou and Ricciardi 1951:425-426). Deane, Causey and

Deane (1948:877) state that in the northeast and Amazon basin regions of Brazil,

galvaoi is restricted to the high Amazon basin. These investigators encountered gal-

vaoi in Rio Branco, Rio Zinho and Brasileia (Acre), and in Guajara Mirim (Rondonia).

Garcia and Ronderos (1962:158) record galvaoi from Paraguay.

Material Examined: 7 specimens: | male, 2 male genitalia, 4 females.

BRAZIL. Acre (1M, 1Mgen,4F): Rio Branco, O. Causey, 1M and 1Mgen (paratype 58037), 1F

(paratype 58039), 1F (paratype); same locality, S.E.S.P. 699, 2F (13428-12 and 13428-42) [JH].

Sao Paulo [?| (1Mgen): Locality not specified, det. A. Galvao, 1Mgen (1085-28).

4. Anopheles (Nys.) noroestensis Galvao & Lane

Figs. 2, 4, 6, 13, 14

1937. Anopheles (Nyssorhynchus) oswaldoi var. noroestensis Galvao and Lane 1937e:220-221.

TYPE: Lectotype male genitalia slide (2177, 10293), adult apparently lost, (possible syn-

type FMSP, 373), Novo Oriente [Pereira Barreto] , near Lussanvira (Sao Paulo), Brazil

[FH; designation by Belkin, Schick and Heinemann 1971:6].

1937. Anopheles (Nyssorhynchus) oswaldoi var. metcalfi Galvao and Lane 1937e:218-220.

TYPE: Lectotype male (96) with genitalia on slide (710210-1), Porto das Caixas (Rio de

Janeiro), Brazil, 29 May 1925, F. M. Root; one of several specimens identified as tarsima-

culatus by Root (1926b:711), on which Galvao and Lane based their metcalfi [|USNM; de-

signation by Belkin, Schick and Heinemann 1971:6]. TRANSFERRED SYNONYMY.

1940. Anofeles (Nyssorynchus) osvaldoi (!) var. ayrozai Unti 1940a:379-383. TYPE. Syntypes

female(s), larva(e), eggs, Vale do Rio Paraiba, Guaratingueta (Sao Paulo), Brazil, Nov 1939-

Aug 1940 [NE; not in FH or SPM, Belkin, Schick and Heinemann 1971:6]. Synonymy

with noroestensis by Galvao, Lane and Unti (1944:39).

1942. Anopheles (Nyssorhynchus) clarki Komp 1943:197-201. TYPE: Holotype dissected male

genitalia mounted on slide, Monteros (Tucuman), Argentina, Jun 1940, C. A. Alvarado

[USNM, 56476; male apparently lost] .

Anopheles (Nyssorhynchus) noroestensis of Galvao and Damasceno (1942:125-129, 131); Barretto

and Coutinho (1943:321); Correa (1943:128); Fonseca and Unti (1943:47-49); Galvao (1943:

143, 148, 149, 151); Ramos (1943:52); Unti and Ramos (1943:28); Causey, Deane and Deane

(1944:2, 4, 5; 1946:26); Correa and Ramos (1944b:130); Galvao, Lane and Unti (1944:37-43);

Lane (1944: 264, 265; 1949:403; 1953:264-266); Deane, L. M., Causey and Deane (1946:7, 9,

10, 12; 1948:878-880); Deane, M. P., Causey and Deane (1946:38, 43, 45); Amaral and Penido

(1947:168, 173-179); Senevet (1948a:278); Levi-Castillo (1949:10, 11, 15, 28, 67, 72, 76, 82,

86); Duret (1950a:371; 1950b:301-302; 1952:347); Carvalho and Rachou (1951:474-477, 480);

Pinotti (1951:670); Rachou and Ferraz (1951:542-543, 547-553); Rachou and Ricciardi (1951:

424-426, 432-437); Gabaldon and Cova Garcia (1952:178, 193, 201); Horsfall (1955:170); Be-

jarano (1956:9, 20; 1957:326-335; 1959:305, 314, 323, 325); Guedes, Amorim and Schreiber

(1957:247); de Andrade (1958b:117-125); Rachou (1958:148); Rachou, Moura Lima, Ferreira

Neto and Martins (1958:417, 421, 423); Castro (1959a:175); Castro and Garcia (1959:600);

Castro, Garcia and Bressanello (1959:549); Schreiber and Guedes (1959b: 128-129; 1960:356-

68 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

357; 1961:657-658); Stone, Knight and Starcke (1959:33); Vargas (1959:370, 383); Cerqueira

(1961:124); Forattini (1962:311, 394-395); Garcia and Ronderos (1962:137-139, 158); Belkin,

Schick and Heinemann (1971:6); Consolim and Galvao (1973:177-179); Knight and Stone

(1977:63).

Anopheles noroestensis of Foote and Cook (1959:29, 32); Schreiber and Guedes (1959a:97-98);

Maciel (1962:473-775); Cova Garcia (1964:200); Mattos and Xavier (1965:272-273); Gorham,

Stojanovich and Scott (1967:17, 47, 58; 1973:113, 138, 147).

Anopheles (Nyssorhynchus) oswaldoi var. noroestensis of Galvis (1938a:52, 54-56, 58; 1940: 432-

437); Galvao and Amaral (1938:14); Galvao and Lane (1938:176-177); Fonseca and Fonseca

(1942:99); Oliveira and Verano (1942:355); Rozeboom (1942a:240).

Anofeles (Nyssorynchus) osvaldoi (!) noroestensis of Unti (1940a:378).

Anopheles oswaldoi noroestensis of Galvao (1938b: 101-103); Russell, Rozeboom and Stone (1943:

51); Vargas (1943:59).

Anopheles (Nyssorhynchus) oswaldoi in part of Oliveira and Verano (1942:353-358).

Anopheles (Nyssorhynchus) oswaldoi var. metcalfi of Galvao and Lane (1938:175-176); Galvao

(1940:428-433; 1943:142-143); Correa and Ramos (1942a:37, 40-44, 46-47; 1942b:381-386);

Galvao and Damasceno (1942:123-125); Rozeboom (1942a: 239).

Anopheles (Nyssorhynchus) oswaldoi var. ayrozai of Rozeboom (1942a:240); Correa and Ramos

(1944b:130).

Anopheles clarki of Russell, Rozeboom and Stone (1943:44); Cova Garcia (1964: 200).

Anopheles evansi of Gabaldon (1949:765).

Anopheles (Nyssorhynchus) tarsimaculatus in part of Root (1922a:322, ?; 1926b:684-700, 709-

711; 1932:779); Davis (1928:549-553); Shannon and Del Ponte (1928:42, 44-56, 53-54); Shan-

non and Davis (1930:488); Shannon (1931:10, 22); Lane (1939:27-29); Pinto (1939:390-398);

Martinez and Prosen (1951:39, ?); Bejarano (1957:333-335); Castro (1959a:173-174).

Anopheles tarsimaculatus in part of Pinto (1930:154, 156); Kumm (1932:1-6); Unti and Ramos

(1942:94, 99-100).

Anopheles (Nyssorhynchus) tarsimaculata in part of Root (1923:271; 1924b:461-463).

Anopheles tarsimaculata in part (?) of Muehlens, Dios, Petrocchi and Zuccarini (1925:262-265).

Cellia tarsimaculata in part (?) of Peryassu (1921b:183); Neiva and Pinto (1922a:321; 1922b:356-

357); Godoy and Pinto (1923:29-33).

Cellia albimana in part (?) of Neiva (1909:69-77); Lutz, Souza Araujo and Fonseca Filho (1918:

162).

Cellia albipes in part (?) of Bourroul (1904:35).

FEMALE (fig. 6). Wing: 3.3 mm. Proboscis: 2.1 mm. Palpus: 2.0 mm. Forefe-

mur: 1.4mm. Abdomen: about 2.7 mm. Head: Integument reddish brown to dark

brown. Proboscis about 1.5 length of forefemur. Long, erect, white, cuneate scales

on vertex. Occiput with predominantly cream to golden scales. Palpal segment 2

with a few lighter scales on dorsal surface; segment 3 usually with a gray to white,

dorsolateral stripe; apex of segment 2 usually with a few light scales; apex of segment

3 with a broad, distinct, white band; segment 4 with a dark basal band and a smaller,

dark, apical band, often dark scales extending along ventral surface to form a stripe;

segment 5 white with dark basal band extending from apex of segment 4. Antenna:

Flagellar segment 1 with numerous, short, white, obovate scales on basomesal, dorsal

and dorsolateral surfaces, and a distinct patch of long setiform scales distally on dor-

sal and dorsomesal surfaces. Flagellar segments 2-13 each with about 7-9 long, silver

setae in a basal whorl. Thorax: Integument as on head. Median anterior promon-

tory area with numerous, long, white, setiform scales extending dorsad onto anterior

of acrostichal line; lateral promontory area with numerous light scales. Silver to

cream scales on scutum large. Prescutellar space small to moderate, subtriangular or

horseshoe shaped. Upper stp with 4, 5 long, dark setae in a horizontal to vertical arc

and a patch of gray to cream scales. Lower stp usually with a long dark seta and a

patch of gray to cream scales. Upper mep with about 4-7 long, brownish setae and

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 69

occasionally 1, 2 small scales. Legs: Light scales on coxae pale gray or cream, usual-

ly not white. Foretarsal segment 2 cream to white in apical 0.26-0.55; segment 3

with a broad, cream to white band in apical 0.55-0.80; segment 4 all dark to 0.5 api-

cally light; segment 5 cream, gray or golden in about apical 0.3-0.5. Midtarsal seg-

ment 2 cream in about apical 0.1; segments 3 and 4 predominantly dark, with a few

gray or golden apical scales; segment 5 often light in about apical 0.5. Hindtarsal

segment 2 with a dark band in basal 0.25-0.35 (0.2-0.4); segment 5 cream to white in

about apical 0.5. Wing: Light spots on wing yellow to cream, not white, at least on

anterior veins. Vein C humeral light spot more than twice, 2.0-4.5 (1.75-4.50),

length of basal dark spot; usually subbasal light spot greater than or equal to subba-

sal dark spot; rarely subbasal light spot absent, or subbasal and presectoral dark spots

fused into one large spot; often presectoral light spot absent and the presectoral and

sectoral dark spots fused; subcostal light spot moderately large, 0.30-0.45 (0.25-

0.46) of subcostal dark spot; preapical light spot moderately large, 0.30-0.43 (0.17-

0.43) of preapical dark spot; apical dark spot small, but variable. R presectoral dark

spot much larger than sectoral dark spot; sectoral dark spot often absent. Dark pre-

apical spot of R, large, conspicuous, 0.45-0.54 (0.40-0.54) of vein. Apical dark spot

of R; large, occasionally broken by a few light scales. M sectoral and preapical dark

spots moderately large to large, conspicuous; preapical dark spot not extending

onto M,,,. Cu, subcostal dark spot usually smaller than subsequent preapical dark

spot. Apical light fringe spot moderately large to large, conspicuous and unbroken

to apex of R4,.; additional small light fringe spots at the apices of Rais, Mis2, Mea,

Cu,, Cu, and A; occasionally a light fringe spot at level of 0.5 distance from base of

A. Abdomen: Caudolateral scale tufts of segments II-VII with moderately large, ob-

ovate scales.

MALE. Wing: about 3.4mm. Proboscis: 2.55 mm. Forefemur: 1.55 mm. Abdo-

men: about 2.75 mm. Essentially as in female except for sexual characters. Head:

Palpal segments 2 and 3 with a dorsal line of cream scales; apex of segment 2 with a

small white band; apex of segment 3 with a large white band; segment 4 with a

brown basal band, rest of segment golden to white; segment 5 usually with a brown

basal band extending along ventral surface to very near apex; dorsally and dorsolater-

ally remainder of segment 5 cream to white. Antenna: About 0.75 length of probos-

cis. Flagellar segment 1 with numerous, elongate, white setiform scales on dorsome-

sal margin. Legs: Forefemur about 0.6 length of proboscis. Basal plantar surface of

foretarsal segment 5 with about 6-8 moderately long, distinct, spiniform setae. Claw

on foreleg long, moderately curved; submedian tooth about 0.25-0.30 length of claw,

sharp and apically recurved; basal tooth subequal to submedian tooth, heavy and de-

curved.

MALE GENITALIA (fig. 13). In general very similar to that of oswaldoi and gal-

vaoi. Segment VIII: Tergite and sternite with narrow, long, obovate scales. Segment

IX: Sternite similar to that of oswaldoi, subtrapezoidal, moderately long, about 0.2

length of sidepiece. Anterior apodeme long, about 0.3 length of sternite and subtra-

pezoidal. Sidepiece: Moderately narrow and subconical. Tergal surface usually with

4, 5 long, submedian tergomedial bristles and 1, 2 subapicolateral bristles; with mod-

erately short bristles mesad of tergomedial bristles, often sparse with only 2, 3 short

bristles present, more basal and apical bristles occasionally longer than rest. Tubercle

of parabasal spine appearing short, less than 0.5 of parabasal spine. Basal apodeme

moderately long, usually 0.25 or slightly less length of sidepiece. Longer, more dor-

sal accessory spine slightly less than 0.5 of sidepiece; more ventral spine about 0.75

of longer spine. Internal spine subequal to shorter accessory spine, moderately to

70 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

strongly retrorse apically. Clasper: Spiniform long and moderately heavy; seta b

about 1.5 of spiniform. Dorsal Claspette: Pedicel moderately broad, sinuous; base

rounded and weakly curved mesad. Leaflets about 0.35-0.45 length of claspette,

broad; basomesal projection of dorsal leaflet very reduced. Ventral Claspette: About

0.35-0.45 length of sidepiece. Lateral margins tapered toward a narrow apex, width

at apex 0.20-0.36 length of claspette and slightly wider than apex of aedeagus. Basal

lobules large, expanded laterally, with long, often reflexed setae; setae on basal mar-

gin about 1.5-1.8 width of aedeagus; setae not concentrated on basomesal margin.

Ventral and lateral surfaces (exclusive of basal lobule) with short setae about 0.5-0.7

width of aedeagus; setae extending to or nearly to apex. Apex with rounded, sclero-

tized lateral margins, separated by a shallow median sulcus. Preapical plate large,

semicircular to oval, occasionally concave basally, heavily sclerotized, located about

0.3 length of claspette basad of apex. Transparent membranous area immediately

basad of preapical plate in shape of inverted U. Refringent structure distinct, in

shape of an inverted V, with or without lateral arms. Phallosome: Aedeagus sube-

qual to length of ventral claspette; apex appearing distinctly truncate; leaflets absent.

PUPA (fig. 13). Abdomen: about 2.6 mm. Trumpet: 0.40 mm. Paddle: 0.75 X

0.55 mm. Cephalothorax: Darkly pigmented. Wing cases often with dark, sclero-

tized, longitudinal stripes. Hair 2-C 1-3 branched, longer than 1,3-C; 1,3-C 2,3

branched. Hairs 4,5-C moderately developed; 4-C 1-3 forked; 5-C 2,3 forked (1-4).

Hair 6-C 1,2 forked, long. Hair 7-C 2,3 branched, long, one branch usually about 1.5

length of other branch(es). Trumpet: Pinna moderately to heavily pigmented; mod-

erately long, about 3.3-4.1 (3.3-4.3) length of meatus; in lateral aspect, appearing

broad medially and tapered toward apex (not as it appears in fig. 13). Meatal cleft

usually basally pointed. Metanotum: Hair 10-C single and strong. Hair 11-C 3-5

branched, 1.0-1.3 length of 10-C. Hair 12-C single to triple, long, about 1.4-2.0

length of 10-C. Abdomen: Chaetotaxy in general very similar to that of oswaldoi.

Hair 2-I 5-8 branched, moderately long. Hair 3-I single, about 0.5-1.0 length of 2-I.

Hair 4-I 4,5 branched. Hair 5-I single or double, long. Hair 6-I single and long. Hair

7-I 2,3 branched, about 0.5 length of 6-I. Hair 9-I single, longer than 7-I and shorter

than 6-I. Hair O-II-VII moderately developed; O-II,III 4-7 branched (3-7); 0-IV-VII 3-

5 branched (2-6). Hair 1-II,III 5-9 (4-9) and 6-10 (5-10) branched respectively,

strongly developed; 1-IV-VII strong and very long, 1.75-2.00 length of segment. Hair

2-IV-VII single or double, rarely triple, long, longer than that of oswaldoi, 0.5-0.7

length of segment. Hair 3-IV 3,4 branched (3-5), moderately long; 3-V 2,3 forked (1-

3), moderately long. Hair 5-III 6-11 branched (5-12), strongly developed; 5-IV 3-5

branched (2-5), median branches longer and stronger than lateral branches; 5-V-VII

heavy and long, usually equal to or slightly longer than segment. Hair 6-II-VI single

or double, long; 6-II about 1.5 length of 7-II. Hair 7-II 2,3 branched (2-5); 7-IH-V

and 8-III-VII moderately short; 7-III 2,3 branched (2-4); 7-IV 1-4 branched (1-5); 7-

V 1-3 branched; 7-VI,VII single, very long, about 0.7 length of segment; 8-III 2-4

branched (2-5); 8-I[V-VI 1-3 branched; 8-VII 3 branched (2-4). Hair 9-II thin, point-

ed, small, unpigmented; 9-III small, 2.5 or less length of 9-II; 9-IV thick, 2.0-3.0

length of 9-III, heavily sclerotized; 9-V heavy, 1.5-2.0 (1.1-2.0) length of 9-IV; 9-VI

strong, thick, weakly curved, 1.0-1.5 length of 9-V; 9-VII thick, curved, acuminate,

1.5-2.0 (1.4-2.0) length of 9-VI; 9-VIII straight, about equal to length of 9-VII; 9-II-

VI less than 0.33 length of segment; 9-VII,VIII 0.3-0.5 length of segment. Hair 10-

III 2,3 forked, moderately long; 10-IV,V single, very long. Hair 4-VIII 2,3 forked (1-

3). Terminal Segments: Male genital lobe heavy, thick at base, with sides sloping

toward apex; apex with distinct mammilliform protuberance. Paddle: Obovate,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 71

slightly emarginate at insertion of hair 1-P. Midrib basally distinct, apically becom-

ing indistinct. External buttress about 0.6 length of paddle. External margin distad

of buttress with very fine, filamentous spicules extending around apex and becoming

very sparse along caudal 0.5 of inner margin. Hair 1-P moderately short; 2-P single or

forked, subequal to 1-P.

LARVA (fig. 14). Head: 0.6 mm. Antenna: 0.28 mm. Anal Saddle: about 0.3

mm. Head: Darkly pigmented, reddish brown. Median tooth of mental plate usual-

ly wider than combined widths of 2 adjacent teeth from one side. Inner and outer

clypeal hairs (2,3-C) single and barbed in about apical half; 3-C about 0.8-0.9 length

of 2-C; hairs 2-C widely spaced, clypeal index about 1.7 (1.0-2.0). Hair 4-C 3,4

branched (2-5), moderately short to moderately long, 0.30-0.45 length of 3-C. Hairs

8,9-C often dendritic, moderately large, length about 1.5-2.0 distance separating hairs

5-C; 8-C 4-6 branched (4-7); 9-C 6-8 branched. Hair 10-C 3,4 branched (2-4); 12-C 4,

5 branched, large, subequal to 9-C. Hair 15-C 4-7 branched, moderately large. Collar

very dark reddish brown, moderately wide to wide dorsomedially. Antenna: Slight-

ly darker than remainder of head. Spicules along mesal margin moderately long and

thin; shorter and fewer on ventral surface. Hair 1-A 4-8 branched, short, about as

long as width of antenna at point of insertion. Thorax: Reddish brown or greenish

brown. Inner submedian prothoracic group (1-3-P) usually with hairs 1,2-P not shar-

ing a common tubercle; palmate hair 1-P with 10-14 moderately narrow, pointed,

lanceolate leaflets; 2-P 13-18 branched (12-18). Hair 14-P 8,9 branched (7-10), large;

branches long (0.3 mm), arising from a moderately short shaft, median branches

longer than lateral and extending beyond anterior margin of thorax. Mesothoracic

hair 1-M strongly plumose, 26-30 branched (26-32). Metathoracic hair 2-T long, ex-

tending beyond caudal margin of thorax. Palmate hair 3-T with 11-15 moderately

long, narrow, semitransparent leaflets. Abdomen: Integument pigmented as thorax.

Hair O-II-VII moderately large to large; O-II,III 6-9 branched (5-10); O-IV-VII 5-7

branched (5-8). Palmate hair 1-I with 15-18 (13-18) semitransparent, narrow, spread-

ing leaflets; 1-II-VII with pigmented, moderately narrow, pointed leaflets; 1-II-V with

21-30 leaflets; 1-VI with 24-25 leaflets; 1-VII with 19-28 leaflets; 1-II,VII slightly

smaller than 1-III-VI. Hair 2-II 6-9 branched (4-9), strongly developed, large; 2-III 4-

6 branched, large, more than 2.0 length of leaflets of 1-III and stronger than 2-II; 2-

IV single, moderately long, about 1.5 length of leaflets of 1-IV; 2-V very long, 3.0-

4.0 length of leaflets of 1-V. Hair 5-I 3-5 branched, moderately short, inserted less

than 0.75 its length from lateral margin of abdomen; 5-II 8-10 branched, moderately

developed. Hair 9-1 4,5 branched, moderately large. Hair 11-I 3,4 branched, large.

Hairs 13-[,IL,III 5-9, 7-9 (6-9) and 8,9 (8-12) branched respectively, moderately devel-

oped; 13-IV 3,4 branched, moderately long, subequal to 2-IV and about 1.5 length of

leaflets of 1-IV; 13-V almost always 3 branched (3-5) at least on one side of larva,

very large, subequal to 2-V, about 3.0-4.0 length of leaflets of 1-V and extending be-

yond caudal margin of segment; 13-VI 8-12 branched (6-12), moderately developed.

Spiracular Lobe: Pecten with 15-20 teeth; arrangement of teeth variable, but similar

to that of oswaldoi; teeth usually as follows beginning ventrally: (1) 1 long, (2) about

4-6 moderate followed by 2-5 mixed moderate to short, (3) 1 long, (4) 1-5 mixed

moderate to short and (5) usually 1 or 2 long occasionally followed by a short tooth;

occasionally with another long tooth interspersed in dorsal half of pecten; serrations

on teeth moderate in length. Lateral arm of spiracular apparatus short to moderately

short, directed caudolaterally. Hairs 8,9-S 4-7 and 5,6 (4-7) branched respectively,

moderately large. Anal Segment: Most of segment covered with fine spicules more

strongly developed apically. Hair 1-X about 1.5-2.0 length of saddle; inserted on sad-

ae: Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

dle near ventral margin, or occasionally on ventral margin of saddle at base of inden-

tation. Anal gills long, usually longer than saddle.

DISCUSSION. An. noroestensis can be distinguished from the other species in the

Oswaldoi Subgroup in the female by the combination of (1) palpal segment 4 often

with a dark ventral stripe, (2) hindtarsal segment 2 with a dark band in basal 0.25-

0.35 (0.2-0.4), (3) foretarsal segment 3 with a light band in apical 0.55-0.80, (4) light

wing spots yellow to cream (at least on anterior veins), and humeral light spot of vein

C usually greater than 2.0 of basal dark spot, (5) subcostal light spot of vein C mod-

erately large, about 0.30-0.45 of subcostal dark spot, (6) sectoral and preapical dark

spots of vein M moderately large to large and (7) apical light fringe spot moderately

large to large and conspicuous; in the male genitalia by the (1) apically truncate aede-

agus subequal to length of ventral claspette, (2) basal lobule of ventral claspette large,

expanded laterally, with setae along basal margin long, about 1.5-1.8 width of aedea-

gus, (3) apex of ventral claspette narrow, 0.20-0.36 length of claspette and (4) pre-

apical plate large, semicircular to oval and heavily sclerotized; in the pupa by the

combination of (1) pinna moderately long, about 3.3-4.1 length of meatus, appearing

broad medially and tapered toward apex, (2) hair 9-V relatively short, 1.5-2.0 (1.1-

2.0) length of 9-IV, (3) hair 9-VII 1.5-2.0 (1.4-2.0) length of 9-VI, (4) hair 9-VII, VIII

longer than in oswaldoi, 0.3-0.5 length of segment, (5) hairs 2-V and 7-VI long, usu-

ally more than 0.5 length of segment, (6) hair 7-C with one branch about 1.5 length

of other branch(es), (7) hair 2-I 5-8 branched and moderately long, 3-I about 0.5-1.0

length of 2-I, (8) hair 6-II about 1.5 length of 7-II and (9) hair 1-P moderately short;

and in the larva by the combination of (1) hairs 2,3-C barbed in about apical half,

hairs 2-C widely spaced, clypeal index about 1.7 (1.0-2.0), (2) hair 4-C 3,4 branched

(2-5), moderately short to moderately long, 0.30-0.45 length of 3-C, (3) hair 14P 8,

9 branched (7-10), large, with long branches, (4) hair 13-IV 3,4 branched, moderate-

ly long, about 1.5 length of leaflets of 1-IV, (5) hair 2-T long, extending beyond cau-

dal margin of thorax, (6) hair 2-IV moderately long and 2-V very long, 3.0-4.0 length

of leaflets of 1-V, (7) hair 5-I moderately short, inserted less than 0.75 its length from

lateral margin, (8) hair 13-V very large, about 3.0-4.0 length of leaflets of 1-V, almost

always 3 branched (3-5) at least on one side of larva and (9) lateral arm of spiracular

apparatus short to moderately short, directed caudolaterally. An. noroestensis is dif-

ficult to diagnose in all stages, except for the male genitalia. The larva is almost iden-

tical with that of rangeli, except for the position of hairs 4-C and 5-I,II, and the rela-

tive lengths of several of the abdominal hairs. The pupa is very similar to those of

strodei, benarrochi and oswaldoi. The adult is like those of strodei and anomalophyl-

lus, except for the coloration of the scales of the thorax, coxae and wing, and the

banding pattern of the foretarsus.

An. noroestensis is most closely allied to oswaldoi and galvaoi, and, to a somewhat

lesser extent, to aquasalis. The male genitalia differ from those of galvaoi and oswal-

doi primarily in the shape of the apex of the aedeagus, and from that of oswaldoi in

the shape of the preapical plate. The basal band on the second hindtarsal segment of

the adult is the main character separating noroestensis from aquasalis, galvaoi and os-

waldoi. But because intraspecific variation is so great, individuals at the extremes

(with either very small or very large second hindtarsal dark bands) cannot be distin-

guished from the adults of the latter 3 species. Ecologically, noroestensis occupies a

different niche than either aquasalis or oswaldoi; it is found in fresh water in second-

ary growth areas, usually not in forested areas or coastal brackish water habitats.

The salivary and autosomal chromosomes were examined by Guedes, Amorim and

Schreiber (1957) and Schreiber and Guedes (1959b; 1960; 1961). These authors

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 73

state that the X chromosomes are metacentric in neuroblasts and acrocentric in the

salivary cells, and that some heterozygous inversions and asynaptic zones have been

found in the autosomes.

An. noroestensis and metcalfi were first described as varieties of oswaldoi by Gal-

vao and Lane at a meeting of the Sociedade de Biologia de Sao Paulo, 15 September

1937. The transcripts of this presentation were not published until 1938 in a fest-

schrift for Lauro Travassos. In the interim, Galvao and Lane (1937e) described, in an

_abbreviated form, these new varieties and included them in keys along with illustra-

tions of the eggs. The description of metcalfi was based on a description of an egg by

Root (1926b:700) from Brazil. In comparing the surface ornamentation of the eggs,

Root stated, “The species of the Nyssorhynchus group seem not to show the elon-

gate hexagonal marking, so conspicuous in the eggs of such species as quadrimacula-

tus or pseudomaculipes. Instead, one finds the whole ventral and lateral portion of

the egg studded with little white stars, each consisting of eight or ten short lines radi-

ating from an imaginary common center.” Because of the statement regarding the

‘little white stars,’ Galvao and Lane assumed that Root was describing a new and

different species, unlike either noroestensis, oswaldoi or tarsimaculatus. In 1940,

Galvao stated that in the state of Sao Paulo metcalfi must be rare or must not exist in

that region, since no eggs had ever been collected with the characteristics described

by Root. Later, Galvao and Damasceno (1942:123-125) reported that the eggs illus-

trated by Root (1926b) could have been those of oswaldoi, noroestensis or tarsima-

culatus, and that it was impossible to tell what species his description referred to.

They wrote, “*. . . julgamos que o A. oswaldoi metcalfi nao pode ser caracterizado pe-

lo descricao de Galvao e Lane (1938) e o consideramos como nomen nudum.”’ By

nomen nudum the authors, I believe, meant nomen dubium, since nomen nudum re-

fers to a name not satisfying Articles 12-16 of the International Code of Zoological

Nomenclature. As Mayr (1969:347) states, ““A nomen nudum has no standing in zoo-

logical nomenclature and is best never recorded, not even in synonymy.”’ Whereas a

nomen dubium refers to a name for which there is insufficient evidence for recogni-

tion of the species to which the name was applied. In the same publication, Galvao

and Damasceno (1942:125-129) raised noroestensis to specific rank based on the egg,

larva, male genitalia and female. In 1971, Belkin, Schick and Heinemann (p 6) desig-

nated a male (96) with its genitalia on a slide (710210-1), collected by Root, 29 May

1925 in Porto das Caixas (Rio de Janeiro), as the lectotype of metcalfi. They stated

that the lectotype was, “1 of several specimens identified as tarsimaculatus by Root

(1926:711), on which Galvao and Lane based their metcalfi (USNM).”’ After careful

examination of this lectotype, Iam synonymizing metcalfi with noroestensis. The

apex of the aedeagus clearly appears truncate, and the shape of the preapical plate

corresponds to that of noroestensis. The dark basal band of hindtarsal segment 2 is

0.35 the length of the segment, also as in noroestensis. All the other characters exa-

mined correlate with those found in noroestensis. Additional specimens were collec-

ted near the type locality of metcalfi by J. N. Belkin, H. C. Barnett, G. K. Bryce and

myself, December 1975 to February 1976 in the state of Rio de Janeiro. After study-

ing these individually reared specimens and correlating the characters in the imma-

tures and the adults, I have identified these specimens as noroestensis.

BIONOMICS. The immatures of noroestensis have been collected in both perma-

nent and temporary water in drainage ditches, small ground pools, potholes and along

stream margins. The immatures are usually found in fresh water, either exposed to

the sun or partially shaded. The water is frequently turbid and brownish. One collec-

tion from the state of Rio de Janeiro was from a flooded grassy yard highly contami-

74 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

nated with pig and chicken feces. The immatures were all collected in regions of sec-

ondary growth or in areas of cultivation, and not in the forest, although they may oc-

cur along the edge of forested areas. Species that have been collected in association

with noroestensis are Anopheles (Nys.) albitarsis, Culex (Cux.) coronator group, Cx.

(Cux.) sp, Cx. (Mel.) sp and Uranotaenia ( Ura.) lowii. Additional larval habitats re-

ported in the literature are muddy puddles, rock holes and wells (Deane, Causey and

Deane 1948:878-880), and marshes and swamps (Galvao 1940:433). Deane, Causey

and Deane (1948) showed that noroestensis was highly sensitive to sodium chloride

and would not develop to the third instar even in a dilute solution of 0.5%. They al-

so reported collecting noroestensis at altitudes greater than 700 m in the state of Ce- .

ara, Brazil. Galvao (1938b:102) reared noroestensis in the laboratory at a mean tem-

perature of 25.7°C and showed that the development from egg to adult required

about 13 days.

Most reports indicate that noroestensis is not very anthropophilic or endophilic.

Of the 1342 females collected in northeastern Brazil by Deane, Causey and Deane

(1948), only 53 or 3.9% were collected in houses. Forattini (1962:394-395) also re-

ported that noroestensis was not domestic or anthropophilic. Contrary to these au-

thors, Correa and Ramos (1942a:37-47) indicated that noroestensis (as metcalfi) was

largely anthropophilic and endophilic. Deane, Causey and Deane (1948) stated that

noroestensis was most abundant during the rainy season (June 1941) in Ceara. In 2

days in June, they collected 447 females on animal bait, representing 96.9% of the

anophelines captured. During the dry season in December over a 2 day period, they

collected only 12 adults representing 35.3% of the anophelines caught. In host pre-

ference studies made between 1730 and 1930 hours in June, Deane, Causey and

Deane (1948) collected 38 females on a horse and only one on a man. Biting peaked

around sundown, and there was another small peak of activity at dawn, although

some individuals could be captured throughout the night.

MEDICAL IMPORTANCE. An. noroestensis does not seem to be important as a

primary vector of malaria. Correa and Ramos (1942a:37; 1942b:385-386) dissected

24 females of “metcalfi’’ which were collected inside houses in Ribeira, Sao Paulo,

Brazil and found 2 (8.3%) with an oocyst infection of Plasmodium sp; however, I am

not certain that the species these authors reported as metcalfi was actually noroesten-

sis. Nevertheless, a photograph of the head capsule of the supposed metcalfi in Cor-

rea and Ramos (1942b:383) corresponds to that of noroestensis, as does the habitat

in which these mosquitoes were collected. Deane, Causey and Deane (1948), after

studying noroestensis in many different localities in the northeast of Brazil, stated that

all indications are that this species is not important in the transmission of malaria.

Forattini (1962:311) reported that it is possibly a secondary vector. An. noroesten-

sis from Guaratingueta and Guaruja, Sao Paulo, Brazil has been experimentally infect-

ed with Plasmodium vivax by Fonseca and Unti (1943:47-52).

DISTRIBUTION (fig. 2). An. noroestensis is distributed throughout central and

southeastern South America. Its northernmost limits are the southern margins of the

Amazon basin and the northeastern states of Brazil (Maranhao, Ceara). In the west,

noroestensis extends to the eastern versant of the Andes. An. noroestensis is reported

to occur as far south as the provinces of Chaco, Corrientes, Formosa, Misiones, Salta,

Santa Fe and Tucuman in Argentina (Garcia and Ronderos 1962:161).

Material Examined: 178 specimens: 22 males, 18 male genitalia, 57 females, 19

pupae, 62 larvae; 19 individual rearings: 9 larval, 7 pupal, 3 incomplete.

ARGENTINA (7 specimens: 2M, 3Mgen, 2F). Tucuman: Concepcion, N. Davis, 3Mgen (clarki)

[JH]. Monteros, Jun 1940, 2M, 2F.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 13

Wa ERE ARI tenia

BOLIVIA (17 specimens: 6M, 4Mgen, 7F). Santa Cruz: Charagua, 16 May 1944, 3153,4F. La

Guardia, nr Santa Cruz, 27 Sep 1943, Torres Munoz, 3083, BOL 21, 4M, 3Mgen. Department and

locality not specified: Carr, No. 2919, 2M, 1Mgen, 2F; same collector, 1F.

BRAZIL. Bahia (47 specimens: 1M, 9F,37L): C. de Maria, 1F [JH]. Escondido, 1F [JH]. Ita-

pebi, 3L [JH]. Itapira, Mar 1931, 1F. Jequie,4F [JH]. Jitauna, 1F [JH]. Pindobacu, 2L [JH].

Poiri, 1L [JH]. Salvador da Bahia, 1972, J. Fowler, BRS 82, 3L; same data except Mar 1972, BRS

101, 1L; same data, BRS 110, 1L; same locality and collector, 1972, BRS 116, 3L; same data, BRS

127,4L; same data, BRS 147, 1L; same data, BRS 149, 1L; same data, BRS 164, 3L; same data,

BRS 171, 2L. Rio Ipitanga, 16 Mar 1972, J. Fowler, BRS 92,3L. Santo Amaro, 1L [JH]. Santo

Antonio, 1F [JH]. Locality not specified, thru M. Boyd, 1M; 8L [JH]. Ceara (1F): Guaramiranga,

det. Deane, 1F [CU]. Maranhao (1Mgen): Locality not specified, IMgen (10.190) [JH]. Pernam-

buco (3M): Locality not specified, D. Lucen, 3M. Rio de Janeiro (86 specimens: 9M, 9Mgen, 27F,

19p, 22L&1; 19 ind rear: 91, 7p, 3inc): Bangu, F. Root, 1Mgen (105) [JH] ; same locality, 3Mgen

(109) [JH]. Itaquai, 19F [JH] ; same locality, BRA 282. Mage, 1F [JH]. Seropedica, BRA 230,

231. Varzea das Mocas, BRA 253, 291. Rio Grande do Norte (2L): Rio Cajupiranga, on road to

San Jose, 6 Jan 1945, H. Dalmat, 1L [CU]. Rio Pirangi, Cajupiranguinha, 1 Nov 1944, H. Dalmat,

1L [CU]. Sao Paulo (1M, 11F): Novo Oriente, 1938, A. Galvao, J. Lane, 1M, 11F. State not spe-

cified (1Mgen, 1L): Manibu, Rio Araray, 18 Oct 1944, H. Dalmat, 1L (21) [CU]. Locality not spe-

cified, F. Root, 1Mgen (109-25) [JH].

5. Anopheles (Nys.) aquasalis Curry

Figs. 2,4, 6, 15, 16

1932. Anopheles (Nyssorhynchus) tarsimaculatus var. aquasalis Curry 1932:566-572. TYPE:

Syntypes male with genitalia slide, female and larva, Atlantic side of Canal Zone (no type

locality cited although Colon and Cristobal mentioned), D. P. Curry et al. [LU] (Belkin,

Schick and Heinemann 1965:44).

1941. Anopheles (Nyssorhynchus) emilianus Komp 1941:794-800. TYPE: Holotype male with

genitalia slide, northern outskirts of Belem (Para), Brazil, reared from larva collected in

grassy pool, 14 Apr 1941, W. H. W. Komp [USNM]. Synonymy with aquasalis by Causey,

Deane and Deane (1945:247-248). |

1942. Anopheles (Nyssorhynchus) oswaldoi var. guarujaensis Ramos 1942:61-63. TYPE: Syn-

types adults, larvae, eggs, Ilha do Guaruja (Sao Paulo), Brazil, Mar 1939, collected in part

by Seccao de Ecologia (A. Carvalho Franco) [LU]. Synonymy with tarsimaculatus by

Galvao and Damasceno (1942:131). Synonymy with aquasalis by Lane (1949:402).

1948. Anopheles aquasalis var. guarauno Anduze 1948:18. TYPE: Syntypes adults, reared from

eggs of the normal aquasalis ‘“‘typicus,”’ Pedernales (Delta Amacuro), Venezuela [NE].

Considered as subspecies by Lane (1953:260).

1948. Anopheles aquasalis var. delta Anduze 1948:18. TYPE: Syntypes adults, reared from eggs

of the normal aquasalis “‘typicus,’”’ Pedernales (Delta Amacuro), Venezuela [NE]. Consid-

ered as subspecies by Lane (1953:260).

1978. Anopheles (Nyssorhynchus) deltaorinoquensis Cova Garcia, Pulido F. and Amarista M.

1978:150-161. TYPE: Syntypes adults, eggs, larvae and pupae, holotype not specified

according to Article 73(b), La Brea, Punta Gorda and Las Parcelas (Sucre), Caribe, Zona de

Maturin and Boca de Guanipa (Monagas), and Capure, Punta de Tolete, Pedennales, Isla de

Guanoco and Boca de Capurito (Delta Amacuro), Venezuela [DERM]. NEW SYNONY-

MY.

Anopheles (Nyssorhynchus) aquasalis of Senevet and Abonnenc (1938:487-493); Gabaldon (1940:

4-5); Gabaldon and Aguilera (1940:66-68, 72-80); Gabaldon, Cova Garcia and Arevalo (1940:

25-28); Gabaldon, Cova Garcia and Lopez (1940:10, 11, 13-16); Gabaldon, Lopez and Ochoa

Palacios (1940:37-38); Galvao (1941 :94; 1943:142-143); Rozeboom (1941:102; 1942a:242);

Rozeboom and Gabaldon (1941:92-98); Galvao and Damasceno (1942:130-131); Komp (1942:

39, 40, 73-74, 81, 122-124, 132, 159-160); Simmons and Aitken (1942:38, 46, 54, 62, 87-90);

Floch and Abonnenc (1943a:1-9; 1943b:1-8; 1945:2, 4-8, 11, 13-16; 1946b:3-5; 1947:1-5:

1951:44-48); Causey, Deane and Deane (1944:2, 4, 5, 6; 1945:243-250; 1946:27); Lane (1944:

263-266; 1949:402; 1953:259-261); Levi-Castillo (1945:105-116; 1949:10, 11, 15, 27, 32, 58,

76 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

67, 72, 76, 81, 85); Deane, L. M., Causey and Deane (1946:2-3, 7-10; 1948:881-894); Deane, M.

P., Causey and Deane (1946:38, 42, 45); Arnett (1947:198); Coher (1948:87-88); Vargas (1948:

(158; 1959:377); van der Kuyp (1949a:67-68); Downs (1950:29-30); Pinotti (195 1:667-668,

673, 681-682); Senior-White (195 1a:293-403; 1951b:465-512); Gabaldon and Cova Garcia

(1952:179, 182, 189); Frizzi and Ricciardi (1955:399-407); Horsfall (1955:178-181); Vargas V.

(1956:29; 1957:1; 1958a; 1958b); Bejarano (1957:326, 327, 329, 333); Rozeboom and Kitz-

miller (1958:244); Schreiber and Guedes (1959b:128-129; 1960:356-357); Stone, Knight and

Starcke (1959:31); Cerqueira (1961:121-122); Fauran (1961 :8-9); Forattini (1962:311, 382-

388); Belkin, Schick and Heinemann (1965:44); Kitzmiller and Chow (1971:65-85); Kreutzer,

Kitzmiller and Rabbani (1975:363-364); Cova Garcia and Sutil O. (1976:31; 1977:18, 47, 63,

88); Cova Garcia, Pulido F. and Amarista M. (1978:155, 157).

Anopheles aquasalis of Rozeboom (1941:102); Kumm and Zuniga (1942:399); Downs, Gillette

and Shannon (1943:33-42); Ross and Roberts (1943:33-34); Russell, Rozeboom and Stone

(1943:26, 29, 30, 31, 42); Eyles (1944: 12-13); Deane and Damasceno (1948:501-508); Deane,

Vernin and Damasceno (1948:793-795, 804-807); Senior-White (1950:5-6; 1954); Floch (1954:

3-5; 1956:45-46); Vargas V. (1958c:1, 3); Foote and Cook (1959:114); Gabaldon and Guerrero

(1959:433-434); Giglioli (1959:280; 1963:131-145); Cova Garcia (1961 :35-36, 69, 87-88, 113,

124-125, 154; 1964:199-214); Fauran (1962:73-76; 1963:44-45; 1964:51-52); Sellers, Bergold,

Suarez and Morales (1965:466-469); Fauran and Courmes (1966:104-105; 1967:70); Stojano-

vich, Gorham and Scott (1966a:12, 19, 30; 1966b:21, 31, 39); Gorham, Stojanovich and Scott

(1967:18, 48, 50; 1973:114, 139, 141); Serie, Kramer and Chatenay (1968:198-200, 202-203,

208, 209); Hamon, Brenques and Chauvet (1970:33); Elliott (1972:757); Panday (1975a:297-

301; 1975b:305-308; 1977:732, 734); Cova Garcia and Sutil O. (1975a:23; 1975b:212).

Anopheles (Nyssorhynchus) emilianus of Bruce, Knigin, Yolles and Graham (1943:441); Galvao

(1943:142); Causey, Deane and Deane (1945:243-250); Cova Garcia and Sutil O. (1976:31;

1977:18, 47, 63, 88); Cova Garcia, Pulido F. and Amarista M. (1978:155, 157).

Anopheles (Nyssorhinchus) (!) emilianus of Cova Garcia (1964: 199-217).

Anopheles emilianus of Russell, Rozeboom and Stone (1943:37, 42, 45); Cova Garcia and Sutil O.

(197 5a: 235 1975b:212).

Anopheles (Nyssorhynchus) tarsimaculatus of Root (1922a:322, in part?; 1926b:684-700, 709-

711, in part; 1932:781, in part); Christophers (1924:39-40, 89, in part); Dyar (1925:187-188,

195, in part?; 1928:439-440); Kumm (1929:101-103, in part); Hill (1930:712); Shannon and

Davis (1930:488, in part?); Shannon (1931:10, 22); Edwards (1932:45, in part?); Senevet

(1932:252, in part?; 1937:358-360; 1938:179-181; 1948c:434-439); Antunes and Lane (1933:

97, in part); Davis (1933:278, 280, in part?); Earle (1936:459-469); Komp (1936a:161-162, in

part); Galvao and Lane (1937a:77-78; 193 7e:213-216; 1938:169, 170, 172-173, in part); Gal-

vao, Lane and Correa (1937:40-41); Galvao and Amaral (1938:13); Root and Andrews (1938:

566-578); Rozeboom (1938a:96, 100-101, 104; 1938b:289, 293); Lane (1939:27-29, in part);

Pinto (1939:398, in part?); Galvao (1940:421-425; 1941:92, in part?; 1943:141-143, 149, 151);

Komp (1941a:93, 95, 97); Senevet and Quievreux (1941:248); Floch and Abonnenc (1942b:2);

Galvao and Damasceno (1942: 129-130); Galvao, Damasceno and Marques (1942:51-111); Bar-

retto and Coutinho (1943:321); Correa (1943:124, 126, 127); Coutinho (1943:65-77); Ramos

(1943:51-62); Correa and Ramos (1944a:105-108; 1944b:131; 1944c:10-11); Galvao, Correa

and Grieco (1944:85-102); Galvao, Lane and Unti (1944:45); Amaral and Penido (1947:164-

165, 168, 173-180); Vargas (1948:157-158); Carvalho and Rachou (1951:474-477, 480); de An-

drade (1953a:95-107; 1953b:145-156; 1958a:17-30); Senevet and Andarelli (1955:339, 342);

Bejarano (1957:333-335, in part); Rachou (1958:161-168).

Anopheles tarsimaculatus of Edwards (1922:75); Root (1922b:392); Cleare (1926:404-406); Hoff-

man (1930:366); Benarrochi (193 1:691-693); Davis (1931:45-47, 49-50, in part); Shannon

(1931:10, 22, in part); Beattie (1932:477-496, in part); Kumm (1932:1-6, in part); Buxton

(1934:491-494); Hill (1934:425-428); Earle (1936:460-465, 467-468); de Verteuil and Spence

(1937:450, 454-455); Kumm and Novis (1938:503); Kumm and Ruiz (1939:438, 441); Kumm,

Komp and Ruiz (1940:388, 389, 391, 412, 419); Kumm (1941a:359); Vargas (1941:118); Unti

and Ramos (1942:93, in part?); Oliveira, de Andrade and Nascimento (1951:151-247).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) Ve

Nyssorhynchus tarsimaculatus in part (?) of Lima (1928:95-98); Townsend (1933b:7-12).

Anopheles (Nyssorhynchus) tarsimaculata in part of Root (1923:276; 1924b:461, 462).

Anopheles tarsimaculata of Dyar and Knab (1906b: 160-161, in part); Busck (1908:58); Knab

(1913:36, 37, in part); Zetek (1915:221-271); LePrince and Orenstein (1916:43-114); Howard,

Dyar and Knab (1917:975-979, in part?); Bonne (1923:127-128, in part); Dyar (1923:185, in

part); Muehlens, Dios, Petrocchi and Zuccarini (1925:252-254, 257, 264-265); Senevet (1931:

79-82, in part?).

Anopheles (Anopheles) tarsimaculata in part of Bonne-Wepster and Bonne (1923:127); Bonne and

Bonne-Wepster (1925:511-515).

Anopheles (Cellia) tarsimaculata in part (?) of Dyar (1918:151).

Cellia tarsimaculata in part (?) of Neiva and Penna (1916:94); Peryassu (1921a:70; 1921b:181-

183); Neiva and Pinto (1922b:356-357); Godoy and Pinto (1923:29-33); Bonne (1924: 133-

137); Boyd (1926:31, 36, 48).

Anopheles albimanus var. tarsimaculata of Evans (1921:460; 1922:213).

Cellia albimana in part (?) of Peryassu (1908: 120-121); Neiva (1909:69-77); Theobald (1910:69-

70); Godoy and Pinto (1923:29-33).

Anopheles albimanus in part of Nicholls (1912:251-268).

Anopheles albipes of Gray and Low (1902:194).

Cellia albipes in part (?) of Theobald (1903:110-113; 1905b:11); Bourroul (1904:33, 63, 75);

Peryassu (1908:61).

Anopheles (Nyssorhynchus) oswaldoi var. metcalfi (2) of Lucena (1940a:107-112; 1940b: 176-177).

FEMALE (fig. 6). Wing: 3.3 mm. Proboscis: 2.1mm. Palpus: 2.0 mm. Forefe-

mur: 1.4mm. Abdomen: 2.85 mm. Head: Integument brown to dark brown. Pro-

boscis about 1.5 length of forefemur. Palpal segment 2 with a few, light, erect scales

on dorsal or dorsolateral surface; segment 3 with a gray to white, dorsal or dorsolater-

al, longitudinal stripe beginning 0.3-0.5 from base and usually ending near apex, with

a white apical band larger than light apical band of segment 2; base and apex of seg-

ment 4 with dark scales occasionally extending onto base of 5; remainder of seg-

ments 4 and 5 white; ventral surface of 4 usually without a distinct stripe of dark

scales. Antenna: Flagellar segment 1 with elongate, oblanceolate, white scales distal-

ly on dorsomesal surface, and often with shorter scales on dorsolateral surface; basal-

ly with short, obovate, white scales on dorsal or dorsolateral surface. Flagellar seg-

ments 2-13 each with basal whorl of 8-10 long, curved, silver setae. Thorax: Integu-

ment dark brown to almost black. Prescutellar space generally horseshoe shaped, oc-

casionally triangular. Ppl with 2 (1-3) long, dark setae. Sp with 3-9 moderately long,

light setae. Pra with 5-10 long setae, and a patch of about 7 long, broad, whitish

scales. Upper stp with about 4 dark setae, and about 8 (6-11) gray to white, broad,

obovate scales distributed in horizontal arc. Lower stp with 1, 2 long setae and a

patch of about 6 gray to white, obovate scales. Upper mep with 3-6 long, dark setae;

scales absent. Legs: Light scales on coxae light cream to grayish, not white. Fore-

tarsal segment 2 cream to white in about apical 0.25 (0.15-0.36); segment 3 highly

variable, light in apical 0.40-0.65 (0.40-0.70) in Central and South America, and light

in apical 0.15-0.40 in the Lesser Antilles; segment 4 all dark or with a few, golden,

apical scales; segment 5 with apical 0.3-0.7 golden to cream or gray. Midtarsal seg-

ment 2 with a cream band in apical 0.1-0.3; segment 3 with a small, apical, golden

band; segment 4 all dark; segment 5 golden in apical 0.3-0.5. Hindtarsal segment 2

with brown band in basal 0.40-0.55 (0.3-0.6). Wing: Highly variable, characters list-

ed below represent the usual conditon and some of the common variations. Dark

spots on posterior veins distinct. Light spots at least on veins C and R light cream to

yellowish, occasionally white on posterior veins. Vein C with humeral light spot 1.8-

3.5 (1.5-3.5) of basal dark spot; basal dark spot usually 0.5 its length from humeral

cross vein; occasionally presectoral light spot absent; very rarely both subbasal and

; 78 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

presectoral light spots absent, so that subbasal, presectoral and sectoral dark spots

fused into one large, dark spot; subcostal light spot about 0.35 (0.3-0.5) of subcostal

dark spot, occasionally longer in specimens from the Lesser Antilles; preapical light

spot about 0.45 (0.3-0.6) of preapical dark spot; apical dark spot moderately small.

Rs-R,,3 usually with an extra subcostal light and dark spot. R3; variable, with 2 or 3

light and dark spots, occasionally apical dark spot absent. R,4,; subcostal and preapi-

cal dark spots moderately small. M sectoral dark spot distinct, usually moderately

long although variable; preapical dark spot not extending onto M,,,. Vein A subcos-

tal dark spot moderately long, about 0.1-0.2 length of A. Apical light fringe spot

small to moderate; additional small light fringe spots at apices of R4.5, Mjs2, M34,

Cu,, Cu, and A; a moderately small, inconspicuous, light fringe spot at level of 0.5

distance from base of A. Abdomen: Sternite I with a few, moderately long setae.

Dark caudolateral scale tufts on tergites and sternites II-VII with scales long, broad

and very conspicuous. Tergite densely covered with narrow, moderately large, light

- gray to golden scales in subtriangular pattern on segments II, III and rectangular pat-

tern on [V-VIII.

MALE. Wing: 3.11 mm. Proboscis: 2.55 mm. Forefemur: 1.5 mm. Abdomen:

about 2.8 mm. Essentially as in female except for sexual characters. Head: Palpal

segment 2 and almost always segment 3 with a light dorsal stripe; segment 4 with

pale golden, cream or white scales mediodorsally and laterally, usually with a basal

and an apical, dark band, and scattered, inconspicuous, dark scales ventrally. Anten-

na: About 0.75 length of proboscis. Flagellar segment 1 with numerous, long, cur-

ving, white, lanceolate scales on dorsomesal surface. Legs: Forefemur about 0.7

length of proboscis. Claw on foreleg curved and pointed; submedian tooth recurved

at apex, about 0.2 length of claw; basal tooth subequal to submedian tooth, strongly

decurved and blunt.

MALE GENITALIA (fig. 15). Segment VIII: Scales on tergite and sternite nar-

row, lanceolate to oblanceolate and predominantly light; scales somewhat narrower

on sternite than on tergite. Segment IX: Sternite long, about 0.2 length of sidepiece,

subtrapezoidal. Anterior apodeme subtrapezoidal and moderately long, about 0.15

length of sternite. Sidepiece: Moderately narrow, conical. Tergal surface usually

with 3, 4 long, submedian tergomedial bristles and one subapicolateral bristle; with

moderately short bristles mesad of basal tergomedial bristles and longer bristles later-

ad of more apical tergomedial bristles. Parabasal spine usually at least 2.0 length of

its tubercle. Basal apodeme about 0.2 length of sidepiece. Longer, more dorsal ac-

cessory spine about 0.5 length of sidepiece; more ventral spine about 0.70-0.75 of

longer spine. Internal spine subequal to shorter accessory spine, moderately to

strongly retrorse apically. Clasper: Spiniform blunt, moderately long, slightly less

than or equal to length of seta b. Dorsal Claspette: Pedicel moderately broad; base

broad and rounded, weakly curved mesad. Leaflets broad; dorsal leaflet with large,

prominent basomesal projection. Ventral Claspette: Lateral margins tapered toward

a moderately narrow apex, width at apex about 0.3 length of claspette. Basal lobule

broad and semispherical, covered basally and laterally with radiating, moderately

short setae; setae along basal margin about equal to or slightly longer than width of

aedeagus; setae shorter on mesal margin and more or less uniformly distributed on

lobule. Lateral and ventral surfaces (exclusive of basal lobule) with short setae; setae

0.5 or less width of aedeagus; setae extending to or nearly to apex. Apex with round-

ed lateral margins and a shallow median sulcus; sulcus with gently sloping sides. Pre-

apical plate moderately small, circular to oval, weakly to heavily sclerotized, located

about 0.3 length of claspette basad of apex. Region immediately basad of preapical

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 79

plate with a moderately broad, transparent, horseshoe shaped membranous area. Re-

fringent structure in shape of inverted V. Phallosome: Aedeagus about 1.20-1.35

length of ventral claspette; apex rounded, slightly wider than long; leaflets absent.

PUPA (fig. 15). Abdomen: about 2.8 mm. Trumpet: 0.5 mm. Paddle: 0.75 X

0.55 mm. Cephalothorax: Dark brown to brownish black. Wing cases often with

longitudinal sclerotized stripes. Hairs 1-3-C 2,3 branched (1-3); 1-C longest. Hairs 4,

5-C 2-4 forked (1-4). Hair 6-C 1-3 forked (1-4). Hair 7-C 2,3 branched, often caudal

branch about 1.5-2.0 length of other branch(es). Hairs 8,9-C subequal in length; 8-C

single; 9-C 1-3 forked (1-4). Trumpet: Pinna pale yellow, with margin moderately

pigmented; moderately long, about 3.5-4.3 (3.5-4.7) length of meatus; in lateral as-

pect, appearing broad medially and tapered toward apex. Meatal cleft long and basal-

ly rounded. Metanotum: Hair 10-C single or occasionally double, short, about 0.50-

0.75 length of 12-C. Hair 11-C 1-5 forked (2-6), equal to or slightly longer than 10-

C. Hair 12-C 1-3 forked about 0.3 from base, long. Abdomen: Pigmented as on

cephalothorax. Hair 2-I 4-8 branched (2-10) near or at base, moderately short, most

branches subequal. Hair 3-I single or double, about 0.5 length of 2-I. Hair 41 4-6

branched (3-8), moderately long. Hair 5-I 1-3 branched, moderately long, slightly

shorter than 6-I. Hair 6-I single and long. Hair 7-I 3-7 branched, 0.50-0.65 length of

6-I. Hair 9-I single, 0.65-0.85 length of 6-I. Hair O-II-VI 2-6 branched (2-8); 0-VII 1-

3 branched (1-5). Hair 1-IL, IT] +9 branched (3-11), moderately long, median branch-

es longer than lateral; 1-I[V-VII very long, 1.50-1.75 length of segment. Hair 3-IV 3-

8 branched (2-8), moderately developed; 3-V 2-4 forked (1-5), moderately long. Hair

5-III 3-6 branched (3-9) and 5-IV 2,3 branched (1-4), median branches longer than

lateral; 5-V-VII single, subequal to length of segment. Hair 6-II single or double,

about 2.0 length of 7-II; 6-III 1-3 forked, moderately long; 7-II 3-7 branched (2-7).

Hairs 7,8-III-V 2-5 branched; 8-III-VII and 7-III,1V small; 7-V-VII 1-4 forked, moder-

ately long. Hair 9-IIT minute, unpigmented; 9-III small, less than 2.0 length of 9-II; 9-

IV usually blunt, always 2.0 or more length of 9-III; 9-V-VII moderately slender, cur-

ved, acuminate, long; 9-V 2.4-4.0 (2.3-5.0) length of 9-IV, slightly less than or equal

to 0.5 length of segment; 9-VI 1.0-1.3 length of 9-V; 9-VII 1.1-1.5 length of 9-VI, us-

ually greater than 0.5 length of segment; 9-VIII subequal to and straighter than 9-VII.

Hair 10-III 2-4 forked (1-4). Hair 4-VIII 1-3 branched (1-4). Terminal Segments:

Apex of male genital lobe with large, distinct, mammilliform protuberance. Paddle:

Light brownish yellow. Large, obovate, moderately broad, emarginate subapically at

insertion of hair 1-P. Midrib not reaching apex. External buttress about 0.6 length

of paddle, serrated apically with very small teeth. External margin distad.of buttress

with fine, short, filamentous spicules extending around apex and becoming shorter

along caudal 0.7 of inner margin. Hair 1-P strong and moderately long; 2-P 1-3

forked (1-4), weak, subequal to 1-P.

LARVA (fig. 16). Head: 0.6 mm. Antenna: 0.27 mm. Anal Saddle: 0.3 mm.

Head: Heavily pigmented, mottled over much of ventral and dorsal surfaces, reddish

brown to dark brown, lighter in ocular region. Inner and outer clypeal hairs (2,3-C)

plumose in apical 0.3-0.7, with 7-12 simple, rarely dendritic, moderately short to

long branches; 3-C occasionally with slightly longer branches than 2-C; hairs 2-C

widely spaced, clypeal index 1.67. Hair 4-C 1-3 branched, short to moderately short,

length usually less than or equal to distance between insertion of hairs 2-C and 3-C;

inserted moderately close to moderately far from anterior margin of head. Hair 8-C

3-5 branched (2-6) near base, slightly longer than distance separating insertion of

hairs 5-C. Hair 9-C dendritic, 3-6 branched (3-8), about 1.3 length of 8-C. Hair 12-C

3-5 branched (3-6), subequal to 9-C. Collar wide dorsomedially, moderately sclero-

80 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

tized, yellowish brown to reddish brown. Antenna: More heavily sclerotized than

remainder of head. Mesal margin with numerous, strongly developed spicules thicker

than in oswaldoi. Hair 1-A 4-7 branched (3-10), moderately short, always less than

2.0 width of antenna at point of insertion. Thorax: Reddish brown to dark reddish

brown or dark greenish brown, often darker medially. Submedian prothoracic group

(1-3-P) with or without hairs 1,2-P sharing a common tubercle; often hair 1-P arising

from very weakly pigmented tubercle; palmate hair 1-P with 10-15 (10-18) moderate-

ly narrow leaflets; 2-P 16-20 branched (13-23), about 4.0 length of leaflets of 1-P.

Hair 14-P 6-10 branched from a short, flattened stalk; lateral branches equal to or

slightly shorter than median. Mesothoracic hair 1-M 25-40 branched, lanceolate in

outline, lateral branches longer than basal and apical. Metathoracic hair 2-T long,

usually extending to about caudal margin of thorax. Palmate hair 3-T with 10-15 (9-

19) semitransparent, moderately long, spreading leaflets. Pro-, meso- and metathora-

cic pleural group spines long, strongly developed. Abdomen: Pigmented as on thor-

ax, sternum lighter than tergum. Hair O-II-VI 3-7 branched (2-8), moderately devel-

oped. Palmate hair 1-I with 11-15 (10-17) semitransparent, moderately narrow leaf-

lets; 1-II, VII with 18-26 broad, pointed leaflets, smaller than 1-III-VI; 1-III-VI with

with 20-30 (20-32) leaflets. Hair 2-II 3-6 branched, strongly developed; 2-III 2-5

branched, large; 2-IV single, moderately long; 2-V 1-3 branched, very long. Hair 5-I

3,4 branched (3-5), branches subequal and moderately long; 5-II 5-9 branched (49).

Hair 9-I 4-7 branched (3-8). Hairs 13-LII,III 4-7 (3-7), 5-9 (5-11) and 5-9 (5-13)

branched respectively, small; 13-IV 4-6 branched (3-8), moderately large; 13-V 4-7

branched (3-7), large; 13-VI 8-13 branched (7-13), moderately small. Spiracular

Lobe: Pecten with 12-19 teeth; median teeth mixed moderate and short. Lateral

arm of spiracular apparatus very short. Hairs 8,9-S 3-5 branched (2-6). Anal Seg-

ment: Covered, except for base, with very fine, short spicules becoming stronger

toward apex. Saddle rusty brown, with irregular ventral margin. Hair 1-X longer

than saddle; inserted on ventral margin of saddle at base of indentation, or on saddle

near ventral margin; inserted about 0.3 cephalad of caudal margin. Anal gills usually

short, about 0.5 length of anal segment, very rarely equal to length of saddle.

DISCUSSION. An. aquasalis can be distinguished from the other species in the Os-

waldoi Subgroup in the female by the combination of (1) palpal segment 4 usually

without a distinct ventral stripe of dark scales, (2) light scales on coxae light cream to

grayish, not white, (3) foretarsal segment 3 highly variable, light in apical 0.40-0.70

in Central and South America and light in apical 0.15-0.40 in the Lesser Antilles, (4)

hindtarsal segment 2 brown in basal 0.40-0.55 (0.3-0.6), (5) vein C with humeral light

spot large, 1.5-3.5 length of basal dark spot, and with basal dark spot usually about

0.5 its length from humeral cross vein, (6) subcostal light spot on vein C usually less

than 0.5 subcostal dark spot, (7) apical light fringe spot small to moderate, (8) dark

spots on posterior veins distinct and (9) light wing spots at least on veins C and R

light cream to yellowish, not white; in the male genitalia by the combination of (1)

apex of ventral claspette moderately narrow, width at apex about 0.3 length of clas-

pette, (2) basal lobule broad with uniformly distributed setae, setae along basal mar-

gin equal to or slightly longer than width of aedeagus, (3) lateral and ventral surfaces

of ventral claspette (exclusive of basal lobules) with short setae 0.5 or less width of

aedeagus and (4) preapical plate moderately small, oval to circular; in the pupa by the

combination of (1) hair 7-C with one branch about 1.5-2.0 length of other branch(es),

(2) pinna of trumpet moderately long, about 3.5-4.3 (3.5-4.7) length of meatus, with

margin moderately pigmented, and with a long, basally rounded meatal cleft, (3) hair

10-C single, about 0.50-0.75 length of 12-C, (4) hair 12-C 1-3 forked, long, (5) hair

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 81

2-I +8 branched (2-10) near or at base, moderately short, (6) hair 3-I about 0.5

length of 2-I, (7) hair 7-I 3-7 branched, 0.50-0.65 length of 6-I, (8) hair 9-III small,

less than 2.0 length of 9-II, and 9-IV always 2.0 or more length of 9-III, (9) hair 9-V

2.4-4.0 (2.3-5.0) length of 9-IV and (10) hair 9-V-VIII usually about 0.5 length of

segment; and in the larva by the combination of (1) hairs 2,3-C plumose, with 7-12

simple, rarely dendritic branches, and hairs 2-C widely spaced, clypeal index about

1.67, (2) hair 4C 1-3 branched, short to moderately short, (3) hair 8-C slightly long-

er than distance separating insertion of hairs 5-C, (4) pecten with median teeth mixed

medium and short, (5) lateral arm of spiracular apparatus very short, (6) saddle with

ventral margin irregular, hair 1-X inserted on ventral margin at base of indentation, or

on saddle near ventral margin and (7) anal gills usually short, about 0.5 length of anal

segment, rarely longer. The adults of aquasalis are easily distinguished in most cases

from all other species in the Oswaldoi Subgroup except for benarrochi. Even so,

aquasalis can usually be distinguished from benarrochi by the characters given in the

above discussion and by its geographical distribution. Because of the paucity of ma-

terial available for study of benarrochi, an attempt has not been made to separate

aquasalis from the former in the key to the adult females. I believe that separation

of these 2 species in the key would lead to unnecessary misidentification.

There does not seem to be any consistent geographical differentiation in aqua-

salis except when the females from Central and South America are compared with

those from the Lesser Antilles, and then only in the banding pattern of the third fore-

tarsal segment. The apical light band of foretarsal segment 3 on the mainland speci-

mens varies from 0.40-0.70 the length of the segment, whereas the apical light band

of the island specimens is in general much less than half (0.15-0.40) the length of the

third foretarsal segment.

The amount of interpopulational and intrapopulational variation is very great in

aquasalis from the Lesser Antilles. On first examination of the material from this re-

gion, | believed, as did Senior-White (195 1a:306-307), that possibly I was looking at

a different subspecies or even species. On further examination, I found that in al-

most every population from which I had a fair number of specimens, there occurred

individuals that were almost identical with those examined from mainland and other

island localities, suggesting a single, although highly polymorphic species. The great-

est diversity was found among the specimens from the island of St. Lucia. In the

adults from the Lesser Antilles, there are considerable differences in size and occa-

sionally in color. Many of the adult males from St. Lucia show a reduction in the

amount of white on palpal segments 4 and 5. In the females, the light banding on

the midtarsi and the length of the wing spots also varies. Vein C occasionally has

large subcostal and preapical dark spots, and vein M may be almost entirely dark.

Some specimens from St. Lucia exhibit a fusion of the subbasal, presectoral and sec-

toral dark spots on vein C to form one large spot. In contrast, several specimens

from Dominica exhibit a large subcostal light spot on the costal vein, often longer

than 0.5 the length of the subcostal dark spot. In the male genitalia from the Lesser

Antilles, the preapical plate varies from dark to light, and from a small rounded plate

to a larger, lighter, more oval, diffuse plate; the refringent structure may be in the

shape of an inverted U or V;; the setae on the ventral claspette vary from short to

moderately long; and the parameres may be short and thick at their bases, or long

and slender. In the larvae and pupae from the Lesser Antilles, most of the variation

occurs in the branching of the hairs; this is included in the data on chaetotaxy in the

descriptions. In general, the pupae demonstrate considerably more variation than do

the other stages; some of the variation includes the length and thickness of hair 9-IV-

82 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

VIII, the extent of branching of hair 1-I and the general pigmentation. In the larvae,

the frontal ecdysial suture may be rounded or pointed. Usually the branches of hairs

2,3-C are long and simple, although occasionally they are dendritic; this is particular-

ly evident in the St. Lucia specimens, while on the other hand, one specimen from

Costa Rica had short clypeal branches. Also in the larvae, hair 4-C is often single and

long in specimens from northern Brazil, Trinidad, Tobago and the Lesser Antilles.

The pecten can vary in the number and the length of the teeth, with the St. Lucia

specimens again demonstrating the greatest variability. The length of the anal gills

varies apparently depending on the type of aquatic habitat; occasionally larvae have

moderately large anal gills, usually when collected in fresh water.

The great amount of diversity within and sometimes among populations of aquasa-

lis on the islands in the Lesser Antilles may possibly be attributed to 3 factors. First,

under island conditions there may be decreased interspecific competition (fewer spe-

cies present) that may result in a shifting and/or expansion of a species’ niche, which

may in turn be reflected in some external morphological character. Second, the

founder effect is possibly an important factor; when only a few individuals colonize

an island, genetic drift can result. And third, species may be subjected to quite dif-

ferent selection pressures on islands than they are on the mainland; the different se-

lection pressures may then result in a phenotype different from that on the mainland.

Although the immatures of aquasalis are found only occasionally in fresh water in

Central and South America, many of the specimens examined from the Lesser Antil-

les have been collected in fresh water, even in rain barrels. This indicates that at least

some niche expansion has occurred on certain of the islands.

The closest allies of aquasalis are oswaldoi, galvaoi and noroestensis. As previously

mentioned (p 72), the male genitalia of these 4 species are very similar, differing pri-

marily in the shape and sclerotization of the preapical plate, the length of setae on

the ventral claspette and, in the case of noroestensis, the shape of the aedeagus. An.

aquasalis does not share as many characters with galvaoi, oswaldoi and noroestensis

as these share with each other; however, the larvae and, to some degree, the pupae of

all 4 are similar.

The first cytogenetic studies of the salivary gland chromosomes of aquasalis were

performed by Frizzi and Ricciardi (1955:403-407) on specimens from Sao Bento and

Duque de Caxias, Rio de Janeiro, Brazil. These workers reported no chromosome ab-

normalities. Later, Kitzmiller and Chow (1971) examined the salivary gland chromo-

somes of aquasalis from 2 populations in Brazil, one collected near Belem and an-

other collected near Rio de Janeiro. They found no inversions and stated, “‘aquasalis,

albimanus and oswaldoi show a close morphological relationship.”” Further, when

comparing the extent of regions of homologous banding, the chromosomes of aqua-

salis were more similar to those of oswaldoi than to those of albimanus. In compar-

ing the chromosomes of nuneztovari with albimanus, aquasalis and darlingi, Kitzmil-

ler, Kreutzer and Tallaferro (1973:443) found the “‘strongest”’ ae homologies

between nuneztovari and aquasalis.

An. aquasalis, for many years prior to its description by Curry in 1932, was re-

ferred to as tarsimaculata (or tarsimaculatus) as is evident in the references listed in

the synonymy (p 76-77). As early as 1915, Zetek recognized this brackish water

mosquito in Panama and considered it a racial variety. Much of the information re-

garding tarsimaculatus has been summarized earlier in the Taxonomic History chap-

ter (p 5-7). In 1941, Komp (1941b) described emilianus from Belem, Brazil, and

stated that it could be separated from aquasalis by the shape of the aedeagus, the

slightly longer setae covering the ventral claspette (both of which he states are sub-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 83

ject to variation and are rather indefinite) and, most easily, by the egg. Causey,

Deane and Deane (1945:247-248) studied 7000 ovipostions from isolated females of

aquasalis and found considerable variation among the eggs even within a single ovipo-

sition. In addition, they noted, “Variations in the male genitalia as great as the dif-

ferences described by Komp between emilianus and aquasalis may also be found

among mosquitoes from the same oviposition.’’ After examining the holotype adult

male and its genitalia, and several other specimens from the type locality of emilian-

us, | agree with Causey, Deane and Deane (1945:247-248) and Floch and Abonnenc

(1945:7) in the synonymy of emilianus with aquasalis. In 1942, Ramos described a

variety of oswaldoi, guarujaensis, from the island of Guaruja in the state of Sao Paulo,

Brazil. Galvao and Damasceno (1942:131) synonymized guarujaensis with tarsimacu-

latus which was later synonymized with aquasalis by Lane (1949:402). As with

many species in the Albimanus Section, mutants have been described as varieties.

Anduze (1948) described 2 varieties of aquasalis from Venezuela based on adults,

guarauno and delta. An. guarauno supposedly has a black apical ring on the fourth

hindtarsal segment, and delta supposedly has 2 additional apical dark rings on hind-

tarsal segments 3 and 4 (the ring on segment 4 being more pronounced than on 3).

Recently, Cova Garcia, Pulido F. and Amarista M. (1978) described deltaorinoqu-

ensis from Sucre, Monagas and Delta Amacuro, Venezuela. I have examined one spe-

cimen each of the male, male genitalia, female, pupa and larva labelled “‘A. deltaorin-

oquensis, 2697,’ which I received from Dr. P. Cova Garcia through Dr. J. N. Belkin.

With the exception of the hindtarsal segments and the wing of the adults, and a few

other characters in the immatures summarized below, all stages closely resemble

those of aquasalis. The observed differences which distinguish these specimens are

in the female (1) hindtarsal segment 2 dark in basal 0.66, (2) hindtarsal segment 3

with a small dark band in basal 0.1, (3) hindtarsal segment 4 with a small dark band

in basal 0.16 and (4) vein C subcostal light spot small, 0.15 length of C subcostal

dark spot; in the pupa (1) hair 2-I long (0.16 mm) and (2) hair 5-III,IV 7,9 and 4-6

branched respectively; and in the larva (1) clypeal hair 3-C with distinctly longer

branches than 2-C, some branches dendritic, (2) hair 14-P 4,5 branched and (3) hair

13-1 8,9 branched. Cova Garcia, Pulido F. and Amarista M. also state that in the lar-

va, the lateral arms of the spiracular apparatus are long, reaching the spiracular open-

ings, and that in the male genitalia, the preapical plate is oval, similar in shape to a

““grano de cafe.” I did not see any significant differences from aquasalis in these lat-

ter characters in the available specimens. These authors report that the immatures of

deltaorinoquensis were collected along the coast or along waterways, in brackish and

fresh water, at elevations of 0-20 m above sea level. The preferred habitat was brack-

ish mangrove swamps of Rhizophora mangle, Laguncularia racemose and/or A vicen-

nia nitida. Additional immatures were encountered in puddles of rainwater and

ground pools bordering rivers, streams and lakes. Most of the collections were from

amber colored water with a high organic content. The immatures were associated

with the following additional aquatic plants: Pistia stratiotes, Eichhornia sp, Lemna

sp, Azolla sp, Sagittaria guianensis, Salvinia sp, Utricularia sp. The pH of the water

tested varied from 6.6 to 8.4 and the salinity (gm/lit NaCl) from 7.5 to 17.4. In the

laboratory at 26-29°C, deltaorinoquensis required 18 days for development from egg

to adult (egg 2 days, larva 14 days, pupa 2 days). They report that of the 1681

adults captured, 1376 were deltaorinoquensis and 305 were emilianus (=aquasalis).

Further, they state that both the immatures and adults of emilianus are commonly

encountered in association with deltaorinoquensis. Nevertheless, they state that the

adults reared from eggs of emilianus never had spotted [hind] tarsi, and, in contrast,

«84 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

the legs of the adults reared from eggs of deltaorinoquensis were never white.

For the present, even though the differences enumerated above exist in the single

specimen of each stage I examined, I do not recognize deltaorinoquensis as a distinct

species. Geographically aqguasalis and deltaorinoquensis are sympatric. Also, both

“‘species’’ occupy a similar ecological niche, occurring primarily in brackish water ha-

bitats in relatively close proximity to the coast. In addition, as shown by Roze-

boom’s experiments (1963) with the mutant forms of albimanus (bisignatus with an

extra dark band on hindtarsal segment 3 and trisignatus with extra dark bands on

hindtarsal segments 3 and 4), a character or characters that “breeds true”’ in the la-

boratory from eggs isolated from a particular phenotype does not prove that this

phenotype represents a separate species (see Discussion section for rondoni, p 134).

Because of the great overall morphological, distributional and ecological similarity

between these nominal taxa, I believe that, until more conclusive evidence becomes

available, deltaorinoquensis should be considered a synonym of aquasalis.

BIONOMICS. As with albimanus, the literature on the bionomics and medical im-

portance of aquasalis is voluminous. I will present only a brief summary.

An. aquasalis is considered to be primarily a brackish water mosquito; however, it

has been collected from a wide variety of habitats in both fresh and brackish water.

It is more or less restricted to the coast and areas that are influenced by the tides, al-

though on certain occasions it has been collected considerable distances inland.

Deane, Causey and Deane (1948:882) collected aquasalis 60 km from the coast in

the state of Pernambuco, Brazil in a brackish water lake, Lagoa Salgada [Salty Lake].

In the Amazon basin, the same authors collected aquasalis 140 km from the coast on

the right and left banks of Rio Para. Recently, adults that resemble aquasalis have

been collected at numerous localities in the Maraba area, Para, Brazil (D. R. Roberts,

personal communication). Since these collections were made about 500 km from

the coast in fresh water, and the immatures have not been studied, I am reluctant to

state that these adults are definitely aquasalis. Interestingly, in this region these adults

are endophilic with an outdoor to indoor ratio of 3.9. Lucena (1946) collected aqua-

salis 252 km from the coast at Jupi in northeastern Brazil, at an altitude of 750 m

and in an area in which the soil had a high salt concentration. In all these instances,

except possibly in the Maraba area, salt seemed to be essential for the presence of

aquasalis inland from the coast.

The immatures have been collected in mangrove swamps, brackish marshes, feed:

ed meadows, rice fields, sugar cane plantations, ditches, fresh water ponds, pools

along stream margins, temporary and semipermanent ground pools, large artificial

containers and crab holes. Although there seems to be a preference for brackish wa-

ter, many of the specimens examined came from fresh water but usually no farther

than 5-8 km from the coast at altitudes of 200-300 m; its occurrence in fresh water

is particularly common in the Lesser Antilles. Most of the collections were from

aquatic habitats in full sun or partial shade, but some were in deep shade, such as in

mangrove swamps and in crab holes. Deane, Causey and Deane (1948:885-886) sta-

ted that most of their collections came from water with a sodium chloride concentra-

tion of 0.2-1.0%, although some came from water with a concentration as high as

1.5%. They also found that the number of pupae and adults from larvae reared in

0.5-1.5% sodium chloride solutions was greater than the number produced when lar-

vae were reared in fresh water. De Verteuil and Spence (1937:454) collected aqua-

salis in Trinidad in 70% sea water. Senior-White (1951a:310) reported that in the

wet season in Trinidad, breeding is in water not exceeding 0.35% sodium chloride,

but that during the dry season breeding occurs in water with a salt concentration of

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 85

2.38%. Causey, Deane and Deane (1945:247) showed that under laboratory condi-

tions, aquasalis can complete development in pure sea water, and, if the water is

allowed to evaporate, larvae continue to grow and pupate in water with a concentra-

tion as high as 7.4% sodium chloride. Floch and Abonnenc (1943a:7) collected lar-

vae in fresh water about 3 km from the beach in French Guiana; these survived when

placed in a 19 gm/lit solution of sodium chloride. They also placed larvae collected

in brackish water into fresh water without any apparent ill effects. De Andrade

(1958a:29) stated that aquasalis is an acidophilic species, and that larval density de-

creased as pH increased from 3 to 7. Buxton (1934:491) reported that larvae are

rarely found in water containing more than 0.3 ppm ammonia. Senior-White (195 la:

312), however, stated that saline or ammonia concentration “‘plays no part in regula-

ting either the oviposition or the larval life of A. aquasalis.’”’ The larvae seem to be

able to tolerate moderate amounts of pollution.

In Brazil (Deane, Causey and Deane 1948:884) and in Suriname (Panday 1975b:

305), the maximum density of aquasalis occurs during the rainy season. An. aquasa-

lis during this season is found in a greater number of habitats than in the dry season

when this species is more restricted to the coastal flats. Notwithstanding, in some

areas aquasalis may be more abundant locally during the dry season. Downs, Gillette

and Shannon (1943:39) found a higher density of aquasalis in Tobago and Trinidad

in the dry season, because at that time the mouths of streams are blocked by sand

bars which dam the streams and provide protected breeding habitats. Giglioli (1963:

132-133) discovered aquasalis to be particularly abundant during a drought in Guy-

ana and noted, “‘During dry periods, tidal salt water invasion of the estuaries and

lower reaches of the rivers extends far beyond the average level; thus, not only the

prevalence of A. aquasalis, but its range as well, is increased.”

The most common species collected in association with aquasalis in both fresh and

brackish habitats are An. albimanus, Culex (Cux.) nigripalpus, Cx. (Mel.) atratus and

Aedes (Och.) taeniorhynchus. In mangrove swamps and flooded meadows, Ae.

(Och. ) tortilis and Cx. (Mel.) madininensis are occasionally associated. Psorophora

(Gra.) cingulata group and Cx. (Cux.) coronator group are commonly associated with

aquasalis in fresh semipermanent ground pools and ditches, and Cx. (Mel. ) idottus

group and Uranotaenia (Ura. ) lowii breed with aquasalis in permanent fresh water

ponds. Occasionally aquasalis has been collected along stream margins in association

with Cx. (Cux.) declarator group, and occasionally in crab holes with Deinocerites

magnus. Other species that have been reported in association with aquasalis are tri-

annulatus (rarely), An. (Ano.) punctimacula, An. (Ano.) pseudopunctipennis, An.

(Ano.) apicimacula and Chagasia bathana. Common plant associates are Typha an-

gustifolia, Cyperus articulatus, Rhizophora sp and Avicennia sp (mangroves), Eleo-

charis mutata, Ipomoea reptans, Chara sp, Naias sp and Utricularia sp. Frequently

aquasalis is associated with mats of algae. |

The adults of aquasalis are crepuscular, usually being active from dusk to a few

hours after sundown. The adult females feed readily on man and on domestic ani-

mals such as cattle, horses, donkeys, dogs, sheep and pigs. They rest inside houses on

the walls normally lower than one meter (Deane and Damasceno 1948:508).

Throughout vast stretches of northeastern Brazil and in some areas of the Guianas

and Lesser Antilles, it is the only anopheline that occurs in large numbers in

houses; however, it is reported that less than 10% stay in the house for 24 hours

(Deane, Causey and Deane 1948:891). Feeding preference studies indicate that aqua-

salis is more zoophilic than anthropophilic, except in the Lesser Antilles where it

may be more anthropophilic. Nevertheless, when domestic animals are removed

. 86 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

from an area, which has happened in many places because of mechanization, man us-

ually becomes the primary host; under these conditions aquasalis becomes a major

health threat (Downs, Gillette and Shannon 1943; Giglioli 1959, 1963; Hamon,

Mouchet et al. 1970:33). The adults of aquasalis are considered to be strong fliers

capable of migrating considerable distances. In marked release and recapture studies

- conducted in Panama by LePrince and Orenstein (1916:114), individuals of tarsima-

culata (=aquasalis) marked with aniline dye traveled from 366 to 1905 meters from

the release station. De Verteuil and Spence (1937:454-455) reported that aquasalis

was able to fly at least 4.8 km in Trinidad; however, this was not experimentally ver-

ified.

MEDICAL IMPORTANCE. An. aquasalis has been a major vector of malaria along

the coast of Brazil south to the state of Parana, and in the Lesser Antilles. It is be-

lieved not to be a vector in Panama, probably because of its zoophilic behavior there.

In Venezuela and the Guianas, aquasalis has played a role in residual malaria trans-

mission (Gabaldon and Guerrero 1959; Floch 1956:45-46; Giglioli 1959:280; Ha-

mon, Mouchet et al. 1970:33). An. aquasalis has been extremely important as a ma-

larial vector in Trinidad, Tobago and in northeastern Brazil. De Verteuil and Spence

(1937:454) reported that 70% of the mortality due to malaria from 1931 to 1933

in Trinidad was a result of the presence of An. tarsimaculatus (=aquasalis) within 4

miles of brackish water. An. aquasalis has been incriminated as the major vector of

malaria in Grenada (Earle 1936; Root and Andrews 1938:577), St. Lucia (Earle

1936) and Guadeloupe (Fauran 1962:76). It has been found to be naturally infected

with Plasmodium sp in Brazil (Deane, L. M., Causey and Deane 1946; Correa and Ra-

mos 1944c; Coutinho 1943:69; Fonseca and Fonseca 1942:102-106, 110, 117),

Trinidad and Tobago (Downs, Gillett and Shannon 1943:35-37), Grenada and St.

Lucia (Earle 1936:463-465). In all the.other areas where it is important as a poten-

tial vector, it is considered to be dangerous only when it occurs in large numbers.

Thus, its vector effectiveness is dependent on adult density (Deane, L. M., Causey and

Deane 1946; Forattini 1962:385; Senior-White 1951b) and to some extent on the

presence of alternate hosts which serve to decrease the number of females feeding on

man.

An. aquasalis does not seem to be important in the transmission of filariasis, al-

though Deane, Causey and Deane (1948:893-894) found a filarial worm, Wuchereria

bancrofti, in the thorax of a female in Belem, Para, Brazil. Venezuelan Encephalitis

virus was isolated from one of 40 lots of aquasalis representing a total of 1537 speci-

mens tested in Venezuela during the VE outbreak in 1962-1964 (Sellers, Bergold et

al. 1965). Aedes taeniorhynchus appeared to be the major carrier during the above

outbreak. An. aquasalis may or may not be capable of transmitting the virus.

DISTRIBUTION (fig. 2). An. aquasalis is primarily restricted to the coast, coastal

lowlands and coastal waterways influenced by tides, except in the islands of the Les-

ser Antilles and possibly in Para, Brazil. It occurs in the state of Parana, Brazil, north

along the coast, west through the Guianas, Venezuela and Colombia, and into Cen-

tral America predominantly along the Atlantic side of Panama and Costa Rica to Nic-

aragua. On the Pacific side of South America, aquasalis is reported to occur as far

south as the province of El Oro, Ecuador on the shores of the Gulf of Guayaquil. An.

aquasalis is present in Trinidad, Tobago and the Lesser Antilles as far north as the

Leeward Islands of Antigua and St. Kitts, but it does not occur in the Greater Antil-

les, Virgin Islands or the Bahama Islands.

Material Examined: 2159 specimens: 432 males, 132 male genitalia, 839 females,

189 pupae, 567 larvae; 150 individual rearings: 77 larval, 52 pupal, 21 incomplete.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 87

ANTIGUA (62 specimens: 55F, 3p, 4L&1; 3 ind rear: 11, 2p). St. George: Fitches Creek, ANT

66,67. St. John: Five Islands, 8 Oct 1948, H. Box, 1F [BM1939-219] ; same locality, ANT 101.

Parish and locality not specified: 1 Dec 1899, 1F on slide [BM] ; 27 Sep 1950, 50F.

BRAZIL. Bahia (35 specimens: 1M, 25F, 9L): Bonfim, 26 Jan 1930, N. Davis, R. Shannon, 1

M, 25F. Salvador da Bahia, Mar 1972, J. Fowler, BRS 89, 3L; same locality and collector, 1972,

BRS 117, 1L; same data, BRS 144, 2L; same data, BRS 180, 1L; same locality and collector, 1972-

73, BRS 499, 2L. Ceara (1M, 2F): Aracati, Beirada, 1M, 2F [CU]. Maranhao (3M): [Sao Luis],

3M [JH]. Para (97 specimens: 20M, 8Mgen, 69F): Belem, Goeldi, KO H-20-28, 4F; same locality,

Apr 1930, N. Davis, 15F; same locality, 13 Apr 1941, W. Komp, BRAK 2, 9M, 6Mgen, 25F; same

locality and collector, 14 Apr 1941, 1M and 1Mgen (holotype of emilianus), 2F (paratypes of emil-

ianus); same locality and collector, Apr 1941, 3M, 3F (paratypes and allotypes of emilianus); same

data, 1M, 2F (paratypes of emilianus) [JH] ; same data, 3M, 15F; same locality, 1944, Causey, 1M;

same locality, Malinha, 1Mgen [JH] ; same locality, 1M (689-4), 1M (889-2) [JH] ; same locality, S.

E.S.P. No. 592, 1F (11.082-4) [JH] ; same locality, S.E.S.P. No. 597, 1F (11.352-1) [JH] ; same lo-

cality, S.E.S.P. No. 626, 1F (11.934-2) [JH]. Pernambuco (10 specimens: 1M, 3F, 6L): Boa Via-

gem, nr Recife, 24 Aug 1944, R. Wenzel, 3L; same data except 4 Sep, 1L; same data except 5 Sep,

1L; same data except 7 Sep, 1L. Recife, Pontazinha, May 1945, E. Coner, 1F [CU]. Recife, Ibura

Field, det H. Dalmat 1944, 1F (27-3-H-3) [CU]. Locality not specified, 1M (1269), 1F (1615).

Rio de Janeiro (26 specimens: 6M, 3Mgen, 2F, 7p, 8L&1; 7 ind rear: 41, 3p): Duque de Caxias,

BRA 227, 228. Itaguai, BRA 289. Itaipu, BRA 248. Rio de Janeiro, 1M (3551) [JH]. Sao Paulo

(3M, 3F): Sao Vincente, 3M [JH]. Locality not specified, A. Lutz (3F). State and locality not

specified (2Mgen): O. Causey, 1Mgen (689-2), 1Mgen (1269) [JH].

COSTA RICA (8 specimens: 2F, 1p, 5L&l; 1 1ind rear). Limon: Puerto Viejo, H. Kumm, No.

179, 1F [BM1938-696]. Westfalia, CR 473.

DOMINICA (105 specimens: 8M, 2Mgen, 31F, 10P&p, 54L&l; 6 ind rear: 4 1, 2inc). St. An-

drew: Portsmouth, 2 Mar 1964, D. Bray, 2F; same locality, 19-21 Oct 1966, A. Gurney, 1F; same

locality, L. Charles, 15L. St. David: Bataka, DOM 173. St. John: Cabrit Swamp, 23 Feb 1965,

W. Wirth, 4M, 20F. Cotton Hill Estate, DOM 97. Picard Estate, DOM 158. Prince Rupert Bay,

DOM 213. St. Joseph: Mero, DOM 77, 78,79. Parish not specified: Canefield Estate, 25 Apr

1944, L. Charles, 10L. Clarke Hall, 21-31 Mar 1965, W. Wirth, 3F. Locality not specified, 1927,

A. Carment, 1M [BM1928-132].

FRENCH GUIANA (109 specimens: 14M, 10Mgen, 51F, 15p, 19L&l; 15 ind rear: 4 LF. |

inc). Guyane: Cabassou (foret de), FGC 3278. Cabassou (Mont), FG 46. Cayenne, 14 Jul 1942,

FGA 72-596, 1F; same locality, Dec 1942, FGA 83, 5M, 4F; same locality, 13 Aug 1947, 1M(852),

1Mgen; same locality, 20 Aug 1947, 1 lp; same locality, 5 Apr 1948, 1 lpF; same locality, FGC

3417; same locality, IMgen. Cayenne, road to Baduel, 11 Jul 1942, FGA 71-10, 1M. Couachi,

FGC 4024. Fort Diamant, FGC 3293. Gallion (foret du), FGC 3381. Iracoubo, 12 Mar 1952,

FGA 184, 1F. La Chaumiere, 2 Jul 1942, FGA 67, 3F. Montjoly, FG 69, FGC 3438. Raban, FG

43. Rorota (Lac du), FG 67.

GRENADA (47 specimens: 16M, 8Mgen, 11F, 5p, 7L&1; 5 ind rear: 3 1, 2p). St. Andrew: Gren-

ville, GRR 12. Pearls Airport, GR 37. Pearls Bay, LAR 43, 44. Pearls Estate, LAR 46. St. George:

Caliviny, GR 18. Point Saline, GR 8. St. Patrick: Mount Rodney Estate, GR 46. Parish not speci-

fied: Pevis Port, LAR 51. Locality not specified, GRR 126; 28 Jun 1929. No. 1, 1b, 2p EI.

GUADELOUPE. BASSE TERRE (1F, 65L). Baie Mahault: Belcourt, FWI 816. La J aille, FWI

871,955. Jarry, FWI 1027. Capesterre: Roseau, FWI 1014. Lamentin: Blachon, FWI 1005.

Petit Bourg: Ste. Claire, FWI 1012. Viard, FWI 944, 988,989. GRANDE TERRE (120 speci-

mens: 4M, 3Mgen, 4F, 12P&p, 97L&1; 9 ind rear: 4 1, Ip, 4inc). Abymes: Besson, FWI 202. Rai-

zet, FWI 704, 851, 855, 927, 951, 959, 1022. Anse Bertrand: Port Louis, FWI 200, 929. Gosier:

Anse Vinaigri, FWI 967. Blanchas, FWI 818. Grande Ravine, FWI 876. Labrousse, FWI 958, 960.

Poucet, FWI 203. St. Feliz, FWI 931. Morne-a-l’Eau: Berville, FWI 865. Blain, FWI 973. Ste.

Anne: Anse du Belley, FWI 1004. MARIE GALANTE (119 specimens: 8M, 7Mgen, 12F, 32P&p,

60L&l; 20 ind rear: 101, 10p). Grand Bourg: Route de Folle Anse, FWI 236. St. Louis: St. Louis,

FWI 264.

-- 88 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

GUYANA (155 specimens: 11M, 7Mgen, 137F). East Berbice-Corentyne: New Amsterdam, 10

Mar 1900, E. Rowland, 1F [BM]; same locality, GUY 1, 17, 20A; same locality, Aiken, 1F. Local-

ity not specified, 12 Feb 1908, Aiken, 3F; 1936, W. Komp, 22F. East Demerara-West Coast Ber-

bice: Georgetown, 1931, W. Beveridge, 1M, 1F [BM1931-210] ; same locality, E. Rowland, 6F.

Hyde Park, 29 Jul 1941, BGR 2, 1M, 1Mgen, 3F; same data, 1F (1D) [JH]. North West: Locality

not specified, Sep 1949, BGR 7, 15F. District and locality not specified: 6 Mar 1941, GUYK 10,

1F; 10 Jul 1941, Fox, 2M [JH] ; Giglioli, GUYK 34, 1M, 1F; no data, 1F [JH].

MARTINIQUE (46 specimens: 8M, 6Mgen, 6F, 13P&p, 13L&l; 13 ind rear: 5 1, 6p, 2 inc).

Anse d’Arlets: Anses d’Arlets, FWIM 1. Ste. Marie: Riviere Charpentier, MAR 116. La Trinite:

Galion Sugar Refinery, MAR 7. Locality not specified: July, A. Busck, IF.

MONTSERRAT (1 IpF). St. Anthony: Foxs Bay, MNT 39.

PANAMA AND CANAL ZONE (327 specimens: 137M, 7Mgen, 166F, 8P&p, 9L). Bocas del

Toro: Punta de Pena, PA 219, 253. Canal Zone: Ancon, 1912, J. Zetek, 1F; same locality, Jul

1941, PAX 181, 1F. Cocolis [Cocoli] , 26 Feb 1943, 1M. Cristobal, 13 Jul 1944, Sterns e¢ al;

ASM 68-1, 2M, 2Mgen, 1F; same data, 3M, 2F, 10L [CU]. France Field, Mar 1941, IM, 5F. Gale-

ta Is., 3 Sep 1930, PAX 17, 5F. Gatun, 28 Feb 1930, D. Curry, PAX 7, 3M, 2F; same locality and

collector, 29 Feb 1930, PAX 8, 6M, 6F; same locality and collector, Jan 1931, PAX 21, OM, 17F;

same locality, 25 Mar 1933, PAX 31, 8M, 22F; same locality, 10 Mar 1937, 2F; same locality, PAR

46, 10M, 2Mgen, 32F; same locality, KO H-19-18, 48M. Gatun Lake, 20 Feb 1934, PAX 36, IM,

3F. Largo Reme [Remo], 1926, D. Curry, 1F. Mindi, 10 May 1941, C. Brown, PAX 176, 31M,

28F. Mojinga Swamp, PA 1165. Mt. Hope, 29 Jan 1944, 5M, 2F. Pedro Miguel Locks, 22 May

1939, PAX 110, 1F. Locality not specified, Feb 1944, 1F. Colon: Colon, 11 May 1930, D. Curry,

PAX 9, 1M, 1F; same locality and collector, 27 May 1930, PAX 10, 1F; same locality and collector,

7 Jun 1930, PAX 12, 1M; same locality and collector, 10 Jun 1930, PAX 13, 4M, 4F. Pina, 27 Jun

1945, 1F. Portobelo, PA 603, 604, 606. Panama: Chepo Road, 19 Nov 1939, PAX 157, 1M.

Province and locality not specified: PAR 136, 1F; from D. Curry, SL [JH] ; no data, 2Mgen [JH].

SAINT KITTS (5F). Conaree Swamp, 22 Nov 1949, 5F.

SAINT LUCIA (192 specimens: 19M, 9Mgen, 23F, 38P&p, 103L&l; 30 ind rear: 24 1, 3p, 3inc).

Anse La Raye: Anse La Raye, LU 71, 72, 127. Roseau Sugar Estate, LU 23. Roseau Sugar Fac-

tory, LU 20, 86. Castries: La Toc, LU 6. Gros Islet: Bois D’Orange, LU 37. Choc Swamp, LU

153, 154. Gros Islet, 24 Feb 1941, 1F; same locality, 29 Jul 1941, 4F; same locality, 2 Aug 1945,

Ingle, 2F. Marisule, LU 39, 40. Union Agricultural Station, LU 150, 151, 157. Laborie: Sapphire

Estate, LU 121. Micoud: Troumassee Estate, LU 118,119. Praslin: Volet Estate, LU 98, 102.

Soufriere: Soufriere, LU 84,85. Quarter and locality not specified: No data, 1L.

SAINT VINCENT (1M). Locality not specified, 1M [BM].

SURINAME (59 specimens: 11M, 8Mgen, 33F, 2p, 5L&l; 2 inc ind rear). Coronie: Friendship,

15 Jun 1946, E. Kuyp, 1Mgen (46-x-17a). Sarahmeer, 17 Jun 1946, E. Kuyp, 1Mgen (46-x-16b).

Zoetwater Kanaal, 14 Jun 1946, E. Kuyp, 1M. Locality not specified, 15 Jun 1946, E. Kuyp, 3M,

1F. Marowijne: Moengo, Sep 1944, Guicherit, 2F. Nickerie: Nieuw Nickerie, 14 Jun 1946, E.

Kuyp, 1Mgen (46-x-21a). Paramaribo: Charlesburg, 9 Jun 1946, E. Kuyp, 1M, 1Mgen (46-x-22a),

1F, 2L; same locality, 11 Aug 1949, D. Geijskes, 4F [BM1949-475]; same locality, SUR 49. Suri-

name: Domburg, SUR 39. Ma Retraite, SUR 70, 72. Morgenstand, 22 Jun 1946, E. Kuyp, 4M, 1

Mgen (46-ix-16b-82), 1Mgen (46-ix-16c-83), 3F, 1 lp (87), 11(88). Meerzorg II, 10 Jun 1946, E.

Kuyp, 1M, 1Mgen (46-x-22b). Ornamibo, 23 Jun 1946, E. Kuyp, 1F, 1 lp. Tweekinderem, 22 Jun

1946, E. Kuyp, 1M, 1Mgen (46-x-22d). District and locality not specified: SUR 117; H. Polah, IF.

TRINIDAD AND TOBAGO. TOBAGO (28 specimens: 3M, 2Mgen, 7F, 6P&p, 1OL&I; 6 ind

rear: 21, 3p, linc). St. Patrick: Bon Accord Estate, 16 Nov 1965, T. Aitken, R. Martinez, A. Guer-

ra, TOB 22, 1 pF; same data, TOB 24, 1 IpM, 1 lpF, 1 IP, 6L. Golden Grove, 19 Nov 1965, T. Ait-

ken, R. Martinez, A. Guerra, TOB 58, 1Mgen, 1 pM; same data, TOB 59, SF. Orange Hill, 30 Nov

1965, R. Martinez, A. Guerra, TOB 142, 1Mgen, 1 pM, 1L. TRINIDAD (463 specimens: 143M, 47

Mgen, 171F, 24p, 78L&l; 20 ind rear: 141, Sp, linc). Caroni: Caroni Swamp, 15 Oct 1964, A.

Guerra, TR 762, 1Mgen, 1 pM, 1 pF, 1 lpM, 1 lpF; same data, TR 763, 1 IpF, 2L. Locality not spe-

cified, Jan 1946, 2F. Mayaro: Guayaguayare Rd., 27 Oct 1964, A. Guerra, TR 797, 4L. St. An-

drew: Cumuto, 11 Jul 1941, TRR 21B, 1M, 1Mgen. Turure Rd. nr Guaico, 9 Apr 1966, A. Guerra,

TR 1492, 1Mgen, 1 lpM, 1 lpF, 1P,3L. St. David: Redhead, 28 Feb 1964, A. Guerra, TR 122,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 89

1 lpF, 3L; same data, TR 124, 1L; same data, TR 125, 1 lpM, 2 lpF. Sans Souci, 5 Mar 1964, A.

Guerra, TR 147, 1Mgen, 2 pF, 3 lpM, 1 IpF, 2L; same data, TR 148, 1 pF, 2L. Toco, 6 Mar 1964,

A. Guerra, TR 162, 1L; same data, TR 166, 2L; same data, TR 167,4L; same data, TR 168, 2L;

same locality, TRK 9, 26M, 1Mgen, 21F. St. George: Blanchisseuse, 27 Mar 1941, W. Komp, KO

111-9, 9M; same locality, 10 Apr 1964, A. Guerra, TR 306, 1 1. Chaguaramas, 29 Jun 1961, T.

Aitken, | lp. Diego Martin River, 8-26 Feb 1941, W. Komp, TRK 35, 24M, 2Mgen, 40F. El Socor-

to, Nov 1945, 4M, 2F. Long Stretch, 5 Feb 1941, TRK 4, 1Mgen. Piarco, Golden Grove, 14 Sep

1954, 1F [BM]; same locality, Sep 1954, 1F [BM]. Port of Spain, 10 Jun 1941, 6Mgen [JH];

same locality, 14 Jun 1941, TRR 7, 14M, SMgen, 7F; same locality, 16 Jun 1941, L. Rozeboom,

4L; same data, 24L [JH]; same locality, 23 Jun 1941, TRR 56, 1M, 1Mgen; same locality, 26 Jun

1941, TRR 57, 28M, 7Mgen, 1F; same locality, 9 Jul 1941, TRR 59, 16M, 4Mgen, 27F. St. Pa-

trick: La Brea, 29 Jul 1966, A. Guerra, TR 1563, 2F, 1 lpF. Lillette Swamp, 9 Jul 1966, T. Ait-

ken, R. Martinez, TR 1552, 2M, 1Mgen, 1F,3p, 5L. County not specified: S. Joseph, 2 Aug 1899,

F. Urich, 2F. Locality not specified, 1915-1928, J. Dickson, 1 1(191) [BM]; 22 Jul 1931, E. de

Verteuil, 1F [BM1931-562] ; 25 Jul 1931, E. de Verteuil, 1M, 1F [BM1931-562] ; Mar 1941, W.

Komp, KO 111-7, 2M, 3F; Jul 1941, 1Mgen [JH]; May 1963, T. Aitken, 2M, 9F; Jun, A. Busck,

36F; W. Downs, 9Mgen [JH]; F. Urich, 1M, 3F; no data, 5M, 5Mgen (750915-23, 24, 25, 26, 27).

VENEZUELA (62 specimens: 14M, 3Mgen, 19F, 12p, 14L&1; 12 ind rear: 9p, 3inc). Anzoate-

gui: Barcelona, 16 Sep 1944, VZK 20, 2M. Aragua: Turiamo, 11 Sep 1944, VZK 3, 2M, 5F;same

locality, Sep 1944, VZK 30, 4M, 1F. Carabobo: Boca de Yaracuy, VZ 259. Puerto Cabello, 1M

gen [JH]. Guarico: El Sombrero, 14 Jul 1945, 1F. San Juan de los Morros, 17 Jan 1945, 1M, 1F.

Monagas: Punta Tigre, Rio Guanaco, 3 Jan 1929, F. Urich, 1F [BM1929-453]. Jusepin, 30 Apr

1945, 2M. Province not specified: Higuerote, Nov 1949, 1L (616-Zone I, Cot. 185).

6. Anopheles (Nys.) ininii Senevet & Abonnenc

| Figs, 2; 4, 6,17, 18

1938. Anopheles (Nyssorhynchus) ininii Senevet and Abonnenc 1938:494-500. TYPE: Syn-

types 2 females with associated larval and pupal skins and genitalia slides, St. Elie mines,

Sinnamary River (Inini), French Guiana, artificial reservoir, 7 Feb 1938 [NE; Belkin

(1968b:53) reported that type was lost or destroyed during Algerian conflict] .

Anopheles (Nyssorhynchus) ininii of Rozeboom and Gabaldon (1941:97-98); Floch and Abonnenc

(1942b:2; 1943a:1-3; 1946a:1-4; 1946b:3-5; 1947:6; 1951:17, 50-51); Senevet (1948a:277-

279; 1948b:431-432; 1948c:436-437); Levi-Castillo (1949:10, 15, 28, 58, 67, 77, 82, 86); Ga-

baldon and Cova Garcia (1952:178, 191); Senevet and Andarelli (1955:339); Stone, Knight and

Starcke (1959:33); Vargas (1959:370); Fauran (1961:10-11); Belkin, Schick and Heinemann

(1965:23); Belkin (1968b:53). |

Anopheles ininii of Russell, Rozeboom and Stone (1943:51); Foote and Cook (1959:28, 29).

FEMALE (fig. 6). Wing: 3.7 mm. Proboscis: 2.3 mm. Palpus: 2.35 mm. Forefe-

mur: 1.6 mm. Abdomen: about 3.1 mm. Dark, very similar to that of oswaldoi.

Head: Integument dark brown with silver pruinose areas. Interocular space moder-

ately wide, about 3-6 ommatidial diameters. Proboscis about 1.4 length of forefe-

mur. High contrast between the integument and white erect scales on vertex, and de-

cumbent scales on interorbital line and upper orbitals. Erect scales of vertex as those

of oswaldoi. Dark scales of palpus dark brown to black; apex of palpal segment 2

with a small white band; apex of segment 3 with a moderately large, white band; dor-

sal surface of 2 and 3 with tan scales, not always conspicuous on 2, but as a longitu-

dinal stripe on 3; segment 4 with a moderately large, dark, basal band and usually a

much smaller, dark, apical ring; dark scales on ventral surface of segment 4 extending

distad from basal band to form a distinct, dark, ventral stripe, usually not reaching

the dark apical ring; mediolateral surface of segment 4 and 5 white. Antenna: Fla-

gellar segment | with long, numerous, very prominent, white scales on dorsomesal

surface. Flagellar segments 2-13 each with 7-9 setae in basal whorl. Thorax: Integu-

#90 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

ment dark brown to brownish black. Short light scales on scutum ivory to silver.

Anterior promontory area with long, white, setiform scales extending dorsad onto

acrostichal line; humeral scale tuft very prominent, with white scales above and nu-

merous, cuneate, brownish black scales below. Prescutellar space large, horseshoe

shaped and very dark. Upper stp with about 3 long setae and 10 gray to white, obo-

vate scales. Lower stp with one long seta and about 8 scales; scales similar to those

of upper stp, occasionally lighter, in a large, conspicuous, diagonal patch. Upper mep

with 5,6 setae; scales absent. Legs: Coxal scales white. Foretarsal segments 3-5 pre-

dominantly cream to white; light apical bands on segments 2 and 3 0.45 (0.35-0.55)

and 0.75 (0.70-0.86) respectively; segment 4 often all light cream to golden brown or

with a small, dark, basal area rarely longer than 0.3 of segment; often dark basal

scales on segments 1-3 and occasionally 4,5 primarily restricted to dorsal, or dorsal

and part of anterior and posterior surfaces so that basal bands appear incomplete ven-

trally; segment 5 all cream to about 0.5 basally dark. Midtarsal segments 2 and 3

with a small cream band in apical 0.08-0.17 and 0.12-0.24 respectively; segment 4

with a cream band in apical 0.15-0.25; segment 5 from completely cream to dark in

about basal 0.5. Hindtarsal segment 2 with a dark band in basal 0.17-0.26; segment

5 cream to white in apical 0.5-0.6. Wing: Light wing spots white to very light cream.

Dark spots dark brown to brownish black. Vein C humeral light spot 2.0-3.3 (1.5-

3.3) of basal dark spot; subcostal light spot 0.35-0.70 of subcostal dark spot; preapi-

cal light spot 0.25-0.50 of preapical dark spot; apical dark spot small. R presectoral

dark spot moderately small; sectoral dark spot small. Rs-R,,3 usually with an extra,

small to moderately large, subcostal dark spot. M highly variable, with a moderately

long to very long, sectoral dark spot; preapical dark spot not extending onto M44».

Cu, subcostal and preapical dark spots small to moderate. Apical light fringe spot

moderately small to large, undivided or divided by a few dark scales between R, and

R;, and/or R; and R4,,; additional light fringe spots as in remaining species. Abdo-

men: Tergite I with a few light scales along caudomedian border. Sternite I with a

few setae. Caudolateral scale tufts large and prominent on segments II-VII. Light

scales of tergites II-VII often overlapping bases of dark caudolateral tufts. Scales on

sternite II-VII as in section description except very distinct and numerous.

MALE. Wing: 3.6 mm. Proboscis: 2.9 mm. Forefemur: 1.6 mm. Abdomen:

about 3.2 mm. Large, essentially as in female except for sexual characters. Head:

Palpal segments 2 and 3 with a dorsal cream stripe; segment 4 with a white band at

base and apex, a small, brown, subbasal band usually present; segment 4 cream to

white mediolaterally and brown ventrally. Antenna: About 0.75 length of proboscis.

Flagellar segment 1 with numerous, long, thin, conspicuous, white, lanceolate scales.

Legs: Forefemur about 0.55 length of proboscis. Basal plantar surface of foretarsal

segment 5 with 7,8 moderately long, spiniform setae; setae longest toward apex.

Claw on foreleg large, curved, acuminate; submedian tooth long, about 0.25 length of

claw, tapering to blunt recurved point; basal tooth long, subequal to submedian

tooth, strongly decurved.

MALE GENITALIA (fig. 17). Segment VIII: Tergite and sternite with moderate-

ly large, obovate scales. Sternite with a few, median, dark scales, and laterally with

cream to white scales. Tergite with dark scales along posterior margin. Segment IX:

Sternite long, about 0.2 length of sidepiece, subtrapezoidal. Anterior apodeme a

sclerotized band across anterior border of sternite, about 0.15 length of sternite, oc-

casionally appearing weakly bilobed. Sidepiece: Subconical. Tergal surface with 4,

5 long, submedian, tergomedial bristles; bristles mesad of tergomedial bristles moder-

ately long. Parabasal spine moderately long, at least 2.0 length of its tubercle. Basal

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 91

apodeme thick, moderately long, about 0.2 length of sidepiece. Longer, more dorsal

accessory spine slightly longer than 0.5 length of sidepiece; more ventral spine 0.65-

0.70 of longer spine. Internal spine subequal to shorter accessory spine, apically re-

trorse. Clasper: Spiniform long and narrow, subequal to seta b. Dorsal Claspette:

Long, 0.55-0.70 length of sidepiece. Pedicel moderately thick; base rounded, occa-

sionally curved mesad. Leaflets long, about 0.5 length of claspette, sharply curved

mesad about 0.5 from base; dorsal leaflet with small basomesal projection. Ventral

Claspette: Strongly conoid; long, about 0.5 length of sidepiece; narrow at apex,

width at apex about 0.3 length of claspette. Basal lobule very large, extending later-

ad, and usually bent at base ventrally so that setae on basal margin projecting caudad;

setae along basal margin very strong and long, about 2.0-3.0 width of aedeagus; setae

distributed uniformly over surface of lobule. Lateral and ventral surfaces (exclusive

of basal lobule) with short setae about 0.5 width of aedeagus; setae extending toward

apex only as far as level of apical margin of preapical plate. Apex without setae, pro-

duced into rounded cone with a very small, median sulcus. Preapical plate distinct,

very large, crescent shaped, heavily sclerotized. Refringent structure rooflike, in

shape of an inverted V with a rounded vertex. Phallosome: Aedeagus about 0.5-0.6

length of sidepiece; apex about as long as wide, moderately rounded, with or without

very small, membranous leaflets as in nuneztovari.

PUPA (fig. 17). Abdomen: about 2.9 mm. Trumpet: 0.5 mm. Paddle: 0.8 X 0.6

mm. In general, setae long and numerously branched. Cephalothorax: Heavily scler-

otized. Wing cases with or without longitudinal stripes. Hair 2-C 2,3 forked (1-3),

longer than 1,3-C; 1-C 2 branched (1-3), shorter than 3-C; 3-C 2,3 branched near base.

Hairs 4,5-C subequal; 4-C 2-4 forked; 5-C 2-5 forked. Hair 6-C 1-3 branched. Hair 7-

C 2,3 branched (1-4) near base, shorter branches subequal to 6-C, with one branch

about 1.25-1.50 length of shorter branch(es). Trumpet: Similar to that of strodei

and aquasalis. Pinna amber; about 3.7-4.4 (3.6-4.6) length of meatus; in lateral as-

pect, appearing broad medially and tapered toward apex. Meatal cleft long, moder-

ately pointed at base. Metanotum: Hair 10-C single or double, strong, moderately

long, slightly less than or equal to 11-C. Hair 11-C 3-5 branched near base. Hair 12-

C 1-3 forked, about 2.0 length of 10-C. Abdomen: Heavily sclerotized. Hair 2-I 5-8

branched near base, moderately long; 3-I single or double, slightly more than 0.5

length of 2-I. Hair 4-I 5-8 branched. Hair 5-I 2,3 forked (1-3), long. Hair 6-I single

and very long. Hair 7-I 3-6 branched, 0.5 length of 6-I. Hair 9-I single, longer than 7-

I and shorter than 6-I. Hair O-II,III, VI, VII moderately large; O0-IV,V large; O-II 4-7

branched (4-10); O-III 5-7 braved (4-7): 0-IV,V 4-6 branched (3-8); 0-VI 47

branched; O-VII 4-6 branched. Hair 1-II,III 8-12 (8-13) and 7-13 (6-14) branched re-

spectively, large; 1-[V-VII very long, 1. 7s to more than 2.00 length of segment. Hair

3-IV 4-6 branched (4-7); 3-V 3-5 forked (2-5). Hair 5-II 4-7 branched (4-8) near base,

moderately small; 5-III numerously, 9-13 (5-13) branched, moderately developed; 5-

IV 2-6 branched (2-8), large, subequal to length of segment; 5-V-VII single, long.

Hair 6-II single or double, long, at least 2.0 length of 7-II. Hair 7-II 4-6 branched; 7-

III,[V about 3-7 branched, moderately developed; 7-V 3,4 branched, longer than 7-

III,1V; 7-VI usually double (1-4 branched), moderately long, about 0.5 or less length

of segment. Hair 8-III-VII moderately developed; 8-III-VI about 2-5 branched; 8-VII

4-6 branched. Hair 9-II very small, unpigmented; 9-III blunt, stout, short, subequal

to or slightly longer than 9-II, pigmented; 9-IV stout, about 2.0 of 9-III; 9-V acumi-

nate, stout, curved, 1.75-2.00 (1.4-2.4) length of 9-IV; 9-VI strong, relatively short,

about 0.25-0.33 length of segment; 9-VII 1.41.8 (1.3-1.8) length of 9-VI; 9-VII, VIII

tapering to a fine point, long, about 0.5 or more length of segment; 9-VII curved;

92 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

9-VIII almost straight. Hair 10-III 2-4 forked, moderately long. Hair 4+VIII 2-4

forked. Sum of branches of hairs 1-III, 5-IIJ and 0-VI 22-27 (22-31). Terminal Seg-

ments: Male genital lobe heavy, thick, with mammilliform protuberance moderately

prominent. Paddle: Large, obovate, weakly emarginate at insertion of hair 1-P. Ex-

ternal buttress 0.55 length of paddle. External margin distad of buttress with moder-

ately long, filamentous spicules extending around apex and becoming minute along

caudal 0.5 of inner margin. Hair 1-P moderately long (0.1 mm); 2-P 1-3 forked, sub-

equal to 1-P.

LARVA (fig. 18). Head: 0.6 mm. Antenna: 0.3 mm. Anal Saddle: 0.3 mm.

Head: Moderately large. Very darkly pigmented, often almost black. Setae strongly

developed and long. Inner and outer clypeal hairs (2,3-C) single and barbed in about

apical half, often appearing simple; hairs 2-C widely spaced, clypeal index about 1.25;

3-C about 0.8-0.9 length of 2-C. Hair 4-C 4,5 branched (3-6), short, not extending to

base of 2-C. Hairs 8,9-C 6-8 branched (5-10), strongly developed; 9-C weakly plu-

mose with moderately long shaft, longer than 8-C, subequal to 2.0 distance separa-

ting insertion of 5-C and 6-C. Hair 10-C 3,4 branched, subequal to 8-C. Hair 12-C +

6 branched, subequal to 8-C. Hair 15-C about 5 branched, strongly developed, long.

Collar heavily pigmented and wide dorsomedially. Antenna: Very heavily pigment-

ed. Mesal margin with strongly developed, stout spicules; ventral surface with numer-

ous, shorter, heavy spicules; dorsal surface with a few small spicules. Hair 1-A den-

dritic, 7-9 branched (6-10), long, about 2.0 width of antenna at point of insertion,

inserted about 0.25-0.33 from base of antenna. Hairs 2,3-A long and narrow. Thor-

ax: Dark, greenish brown, heavily pigmented. Submedian prothorax group (1-3-P)

often with 1,2-P sharing common tubercle, occasionally tubercle appearing more

sclerotized at insertion of 2-P than at insertion of 1-P; palmate hair 1-P with 11-16

(10-18) long, moderately narrow leaflets; 2-P 14-17 branched (10-17), long, almost

4.0 length of 1-P. Hair 14-P 7-10 branched (7-13) from moderately long shaft. Meso-

thoracic hair 1-M 28-33 branched, apical branches shorter than lateral. Metathoracic

hair 2-T long, extending to or beyond caudal margin of thorax. Palmate hair 3-T

with 12-18 (10-18) large, weakly pigmented leaflets. Prothoracic pleural group spine

larger than mesothoracic spine; metathoracic pleural spine smallest. Abdomen: Hair

O-II-VI large; O-II,III,V,VI about 6-10 branched (5-10); O-I[V 5-8 branched. Palmate

hair 1-I with 14-17 (11-20) weakly pigmented, moderately long, narrow, spreading

leaflets; 1-III-VI with 23-30 (20-31) long, narrow, lanceolate leaflets; 1-I[,II, VII short-

er than 1-ITI-VI. Hair 2-II 4-6 branched (3-6), moderately large; 2-III 3-4 branched,

large; 2-IV single, slightly longer than length of leaflets of 1-IV; 2-V single, very long

(0.3 mm). Hair 3-I single, moderately long. Hair 5-1 4,5 branched (3-6), moderately

developed; 5-II 8-10 branched, medium. Hair 9-I about 4-7 branched. Hair 13-I,II,

III 46 (4-9), 7-11 (6-11) and 8-10 (6-14) branched respectively, small; 13-IV 3,4

branched, larger than 13-I-III; 13-V large, extending beyond caudal margin of seg-

ment; 13-VI 7-12 branched. Spiracular Lobe: Pecten with 15-22 teeth; many (8-11)

of median teeth subequal. Lateral arm of spiracular apparatus long, directed laterally,

appearing truncate on lateral margin at point of contact with spiracular opening.

Anal Segment: Covered with very fine spicules becoming very short and inconspicu-

ous basally. Saddle reddish brown. Hair 1-X longer than saddle; inserted on saddle

near ventral margin, never on lateral margin at base of indentation; inserted about 0.3

cephalad of caudal margin. Anal gills very long, usually 2.0 or more length of saddle.

DISCUSSION. An. ininii can be distinguished from the other species in the Oswal-

doi Subgroup in the female by the combination of (1) palpal segment 4 with a dark

basal band extending distad on the ventral surface to form a distinct, dark, ventral

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 93

stripe usually not reaching apical dark ring, (2) foretarsal segments 3-5 predominant-

ly cream to white with dark scales often present only on dorsobasal surface of seg-

ment, segments 2 and 3 light in apical 0.45 (0.35-0.55) and 0.75 (0.70-0.86) respec-

tively, segment 4 often all light cream to golden brown or with a small, dark, basal

area rarely longer than 0.3 length of segment, segment 5 from about 0.5 basally dark

to all cream, (3) midtarsal segment 4 with a cream band in apical 0.15-0.25, and seg-

ment 5 from completely cream to 0.5 basally dark, (4) hindtarsal segment 2 with a

dark band in basal 0.17-0.26, (5) wing spots white to very light cream, (6) humeral

light spot on vein C about 2.0-3.3 (1.5-3.3) of basal dark spot and (7) subcostal light

spot on vein C 0.35-0.70 of subcostal dark spot; in the male genitalia by the combi-

nation of (1) ventral claspette strongly conoid, narrow and rounded at apex, width

at apex about 0.3 length of claspette, (2) basal lobule very large, extending laterad

and usually bent at base ventrally so that setae on basal margin projecting caudad, (3)

setae along basal margin of basal lobule very strong and long, 2.0-3.0 width of aedea-

gus, setae distributed uniformly over surface of lobule, (4) setae on lateral and ven-

tral surfaces of ventral claspette (exclusive of basal lobule) short, about 0.5 width of

aedeagus, extending toward apex only as far as level of apical margin of preapical

plate, (5) preapical plate heavily sclerotized, very large, crescent shaped and (6) apex

of aedeagus with or without very small, membranous leaflets; in the pupa by the

combination of (1) setae in general long and numerously branched, (2) hair 7-C 2,3

branched (1-4), shorter branches subequal to 6-C, with one branch about 1.25-1.50

length of shorter branch(es), (3) hair 5-II 4-7 branched (4-8) near base, moderately

small, (4) hair 9-V 1.75-2.00 (1.4-2.4) length of 9-IV, (5) hair 9-VIT 1.4-1.8 (1.3-1.8)

length of 9-VI and about 0.5 or more length of segment, tapering to a fine point, (6)

hair 2-I 5-8 branched near base, moderately long, hair 3-I slightly more than 0.5

length of 2-I, (7) hair 6-II single or double, at least 2.0 length of 7-II and (8) sum of

branches of hairs 1-III, 5-II] and 0-VI 22-27 (22-31); and in the larva by the combina-

tion of (1) hairs 2,3-C single and barbed in about apical half, hairs 2-C widely spaced,

clypeal index about 1.25, hair 3-C about 0.8-0.9 length of 2-C, (2) hair 4-C short, not

extending to base of 2-C, (3) hairs 8,9-C 6-8 branched (5-10), strongly developed, 9-

C weakly plumose with moderately long shaft, subequal to 2.0 distance separating in-

sertion of hairs 5-C and 6-C, (4) hair 1-A dendritic, 7-9 branched (6-10), long, about

2.0 width of antenna at point of insertion, (5) hair 14-P 7-10 branched (7-13) from

moderately long shaft, (6) hair 13-IV 3,4 branched, larger than 13-I-III, (7) lateral

arm of spiracular apparatus long, directed laterally, appearing truncate on lateral mar-

gin, (8) hair 1-X inserted on saddle near ventral margin, never on ventral margin at

base of indentation and (9) anal gills very long, usually 2.0 or more length of saddle.

An. ininii is a highly derived species, but it still retains a few ancestral characters

_ which it shares with other species in the Oswaldoi Group and with some species in

the Argyritarsis Section. These ancestral characters are in the larva (1) hair 13-IV

large, with few branches and (2) hair 1-X inserted on saddle near but not on ventral

margin; and in the male genitalia (1) the frequent presence of very small leaflets on

the apex of the aedeagus and (2) possibly the shape of the ventral claspette. Al

though ininii often has very small leaflets on the aedeagus as do nuneztovari and rang-

eli, it is probably more closely related to oswaldoi, galvaoi, noroestensis and aquasalis.

An. oswaldoi occasionally has leaflets on the aedeagus, and in general its ventral clas-

pette is similar to that of ininii. The preapical plates are almost identical in oswaldoi

and ininii, and the setae are distributed similarly on the basal lobules of the ventral

claspette. The females of oswaldoi and ininii also are very much alike in all charac-

ters except for the fourth palpal segment and the banding patterns of the foretarsus.

94 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

It is impossible to differentiate these 2 species on the basis of the wing or the second

hindtarsal segment. Likewise, the larvae of oswaldoi and ininii share many similar

characters, such as the length of the lateral arm of the spiracular apparatus, the devel-

opment of hairs 8,9-C and the arrangement of teeth on the pecten; these characters

are usually more strongly developed (i.e. longer, larger) in ininii, except for the

branching of hairs 2,3-C which has been strongly reduced in ininii as compared to os-

waldoi. Ecologically, ininii occurs sympatrically with oswaldoi, the immatures being

found in many of the same habitats, although it usually occurs in more open areas

than oswaldoi. Speciation of ininii may have occurred in a geographically isolated

area (refugium) in northern central South America.

BIONOMICS. Very little is known about the bionomics of ininii, particularly re-

garding the adults. Almost all the information relating to the larval habitats discus-

sed below came from data recorded on collection forms by John F. Reinert while in

Para, Brazil in 1974. The immatures of ininii are found along grassy margins of

ponds, lakes, reservoirs, temporary or permanent ground pools, and animal or wheel

tracks, in full sun or occasionally in partial shade. The water is fresh, clear, and with

algae and emergent vegetation, although vegetation is usually scanty in the larval ha-

bitat. The general habitat in Para was characterized as secondary scrub. The 3 spe-

cies in the Albimanus Section collected in association with ininii were oswaldoi, nu-

neztovari and triannulatus; the last 2 species were by far more common. The follow-

ing 11 species have also been found in association with ininii: Anopheles (Ano.) sp,

Culex (Cux.) coronator group, Cx. (Cux.) mollis, Cx. (Mel.) bastagarius, Cx. (Mel.)

dunni group, Cx. (Mel.) ?rachoui, Cx. (Mel.) zeteki group, Cx. (Mel.) sp, Cx. (Ads.)

amazonensis, Uranotaenia (Ura.) geometrica and Ur. (Ura.) pulcherrima. The type

material of ininii was collected in an artificial reservoir in St. Elie, French Guiana

(Senevet and Abonnenc 1938:501-511). Floch and Abonnenc (1947:6; 1951:50-51)

reported that ininii occurs in dense vegetation with An. (Nys.) braziliensis, darlingi,

nuneztovari, oswaldoi and triannulatus. Floch and Abonnenc (1951:17) reported

that in 9 years a total of 6 specimens were collected.

MEDICAL IMPORTANCE. Nothing is known about the adult behavior or the me-

dical importance of ininii.

DISTRIBUTION (fig. 2). An. ininii has been collected only in French Guiana and

in the state of Para, Brazil.

Material Examined: 192 specimens: 32 males, 14 male genitalia, 57 females, 66

pupae, 23 larvae; 66 individual rearings: 18 larval, 48 pupal.

BRAZIL. Para (192 specimens: 32M, 14Mgen, 57F, 66p, 23 L&I; 66 ind rear: 181, 48p). Alta-

mira, 150 km W of, 5 Nov 1974, J. Reinert et al., coll. 77, 1 pF; same data, coll. 79, 3Mgen, 7 pM,

7 pF; same data, coll. 80, 2 pM, 1 pF; same data except 163 km W of Altamira and 6 Nov, coll. 85,

5Mgen, 4 IpM, 5 IpF, 4 pM, 1 pF, 5L; same data except 158 km W of Altamira and 9 Noy, coll. 111,

1 pM. Bacuri, Gleba 29, Lote 03, 20 Oct 1974, J. Reinert et al., coll. 5, 1M, 1Mgen; same data ex-

cept Gleba 36, Lote 02 and 23 Oct, coll. 33,3 IpF; same data except Gleba 29, Lote 03 and 25 Oct,

coll. 44, 2Mgen, 2 IpM, 2 lpF, 1 pM; same data except Gleba 36, Lote 02 and 26 Oct, coll. 50, 1M

gen, 2 pM, 1 pF; same data except Gleba 34, Lote 02 and 28 Oct, coll. 62, 2Mgen, 2 pM, 1 pF;

same data except Gleba 38, Lote 02 and 29 Oct, coll. 64, 21pM, 2 pM, 2 pF. Belem, APEG Forest,

22 Nov 1974, J. Reinert et al., coll. 140, 1 pF. Jatobal, 18 Feb 1976, Moramay, 1102, 3 pF; same

data except 19 Feb, 1104, 2 pM, 7 pF. Maraba area, 3 Jun 1976, 1F (R-5); same locality, 15 Jun

1976, D. Roberts, 1F (A-1); same locality, 23 Jun 1976, 5F (A-2); same locality, 24 Jun 1976, D.

Roberts, 11F (A-1); same locality, 21 Oct 1976, 2F (1); same locality, 8 Nov 1976, 1F (1); same

locality, 9 Nov 1976, 1F (2).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 95

7. Anopheles (Nys.) anomalophyllus Komp

Figs. 2, 4, 6, 19, 20

1936. Anopheles (Nyssorhynchus) anomalophyllus Komp 1936:163-164. TYPE: Holotype

male with genitalia slide, Wenham’s Farm, near Almirante (Bocas del Toro), Panama, bred

from pupa, Feb 1929, W. H. W. Komp [USNM, 52018].

Anopheles (Nyssorhynchus) anomalophyllus of Lane (1939:21; 1944:262; 1949:402; 1953:263-

264); Komp (1941a:90, 93, 96; 1942:39, 43, 75, 80, 125-127, 132, 161-162); Rozeboom and

Gabaldon (1941:92-98); Galvao and Damasceno (1942:132); Simmons and Aitken (1942:39,

46, 52, 62, 87); Levi-Castillo (1949:9, 15, 27, 58, 67, 72, 76, 85); Gabaldon and Cova Garcia

(1952:182); Vargas V. (1956:29; 1957; 1958a; 1958b); Bejarano (1957:326, 328, 330, 331);

Stone, Knight and Starcke (1959:31); Vargas (1959:376); Belkin, Schick and Heinemann (1965:

43); Cova Garcia, Pulido F. and Amarista M. (1978:157).

Anopheles anomalophyllus of Kumm and Ruiz (1939:438); Kumm, Komp and Ruiz (1940:389,

408); Kumm (1941b:95-97); Rozeboom (1941:103); Russell, Rozeboom and Stone (1943:26,

29, 31); Vargas V. (1958c:1, 3); Cova Garcia (1964:200); Stojanovich, Gorham and Scott

(1966a:10, 19, 30).

FEMALE (fig. 6). Description based primarily on 4 specimens from Costa Rica.

Wing: 3.1 mm. Proboscis: 1.9mm. Palpus: about 1.9 mm. Forefemur: 1.2 mm.

Abdomen: about 2.7 mm. Generally very similar to strodei. Head: Integument dark

brown. Proboscis about 1.6 length of forefemur. Vertex with very elongate, cuneate

or slightly forked, cream to white scales. Dorsal surface of palpal segment 2 with a

very few, light, erect scales; dorsal surface of segment 3 with light scales forming a

longitudinal stripe; segments 2 and 3 with light apical band, apical band of 2 much

smaller than apical band of 3; segment 4 with a basal and an apical dark band, light

scales on ventral surface darker than on lateral surface, apparently with light brown

to brown scales forming an indistinct ventral stripe; segments 4 and 5 predominantly

white; base of 5 with a few dark scales extending from apex of 4. Antenna: Flagel-

lar segment 1 with moderately long to long, white, oblanceolate scales distally on

dorsomesal surface; base of 1 with small, gray to white, obovate scales. Thorax: In-

tegument light brown to reddish brown. Elongate, white anterior promontory scales

extending onto acrostichal line. Prescutellar space horseshoe shaped. Upper stp with

3,4 long, light brown to dark brown setae and a horizontal row of light scales. Low-

er stp with one long brown seta and a patch of light scales. Upper mep with about 4,

5 long, cream to light brown setae; scales not visible in specimens examined. Legs:

Integument predominantly reddish brown. Light scales on coxae white. Midfemur

with long, narrow, cream anteroapical spot and a small knee spot. Foretarsal seg-

ments 2 and 3 with a white band in apical 0.35 (0.29-0.42) and 0.51 (0.23-0.78) re-

spectively; segment 4 all brown; segment 5 cream in about apical 0.5. Midtarsal seg-

ments | and 2 with a small, inconspicuous, white apical band; segment 3 with a

cream apical ring (inconspicuous in one specimen); segment 4 all dark; segment 5

with a cream band in about apical 0.5. Hindtarsal segment 2 with brown band in ba-

sal 0.4 (0.30-0.45); segment 5 cream in apical 0.5-0.7. Wing: Slightly narrower and

more pointed at apex than that of aquasalis. Light spots of wing white except for

vein C and possibly R. Vein C humeral light spot about 2.5 (2.3-4.0) of basal dark

spot; subcostal light spot 0.23-0.60 of subcostal dark spot; preapical light spot about

0.5 of preapical dark spot; 2 specimens with presectoral and sectoral dark spots fused.

R presectoral dark spot moderately short to long; sectoral dark spot very small in 2

specimens and absent in the other 2. R,,; subcostal and apical dark spots small to

moderately small. M sectoral dark spot small to moderate, in 2 specimens absent or

indistinct. Apical light fringe spot moderately large, distinctly broken by dark scales

96 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

~ between R; and Ry,;; additional, small, inconspicuous, light fringe spots at apices of

all veins posterior of, and including, Ry,,; a moderately large, inconspicuous, light

fringe spot at level of 0.5 distance from base of A. Abdomen: Dark scales of caudo-

lateral tuft on tergites and sternites relatively small, about 0.07 mm. Remainder of

abdomen as for that of aquasalis.

MALE. Wing: 2.8 mm. Proboscis: 2.1mm. Forefemur: 1.45 mm. Abdomen:

about 2.6 mm. Head: Palpal segments 2 and 3 with cream dorsal stripe; segment 4

predominantly cream, with a basal and an apical white band, and with or without a

subbasal and a subapical dark ring. Antenna: About 0.7-0.8 length of proboscis.

Flagellar segment 1 with moderately long, white, lanceolate scales. Legs: Forefemur

0.7 length of proboscis. Basal plantar surface of foretarsal segment 5 with about 7

moderately long, spiniform setae. Claw on foreleg large, curved, acuminate; subme-

dian tooth long, tapering to a blunt, slightly recurved apex; basal tooth strongly de-

curved. i 3

MALE GENITALIA (fig. 19). Description based on 5 damaged specimens, includ-

ing holotype and 2 paratypes. Segment IX: Sternite long, about 0.2 length of side-

piece, curved, subtrapezoidal. Anterior apodeme on anterior border moderately de-

veloped, subtrapezoidal. Sidepiece: Tergal surface with 4,5 long, submedian, tergo-

medial bristles, basal 3 approximated; bristles mesad of tergomedial bristles moder-

ate in length. Parabasal spine tapering, slender, moderately long, about 2.0 length of

its tubercle. Basal apodeme moderately thick, 0.18 length of sidepiece. Longer,

more dorsal accessory spine slightly less than 0.5 length of sidepiece; more ventral

spine about 0.8 of longer spine. Internal spine subequal to shorter accessory spine,

apically retrorse. Clasper: Spiniform acuminate and moderately long, slightly less

than length of seta b (one specimen). Dorsal Claspette: About 0.6 length of side-

piece. Pedicel moderately broad; internal apodeme distinct. Leaflets broad, as in

section description. Ventral Claspette: On holotype slide, about 0.87 length of aede-

agus; width at apex about 0.2 length of claspette. Basal lobule large, expanded at

base; setae along basal margin long, 1.5-2.0 width of aedeagus. Lateral and ventral

surfaces (exclusive of basal lobule) with short setae extending to or nearly to apex.

Apex striated, rounded and appearing domelike. Preapical plate moderately large,

oval to semicircular, moderately sclerotized, located about 0.3 length of claspette ba-

sad of apex. Refringent structure not apparent. Phallosome: Aedeagus with apex

rounded, longer than wide; with a pair of long, pointed, sclerotized leaflets about 0.4

length of aedeagus, leaflets directed basolaterally and serrated with 4,5 small teeth

apically; subtriangular median sclerotizations at base of apex directed caudomesad

instead of mesad as in remaining species.

PUPA (fig. 19). Abdomen: about 2.8 mm. Trumpet: 0.48 mm. Paddle: 0.8 mm.

Description based on 2 laterally oriented specimens; outline of drawing based on stro-

dei, many of setal lengths approximate. Cephalothorax: Very dark brown. Hairs 4,

5-C 3 and 2 branched respectively near base. Hair 6-C 3 forked. Hair 7-C 2 branched.

Trumpet: Pinna long, about 4.5 length of meatus. Meatal cleft long, basally round-

ed. Metanotum: Hair 10-C single, short (0.1 mm). Hair 11-C slightly shorter than

12-C, 4 branched near base, median branch longer than lateral. Hair 12-C single,

about 2.0 length of 10-C. Abdomen: Very darkly pigmented brownish black. Hair

2-I 6 branched 0.25 from base, long (0.16 mm), about 2.0 length of 3-I. Hair 41 4

branched, short. Hair 5-I single and moderately long. Hair 6-I single and long. Hair

7-1 3 branched near base, slightly longer than 0.5 length of 6-I. Hair 9-I single, slight-

ly shorter than 6-I. Hair O-II,III 3 and 4 branched respectively, moderately long; O-

IV-VII 3-5 branched. Hair 1-II 9 branched, strongly developed; 1-III 7,9 branched,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 97

strong; 1-II,]I with median branches longer than lateral; 1-[V-VII single, very long,

1.5-1.8 length of segment, longer on anterior segments than on posterior. Hair 3-IV

5 branched, moderately long; 3-V 2 forked 0.5 from base, long. Hair 5-III 9

branched, subequal to 1-III; 5-IV 5 branched, lateral branches shorter than median;

5-V-VII single, long, subequal to length of segment. Hair 6-II single, probably slight-

ly longer than 7-II; 6-III single, long. Hair 7-II 3 branched; 7-III 2 forked, moderate-

ly developed; 7-V double, longer than 7-IV; 7-VI,VII single, long. Hair 8-III 2 forked,

moderately long; 8-IV 1,2 forked, moderately long; 8-V single, moderately long; 8-

VI, VII 2 and 3 branched respectively. Hair 9-II very small; 9-III strongly developed,

more than 2.0 length of 9-II, heavily pigmented; 9-IV stout, 2.0 of 9-III; 9-IV-VII

thick, acuminate, slightly curved, progressively longer from IV to VII, relatively

short, less than 0.33 length of segment; 9-VIII subequal to 9-VII. Hair 10-III 3

forked, moderately long. Hair 4-VIII 2,3 forked, moderately long. Terminal Seg-

ments and Paddle not clearly discerned because of lateral orientation of specimens.

Buttress about 0.7 length of paddle, serrated distally with small teeth. Hair 1-P mod-

erately short (0.05 mm).

LARVA (fig. 20). Head: 0.6 mm. Antenna: 0.25 mm. Anal Saddle: 0.3 mm.

Head: Moderately pigmented. Median tooth of mental plate moderately wide, less

than combined width of 2 adjacent teeth from one side. Inner and outer clypeal

hairs (2,3-C) single and barbed, with 8-10 and 4-9 barbs respectively; barbs of 3-C

longer than those of 2-C; 3-C about 0.6-0.7 length of 2-C; hairs 2-C closely approxi-

mated, clypeal index about 2.5. Hair 4-C 2,3 branched, short, not reaching base of

2-C; distance between insertion of 4-C and 2-C about 1.5 distance between insertion

of 2-C and 3-C. Hairs 8,9-C 3-5 branched (2-6), large; 8-C slightly shorter than 9-C.

Hair 12-C 47 branched, large, subequal to 9-C. Hair 15-C single or forked, long.

Collar wide dorsomedially, dark, heavily pigmented. Antenna: Darkly pigmented.

Mesal margin with numerous, strong, long spicules; ventral surface with few, moder-

ately strong, short spicules. Hair 1-A 5,6 branched (3-6), short, slightly longer than

width of antenna at point of insertion, inserted about 0.3 from base of antenna.

Thorax: Darkly pigmented. Submedian prothoracic group (1-3-P) with 1,2-P sharing

a common sclerotized tubercle; palmate hair 1-P with 9-12 long, moderately broad to

broad, pointed, lanceolate leaflets; 2-P 12-14 branched (12-17), about 3.0 length of

1-P. Hair 14-P 7-10 branched from a short, flattened stalk, lateral branches shorter

than median. Mesothoracic hair 1-M 20-30 branched (20-32). Metathoracic hair 2-T

moderately short. Palmate hair 3-T with 10-14 (10-16) moderately short, semitrans-

parent, lanceolate leaflets. Abdomen: Hair 0-II-VII moderately large to large; O-II,

III, V-VII 47 branched; 0-IV 6,7 branched. Palmate hair 1-I with 12-16 narrow, mod-

erately long, semitransparent leaflets; 1-II-VII with long, moderately narrow, pig-

mented leaflets; 1-II, VII smaller than 1-III-VI; 1-III,VI slightly smaller than 1-IV,V.

Hair 2-II 4-7 branched, large; 2-III 3-6 branched (3-7), moderately long; 2-IV single,

medium, about 2.0 length of leaflets of 1-IV; 2-V single, long. Hair 5-I 3-5 branched

(3-8); 5-II 8-11 branched, moderately developed. Hair 9-I 5-9 branched, moderately

large. Hair 11-I 2-4 branched, moderately large. Hair 13-I,II,III 5-7, 7-10 (5-12) and

6-10 branched respectively, moderately small; 13-IV 4,5 branched, moderately large;

13-V 3,4 branched (3-7), moderately large to large. Spiracular Lobe: Pecten with 13-

17 teeth; many of median teeth subequal, with a few mixed long and short. Lateral

arm of spiracular apparatus short, similar to that of aquasalis but narrower. Hairs 8,

9-S 3-5 branched, medium. Anal Segment: Hair 1-X longer than saddle; inserted on

ventral margin of saddle at base of indentation, or less often on saddle near ventral

margin. Anal gills slightly shorter than or about equal to length of saddle.

98 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

DISCUSSION. The female of anomalophyllus is very similar in appearance to

those of noroestensis and strodei. It is not possible to separate anomalophyllus from

the 2 latter species, particularly strodei, except by careful correlation of the charac-

ters given below and by comparing them with the descriptions of noroestensis and

strodei. Locality data are also extremely useful in identifying anomalophyllus because

of its limited distribution in Panama and Costa Rica. Due to insufficient material,

anomalophyllus is not included in either the key to females or the key to pupae. An.

anomalophyllus can be distinguished from the other species in the Oswaldoi Sub-

Group, except for strodei, in the female by the combination of (1) palpal segment 4

apparently with an indistinct brown ventral stripe, (2) prescutellar space horseshoe

shaped, (3) foretarsal segment 2 0.35 (0.29-0.42) apically white and segment 3 0.51

(0.23-0.78) apically white, (4) foretarsal segment 5 with a cream band in about apical

0.5, (5) midtarsal segment 5 cream in about apical 0.5, (6) hindtarsal segment 2 with

brown band in basal 0.4 (0.30-0.45), (7) light spots of wing white except for vein C

and possibly R, (8) humeral light spot of vein C about 2.5 (2.3-4.0) length of basal

dark spot and subcostal light spot 0.23-0.60 length of subcostal dark spot and (9)

caudolateral dark scale tufts of tergites and sternites of abdomen relatively small,

about 0.07 mm; in the male genitalia by (1) aedeagus with long serrated leaflets about

0.4 length of aedeagus, (2) basal lobule large, expanded at base, with long setae along

basal margin 1.5-2.0 width of aedeagus and (3) preapical plate moderately large, oval

to semicircular, moderately sclerotized; and in the larva by the combination of (1)

hairs 2,3-C single and barbed, hairs 2-C closely approximated, clypeal index about

2.5, (2) collar wide dorsomedially, dark, heavily pigmented, (3) hairs 1,2-P sharing a

common tubercle, (4) palmate hair 1-P with 9-12 long, moderately broad to broad,

pointed, lanceolate leaflets, and 2-P 12-14 branched (12-17), (5) hair 1-M 20-30

branched (20-32), (6) hair 13-V moderately large to large and (7) hair 1-X inserted on

ventral margin of saddle at base of indentation, or less often on saddle near ventral

margin. |

It is unclear exactly what species are the closest allies of anomalophyllus. | am

placing anomalophyllus in the Oswaldoi Complex on the basis of the similarities in

the male genitalia. In general appearance, the ventral claspette is similar to those of

the other species in the Oswaldoi Complex, including the presence of setae on the

ventral and lateral surfaces of the fused portion of the claspette. An. anomalophyllus

retains long serrated leaflets on the aedeagus, an ancestral character it shares with no

other species in the Albimanus Section. Since phylogeny cannot be demonstrated on

the basis of ancestral characters, the presence of the leaflets does not aid in establish-

ing phylogenetic relationships. However, the female and particularly the larva of ano-

malophyllus are very similar to those of strodei. The larva is so much like that of

strodei, that identification is possible only when all diagnostic characters given in the

Key are correlated. It is probable that very early in the divergence of the species in

the Oswaldoi Subgroup, a population of an ancestral species became isolated in the

Panamanian Isthmus and speciated into what is now recognized as anomalophyllus.

Therefore, anomalophyllus, sharing characters with both the Oswaldoi Complex and

Strodei Complex, represents a separate intermediate phylogenetic line. However, at

this time I do not feel that enough information and material are available to justify

the establishment of a separate monotypic taxon of higher category for anomalophyl-

lus.

BIONOMICS. An. anomalophyllus is a rare mosquito of limited distribution

known from very few collections. All collections of anomalophyllus have been from

running streams at elevations from less than 100 m to about 600 m. Komp (1936a:

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 99

160-161) stated that the stream at the type locality was shaded. Kumm, Komp and

Ruiz (1940:391, 408) took one larva from a sunlit pooled stream at Finca Canada,

near Turrialba, Costa Rica. In 1971, D. A. Schroeder collected one fourth instar lar-

va along a slow moving, clear, fresh, semipermanent, grassy stream in a cacao planta-

tion outside of Zent, Costa Rica.

MEDICAL IMPORTANCE. An. anomalophyllus is of no medical importance.

DISTRIBUTION (fig. 2). An. anomalophyllus is a rare species found only in

southern and central Costa Rica and on the Atlantic coast of western Panama.

Material Examined: 36 specimens: 4 males, 5 male genitalia, 8 females, 2 pupae,

17 larvae; 2 individual rearings: 1 larval, 1 incomplete.

COSTA RICA (27 specimens: 3M, 2Mgen, 8F, 2P&p, 12L&l; 2 ind rear: 11, linc). Cartago:

Turrialba, Finca Canada, [H. Kumm], No. 33, 1M. Limon: Amubre [Amubri], [H. Kumm], No.

1496, 1 lpF [JH], 4F, 41; same locality, 2F. Chase, Jan 1930, W. Komp, 1M, 1Mgen; same data

except Feb 1932, 1L, 11. Suerre, [H. Kumm], No. 1426, 1Mgen; same data except No. 1427, 1L

[JH], 11P,1L. Zent, CR 456, 1L. Province not specified: Margarita, 1 1 [JH]. Volcan, [H.

Kumm], No. 1071, 1M, 1F.

PANAMA (9 specimens: 1M, 3Mgen, 5L&l). Bocas del Toro: Almirante, 19 Dec 1928, W.

Komp, | 1; same data except 1 Jan 1929, 3 1; same data except Feb 1929, 1M and 1Mgen (holo-

type), 2Mgen (paratypes); same data except date not specified, 1L.

8. Anopheles (Nys.) rangeli Gabaldon, Cova Garcia & Lopez

Figs: 3. 4,6; 21,22

1940. Anopheles (Nyssorhynchus) rangeli Gabaldon, Cova Garcia and Lopez 1940:13-20. TYPE:

Holotype male (309.5), Puerto Cabello (Carabobo), Venezuela [MDM].

Anopheles (Nyssorhynchus) rangeli of Gabaldon and Aquilera (1940:69-73, 76); Galvao (1941:93;

1943:149, 150, 151); Rozeboom and Gabaldon (1941:91-98); Galvao and Damasceno (1942:

121-122); Simmons and Aitken (1942:39, 46, 52, 61, 96); Cerqueira (1943:19; 1961:126); Gal-

vis (1943:91-92); Causey, Deane and Deane (1944:2; 1946:27); Lane (1944:263; 1949:403;

1953:266-268); Deane, L. M., Causey and Deane (1946:7, 9, 12; 1948:897-899); Deane, M. P.,

Causey and Deane (1946:38, 43, 45); Gabaldon and Cova Garcia (1946b:114-118; 1952:178,

195-196); Senevet (1948a:278); Bates and de Zulueta (1949:134-139); Levi-Castillo (1949: 10,

15, 28, 72, 76, 86); Rey and Renjifo (1950:534, 537); Bejarano (1957:326, 329, 331); Stone,

Knight and Starcke (1959:34); Vargas (1959:377, 383); Forattini (1962:395-396); Belkin,

Schick and Heinemann (1965:72); Morales-Ayala (1971:139); Cova Garcia and Sutil O. (1976:

28; 1977:23, 46, 64, 85); Cova Garcia, Pulido F. and Amarista M. (1978:157).

Anopheles rangeli of Gabaldon, Cova Garcia and Arevalo (1940:28-29); Cova Garcia and Marcano

(1941:57-58); Rozeboom (1941:103); Russell, Rozeboom and Stone (1943 :26, 30, 41, 49);

Renjifo and de Zulueta (1952:600-601); Komp (1956:39-40); Foote and Cook (1959:12, 13,

23, 26, 29, 32); Cova Garcia (1961:37-38, 68, 90-91, 114, 125-126, 153); Vincke and Pant

(1962:2-4, 6, 7, 10, 12, 13); Mattos and Xavier (1965:273); Stojanovich, Gorham and Scott

(1966b:22, 30, 44); Gorham, Stojanovich and Scott (1967:60; 1973:112, 138, 150); Elliott

(1972: 757): Cova Garcia and Sutil O. (1975a:22; 1975b:213).

FEMALE (fig. 6). Wing: 3.3 mm. Proboscis: 2.0 mm. Palpus: 2.0 mm. Porter:

mur: 1.2mm. Abdomen: about 2.6 mm. Head: Integument light brown to dark

brown with light pruinose areas. Interocular distance 3,4 ommatidial diameters. Pro-

boscis about 1.7 length of forefemur. Erect scales on vertex as those of oswaldoi,

long, narrow, tapering at base into threadlike stalk. Palpal segment 2 occasionally

with many, light, dorsal scales; segment 3 with an indistinct to conspicuous, cream-

ish, dorsolongitudinal stripe, apex with a moderately large, white band; segments 4

and 5 predominantly white; base of 4 with a moderately small, dark band and apex

with a small to indistinct, dark band; segment 5 occasionally with a few, dark, basal

scales; segment 4 usually without brown scales forming a distinct ventral stripe.

100 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Antenna: Dorsomesal and mesal surfaces of flagellar segment | with numerous,

elongate, white scales. Flagellar segments 2-13 each with 8-10 silver setae in basal

whorl. Thorax: Integument extremely variable, tan to very dark brown. Scutum

largely reddish cream, tan or silver, particularly in pruinose areas. Pleuron often with

integument very light so that pruinose light regions hard to distinguish from nonprui-

nose areas. Anterior promontory scales extending dorsad onto acrostichal line. Pre-

scutellar space usually large and subtriangular. Sp with 4-8 setae and a few, elongate,

light scales. Pra with 6-10 elongate, light, setiform scales, and a patch of elongate

lanceolate scales, with some shorter obovate scales interspersed. Upper stp with 4

(3-5) long setae and a patch of about 10 cream to white, small, obovate to lanceolate

scales in a horizontal arc. Lower stp with one long, usually light seta, and 4-8 cream

to white scales in diagonal patch. Upper mep with about 6 long, dark setae; anterior-

ly often with a setiform scale shorter than setae; often with 1-4 light, moderately

small, obovate scales, either dorsal or anterior to setae; often scales rubbed off or dif-

ficult to see. Legs: Light scales on coxae white, grayish white or very light cream.

Foretarsal segments 2 and 3 with a cream to white band in apical 0.30-0.45 and 0.7

(0.60-0.85) respectively ; segment 4 predominantly dark, with or without apical cream

ring; segment 5 light in about apical 0.5. Midtarsal segment 2 with light band in apical

0.10-0.35; segment 3 with small light apical band; segment 4 with or without light

apical scales; segment 5 light in about apical 0.5. Hindtarsal segment 2 with a brown

band in basal 0.28 (0.24-0.35); segment 5 cream to white in about apical 0.5. Wing:

Light spots of vein C very light cream, remaining veins with light spots white. Wing

characterized by several large light spots. Vein C humeral light spot about 1.8-3.5

(1.0-3.7) of basal dark spot; subcostal light spot usually greater than 0.5 (0.45-1.00)

of subcostal dark spot; preapical light spot large, 0.40-0.65 of preapical dark spot. R

presectoral and sectoral dark spots usually small, occasionally sectoral dark spot ab-

sent. Rs-R»,3 highly variable. R,,3; and R, preapical light spots usually large. M pre-

dominantly light, sectoral dark spot often indistinct; preapical dark spot not extend-

ing onto M,,,5. Cu, subcostal and preapical light spots small to moderately long; oc-

casionally subcostal spot absent. Apical light fringe spot large, conspicuous and un-

divided, or with a few dark scales anterior to apex of Ry,;; additional, moderate size,

light fringe spots at apices of Rais, Mys2, M34, Cu,, Cu, and A; occasionally a con-

spicuous, fairly large, light spot between base of wing and A. Abdomen: Sternite I

with a few, moderately long to long, dark setae, and usually a few, small, inconspicu-

ous, obovate to moderately lanceolate, light scales in basal 0.5. Caudolateral scale

tufts of segments II-VII with scales large, dark brown, obovate and conspicuous.

MALE. Wing: 3.2 mm. Proboscis: 2.45 mm. Forefemur: 1.4mm. Abdomen:

about 3.0 mm. Essentially as in female except for sexual characters. Head: Palpal

segments 2 and 3 with a light, dorsal stripe; lateral surface of segment 3 speckled with

cream and dark scales; segment 4 extensively light, occasionally with a subbasal and a

subapical, small, brown band; base and apex of 4 usually white, medially cream with

a few, scattered, dark scales. Antenna: About 0.8 length of proboscis. Flagellar seg-

ment 1 with numerous, conspicuous, long, white, lanceolate scales on dorsomesal sur-

face. Legs: Forefemur about 0.6 length of proboscis. Basal plantar surface of fore-

tarsal segment 5 with about 6 moderately short, heavy, spiniform setae; 2 apical spin-

iform setae longer than others. Claw on foreleg similar to that of aquasalis, except

submedian tooth slightly more attenuate toward apex.

MALE GENITALIA (fig. 21). Segment VIII: Tergite and sternite with narrow

lanceolate scales; scales broader on sternite than tergite. Segment IX: Sternite long,

about 0.25 length of sidepiece, subtrapezoidal. Anterior apodeme long, subtriangular,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 101

may not extend entire width of sternite. Sidepiece: Moderately narrow, conical.

Tergal surface usually with 3,4 long, submedian, tergomedial bristles and one or 2

subapicolateral bristles; with moderately short bristles mesad of tergomedial bristles,

except for occasionally a stronger, longer, basomesal bristle. Parabasal spine slender,

about 2.0 length of its tubercle. Basal apodeme long and narrow, about 0.25 length

of sidepiece. Longer, more dorsal accessory spine about 0.5 length of sidepiece;

more ventral spine about 0.7-0.8 of longer spine. Internal spine subequal to shorter

accessory spine, strongly retrorse apically. Clasper: Spiniform moderately developed,

acuminate, subequal to seta b. Dorsal Claspette: Pedicel moderately broad; base

rounded and usually curved mesad. Dorsal leaflet with a moderately developed baso-

mesal projection. Ventral Claspette: Large, about 0.5 length of sidepiece; width at

apex 0.4-0.5 length of claspette. The nonfused dorsal lobe may appear to project be-

yond apex of fused ventral lobe, depending on angle of observation. Basal lobule

very large, expanded laterally at base and usually curved ventrally; setae along basal

margin strong, reflexed caudally and long, about 1.5 length of aedeagus; basomesal

margin with a concentration of long setae directed caudally into mesal cleft. Ventral

and lateral surfaces (exclusive of basal lobule) with short setae about as long as width

of aedeagus; setae extending to or nearly to apex. Apex truncate; lateral margins ab-

ruptly angled, rounded, sclerotized; apex medially membranous and weakly emargi-

nate. Preapical plate small, oval, heavily sclerotized; located about 0.15 length of

claspette basad of apex; convex apically, flat or slightly concave basally, occasionally

with small basomesal projection, often produced basolaterally into short, caudolater-

ally directed, sharp points. Membranous area transparent, horseshoe shaped, immedi-

ately basad of preapical plate, extending basad bordering mesal cleft. Refringent

structure in shape of an inverted horseshoe, often indistinct, without lateral arms.

Phallosome: Aedeagus about 1.0-1.2 length of ventral claspette; apex rounded with

tapering sides, slightly longer than wide; with or without very small membranous or

weakly sclerotized leaflets.

PUPA (fig. 21). Abdomen: 2.5 mm. Trumpet: 0.45 mm. Paddle: 0.7 X 0.5 mm.

Cephalothorax: Hairs 1-3-C subequal; 1,2-C 2,3 branched; 3-C 1-3 branched. Hairs

4,5-C 2-4 forked (1-4); 4C slightly shorter than 5-C. Hair 6-C single or 2,3 forked in

apical 0.5, subequal to 7-C. Hair 7-C 2,3 branched, branches subequal or one branch

slightly longer than other(s). Trumpet: Very similar to that of aquasalis. Pinna

weakly pigmented; moderately long, 3.8-5.4 (3.6-5.4) length of meatus; in lateral as-

pect, appearing broad medially and tapered toward apex. Meatal cleft long, open,

basally rounded. Metanotum: Hair 10-C single, heavy, moderately short. Hair 11-C

4-6 branched (2-6), subequal to or slightly longer than 10-C. Hair 12-C single or dou-

ble, about 2.0 length of 10-C. Abdomen: Hair 2-I 5-8 forked (2-8) 0.20-0.33 from

base, occasionally dendritic, long; 3-I single or double, subequal to 2-I. Hair 4-I 46

branched (3-6), moderately long. Hair 5-I 1-3 branched, moderately long. Hair 6-I

single or occasionally double, long, about 2.0 length of 7-I. Hair 7-1 3,4 branched (2-

5) near base. Hair 9-I single, slightly shorter than 6-I. Hair O-II-IV 47 branched (3-

7), moderate; sum of branches of O-II and O-III ranging from 8 to 14; 0-V,VI 3-5

branched, smaller than O-II-IV; 0-VII 2-4 branched. Hair 1-II, III strongly developed;

1-II 5-9 branched (3-9); 1-III 4-7 branched (2-10); 1-IV-VII strong, very long, 1.7-2.0

length of segment. Hair 3-IV,V 3-6 (2-6) and 2-4 forked respectively, moderately

long. Hair 5-III 4-8 branched (3-8), strong; 5-IV 1-4 branched, median branch much

longer than lateral; 5-V-VII single, long, about length of segment. Hair 6-II single or

occasionally double, about 2.0 length of 7-II; 6-III single, long. Hair 7-II 3,4

branched (2-5); 7-ITI,IV 2-4 branched (1-4), small; 7-V single to triple; 7-VI,VII single

102 | Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

or rarely double, long. Hair 8-III 3-5 branched (2-5); 8-1V, VII 2-4 branched (1-4); 8-V,

VI 1-3 branched (1-4). Hair 9-II minute, unpigmented; 9-III about 2.0 length of 9-II,

unpigmented or occasionally pigmented; 9-IV-VII strongly pigmented; 9-IV moder-

ately pointed, at least 2.0 of 9-III; 9-V moderately thin, curved, acuminate, usually

2.3 (2.0-4.5) length of 9-IV; 9-VI-VIII thin, acuminate, subequal to 0.5 length of seg-

ment; 9-VI,VII more curved than 9-VIII; 9-VI slightly shorter than 9-VII, VII; 9-VII

1.1-1.5 (1.1-1.7) length of 9-VI. Hair 10-III 2-4 forked (1-4) in basal 0.3-0.5, moder-

ately developed. Hair 4-VIII 2,3 forked (2-4) about 0.5 from base, subequal to 9-

VIII. Terminal Segments: ‘Male genital lobe as that of aquasalis only broader at base,

and sides tapering more medially toward apex. Paddle: Broad distally, obovate, very

weakly emarginate at insertion of hair 1-P. External buttress 0.7 length of paddle,

serrated apically. Spicules on external and internal margin as in aqguasalis. Hair 1-P

moderately long; 2-P single or double, subequal to 1-P.

LARVA (fig. 22). Head: 0.6 mm. Antenna: 0.25 mm. Anal Saddle: 0.25 mm.

Head: Moderately pigmented. Median tooth of mental plate as broad as combined

widths of 2 adjacent teeth from one side, tapering abruptly to a point. Inner and

outer clypeal hairs (2,3-C) single, and simple or barbed; hairs 2-C widely spaced, cly-

peal index about 1.67; 3-C slightly shorter than 2-C. Hair 4-C 2-3 branched (1-4),

short, not extending to base of 2-C. Hairs 8,9-C 4-6 (2-6) and 5-7 branched respec-

tively, dendritic, moderately large. Hair 12-C 3-5 branched, moderately large, sub-

equal to 9-C. Hair 15-C 2-4 branched. Collar wide dorsomedially, dark, strongly pig-

mented. Antenna: Moderately pigmented. Mesal margin with long, moderately thin

spicules extending almost to base. Hair 1-A 46 branched, short, slightly longer than

width of antenna at point of insertion, inserted in basal 0.25 or less of antenna.

Thorax: Moderately pigmented. Submedian prothoracic group (1-3-P) with or with-

out 1,2-P sharing a common tubercle; palmate hair 1-P with 9-13 (8-14) long, broad,

lanceolate leaflets; 2-P 13-17 branched (10-19), about 4.0 length of leaflets of 1-P.

Hair 14-P 6-9 branched (5-9) from a short to moderately short, flattened stalk;

branches long, lateral branches slightly shorter than median. Mesothoracic hair 1-M

28-37 branched, lateral branches equal to or slightly longer than apical branches.

Metathoracic hair 2-T moderately long, reaching or not reaching posterior margin of

thorax. Palmate hair 3-T with 12-18 semitransparent, moderately long, moderately

broad to broad, spreading leaflets. Abdomen: In general, tergal hairs shorter in

length and inserted closer to midline than in other species in Oswaldoi Complex.

Hair 0-II-VI moderately developed; 0O-II,III 6-9 branched; 0-IV,V 4-8 branched (3-10);

0-VI 4-6 branched (4-7). Palmate hair 1-I-VII with moderately narrow to broad, acu-

minate, lanceolate leaflets; 1-I-VII longer than those of other species in Oswaldoi

Complex; 1-I with 13-16 (13-17) weakly pigmented leaflets; leaflets of 1-I,H, VII

slightly shorter than those of 1-III-VI. Hair 2-II 5-8 branched (5-10); 2-III 3-5

branched (3-6), relatively short, 1.5-2.0 length of leaflets of 1-III; 2-IV single, moder-

ately short, slightly longer than leaflets of 1-IV; 2-V single, about 3.0-4.0 length of

leaflets of 1-V. Hair 5-I 3-5 branched, short, inserted 0.75-1.00 its length from later-

al margin of abdomen; 5-II 5-9 branched (5-12), short, inserted as 5-I. Hair 9-I 4-6

branched (3-6), moderately small. Hair 11-I 2 branched (2-4), moderately large.

Hair 13-I, 11,1 4-7 (3-7), 6-10 (5-10) and 7-9 (6-10) branched respectively, small; 13-

IV 3 branched (3-5), moderately short, slightly longer than leaflets of 1-I[V; 13-V 3-5

branched, moderately long, subequal to 2-V; 13-VI 6-11 branched, small. Spiracular

Lobe: Pecten with 15-19 teeth; many of median teeth subequal in length. Lateral

arm of spiracular apparatus short as in aquasalis, directed caudolaterally. Hairs 8,9-S

3-5 forked (3-8). Anal Segment: Covered with very fine spicules over about apical

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 103

0.7 of segment; fewer spicules in basal 0.5 of membranous portion of segment. Sad-

dle tan to brown. Hair 1-X longer than saddle; inserted on saddle near ventral mar-

gin of saddle or on ventral margin at base of indentation, rarely inserted just ventrad

of ventral margin; inserted about 0.3 cephalad of caudal margin. Anal gills about

0.35-0.40 mm long, much longer than saddle.

DISCUSSION. An. rangeli can be distinguished from the other species in the Os-

waldoi Subgroup in the female by the combination of (1) upper mep often with 1-4

light obovate scales, (2) foretarsal segment 2 with a light band in apical 0.30-0.45

and segment 3 with light band in apical 0.7 (0.60-0.85), (3) hindtarsal segment 2

with a brown band in basal 0.28 (0.24-0.35), (4) wing with several large light spots,

humeral light spot of vein C about 1.8-3.5 (1.0-3.7) length of basal dark spot, (5)

subcostal light spot of vein C usually greater than 0.5 (0.45-1.00) length of subcostal

dark spot, (6) preapical light spot of vein C 0.40-0.65 of preapical dark spot, (7) pre-

sectoral and sectoral dark spots of vein R usually small, (8) vein M predominantly

light with sectoral dark spot often indistinct and (9) apical light fringe spot large,

conspicuous and usually undivided; in the male genitalia by (1) ventral claspette

large, apex truncate and wide, width at apex 0.4-0.5 length of claspette, apex with

abruptly angled, rounded lateral margins, (2) basal lobule very large, usually curved

ventrally, with long setae along basal margin about 1.5 width of aedeagus, (3) long

setae along basomesal margin of basal lobule concentrated and directed caudally into

mesal cleft, (4) preapical plate small, oval, heavily sclerotized, convex apically, flat or

slightly concave basally, occasionally with small basomesal projection and (5) apex

of aedeagus with or without very small, membranous or weakly sclerotized leaflets,

apex slightly longer than wide; in the pupa by the combination of (1) hair 7-C with

branches subequal or one branch slightly longer than other(s), (2) pinna moderately

long, about 3.8-5.4 length of meatus, in lateral aspect appearing broad medially and

tapered toward apex, (3) hair 11-C subequal to or slightly longer than 10-C, (4) hair

2-I 5-8 forked (2-8) 0.20-0.33 from base, occasionally dendritic, long, subequal to

3-I, (5) hair O-II moderate, 5-7 (4-7) branched, (6) sum of branches of hairs O-II and

Q-III ranging from 8 to 14 and (7) hair 9-III about 2.0 length of 9-II, unpigmented or

occasionally pigmented, 9-V usually 2.3 (2.0-4.5) length of 9-IV, 9-VII thin, acumi-

nate, about 0.5 length of segment, 1.1-1.5 (1.1-1.7) length of 9-VI; and in the larva

by the combination of (1) hairs 2,3-C single, and simple or barbed, hairs 2-C widely

spaced, clypeal index about 1.67, (2) hair 4-C short, not extending to base of 2-C,

(3) hairs 8,9-C 4-6 (2-6) and 5-7 branched respectively, dendritic, moderately large,

(4) palmate hair 1-P with 9-13 (8-14) long, broad leaflets, (5) hair 2-IV single, moder-

ately short, (6) hair 5-I,II short, inserted 0.75-1.00 its length from lateral margin of

abdomen, (7) hair 13-IV 3 branched (3-5), moderately short, (8) lateral arm of spira-

cular apparatus short and (9) hair 1-X inserted on saddle near ventral margin of sad-

dle or on ventral margin at base of indentation, rarely inserted just ventrad of ventral

margin.

An. rangeli forms the sister group of nuneztovari and trinkae. It differs from these

2 species in the female usually by (1) the absence of ventral dark line on palpal seg-

ment 4, (2) presence of a few light upper mep scales and (3) presence of a small basal

dark spot and a large subcostal light spot on vein C. The male genitalia of these 3

species are very similar, rangeli differing only in the (1) presence of a concentration

of long setae along the basomesal margin of the basal lobule, (2) development of the

preapical plate and (3) structure of the apex of the ventral claspette to some degree.

The pupa of trinkae and the larva of noroestensis are very much like those of rangeli

and can be distinguished only by the characters given in the keys. Locality data are

104 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

extremely useful in distinguishing the larva of rangeli from noroestensis, since the

distributions of the 2 species do not overlap, except possibly in Bolivia and south-

western Brazil.

There seems to be no significant variation between the populations I have studied.

Intrapopulational variation seems as great as or greater than interpopulational varia-

tion. Gabaldon and Aguilera (1940:69-72, 76) examined material from Maracay,

Dividive, Pampanito and Pampan, Venezuela, and could find no significant differen-

ces, although the specimens from Trujillo had a tendency to be darker and smaller.

They also studied progeny rearings and found considerable variation between sib-

lings; in one family they studied (LV4), the dark band on hindtarsal segment 2 var-

ied from 28.3 to 42.0%. Also, as in all the species in this section, there is consider-

able variation in the relative lengths of the wing spots and in the tarsal banding.

BIONOMICS. The immatures of rangeli occur in marshy depressions, temporary

ground pools, animal and wheel tracks, semipermanent ditches, stream margins, and

lakes. They are usually found in full sun or partial shade, and usually not in the for-

est proper but in open glades, meadows, and scrub or grassland areas. There is usual-

ly abundant vegetation in the breeding site, and occasionally the aquatic habitat has

decaying vegetation or is contaminated with feces. The immatures are always found

in fresh water. Gabaldon, Cova Garcia and Arevalo (1940:28-31) reported that the

larvae of rangeli cannot tolerate sodium chloride in concentrations as low as 0.05%.

The time of development from egg to adult averaged 24 days for specimens from La

Victoria (Aragua), Villa de Cura (Aragua) and Puerto Cabello (Carabobo), Venezuela;

however, the temperature at which the immatures developed was not reported. The

average number of eggs deposited at one oviposition varied from 23 to 217 with a

mean of 80.3. In Colombia, the seasonal peak in the populations of rangeli occurs in

June at the beginning of the rainy season, and rangeli continues to be very abundant

until December or January (Bates and de Zulueta 1949:136, 137). The larvae and

pupae are commonly associated with 3 species in the Albimanus Section, trinkae, tri-

annulatus and, in some areas, strodei. They have also been collected with oswaldoi

(Gabaldon 1940). Other species found in association with rangeli are An. (Nys.) ar-

gyritarsis, An. (Ano.) punctimacula, Culex (Cux.) corniger, Cx. (Cux.) coronator

group, Cx. (Cux.) declarator group, Psorophora (Gra.) confinnis group and Uranotae-

nia (Ura.) lowii. An. rangeli occurs in lowlands and at higher elevations; our highest

collection was made at 950 m above sea level.

The adults are predominantly exophilic. Rey and Renjifo (1950:537) collected

2414 anophelines inside houses from September to November near Cucuta, Colom-

bia, of which only 0.62% were rangeli. Of the 3722 adults collected by Deane, Cau-

sey and Deane (1948:899), only 25 or 0.7% were from inside houses. In 131 hours

of collecting inside houses, they found only 5 specimens of rangeli; however, in less

than 41 hours collecting outside, they captured 1391 individuals. The majority of

the specimens were collected on horses or bulls, although many were biting man. In

Boca de Acre, in a collection made at sundown simultaneously from a cow and a

man, 5 females of rangeli were collected from the cow and 8 from the man. Elliott

(1972:757) reported that in Peru the peak hours of biting by rangeli were 1800-2000

h and 0400-0600 h.

MEDICAL IMPORTANCE. Very little is known about the vector capacity of

rangeli. It does not seem to be a vector of malaria, although Forattini (1962:396)

stated that it has been suspected of transmitting malaria in Ecuador. Deane, Causey

and Deane (1948:899) dissected 363 females from Rio Branco (Acre), Brazil, and

found none infected with Plasmodium sp. Rey and Renjifo (1950:537) did not find

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 105

rangeli naturally infected in the Cucuta area of Colombia during a malaria epidemic

in which Plasmodium falciparum (18%), P. vivax (55%) and P. malariae (27%) were

present in the human population.

DISTRIBUTION (fig. 3). An. rangeli occurs in the upper Amazon and Orinoco

basins, Colombia, Venezuela, Ecuador (Morona Santiago, Napo, Pastaza), and south

through eastern Peru and into northern Bolivia (Beni, Santa Cruz).

Material Examined: 1181 specimens: 196 males, 113 male genitalia, 437 females,

267 pupae, 168 larvae; 126 individual rearings: 53 larval, 44 pupal, 29 incomplete;

11 progeny rearings.

BOLIVIA (7 specimens: 2M, 2Mgen, 3F). Beni: San Ignacio, 18 Sep 1943, Carr, 3F. Todos

Santos, Jun 1946, E. Soracho, BOL 36, 2M, 2Mgen.

BRAZIL (1Mgen, 1F). Acre: Rio Branco, 1949, A. Galvao, 1Mgen [JH]. Amazonas: Boca do

Acre, det. Deane, 1F [CU].

COLOMBIA (780 specimens: 130M, 26Mgen, 317F, 186p, 119L&l; 43 ind rear: 17 1, 26 inc;

11 progeny rearings). Boyaca: Puerto Boyaca, COM 650. Caldas: La Dorada, 22 Jun 1943, KO

120A-2, 1M; same locality, 23 Jun 1942 [1943], 1M, 1F; same locality, 25 Jun 1943, W. Komp,

KO 120A-11, 3M; same locality, 26 Jun 1943, KO 120A-5, 1M, 1F. Caqueta: Tres Esquinas, COM

490, 491, 493. Florencia, H. Kumm, 1F [BM]. Meta: Restrepo, Jul 1935, KO 120A-4, 1M, IF;

same locality, Sep 1935, W. Komp, KO H-19-8, 1M, 1F; same locality, 13 Jun 1937, 1M, 4F; same

locality, COB 50; same locality, IMgen. Vega Grande, 24 Nov 1936, COR 147, 11F. Villavicencio,

26 May 1941, M. Bates, CV 265, 2M, 1Mgen; same locality and collector, 30 May 1941, CV 267,

3M, 1Mgen, 1F; same locality and collector, 26 Jun 1941, 1Mgen; same locality, 1 Jun 1942, 1F;

same locality, 3 Jun 1942, W. Komp, 1M; same locality and collector, Jun 1942, KO 120A-3, 1M;

same data, KO 120A-21, 6M, 2F; same locality and collector, 71942, KO 120A-23, 14M, 2F; same

locality, 15 Jul 1943, KO 120A-8, 20F; same locality, 15 Aug 1943, KO 120A-6, 7F; same locality,

22 May 1944, KO 120A-13, 28F; same locality and date, KO 120A-16, 1M; same locality, 23 May

1944, W. Komp, KO 120A-14, 27F; same locality and collector, 8 Jun 1944, 40F; same data, KO

120A-17, 18F; same locality, 20 Jun 1944, COK 47,37F; same locality, 1944, M. Bates, 2 1; same

data, CV 90, 1F; same data, CV 91, 1F; same data, CV 93, 1M; same locality, 3 May 1947, 3Mgen

[JH] ; same locality, 3 May 1947, L. Rozeboom, CV-P | (progeny rearing), 18p, 7L, 4 1; same data,

CV-P 2 (progeny rearing), 17p, 3L, E; same data, CV-P 3 (progeny rearing), 5M, 5Mgen, 3F, 14p,

3L; same data, CV-P 4 (progeny rearing), 15p, 5L; same data, CV-P 5 (progeny rearing), 8 lp, 14p,

1 1; same data, CV-P 6 (progeny rearing), 16p, 3L, 1 1; same data, CV-P 7 (progeny rearing), 15p, 8

L, 11; same data, CV-P 8 (progeny rearing), 12p, SL, 3 1; same data, CV-P 9 (progeny rearing), 13p,

61; same data, CV-P 10 (progeny rearing), 5p, 3 1;same data, CV-P 11 (progeny rearing), 10 lp;

same locality and collector, 6 May 1947, CV-P 17, 1F; same data, CV-P 18, 1F; same data, CV-P

27, 1F; same locality, 9 May 1947, 1Mgen [JH], 3L; same data, [CV] 370.10, 3L, 1 1 [JH]; same

locality, 14 May 1947, CV 379, 1F; same locality, 23 May 1947, M. Bates, Cabo, L. Rozeboom,

CV 389, 1M; same data, CV 391, 3M, 1F; same locality and date, 4p, 4L [JH] ; same locality, 27

May 1947, M. Bates, Cabo, L. Rozeboom, CV 394, 2F; same data, CV 395, 3M, 3F; same locality,

May 1947, CV 396A, 1F; same locality, 10 Jun 1943 [1947], L. Rozeboom, Cabo, [CV] 423.2,

1 lp [JH] ; same locality, 28 Jul 1948, [CV] C595.104, 1Mgen [JH]; same locality, CV 732, 7 Ip,

SL; same locality, COB 39, 42, 53, 60, 61, 63, 70, COM 607; same locality, 27 Jul 1974, J. Kitz-

miller, COZ 41, 62M, 77F; same locality, 4M, 2F. Locality not specified, 24 Nov 1936, COR 147

A, 8F. Santander: Barranca[bermeja] , 8 Sep 1943, D. Fierro, COT 68, 2M, 1Mgen.

ECUADOR (182 specimens: 35M, 14Mgen, 33F, 65p, 35L&l; 67 ind rear: 33 1,34p). Morona

Santiago: Sucua, 6Mgen. Locality not specified, Zamosa, 2Mgen. Napo: Limoncocha, 3 Jun

1977, Y. Huang, coll. 45, 1 pM, 1 pF, 1 lpF; same data except 8 Jun, coll. 57, 4 pM, 4 pF, 7 IpM.

Tena, 1.5 km S of, 13 May 1977, Y.Huang, coll. 26, 5 pF, 2 lpF; same data except 32 km S of

Tena, coll. 27, 3 lpF; same data except 12.5 km SE of Tena and 24 May, coll. 38, 1Mgen, 1 pM, 3

pF, 2 lpM, 4 lpF; same data except 11.7 km SW of Tena and 25 May, coll. 40, 1Mgen, 3 pM, 6 lpM,

3 IpF; same data, coll. 41, 1 IpF; same data except 1.5 km S of Tena and 27 May, coll. 44, 1F, 1

pM, | IpF. Pastaza: Puyo, 39 km N of, 6 May 1977, Y. Huang, coll. 9, 1Mgen, 1 pM, 1 lpF; same

data except 4 km NE of Puyo and 7 May, coll. 10, 2 pF; same data except 5.3 km NE of Puyo,

106 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

coll. 14, 1Mgen, 1 pM, 1 lpF, 1 1; same data except 16 km NE of Puyo, coll. 15, 1 IpF; same data

except 1.5 km S of Puyo and 19 May, coll. 32, 1 pF, 1L; same data, coll. 34, 1Mgen, 2 pM, 1 pF;

same data except 23 km SE of Puyo, coll. 35, 1Mgen, 3 pM.

PERU (35 specimens: 10M, 23F, 2L). Huanuco: Cochicote, 12, 13 Oct 1965, J. Hitchcock, 14

F. Tingo Maria, Apr 1946, A. Valde Rerma [?], 1M, 2F; same locality, E. Viale, 2L; same locality,

E. Hambleton, 6M, 7F. Province not specified: Rio Perene, 1939, Kuczunski, Godard, 3M.

VENEZUELA (177 specimens: 19M, 70Mgen, 60F, 16p, 12L&l; 16 ind rear: 3 1, 10p, 3inc).

Aragua: Cagua, VZ 99. Maracay, 3 Sep 1926, M. Nunez Tovar, 28F; same locality and collector,

11 Sep 1926, 1F; same locality and collector, 9 Oct 1926, 2F; same locality, 9 May 1927, 1Mgen

(H36a), 1 lp [JH]; same locality, 11 May 1927, 1F (H38a); same locality, 27 May 1927, 1Mgen

(40a) [JH] ; same locality, 10 Jun 1927, 2Mgen (63btc), 2 lp [JH] ; same locality, 20 Aug 1928, R.

Hill, 1Mgen [JH]; same locality and collector, Aug 1928, 1Mgen (20), 1Mgen (23) [JH] ; same lo-

cality and collector, 25 Sep 1928, 3Mgen (38) [JH] ; same locality and collector, 1928, 13Mgen

[JH] ; same locality, 2 May 1929, 3Mgen [JH] ; same locality, Jun 1929, 1Mgen (57) [JH] ; same lo-

cality, 22 Jul 1929, 2Mgen (49) [JH] ; same locality, 1929, [R. Hill] , 22Mgen [JH] ; same locality,

20 Aug 1942, VZK 50, 1F; same locality, VZ 130, 131. Turmero, VZ 45, 336. Carabobo: Puerto

Cabello, 26 Dec 1938, 1L; same locality, 28 Feb 1941, 1M; same locality, 2 Aug 1941, 1M, 1F;

same locality, 1Mgen [JH]. Cojedes: San Carlos, 8-15 Jun 1941, 2M, SF; same locality, 16-22 Jun

1941, 2M, 3F; same locality, VZ 115. Tinaquillo, VZ 119. Distrito Federal: Caracas, 2Mgen [JH].

Monagas: Caripito, 11 Jun 1935, 1M. Portuguesa: Acarigua, 13 Jan 1941, 1Mgen (23) [JH] ; same

data, VZK 49, 1M, 1F. Trujillo: Pampanito, 1-7 Aug 1938, 2M, 7F. State not specified: Trinidad,

1Mgen (130-8), 1Mgen (49-11) [JH]. Locality not specified, 15 Jul 1929, [R. Hill] , 2Mgen (40)

[JH] ; 1929, SMgen (13) [JH] ; 1949, 1Mgen, 2L; no data, 3M, 2F.

9. Anopheles (Nys.) trinkae Faran

Figs. 3, 4, 7, 23, 24

1979. Anopheles (Nyssorhynchus) trinkae Faran 1979:26-39. TYPE: Holotype male (MEP Acc.

638, coll. 29-9) with associated genitalia and larval and pupal exuviae on slides, 1.5 km S

of Puyo (Pastaza), Ecuador, large temporary ground pool, 15 May 1977, Yiau-Min Huang

[USNM, 76123].

Anopheles (Nyssorhynchus) nuneztovari in part of Simmons and Aitken (1942:39, 46, 53, 95);

Galvao (1943:146); Galvis (1943:88-89); Gabaldon and Cova Garcia (1952:193); Lane (1949:

403; 1953:268-269); Stone, Knight and Starcke (1959:33); Vargas (1959:376, 382); Forattini

(1962:396-400); Morales-Ayala (1971:139); Kitzmiller, Kreutzer and Tallaferro (1973:435-455,

?in part); Kreutzer, Kitzmiller and Rabbani (1975:363-364, ?in part); Knight and Stone (1977:

63).

poe nuneztovari in part of Russell, Rozeboom and Stone (1943:37, 47); Foote and Cook

(1959:24, 26, 29, 32); Stojanovich, Gorham and Scott (1966b:22, 30, 43); Gorham, Stojano-

vich and Scott (1967:15, 47, 58; 1973:111, 138, 147-148); Elliott (1968:248-252; 1972:756-

763, ?in part).

Anopheles (Nyssorhynchus) goeldii (?) of Cerqueira (1943:19).

FEMALE (fig. 7). Wing: 3.4 mm. Proboscis: 2.2 mm. Palpus: 2.1mm. Forefe-

mur: 1.5mm. Abdomen: about 3.0 mm. Head: Integument reddish brown to dark

brown. Proboscis about 1.4 length of forefemur. Vertex with long, white, spatulate

or setiform scales becoming shorter, darker and more cuneate caudolaterally on occi-

put; occipital scales golden; postoccipital scales reddish brown to brownish black.

Apex of palpal segment 2 with a small white band; apex of segment 3 with a large

white band; segment 2 with a few, erect, cream to white scales; segment 3 with a

cream to white, dorsal or dorsolateral stripe; segment 4 with a moderately large, dark,

basal band and a smaller, dark, apical band, usually with a single row of dark scales

on ventral surface, not always visible; segment 5 white, occasionally with a few, dark,

basal scales extending from apex of 4. Antenna: Flagellar segment 1 with numerous,

long, oblanceolate scales on dorsomesal and mesal surfaces in apical 0.5; with shorter

scales on dorsolateral surface and small obovate scales at base. Flagellar segments 2-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 107

13 each with a basal whorl of about 8,9 long, silver setae. Thorax: Integument of

scutum reddish tan to dark brown, and extensively pruinose. Anterior promontory

area with setiform scales extending a short distance dorsad onto acrostichal line.

Prescutellar space moderately large, horseshoe shaped. Pleuron light brown to brown.

Sp with 7 (3-9) light setae. Upper stp with 4 (2-4) long, dark setae in horizontal arc,

and about 6-15 cream, obovate scales. Lower stp with 1-3 long setae, and a diagonal

patch of 5-11 cream, obovate scales. Upper mep with 7-9 long, cream setae, and oc-

casionally one or 2 cream, obovate scales. Legs: Dark scales reddish brown. Light

scales on coxae white. Midfemur with both anteroapical spot and knee spot distinct.

Foretarsal segments 2 and 3 with a white band in apical 0.3 1-0.46 and 0.6-0.9 respec-

tively, usually longer than in nuneztovari; segment 4 golden to white in apical O03

(0.1-0.5); segment 5 cream to white in about apical 0.5. Dorsal surface of midtibia

and midtarsal segments 1,2 and usually 3 with a cream stripe. Midtarsal segments 1-3

with a small, cream, apical band, band largest and most distinct on 2; segment 4 all

dark; segment 5 cream in apical 0.3-0.7. Hindtarsal segment 2 with a dark band in

basal 0.3-0.4 (0.30-0.43); segment 5 light cream to white in about apical 0.5. Wing:

Light wing spots white or very light cream. Dark wing spots not usually as extensive

as in nuneztovari. Vein C humeral light spot about 1.3-2.5 (1.3-4.0) of basal dark

spot, usually 2.0 or less of basal dark spot; basal dark spot usually not reaching hu-

meral crossvein; subbasal dark spot equal to or longer than subbasal light spot, often

2.0 of subbasal light spot; subcostal light spot 0.25-0.43 (0.25-0.65) of subcostal

dark spot; preapical light spot about 0.31-0.46 of preapical dark spot; apical dark

spot moderate and distinct. R presectoral dark spot moderately large; sectoral dark

spot relatively small, rarely absent. R. preapical dark spot about 0.5 of vein. Ra,s

subcostal and apical dark spots moderately small. M sectoral dark spot variable,

small to large, lighter than dark spots on C, often with interspersed light scales; pre-

apical dark spot not extending onto M,,,. Cu sectoral dark spot moderately large

and distinct. Cu, subcostal spot usually as large as or larger than preapical dark spot.

Vein A subcostal dark spot moderately short to moderately long. Apical light fringe

spot moderately small and divided by small dark spots between R, and R3, and R3

and Ry,<; additional light fringe spots as those of nuneztovari. Abdomen: As that of

nuneztovari, except caudolateral and sternomedian scale tufts on segments II-VI

small.

MALE. Wing: 3.2 mm. Proboscis: 2.5 mm. Forefemur: 1.45 mm. Abdomen:

2.45 mm. Essentially as in female except for sexual characters. Head: Palpal seg-

ments 2 and 3 with a conspicuous, cream, dorsal stripe; apex of segment 2 and base

of segment 3 with a small white band; apex of segment 3 with a moderately broad,

white band; segment 4 with a basal and an apical brown band, occasionally with

white scales interspersed in small basal band, and usually with a few, white, subapical

scales; segment 4 with mediolateral surface predominantly light cream, ventral sur-

face usually with inconspicuous dark scales. Antenna: About 0.70-0.75 length of

proboscis. Flagellar segment 1 with numerous, long, white, oblanceolate scales on

dorsomesal margin. Legs: Forefemur about 0.6 length of proboscis. Basal plantar

surface of foretarsal segment 5 with about 8 moderately long to long, spiniform se-

tae; longest setae about 0.3 length of segment. Claw on foreleg moderately large,

weakly curved, acuminate; submedian tooth 0.25-0.33 length of claw, moderately

thin and apically recurved; basal tooth moderately long, heavy and decurved.

MALE GENITALIA (fig. 23). Segment VIII, Segment IX, Sidepiece, Clasper and

Dorsal Claspette essentially the same as those of nuneztovari. Ventral Claspette:

Very similar to that of nuneztovari except for the following. Moderately long, 0.40-

i 108 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

0.50 length of sidepiece; apex moderately broad, width at apex 0.43-0.50 (0.38-0.54)

length of claspette. Preapical plate moderately small, semicircular to oval, weakly to

moderately sclerotized. Phallosome: Aedeagus about 0.5 length of sidepiece and

1.00-1.20 (1.00-1.33) length of ventral claspette; apex rounded, about as wide as or

slightly wider than long; leaflets not visible on any specimen examined.

PUPA (fig. 23). Abdomen: about 2.5 mm. Trumpet: 0.5 mm. Paddle: 0.75 X

0.55 mm. Only exuviae examined. Cephalothorax: Wing cases with moderately pig-

mented longitudinal stripes. Hair 1-C 2-4 branched; 2,3-C 2,3 branched. Hairs 4,5-C

2-5 forked; 4C slightly shorter than 5-C. Hair 6-C 2,3 forked (1-3), usually as long

as or slightly longer than short branch(es) of 7-C; 7-C 2-4 branched, with one branch

about 1.5 length of other branch(es). Trumpet: Pinna moderately pigmented; long,

about 3.6-4.0 (3.5-4.1) length of meatus; in lateral aspect, not appearing to taper

toward apex. Meatal cleft basally rounded. Metanotum: Hair 10-C single or double,

subequal to 11-C. Hair 11-C 2,3 branched. Hair 12-C 1-3 branched (1-4), about 1.5-

2.0 length of 10-C. Abdomen: Hair 2-I 4-6 forked (2-6), moderately long to long,

branches arising at least 0.20 distad of base. Hair 3-I single, about 0.5-0.9 length of

2-I. Hair 4-I 46 branched (3-6), moderately long. Hair 5-I 1-3 branched. Hair 6-1

single, long, about 2.0 length of 7-I. Hair 7-I 4-7 branched (3-7). Hair 9-I single,

about 0.7 length of 6-I. Hair 0-II 1-3 branched (1-4), small; O-III-VII moderately de-

veloped; O-III 3-6 branched (2-6); sum of branches of 0-II and O-III 47 (3-9); O-IV 4,

5 branched (2-5); 0-V 2-5 branched; 0-VI 2,3 branched (2-4); O-VII 1-3 branched.

Hair 1-II,III strongly developed; 1-II 6-10 branched (4-13); 1-III 3-9 branched (1-10);

1-IV-VII single, long, 1.75-2.00 length of segment. Hair 3-IV dendritic, 3-6 branched

(1-6), moderately developed; 3-V 3-5 forked (1-5). Hair 5-III 5-7 branched (5-9),

strongly developed; 5-IV 2-5 branched (1-5), moderately long, median branch often

longer than lateral; 5-V-VII single (1-3 branched), long, subequal to length of seg-

ment. Hair 6-II single or double, long, about 2.0 length of 7-II. Hair 7-II 3-7

branched (3-8); 7-IIL,IV 3-5 branched, short; 7-V 2-4 forked, longer than 7-III,IV; 7-

VI,VII single or double, long. Hair 8-III,[V 2-6 branched (2-7); 8-V,VI 2-4 branched

(1-4); 8-VII 2-4 branched, moderately short, slightly longer than 8-III-VI. Hair 9-I]

small, unpigmented, thinner and more pointed than 9-III; 9-III small, less than or oc-

casionally equal to 2.0 length of 9-II; 9-IV heavy, 1.6-3.6 length of 9-III, strongly

pigmented; 9-V moderately thin, acuminate, long, 2.5-4.5 (2.5-4.9) length of 9-IV;

9-VI equal to or slightly longer (1.0-1.3) than 9-V; 9-VII 1.05-1.25 (1.05-1.30)

length of 9-VI; 9-VIII usually straighter than 9-V-VII, subequal to 9-VII; 9-V-VIII

about 0.5 length of segment. Hair 10-III 3,4 forked (2-6), long. Hair 4-VIII 3-5

forked (2-5), about 0.65-0.70 length of 9-VIII. Terminal Segments: Male genital

lobe large, heavy, with a very prominent mammilliform protuberance. Paddle:

Large, obovate, emarginate at insertion of hair 1-P. External buttress 0.6 (0.53-0.68)

length of paddle, serrated apically. External margin distad of buttress with fine,

short, filamentous spicules extending around apex and becoming shorter and fewer

along caudal 0.3-0.5 of inner margin. Hair 1-P moderately long; 2-P subequal to 1-P.

LARVA (fig. 24). Head: 0.6 mm. Antenna: 0.25 mm. Anal Saddle: 0.3 mm.

Head: Heavily pigmented, dull reddish brown. Median tooth of mental plate moder-

ately broad, less than combined width of 2 adjacent teeth from one side, tapering to

blunt point. Inner and outer clypeal hairs (2,3-C) single, and simple or with very

small barbs; hairs 2-C widely spaced, clypeal index about 1.25; 3-C 0.5-0.8 length of

2-C. Hair 4C single or 2 forked, long, 0.7-1.0 length of 3-C and usually extending to

beyond base of 2-C. Hair 8-C 2 branched (2,3) near base, occasionally forked, length

about 2.0 distance separating insertion of 2-C and 3-C. Hair 9-C weakly dendritic, 4

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 109

6 branched (4-7), equal to or slightly longer than 8-C. Hair 12-C 2-5 branched, mod-

erately long. Hair 15-C 2-4 forked. Collar wide dorsomedially, heavily pigmented.

Antenna: Pigmented as remainder of head. Mesal margin with stout, moderately

long spicules; ventral surface with fewer and shorter spicules. Hair 1-A 4-6 branched,

short, inserted in basal 0.25 of antenna. Thorax: Darkly pigmented, reddish brown.

Submedian prothoracic group (1-3-P) with or without hairs 1,2-P sharing a common

tubercle; 3-P occasionally arising from the same tubercle as 2-P; palmate hair 1-P with

11-15 (10-18) moderately narrow to narrow, pointed, pigmented, lanceolate leaflets;

2-P 15-19 branched (12-20), 3.0-4.0 length of 1-P. Hair 14-P 7-10 branched (6-11)

from a moderately short shaft. Mesothoracic hair 1-M 28-37 branched (26-37). Met-

athoracic hair 2-T single, moderately long but not reaching posterior margin of thor-

ax. Palmate hair 3-T with 9-13 (9-17) semitransparent, moderately narrow, pointed,

lanceolate leaflets. Pleural group spines as those of nuneztovari. Abdomen: Hair 0-

II 1-3 branched, very short, about 0.5 or less length of leaflets of 1-II, inconspicuous;

O-IJI-VII 1-3 branched (1-4), small to moderately small. Palmate hair 1-I moderately

small, with 12-16 (10-17) narrow, semitransparent, spreading, lanceolate leaflets; 1-

II-VII with strongly pigmented, moderately narrow, pointed, lanceolate leaflets; 1-II

23-27 branched (19-30); 1-III 24-32 branched; 1-IV 22-29 branched; 1-V 23-30

branched (22-30); 1-VI 21-27 branched (20-30); 1-VII 20-26 branched; 1-II, III, VU

slightly shorter than 1-IV-VI. Hair 2-I 2 branched (1-3), very small; 2-II 6,7 branched

(4-9), moderately large; 2-III 3-5 branched (2-6), large; 2-IV single or rarely double,

about 1.5 length of leaflets of 1-I[V; 2-V single, very long. Hair 3-I single (1,2

branched), moderately long. Hair 5-I 3,4 branched (3-5), small; 5-II 6-11 branched

(5-11), small. Hair 9-I 5,6 branched (4-6). Hair 13-I,I1,III 4-6 (3-7), 7-9 (7-10) and

6-8 (6-9) branched respectively, small; 13-IV 3,4 branched, moderately large, 1.5-2.0

length of leaflets of 1-IV; 13-V 3-5 branched (2-6), very large; 13-VI 8-10 branched

(6-10), moderately small. Spiracular Lobe: Pecten with about 15-17 teeth; median

teeth subequal, occasionally mixed short to moderate, with 2-4 long interspersed;

serrations on teeth moderately long. Hair 8-S 3-4 branched (3-5), moderately long,

subequal to 9-S; 9-S 3-5 branched. Lateral arm of spiracular apparatus short. Anal

Segment: Apically with short thin spicules. Saddle reddish brown. Hair 1-X longer

than saddle; inserted on saddle near ventral margin of saddle, or rarely on ventral

margin at base of indentation. Anal gills longer than saddle.

DISCUSSION. An. trinkae can be distinguished from the other species in the Os-

waldoi Subgroup in the female (except occasionally nuneztovari and rangeli) by the

combination of (1) palpal segment 4 usually with a single ventral row of dark scales,

(2) prescutellar space moderately large and horseshoe shaped, (3) foretarsal segment

2 with a large white band in apical 0.31-0.46 and segment 3 with a very large white

band in apical 0.6-0.9, (4) foretarsal segment 4 golden to white in apical 0.3 (0.1-0.5),

(5) hindtarsal segment 2 with a dark band in basal 0.3-0.4 (0.30-0.43), (6) light wing

spots white or very light cream, (7) humeral light spot of vein C 1.3-2.5 (1.3-4.0)

length of basal dark spot and (8) subcostal light spot of vein C 0.25-0.43 (0.25-0.65)

length of subcostal dark spot; in the male genitalia by the same characters as in nu-

neztovari except that (1) ventral claspette long, 0.40-0.50 length of sidepiece and

length of aedeagus 1.00-1.20 (1.00-1.33) length of claspette, (2) width of apex of

ventral claspette 0.43-0.50 (0.38-0.54) length of claspette and (3) apex of aedeagus

rounded, about as wide as or slightly wider than long and without apparent leaflets;

in the pupa by the combination of (1) hair 7-C with one branch about 1.5 length of

shorter branch(es), (2) pinna long, about 3.6-4.0 (3.5-4.1) length of meatus, not ap-

pearing to taper toward apex in lateral aspect, (3) hair 11-C 2,3 branched, subequal

ee 8, Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

to 10-C, and hair 12-C 1-3 branched (1-4), about 1.5-2.0 length of 10-C, (4) hair 2-I

4-6 forked (2-6), moderately long to long, and 3-I about 0.5-0.9 length of 2-I, (5)

hair O-II 1-3 branched (1-4), small, (6) sum of branches of hairs O-II and O-III usually

less than 8 (3-9) and (7) hair 9-III less than or equal to 2.0 length of 9-II, 9-V 2.5-4.5

(2.5-4.9) length of 9-IV, 9-VITI 1.05-1.25 (1.05-1.30) length of 9-VI and 0.5 or more

length of segment; and in the larva by the combination of (1) hairs 2,3-C single, and

simple or with very small barbs, hairs 2-C widely spaced, clypeal index about 1.25,

and 3-C 0.5-0.8 length of 2-C, (2) hair +C single or 2 forked, long, 0.7-1.0 length of

3-C, usually extending to beyond base of 2-C, (3) hairs 1,2-P sharing or not sharing a

common tubercle, 3-P occasionally arising from same tubercle as 2-P, palmate hair 1-

P with 11-15 (10-18) moderately narrow to narrow, pointed, lanceolate leaflets, (4)

hair O-II 1-3 branched, very short, about 0.5 or less length of leaflets of 1-II, incon-

spicuous, and Q0-III-VII 1-4 branched, small to moderately small, (5) hair 13-IV mod-

erately large, 1.5-2.0 length of leaflets of 1-I[V and (6) hair 1-X inserted on saddle

near ventral margin, or rarely on ventral margin at base of indentation.

Superficially, the larva and male genitalia of trinkae appear similar to those of nu-

neztovari, and the pupa is much like that of rangeli; however, these stages can readily

be distinguished by the characters above. Unfortunately, the key features of the

adult female can easily be confused with those of either nuneztovari or rangeli, and

occasionally (when the humeral light spot of the costal vein is greater than 2.0 the

length of the basal dark spot) the adults can be mistaken for those of strodei. To al-

leviate this problem, trinkae keys out twice in the key to adult females. Because the

adult female is the most unreliable stage for identification, careful attention should

be given to the male genitalia, larva and pupa for positive species determination.

In the adult female, the wings of specimens from Ecuador differ from those of spe-

cimens from Colombia by possession of a proportionally smaller basal dark spot and

a larger subcostal light spot on the costal vein. The immatures from Ecuador have 1-

5 fewer branches on several of the abdominal hairs than those examined from Colom-

bia, and palmate hair 1-P of the larva from Ecuador has more numerous (14-18) and

narrower branches.

Since rangeli and trinkae occur sympatrically (the immatures have been collected

from the same aquatic habitats), it is reasonable to assume that they are genetically

isolated. With respect to nuneztovari with which trinkae shows the greater affinity,

it is unknown if trinkae and nuneztovari are sympatric over any part of their ranges.

Therefore, it is impossible to state with absolute certainty that these latter 2 species

have evolved sufficiently to maintain separate gene pools if they were to associate in

nature. However, after having examined numerous individual rearings and one large

progeny rearing of trinkae from several different localities, I feel confident that

trinkae represents a species distinct from both rangeli and nuneztovari.

BIONOMICS. The data on the bionomics of trinkae were extracted from the col-

lection records compiled by Yiau-Min Huang while she was in Ecuador. Additional

information came from field notes on one collection from Colombia (COB 42).

The immatures of trinkae have been collected in temporary and permanent, small

to large ground pools (43%), wheel tracks (19%), ponds and lakes (19%), streamside

pools (14%), and ditches (5%). All collections were from either clear or slightly col-

ored fresh water exposed to the sun. Several collections came from the sides of roads

and one was from a sugar cane plantation. Many of the ground pools were in grassy

areas so that emergent and submerged vegetation was abundant. Green algae fre-

quently occurred in the aquatic habitats. Most of the collections were made near hu-

man habitations, usually within 1 km of the nearest house. The collections came

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 111

from localities ranging in elevation from 82 to 950 m above sea level, and all were

from the eastern slope of the Andes. Huang characterized the general habitat in Ec-

uador as “‘scrub with scattered trees.”’ Hueck and Seibert (1972) recognize these re-

gions of Colombia and Ecuador where trinkae occurs as the “Andes septentrionales y

Cordilleras,”’ the “‘bosques tropicales y subtropicales, deciduos y mesofiticos de Co-

lombia y Venezuela” and the “‘Jaderas orientales de los Andes medios.” The imma-

tures of trinkae occurred with 2 other species in the Albimanus Section, rangeli and

rarely triannulatus. Other species that have been found associated with trinkae are

Culex (Mel.) bastagarius, Cx. (Mel.) chrysonotum and Cx. (Mel.) pilosus.

Because of the uncertainty of the geographical range of trinkae and its morpholo-

gical similarities with rangeli and nuneztovari, there is some confusion as to which of

these 3 species is referred to in much of the literature on bionomics and distribution.

For example, Elliott (1968, 1972) studied the adult behavior of ‘“nuneztovari’’ in 5

localities in Colombia and contrasted this with the behavior of nuneztovari in Brazil.

There is a possibility, although an unlikely one, that at some of the localities Elliott

worked not on the behavior of nuneztovari but on the behavior of trinkae. Never-

theless, without voucher specimens it is impossible to be certain. Similarly, the dis-

tribution records for nuneztovari and rangeli from Ecuador and parts of Colombia

may really refer to trinkae. For these reasons many of the references listed in the

synonymy of trinkae should be considered tentative.

MEDICAL IMPORTANCE. Nothing is known about the vector potential of

trinkae. As with the literature on bionomics and distribution, many of the referen-

ces to either rangeli or nuneztovari concerning transmission of malaria could refer to

trinkae. Huang (personal communication) states that in the Ecuadorian provinces of

Pastaza and Napo, rangeli was thought to be the vector of malaria. Likewise, Forat-

tini (1962:393) reports that rangeli has been suspected as a carrier of malaria in Ec-

uador; but in no other country has rangeli ever been thought to be important as a

vector. Since trinkae is often easily confused with rangeli in the adult female, it is

possible that trinkae is transmitting malaria in Ecuador.

DISTRIBUTION (fig. 3). An. trinkae is known to occur along the eastern slope of

the Andes from Villavicencio, Colombia south to Puyo, Ecuador. It is not known

how much farther north or south the range of this species extends. Cerqueira (1943:

19) reports goeldii from several localities in Bolivia based on examination of adult fe-

males and 2 male genitalia. I do not know if this represents nuneztovari or trinkae.

The following corrections in the ““Material Examined” section of the original de-

scription of trinkae (Faran 1979:33-34) should be noted: (1) collections CV 737,

738 and 739 are not progeny rearings, (2) on page 34, line 4, ‘‘Acarias, 3 Jun 1949”

should be changed to “Acacias, 3 Jun 1947” and listed under the department of Me-

ta in Colombia, (3) Ecuador coll. 26 (page 34, line 5-6) is not trinkae, but rangeli and

(4) Ecuador colls. 32 and 33 (page 34, line 19-21) were made 1.5 km S of Puyo, not

4.3 km E Puyo as stated.

Material Examined: 385 specimens: 73 males, 18 male genitalia, 60 females, 127

pupae, 107 larvae; 127 individual rearings: 56 larval, 48 pupal, 23 incomplete; 1 pro-

geny rearing.

COLOMBIA (102 specimens: 12M, 6Mgen, 17F, 24p, 43L&1; 24 ind rear: 3 1, 21 inc; 1 progeny

rearing). Meta: Acacias, 3 Jun 1947, [CV] 408.1, 1 lp [JH]. Villavicencio, 26 May 1947, L.

Rozeboom, CV-P 28 (progeny rearing), 12M, 6Mgen, 15F, 8 lp, 12L; same locality, 14 Feb 1948,

[CV] 537.1, 1 lp [JH] ; same locality, CV 737, 3 lp, 4L; same locality, CV 738, 4 lp, 3L; same lo-

cality, CV 739, 5 lp, 4L; same locality, COB 42.

ECUADOR (283 specimens: 61M, 12Mgen, 43F, 103P&p, 64L&1; 103 ind rear: 53 1, 48p, 2inc).

Napo: Tena, 11.7 km SW of, 25 May 1977, Y. Huang, coll. 40, 4 pM, 1 pF, 2 lpM, 2 IpF; same

Peak 510: Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

data except 13 km SW of Tena, coll. 43, 1 lpM. Pastaza: Puyo, 5 May 1977, Y. Huang, coll. 4, 1

Mgen, 2 pM, 2 pF, 1L; same data, coll. 5, 2L; same data, coll. 6, 1 lpM; same data except 20 km N

of Puyo and 6 May, coll. 8, 1Mgen, 1 pM, 1 pF, 1 lpM, 1 IpF; same data except 39 km N of Puyo,

coll. 9, 2 pM, 1 pF, 1 lpF, 1 IP; same data except 4 km NE of Puyo and 7 May, coll. 11, 1 pF, 1

IpM, 1 IpF, 1 IP, 2L; same data except 16 km NE of Puyo, coll. 15, 1Mgen, 1 IpM, 1 lpF; same data,

coll. 16, 1Mgen, 1 pF, 2 IpM, 1 IpF; same data except at Puyo and 8 May, coll. 17, 1Mgen, 5 pM, 3

pF, 3 lpF; same data, coll. 18, 1Mgen, 3 pM, 1 pF, 1 IpF; same data except 31 km W of Puyo and 9

May, coll. 20, 3L; same data except 12 km W of Puyo, coll. 23, 1Mgen, 1M; same data except 1.5

km S of Puyo and 15 May, coll. 29, 1Mgen, 2 pM, 1 pF, 4 1pM, 5 lpF; same data, coll. 30, 1Mgen,

4 pM, 3 pF, 6 lpM, 2 IpF, 1F, 1 1; same data except 4.3 km E of Puyo and 17 May, coll. 31, 1Mgen,

2 pM, 2 lpM, 4 lpF; same data except 1.5 km S of Puyo and 19 May, coll. 32, 1Mgen, 5 pM, 7 IpM,

3 lpF; same data, coll. 33, 1Mgen, 2 pM, 1 pF, 1F.

10. Anopheles (Nys.) nuneztovari Gabaldon

Figs. 3, 4, 7, 25, 26

1940. Anopheles (Nyssorhynchus) nuneztovari Gabaldon 1940:5-6. TYPE: Holotype male, San

Carlos (Cojedes), Venezuela [MDM].

1941. Anopheles (Nyssorhynchus) goeldii Rozeboom and Gabaldon 1941:89-91. TYPE: Holo-

type male with associated genitalia slide, labelled ““N. gorgasi no. 6,” Boa Vista, Rio Tapa-

jos (Para), Brazil, C. H. T. Townsend [USNM].

1945. Anopheles (Nyssorhynchus) dunhami Causey 1945:231-234. TYPE: Holotype male with

associated genitalia slide, Tefe (Amazonas), Brazil, captured on animal bait [|USNM,

58036].

Anopheles (Nyssorhynchus) nuneztovari of Gabaldon and Aguilera (1940:68, 69); Gabaldon, Cova

Garcia and Lopez (1940:9); Rozeboom and Gabaldon (1941 :91-98); Komp (1942:5, 26, 34);

Simmons and Aitken (1942:39, 46, 53, 95); Galvao (1943:146); Floch and Abonnenc (1946b:

1, 3-5; 1947:6; 1951:57-58); Gabaldon and Cova Garcia (1946b:98-99; 1952:178, 193, in part);

Deane, Causey and Deane (1948:894-897); Senevet (1948a:278; 1948c; 434-438); Vargas (1948:

158; 1959:376, 382, in part); vander Kuyp (1949a:67, 68); Lane (1949:403; 1953:268-269, in

part); Rey and Renjifo (1950:534-538); Bejarano (1957:326, 329, 330, 331); Giglioli (1959:

280); Stone, Knight and Starcke (1959:33, in part); Cerqueira (1961:125-126); Fauran (1961:

11); Forattini (1962:396-400, in part); Hamon, Mouchet et al. (1970:32-33); Belkin, Schick and

Heinemann (1971:6); Morales-Ayala (1971:139); Gabaldon (1972:634-639); Kitzmiller, Kreut-

zer and Tallaferro (1973:435-455); Kreutzer, Kitzmiller and Rabbani (1975:363-364); Cova

Garcia and Sutil O. (1976:27; 1977:23, 48, 64, 84); Cova Garcia, Pulido F. and Amarista M.

(1978: 157); Kitzmiller (1979: 26-31); Faran (1979: 26, 27, 30, 31, 32, 33).

Anopheles nuneztovari of Russell, Rozeboom and Stone (1943:37, 47); Floch and Abonnenc

(1946a:2); Foote and Cook (1959:24, 26, 29, 32, in part); Gabaldon and Guerrero (1959:433-

434); Cova Garcia (1961:37-69, 89-90, 114, 127-128, 152); Vincke and Pant (1962:2-18, 28, 31,

35-42); Stojanovich, Gorham and Scott (1966b:22, 30, 43, in part); Gorham, Stojanovich and

Scott (1967:15, 47, 58; 1973:111, 138, 147-148, in part); Elliott (1968:248-252 ; 1972:756-

763, ?in part); Cova Garcia and Sutil O. (1975a:24; 1975b:213); Perez de Valderrama and Scorza

(1976: 212-220); Scorza, Tallaferro and Rubiano (1976:129-130); Panday (1977: 728-737;

1979b:57); Panday and Panday-Verheuvel (1979:36).

Anopheles nunes-tovari (!) of Mattos and Xavier (1965:273).

Anopheles (Nyssorhynchus) dunhami of Deane, Causey and Deane (1948: eae Senevet (1948a:

278); Lane (1953:268); Elliott (1972:758, 762).

Anopheles (Nyssorhynchus) goeldii of Gabaldon and Aguilera (1940:72); Gabaldon, Aguilera and

Arevalo (1941:60); Galvao and Damasceno (1942:122-123); Simmons and Aitken (1942:39, 46,

53, 61, 94-95); Cerqueira (1943:19, ?); Galvao (1943:149, 151); Causey, Deane and Deane

(1944:2, 5, 6; 1946:27); Floch and Abonnenc (1944b:2); Lane (1944:263; 1949:403); Deane,

L. M., Causey and Deane (1946:6, 7, 9, 10, 12); Deane, M. P., Causey and Deane (1946:42, 45);

Senevet (1948a:278); Levi-Castillo (1949:11, 15, 28, 67).

Anopheles goeldii of Russell, Rozeboom and Stone (1943:37, 42, 46).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 113

Anopheles (Nyssorhynchus) tarsimaculatus in part of Shannon (1933:124-132).

Nyssorhynchus tarsimaculatus in part of Townsend (1933b:7-12).

Nyssorhynchus albimanus in part (?) of Townsend (1934:493-494).

Anopheles (Nyssorhynchus) gorgasi in part of Townsend (1933a:101-102).

Anopheles (Nyssorhynchus) sp, goeldii series of Bruce, Knigin, Yolles and Graham (1943:442).

Cellia albimana in part (?) of Neiva (1909:69-77).

FEMALE (fig. 7). Wing: 3.2 mm. Proboscis: 2.1mm. Palpus: about 2.1 mm.

Forefemur: 1.5mm. Abdomen: about 2.9mm. Moderate size. Integument dark

brown to very dark brown, with considerable contrast between light and dark re-

gions. Head: Proboscis about 1.4 length of forefemur. Palpal segments 1 and 2 and

base of 3 with predominantly dark, outstanding scales; segments 2 and 3 with asmall,

white, apical band and a small to moderate, white, apical band respectively; segments

2 and 3 with or without a dorsal or lateral speckling of light scales; segment 4 with

base and usually apex dark scaled, usually ventral surface with row of dark scales;

segment 5 usually completely white, occasionally with a few, dark, basal scales. An-

tenna: Flagellar segment 1 with numerous, white, oblanceolate setae distally on dor-

somesal and mesal surfaces; basally with shorter obovate scales. Flagellar segments 2-

13 each with a basal whorl of about 8-10 tan to silver setae. Thorax: Anterior pro-

montory area with long, white, setiform scales usually not extending far dorsad onto

acrostichal line. Prescutellar space subtriangular or horseshoe shaped. Sp with 4-8

moderately long, light setae and a few obovate scales. Upper stp with 3-5 long, dark

setae above a patch of light scales in horizontal arc. Lower stp with 1,2 long setae

and a diagonal patch of light scales. Upper mep with 7 (5-9) long, cream to brown

setae; scales absent. Legs: Light scales on coxae white to pale cream. Midfemur

with both anteroapical spot and knee spot distinct. Foretarsal segments 2 and 3

cream to white in apical 0.2-0.4 (0.15-0.44) and 0.55 (0.40-0.75) respectively; seg-

ment 4 with a small, cream, apical band; segment 5 golden to cream in about apical

0.5. Midtarsal segments 1 and 2 with a small, golden, apical band; segments 3 and 4

all dark, occasionally with a few, lighter, apical scales; segment 5 with apical 0.25-

0.50 lighter than base. Hindtarsal segment 2 brown to dark brown in basal 0.25-0.32

(0.20-0.32); segment 5 dark in about basal 0.5, apical 0.5 cream to white. Wing:

Light wing spots usually cream, not white, at least on anterior veins. Large dark wing

spots distinct. Vein C humeral light spot about 0.7-1.3 (0.7-1.7) of basal dark spot;

basal dark spot often extending to humeral crossvein; subbasal dark spot usually long-

er than subbasal light spot; sectoral dark spot occasionally absent; subcostal light spot

0.20-0.55 of subcostal dark spot, usually less than 0.5; preapical light spot about

0.36 (0.2-0.5) of preapical dark spot; apical dark spot moderately long and distinct.

R presectoral dark spot moderately large, usually much larger than sectoral dark spot.

R, preapical dark spot large, about 0.3-0.5 of R,. R4,;subcostal and apical dark spots

moderate, very dark and conspicuous. M sectoral dark spot variable, moderately long

to long, 0.25-0.70 of M; preapical dark spot not extending onto M,,,. Dark spots of

M,.. and M3,4 very dark and conspicuous. Cu, subcostal, preapical and apical, and

Cu, preapical dark spots moderately long and distinct. Vein A subcostal dark spot

moderately long, about 0.2 (0.15-0.30) of A. Apical light fringe spot moderately

small (to large in Venezuelan specimens) and divided or undivided by a few dark

scales between R, and R3, and R; and Ry,;; additional, and in most cases conspicu-

ous, small, light fringe spots at apices of R4.5, My.2, Cu,, Cu, and A; a moderately

conspicuous to inconspicuous, light fringe spot between base of wing and A. Abdo-

men: Sternite I with a few, moderately long to long setae, longer setae sternomedial-

ly and shorter setae laterally; with occasionally a few, scattered, inconspicuous, light,

-- 114 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

lanceolate to obovate scales, usually only visible on slide preparations. Caudolateral

and sternomedian dark scale tufts on segments II-VII with scales conspicuous, large

and broad.

MALE. Wing: 3.4 mm. Proboscis: 2.75 mm. Forefemur: 1.6 mm. Abdomen:

about 3.2 mm. Essentially as in female except for sexual characters. Head: Palpal

segments 2 and 3 with a stripe of light scales along dorsal surface; segment 4 with a

basal and an apical, white band, and a subbasal and a subapical, smaller, dark band;

mediolateral surface of segment 4 largely speckled with cream scales with a few inter-

mingled dark scales; ventral surface of 4 inconspicuously dark scaled. Antenna:

About 0.75 length of proboscis. Flagellar segment 1 with conspicuous, long, white,

lanceolate, dorsomesal scales. Legs: Forefemur about 0.6 length of proboscis. Basal

plantar surface of foretarsal segment 5 with about 7 long, stout, spiniform setae.

Claw on foreleg long, weakly curved; submedian tooth long, about 0.5 length of claw,

and moderately thick at base, tapering to a blunt, very slightly recurved tip; basal

tooth thick, strongly decurved and blunt, subequal to submedian tooth.

MALE GENITALIA (fig. 25). Segment VIII: Tergite and sternite with narrow to

moderately broad, obovate scales. Tergite usually with darker scales on caudal and

caudolateral margins. Sternite with median dark scales and lateral, cream to white

scales. Segment IX: Sternite long, about 0.25 length of sidepiece, subtrapezoidal.

Anterior apodeme moderately short, subrectangular to subtriangular, extending

across anterior border of sternite. Sidepiece: Tergal surface with 4,5 long, submedi-

an tergomedial bristles and one subapicolateral bristle; bristles mesad of tergomedial

bristles moderately long. Parabasal spine long, at least 2.0 length of its tubercle. Ba-

sal apodeme moderately thick, pointed, long, about 0.20-0.25 length of sidepiece.

Longer, more dorsal accessory spine slightly less than 0.5 length of sidepiece; more

ventral spine about 0.75 of longer spine. Internal spine subequal to shorter accessory

spine, apically retrorse. Clasper: Spiniform moderately developed, acuminate, sub-

equal to seta b. Dorsal Claspette: Pedicel long, moderately narrow to moderately

broad; base rounded, curved mesad; internal apodeme conspicuous. Leaflets broad

and less than 0.5 (0.37-0.47) length of claspette; dorsal leaflet with prominent, fairly

large basomesal projection. Ventral Claspette: Moderately short, 0.25-0.40 length of

sidepiece; lateral margins not tapering appreciably medially toward apex; apex broad,

width at apex about 0.50-0.60 (0.50-0.64) length of claspette. Basal lobule moder-

ately expanded laterally, not appreciably wider than apex of claspette; setae along ba-

sal margin moderately long, slightly longer than to 1.5 width of aedeagus but never

2.0 width; setae distributed evenly over basal surface and radiating in different direc-

tions; no concentration of long setae on basomesal margin directed caudad toward

vertex of mesal cleft. Ventral and lateral surfaces (exclusive of basal lobule) with

short setae about 0.5 width of aedeagus, setae extending to or nearly to apex. De-

pending on angle of observation, apex of ventral claspette appearing either truncate

or with a variously developed median sulcus with gradual to steeply sloping sides;

apex with abruptly angled, rounded, sclerotized, lateral margins. Preapical plate

moderately small, semicircular to oval, weakly to moderately heavily sclerotized, lo-

cated about 0.2 length of claspette basad of apex. Transparent membranous area ba-

sad of preapical plate as in rangeli except slightly more pointed at vertex. Refringent

structure inconspicuous to moderately conspicuous, in shape of inverted V, with

rounded vertex; lateral arms occasionally apparent. Phallosome: Aedeagus about 0.5

length of sidepiece and 1.33-1.60 (1.31-1.89) length of ventral claspette; apex moder-

ately rounded, usually wider than long; with or without very small, membranous,

nonserrated, pointed, basolaterally directed leaflets.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 115

PUPA (fig. 25). Abdomen: about 2.8 mm. Trumpet: 0.45 mm. Paddle: 0.75 X

0.55 mm. Only exuviae examined. Cephalothorax: Moderately to heavily pigment-

ed. Wing cases with heavily sclerotized stripes. Hairs 1-3-C 2,3 branched (2-4); 2-C

shorter than 1,3-C. Hair 4-C 2-4 branched, about 0.7 length of 5-C. Hair 5-C 2,3

branched (2-4). Hair 6-C 2 forked (2,3) 0.25-0.70 from base, moderately short,

slightly longer than 5-C. Hair 7-C 2,3 branched (1-3) near base, long, subequal to 6-

C, one branch about 1.25 length of shorter branch(es). Trumpet: Pinna moderately

pigmented; moderately long, about 3.5-4.5 (3.0-4.8) length of meatus; in lateral as-

pect, appearing broad medially and tapered toward apex. Meatal cleft long and basal-

ly rounded. Metanotum: Hair 10-C 1-3 branched, less than 0.5 length of 12-C and

about 0.8 length of 11-C. Hair 11-C 3-5 branched (2-7) usually near base. Hair 12-C

1-3 branched, very long. Abdomen: Hair 2-I dendritic, 5-8 branched (3-9) near vase,

large, with long branches; 3-I single or double, long, 0.7-1.0 length of 2-I. Hair 4-I 4-

6 branched (4-7), moderately short. Hair 5-I 1-3 branched, long. Hair 6-I single or

occasionally double, very long, usually more than 2.0 length of 7-I. Hair 7-I 3-5

branched (2-5) near base. Hair 9-I single or rarely double, about 0.7 length of 6-1.

Hair O-II-V 4-7 branched (3-8), moderately large; O-VI, VII 2-5 branched, moderately

short. Hair 1-II,III 6-8 (6-13) and 5-9 (4-11) branched respectively, strongly devel-

oped, median branches longer than lateral; 1-[V-VII very long, 1.6-2.0 length of seg-

ment. Hair 3-IV 4-6 branched (3-9) near base, moderately large; 3-V 2-4 forked (1-4).

Hair 5-HI,[V 411 and 3-5 (2-5) branched respectively, strongly developed; 5-V-VII

single, as long as or slightly longer than segment. Hair 6-I] 1-3 branched, 1.4-1.8

length of 7-II or occasionally longer; 6-III 1-3 branched, long. Hair 7-II 3-5 branched

(2-6); 7-III 3-5 branched (1-6); 7-IV 4-6 branched (2-6); 7-V 2-4 branched, longer

than 7-III,IV; 7-VI,VII single or rarely double, long. Hair 8-III,VII 3-5 branched (2-

5); 8-IV 3 branched (2-6); 8-V,VI 2-3 branched (2-4). Hair 9-II small, unpigmented;

9-III pointed, almost always less than 2.0 length of 9-II, pigmented; 9-IV thick, point-

ed, usually 2.0 or more of 9-III; 9-V-VIII thick, curved, acuminate, short, about 0.33

length of segment; 9-V,VI slightly shorter than 9-VII, VIII; 9-VIII straighter than 9-V-

VII. Hair 10-III 3-5 branched (2-5), moderately long. Hair 4-VIII 3,4 branched (3-5),

subequal to 9-VIII. Terminal Segments: Male genital lobe as in aquasalis, with a very

prominent mammilliform protuberance. Paddle: Obovate with moderately broad,

rounded apex, weakly emarginate at insertion of hair 1-P. External buttress 0.65-

0.75 length of paddle, with distinct spiculiform serrations near apex. Spicules on ex-

ternal margin conspicuous, continuing around apex and along inner margin to about

0.3 from base. Hair 1-P single, moderately long; 2-P 2,3 branched (1-3), about 0.5

length of 1-P.

LARVA (fig. 26). Head: 0.6 mm. Antenna: 0.3 mm. Anal Saddle: 0.3 mm.

Head: Moderately to very heavily pigmented. Median tooth of mental plate blunt,

as broad as combined widths of 2 adjacent teeth from one side. Inner and outer cly-

peal hairs (2,3-C) single and barbed in apical 0.5-0.7, barbs usually longer on 3-C;

hairs 2-C widely spaced, clypeal index 1.0-1.3; 3-C slightly shorter (0.75-0.90) than

2-C. Hair 4-C single or 2-4 forked, moderately long, 0.30-0.60 length of 3-C and usu-

ally extending to near or beyond base of 2-C. Hair 8-C 3-5 branched (2-7) near base,

moderately large, length greater than 2.0 distance separating insertion of hairs 2-C

and 3-C. Hair 9-C dendritic, 5-8 branched (4-8), slightly longer than 8-C. Hair 12-C

3,4 branched (2-5), moderately long. Hair 15-C 2-6 branched, moderately long. Col-

lar wide dorsomedially, heavily pigmented. Antenna: Usually lighter than remainder

of head. Mesal margin with moderately short, thin spicules; fewer and shorter spic-

ules on ventral surface. Hair 1-A 5-7 branched (5-8), short, about as long as width of

. 116 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

antenna at point of insertion, inserted about 0.25 from base of antenna. Thorax:

Darkly pigmented. Submedian prothoracic group (1-3-P) with or without 1,2-P shar-

ing a common sclerotized tubercle, often base of 1-P not sclerotized; palmate hair 1-

P with 9-12 (9-15) pigmented, moderately broad leaflets; 2-P 12-18 branched (1 1-

20); 3-P not sharing tubercle with 2-P. Hair 14-P 6-9 branched (5-11) from a moder-

ately short, flattened shaft. Mesothoracic hair 1-M 27-31 branched (25-35). Meta-

thoracic hair 2-T 1-3 forked, moderately long. Palmate hair 3-T with 11-15 (9-18)

semitransparent, moderately long, narrow leaflets. Mesothoracic pleural group spine

large, subequal to prothoracic spine; metathoracic spine smallest. Abdomen: Hair 0-

II-VII 5-8 branched (4-10), large, very conspicuous; O-II subequal to or longer than

length of leaflets of 1-IJ. Palmate hair 1-I with 13-16 semitransparent, spreading, nar-

row, pointed leaflets; 1-III-VI with 23-31 long, pointed, moderately narrow to mod-

erately broad leaflets; 1-II, VII with 20-27 leaflets, slightly shorter than 1-III-VI. Hair

2-I 3-5 branched (2-6), small; 2-I] 6-9 branched (4-10), large; 2-II] 4,5 branched (3-

6), large; 2-IV single or occasionally double, moderately long; 2-V single, very long.

Hair 3-I single or occasionally double, moderately long. Hair 5-I 4,5 branched (3-5),

moderately small; 5-II 8-11 branched (6-13), small. Hair 9-I 5-7 branched (4-7). Hair

13-I 5-7 branched (4-8), small; 13-II,III 8-10 branched (7-11), small; 13-1V 4-6

branched (3-6), moderate, equal to or slightly longer than leaflets of 1-IV; 13-V 4-6

branched (4-7), large; 13-VI 7-10 branched (6-10), moderately small. Spiracular

Lobe: Pecten with 15-19 (14-19) teeth; length of teeth highly variable, usually medi-

an teeth mixed moderate and short, except for one long tooth in middle; dorsally

pecten usually terminating with 2,3 long teeth; serrations on teeth moderately long,

conspicuous. Hairs 8,9-S 3-6 branched; 8-S subequal to 9-S. Lateral arm of spiracu-

lar apparatus short, directed caudolaterally. Anal Segment: Covered with short thin

spicules. Saddle dark reddish brown, with irregular ventral margin. Hair 1-X longer

than saddle; inserted on saddle near ventral margin, on ventral margin at base of in-

dentation, or just ventrad of ventral margin. Anal gills subequal to length of saddle.

DISCUSSION. An. nuneztovari can be distinguished from the other species in the

Oswaldoi Group in the female (except occasionally rangeli and often trinkae) by the

combination of (1) integument dark brown to very dark brown with considerable

contrast between light and dark regions, and light wing spots usually cream at least on

anterior veins, (2) usually ventral surface of palpal segment 4 with row of dark scales,

(3) foretarsal segment 2 with a light band in apical 0.2-0.4 (0.15-0.44) and segment 3

with a light band in apical 0.55 (0.40-0.75), (4) foretarsal segment 4 with a small, api-

cal, cream band, (5) hindtarsal segment 2 with a dark band in basal 0.25-0.32 (0.20-

0.32), (6) large dark wing spots distinct, humeral light spot of vein C usually smaller

than in trinkae, 0.7-1.3 (0.7-1.7) of basal dark spot, (7) subbasal dark spot of vein C

usually longer than subbasal light spot, (8) subcostal light spot of vein C 0.20-0.55 of

subcostal dark spot, usually less than 0.5 and (9) subcostal and apical dark spots of

vein Ry,5 and dark spots of M,,, and M3,, all very dark and conspicuous; in the male

genitalia by the combination of (1) ventral claspette moderately short, 0.25-0.40

length of sidepiece, width at apex about 0.50-0.60 (0.50-0.64) length of claspette,

and apex appearing either truncate or with median sulcus, (2) basal lobule moderate-

ly expanded laterally, setae along basal margin moderately long, slightly longer than

to 1.5 width of aedeagus but never 2.0 width, setae distributed evenly over basal sur-

face and radiating in different directions, not concentrated along basomesal margin

as in rangeli, (3) ventral and lateral surfaces of ventral claspette (exclusive of basal

lobule) with short setae about 0.5 width of aedeagus, (4) preapical plate semicircular

to oval, and weakly to moderately heavily sclerotized and (5) length of aedeagus

Faran: Albimanus Section of Anopheles (Nyssorhynchus) VL?

1.33-1.60 (1.31-1.89) length of ventral claspette, apex of aedeagus usually wider

than long, moderately rounded and with or without very small, membranous leaflets;

in the pupa by the combination of (1) hair 7-C subequal to 6-C, one branch about

1.25 length of other branch(es), (2) pinna moderately long, about 3.5-4.5 (3.0-4.8)

length of meatus, appearing broad medially and tapered toward apex in lateral aspect,

(3) hair 10-C less than 0.5 length of 12-C and about 0.8 length of 11-C, (4) hair 2-I

dendritic, 5-8 branched (3-9), large, 3-I 0.7-1.0 length of 2-I, (5) hair 6-I usually

more than 2.0 length of 7-I and (6) hair 9-II small, unpigmented, 9-III pointed, al-

most always less than 2.0 length of 9-II, pigmented, 9-IV usually 2.0 or more length

of 9-III, and 9-V-VIII thick, curved, acuminate, short, about 0.33 length of segment;

and in the larva by the combination of (1) hairs 2,3-C single and barbed, hairs 2-C

widely spaced, clypeal index 1.0-1.3, (2) hair 3-C 0.75-0.90 length of 2-C, (3) hair 4-

C single or 2-4 forked, moderately long, 0.30-0.60 length of 3-C, usually extending

to near or beyond base of 2-C, (4) palmate hair 1-P with 9-12 (9-15) pigmented,

moderately broad leaflets, (5) hair O-II-VII 5-8 branched (4-10), large, very conspicu-

ous, O-II subequal to or longer than leaflets of 1-II, (6) hair 13-1V 4-6 branched (3-6),

moderate, equal to or slightly longer than leaflets of 1-[V, 13-V 4-6 branched (4-7)

and (7) hair 1-X inserted on saddle near ventral margin, on ventral margin at base of

indentation, or just ventrad of ventral margin.

In the specimens from Venezuela as compared to elsewhere, the immatures usually

have hairs with somewhat fewer branches, while the adults have the light apical band

on foretarsal segment 2 usually wider, the apical light fringe spot and subcostal light

spot on vein C usually larger, and the sectoral dark spot on vein C often absent.

An. nuneztovari and trinkae are sister species based on shared derived (synapo-

morphic) characters in the larvae and male genitalia, and together these species form

the sister group of rangeli. The male genitalia of the 2 former species are so similar

that it is possible to distinguish them only when comparing the lengths of the aedeagi

relative to the lengths of their corresponding ventral claspettes. The larvae of nunez-

tovari and trinkae share the derived feature of the long hair 4-C and are easily sepa-

rated by the characters enumerated in the key. Unfortunately, the adult females of

all 3 of the above species can sometimes be confused due to intraspecific variability

and the paucity of reliable differentiating characters. It is very important when there

is some doubt, to examine as many specimens as possible, and to correlate all the

characters given in the key.

Based on the male genitalia, Gabaldon described nuneztovari from San Carlos, Co-

jedes, Venezuela in 1940. Rozeboom and Gabaldon (1941) described a similar form

of Nyssorhynchus from the Amazon basin which they called goeldii and stated that

goeldii and nuneztovari were distinguished by differences in the ““mesosome”’ (phal-

losome) and the “fused dorsal lobes’’ (ventral claspettes) of the male genitalia. Again

based on the apex of the mesosome of the male genitalia, which was said to be with-

out leaflets and broader than in goeldii, Causey (1945) described dunhami from Tefe

in the upper Amazon basin. Floch and Abonnenc (1946b:1) synonymized goeldii

with nuneztovari after comparison of the male genitalia of goeldii with several speci-

mens of nuneztovari. Lane in his “Neotropical Culicidae”’ (1953:268) listed dunhami

as a synonym of nuneztovari, so nuneztovari became the valid name for the above 3

nominal species.

Elliott (1968: 248-252) studied the relationship of the indoor biting activity of nu-

neztovari with regard to human activity in 5 Colombian localities. In 1972 (p 758),

Elliott stated that 2 species were confused under the name nuneztovari: ‘In Brazil

and Surinam, in Ecuador, and almost certainly in Bolivia and Peru, there is a species

- 118 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

that bites at sunset, and therefore mainly outside houses, and preferentially on an-

imals. In northern Colombia and western Venezuela, a closely similar but slightly

smaller species bites later in the night, and therefore mainly inside houses.”’ Accord-

ing to Elliott, the possible sympatry of the 2 types in Venezuela and Colombia could

have lead to confusion, such that the behavior of the nonvector has been attributed

to the vector of malaria in this region. Whether or not Elliott was dealing only with

nuneztovarl, or with nuneztovari and trinkae,is not known to me.

Although I have not examined the holotype of nuneztovari, | have studied the ho-

lotype and several paratypes of goeldii, the holotype of dunhami, and numerous

other specimens from Brazil (Amazonas, Para, Rondonia), the Guianas and Venezue-

la (Aragua, Zulia, Barinas and Cojedes, including 3 male genitalia from the type local-

ity of nuneztovari). I have been unable to find any apparently significant differences

among any of these specimens and therefore am considering them to be all nunezto-

vari. However, when more information and material becomes available which can be

correlated with the morphological differences I have observed and with those behav-

ioral and chromosomal differences reported by others, it is possible that goeldii will

be resurrected to specific status.

Kitzmiller, Kreutzer and Tallaferro (1973) studied the salivary chromosomes of

nuneztovari from 3 populations in Brazil, 2 in western Venezuela and one in

northern Colombia. They reported that the Colombian and Venezuelan populations

consistently differ from the Brazilian populations by a homozygous inversion in the

X-chromosome. They stated that the 2 “‘types” could be easily distinguished by this

fixed homozygous inversion. Also, they compared the chromosomal banding pat-

terns of nuneztovari with those of An. (Nys.) darlingi, aquasalis, albimanus and later

(Kreutzer, Kitzmiller and Rabbani 1975:363-364) An. (Nys.) argyritarsis. The simi-

larities between nuneztovari and albimanus were not as great as between darlingi and

aquasalis. ‘The autosomal homologies were strongest between aquasalis and nunezto-

vari, and aquasalis shared more homologous regions with argyritarsis than did nunez-

tovari.

BIONOMICS. The immatures of nuneztovari are found in open marshy areas,

ponds and lakes often in the grassy margins, small or large permanent or temporary

ground pools, animal or wheel tracks, and along stream margins; they are found in

fresh water that is in full sun or partial shade. Aquatic vegetation may be abundant

and algae are often present. An. nuneztovari is collected in the interior or in clear-

ings within the forest, and in areas of secondary growth (scrub) such as around vil-

lages. In the laboratory, the larvae cannot tolerate a sodium chloride concentration

of even 0.5%. An. triannulatus and especially ininii are frequently associated with

this species in ponds and lakes; occasionally oswaldoi is also collected with nunezto-

vari, Other species associated with nuneztovari are Culex (Mel.) bastagarius, Cx.

(Mel.) dunni group, Cx. (Mel.) ensiformis, Cx. (Mel.) rachoui (2), Cx. (Mel. ) zeteki

group, Cx. (Cux.) coronator group and Uranotaenia (Ura.) geometrica.

Considerable work has been done on the adult behavior of nuneztovari from Co-

lombia by Elliott (1968:248-252; 1972) which I will briefly summarize. Elliott stud-

ied the relationship between the biting activity indoors and outdoors with regard to

human activity in 5 localities in Colombia: El Pescado, Rio Fuego, Puerto Reyes,

Turbo and Las Aranas. At Rio Fuego he found the months nuneztovari was most

abundant were April and May; biting occurred the first 2 hours of the evening out-

doors, the next 2 hours indoors and over the entire night with an outdoor to indoor

ratio of 1:1. At all the other stations he reported the same pattern. In the months

of highest density, biting reached a peak shortly before midnight, indoor biting being

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 119

equal to or slightly greater than outdoor. In the months of low density, biting

peaked an hour or so earlier than during the high density season, biting being slightly

greater outdoors than indoors. He states, “there are indications of a negative correla-

tion between the seasonal density and the proportion of outdoor biting...” An. nu-

neztovari in Colombia also showed earlier peaks of activity during the light phases of

the moon (Elliott 1972:756). Biting activity was unimodal, not showing early eve-

ning or morning peaks. An. nuneztovari collected outside in resting places equidis-

tant between animals and houses had a human blood index (HBI) of less than 10%.

This species usually rests one meter high or less on the walls inside houses, although

occasionally (15%) above this height. Because it feeds inside while most people

sleep, it is effectively endophilic.

In 3 months (September to November), Rey and Renjifo (1950) collected 3301

anophelines inside houses and in stables in northern Colombia of which 14.4% were

identified with certainty as nuneztovari and 67.2% as a mixture of nuneztovari and

rangeli, Of the total of 2722 nuneztovari and rangeli collected, a study of the eggs

from 501 of these females revealed that 476, or 95%, were nuneztovari. These auth-

ors therefore concluded that the majority of females collected inside were actually

nuneztovari. The principal time nuneztovari entered houses was 2200-2400 h.

Gabaldon (1972:635) stated that before spraying the inside walls of houses with

DDT in Venezuela, nuneztovari was very endophilic, remaining in the houses and rest-

ing on the walls and ceiling after taking a blood meal. Spraying of insecticides, how-

ever, selected for “‘intense exophilism.”’ An. nuneztovari is still anthropophilic, but,

immediately after taking a blood meal, the females leave the house, thereby avoiding

a lethal dose of insecticide by not resting on the walls. Gabaldon stated that, even

though strongly exophilic, “A. nuneztovari in Venezuela, for example, maintains a

human blood preference of around 80%, and a man-biting rate of more than 100 dur-

ing a night indoors.” Gabaldon believes that this intense exophilism has been largely

responsible for refractory malaria in Venezuela.

Panday (1977:728-737) reported a unimodal distribution of biting activity of nu-

neztovari in Suriname occurring 1800-1900 h, the peak beginning at the termination

of twilight (1830 h, sunset was fixed at 1800 h). His recent surveys showed a tre-

mendous increase in the numbers of nuneztovari in the “hilly and mountainous for-

est region” in the interior of Suriname. He believes that to a large extent this in-

crease is due to the construction of Afobaka dam, which has been responsible for the

formation of Brokopondo storage lake. From daily collections 15 m inside the forest

near the lake, 1800-1900 h, on human bait, July-December 1976, 13,824 females

were captured. He stated that “‘An. nuneztovari seems to be the dominant anthropo-

philic Anopheles species” in this region and implicated it as the primary vector of

Plasmodium falciparum. In laboratory studies on the life cycle of this species, Pan-

day found the egg stage lasts one day, larval stages 7 days and pupal stage one day.

The first gonadotrophic cycle requires 5 days, whereas all subsequent cycles require

4 days. The maximum parous rate during the above period was 0.69 with a mini-

mum of 0.14 and a mean of 0.34. He also reported that grassy vegetation seems es-

sential for oviposition. Contrary to what is observed in Venezuela, the adults are sus-

ceptible to DDT, dieldrin and malathion.

In Para, Brazil, the females seem to be primarily exophagous, unlike nuneztovari

in Colombia and Venezuela. Deane, Causey and Deane (1948:895-896) reported

that of the 21,967 females of nuneztovari collected, only 411 or 1.9% were captured

inside houses. In 29 hours collecting inside houses in Maraba, Para, one specimen of

nuneztovari was found; however, in 31 hours outside on animal bait, 978 females

36° Contrib. Amer Entilnst., vol 13, 007, 1980

were captured. Feeding preference studies, comparing a horse and a man as bait, in-

dicate that nuneztovari feeds freely on man outdoors. Recent studies, conducted

from March 1975 to April 1976 by the U. S. Army Medical Research Unit-Belem

(USAMRU) in Palestina (100 km SW of Maraba, Para), also indicate that nuneztovari

is exophilic and most active at sunset. In this region, nuneztovari was the dominant

anopheline captured in landing and resting collections.

Scorza, Tallaferro and Rubiano (1976: 130-131) conducted precipitin tests to de-

termine what hosts females of nuneztovari were feeding on in Santa Barbara, Barinas,

Venezuela. Of those blood fed, 75% tested positive for the immune sera used in this

study and 25% did not react; 74.2% (289) had fed on cattle, 13% (50) on dogs, 7.4%

(29) on humans, 4.5% (19) on chickens, 0.7% (3) on horses, 0.2% (1) on cats and

none on pigs. : |

MEDICAL IMPORTANCE. An. nuneztovari is a major vector of malaria in west-

ern Venezuela and northern Colombia. It was first discovered naturally infected with

Plasmodium sp by Rey and Renjifo (1950). Gabaldon and Guerrero (1959:434) sta-

ted that in some areas where nuneztovari was transmitting malaria the spleen indices

were close to 100%. They also found that in areas not close to the jungle, malaria

disappeared when the local inhabitants took chloroquine; however, in districts near

forests chloroquine failed to stop transmission. Hamon, Mouchet et al. (1970:32-33)

also indicated that the importance of nuneztovari depends on the amount and densi-

ty of vegetation around houses. Vector density is reduced in areas where vegetation

has been cleared around houses.

In Suriname, Panday (1977: 728-737) reported that nuneztovari may have been

the principal vector of Plasmodium falciparum in recent epidemics. He stated that

An. (Nys.) darlingi, thought to be the primary vector of malignant malaria, has not

been captured in the epidemic regions. In these same areas nuneztovari has been col-

lected in great numbers. Since nuneztovari is the dominant anthropophilic mosquito

and has a fairly high parous rate (mean 0.34), Panday states that this species in the

Brownsweg area, ‘“‘may well act as a good vector of pathogens.”

An. nuneztovari has not been reported to be an important vector of malaria in the

Amazon basin. Deane, Causey and Deane (1948) dissected 405 adults and found all

to be negative for Plasmodium sp. Recently, in the Maraba area of Para, Brazil, Dix-

on and Roberts (personal communication) found both P. falciparum and P. vivax in

the local human population in the absence of recognized primary or secondary vec-

tors. Dixon and Roberts stated, ‘‘In August 1975, 4 teenage members of a colonist

family in Gleba 36 [a small village] became ill with malaria (3 with falciparum and 1

vivax; all denied recent travel from their lote).’” Furthermore, a high percentage of

the transient workers who are initially negative for Plasmodium sp, contract malaria

after working a short time in this region. As in Suriname, darlingi, thought to be the

vector of malaria along waterways, is not collected in the Maraba area. Although

nuneztovari occurs in high densities in this region, the entomological and/or epidemi-

ological data are insufficient to implicate this species as a vector. Several other ano-

phelines have been collected in this general locality, such as ininii, oswaldoi, strodei,

triannulatus, ?aquasalis, An. (Nys.) allopha, An. (Nys.) braziliensis, An. (Ano.) ?mat-

togrossensis and An. (Ano.) sp. None of these species has been considered a primary

vector with the exception of ?aquasalis which occurs in relatively low numbers. For

these reasons, nuneztovari, a known or suspected vector in other countries of north-

ern South America, should be carefully investigated as a potential vector in Para,

Brazil.

Scorza, Tallaferro and Rubiano (1976: 129-136) attempted to infect nuneztovari

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 121

experimentally with P. falciparum and P. vivax and found that this species is highly

susceptible to the former. The results for P. vivax were unexpected. Mosquitoes

that had shown a high rate of early infection with P. vivax (55.9% with young oo-

cysts) when dissected after 20 days had neither mature oocysts nor sporozoites.

They explained this by the fact that the patient used to infect the mosquitoes had in-

gested the sporocide pyrimethamine which resulted in suppression of the develop-

ment of P. vivax. The strain of P. falciparum used in the experiment was resistant to

chloroquine and pyrimethamine.

DISTRIBUTION (fig. 3). An. nuneztovari occurs throughout much of the Ama-

zon basin; it is also found in the Guianas, northern Colombia and Venezuela, and

eastern Panama. It is not known how far south it occurs in Colombia and Venezuela,

nor how far west in the Amazon basin. Cerqueira’s report (1943:19) of goeldii from

Bolivia may refer to trinkae or nuneztovari.

Material Examined: 529 specimens: 106 males, 61 male genitalia, 153 females,

115 pupae, 94 larvae; 114 individual rearings: 70 larval, 42 pupal, 2 incomplete.

BRAZIL. Amazonas (30 specimens: 9M, 4Mgen, 15F, 1 lp): Manaus, May 1901, Durham, 2F;

same locality, Jun 1931, R. Shannon, 7M, 2Mgen; same data, 1F [BM]; same locality and date, 1M,

11F; same locality, 1942, A. Galvao, M.286-2A, 1 lp, 1Mgen. Tefe, 1945, O. Causey, 1M and 1M

gen (holotype of dunhami), 1F. Mato Grosso (6F): Westborder, May 1931, R. Shannon, 6F. Para

(324 specimens: 63M, 21Mgen, 101F, 89p, 501; 89 ind rear: 501, 39p): Altamira, 163 km W of,

6 Nov 1974, J. Reinert et al., coll. 85, 1 lpM, 1Mgen. Bacuri, Gleba 36, Lote 05, 23 Oct 1974, J.

Reinert et al., coll. 31, 1 pM, 1Mgen; same data except Lote 02, coll. 33, 3Mgen, 11 lpM, 6 IpF, 5

pM, 2 pF; same data except 26 Oct, coll. 50, 1Mgen, 4 lpM, 7 lpF, 2 pF; same data except Gleba

34, Lote 05, coll. 51, 1Mgen, 1 lpM; same data except Gleba 29, Lote 09 and 28 Oct, coll. 60, 1

pM; same data except Gleba 34, Lote 02, coll. 62, 5Mgen, 1 IpM, 6 lpF, 13 pM, 6 pF; same data ex-

cept Gleba 38, Lote 02 and 29 Oct, coll. 64, 2Mgen, 6 lpM, 7 lpF, 3 pM, 6 pF. Belem, 1944, O.

Causey, 3M, ?2F; same locality, 6 Nov 1968, T. Aitken, 4M, 4Mgen; same locality, 29-30 Sep 1970,

T. Aitken, A. Toda, 2M, 2Mgen, 2F; same locality, BRB 43. Boa Vista, C. Townsend, 6M, 6Mgen,

11F (including holotype and paratypes of goeldii). Curralinho, 1935, H. Kumm, 2F (K.91) [BM

1936-319]; same data, Rio Aracaca, 1F (K.256) [BM1936-319]. Fordlandia, Jun 1931, R. Shan-

non, 2M, 1Mgen, 8F; same data, 2F [BM]; same locality, 2M, 1F. Itupiranga, 17 Feb 1976, Mora-

may, 3F (1050). Maraba area, 4 May 1976, SF (M-6); same locality, 6 May 1976, 1F (M-8); same

locality, 2 Jun 1976, 6F (R-3); same locality, 3 Jun 1976, 1F (R-5); same locality, 4 Jun 1976, 2F

(M-6); same locality, 6 Jun 1976, 1F (M-8); same locality, 9 Jun 1976, 1F (M-3); same locality, 17

Jun 1976, 3F (A-3); same locality, 23 Jun 1976, 1F (M-11); same locality, 7 Oct 1976, 1M (1261);

same locality, 8 Oct 1976, 1M (1262); same locality, 1976, D. Roberts, 10F. Palestina, 100km SW

of Maraba, 21 Sep 1976, Moramay, 1F (N-1252). Rondonia (1Mgen, 4F): Guajara Mirim, May

1931, [R. Shannon], 1F. Porto Velho, May 1931, R. Shannon, 1Mgen (43.11.17), 3F.

COLOMBIA (1F). Atlantico: Barranquilla, L. Dunn, 1F.

FRENCH GUIANA (2Mgen). Inini: Oyapock, 1Mgen [JH]. Saut Tigre, 30 Oct 1945, 1Mgen

(No 778 23 A).

GUYANA (2M, 6F). East Berbice-Corentyne: Kwakwani, 18 Jan 1942, G. Giglioli, KO 111-18,

1F; same locality and collector, 13 Jan 1943, KO 111-20, 4F. West Demerara-Essequibo Coast:

MacKenzie, Sep 1945, KO 111-29, 1M. District not specified: 1942, G. Bevier, KO 111-8, 1M, IF.

PANAMA (22 specimens: 5M, 4Mgen, 4F, 5p, 4L&l; 5 ind rear: 1 1, 3p, linc). Darien: Pucro,

PA 630. Rio Tuira, GG 1-159, 43.

SURINAME (35 specimens: 19M, 11Mgen, 5F). Brokopondo: Brownsweg, Prof. Dr. Ir. W. J.

van Blommestein Meer, 21 Oct 1975, R. Panday, 4F; same locality and collector, 31 Jul 1976, 3M,

1Mgen; same locality and collector, 9 Aug 1976, 6M, 3Mgen; same locality and collector, 7M, 5M

gen. Marowijne: Moengo, 6 Jun 1945, Guicherit, 1M. Paramaribo: Paramaribo, 5-9 Jul 1945,

Tillema, 2M, 1Mgen (46-v-8a), 1Mgen (46-v-8b), 1F.

VENEZUELA (96 specimens: 8M, 18Mgen, 11F, 20p, 39L&l; 19 larval ind rear). Aragua: Mar-

acay, 1 Aug 1929, R. Hill, 1Mgen (56) [JH]; same locality, 3 Sep 1926, M. Nunez Tovar, 6F.

‘(1 | Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Barinas: Andres Eloy Blanco, Aug 1977, [P. Cova Garcia] , 8lpM, 11 lpF, 15 L. Locality not spe-

cified, Jan 1943, Iriarle [?], 1F. Cojedes: San Carlos, 2Mgen; same locality, 1Mgen [JH]. Zulia:

Mene Grande, 2Mgen (no 3) [JH] ; same locality, 6Mgen [JH]. State and locality not specified:

27 Dec 1939, A. Gabaldon, 1Mgen [JH] ; same collector, [1977], 1 p, 3L, 3Mgen; same collector,

1Mgen [JH] ; no data, 1L (1949), 1L (1950), 1Mgen.

STRODEI COMPLEX

FEMALES. No diagnostic characters distinguishing the adult females of this com-

plex from the species in the Oswaldoi Complex are apparent.

MALES. Essentially as in females except for sexual characters.

MALE GENITALIA. Segment IX: Sternite moderately long to long. Anterior

apodeme short to moderately long. Sidepiece: Parabasal spine moderately long to

long, often reaching lateral margin of aedeagus on other side. Ventral Claspette: Se-

tae on lateral and ventral margins extending only a short distance toward apex, not

as far as level of apical margin of preapical plate, absent from apex. Apex truncate,

rugose or deeply striated, expanded laterally into rounded or pointed lobes. Preapt-

cal plate oval to circular, distinct. Phallosome: Aedeagus rounded apically, without

leaflets.

LARVAE. Moderately large to large. Head: Inner and outer clypeal hairs (2,3-C)

plumose with moderately long branches, or single and barbed; inner clypeals (2-C)

widely spaced in benarrochi and closely approximated in strodei. Hair 4-C short and

branched. Collar moderately narrow to narrow dorsomedially, heavily pigmented.

Antenna: Hair 1-A large in benarrochi, small in strodei. Hairs 2,3-A moderately de-

veloped. Thorax: Submedian prothoracic group (1-3-P) usually without 1,2-P shar-

ing a common tubercle. Hair 14-P with a short to moderately short, flattened shaft.

Abdomen: Hair 0-II moderately small. Hair 5-I moderately large and close to lateral

margin of abdomen. Hair 13-I small, numerously branched; 13-I[V moderately large

in strodei, small in benarrochi. Spiracular Lobe: Lateral arm of spiracular apparatus

short in strodei, moderately long and heavy in benarrochi. Anal Segment: Hair 1-X

short in benarrochi, long in strodei; inserted on saddle. Anal gills moderately long to

long.

DISCUSSION. The Strodei Complex is characterized in the male genitalia by the

apex of the ventral claspette being laterally expanded, rugose and without setae.

This complex is composed of 3 species. Two of these, strodei and rondoni, are ex-

tremely similar and represent a clearly defined phyletic line. These 2 species share

several derived characters. The male genitalia are almost identical, and the larvae re-

portedly differ only in the number of branches of a few setae. An. strodei is widely

distributed throughout Central and South America, often being found at relatively

high altitudes. An. rondoni is limited to the interior of southern South America on

the eastern side of the Andes extending south to the northern provinces of Argentina.

An. rondoni is highly derived in the adult female, possessing a dark basal band on

hindtarsal segment 3, and unique and extensive dark spots on the wing.

An. benarrochi is the most derived species in the complex. Partly because it has so

many unique characters, its phylogenetic relationship to the rest of the species in the

Oswaldoi Subgroup is not clearly understood. On the basis of the male genitalia, I

~am placing benarrochi in this complex, but it may be shown later to be more closely

allied with the Oswaldoi Complex. The distribution of benarrochi is similar to that

of rangeli, these species occurring principally in the Orinoco basin and extending

south into Brazil and Peru. The adult of benarrochi is very similar to that of aquasa-

lis, making species identification based on external morphology very difficult. The

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 123

larva of benarrochi is very different from that of strodei, having unique characters

such as a large hair 1-A of the antenna and a small, numerously branched hair 13-IV.

11, Anopheles (Nys.) strodei Root

Fis, 1 47,27, 28

1926. Anopheles (Nyssorhynchus) strodei Root 1926b:711-714. TYPE: Lectotype male (64)

with associated genitalia on slide (750923-29), a few miles from Agua Limpa, near Juiz de

Fora (Minas Gerais), Brazil, 27 Mar 1925 [USNM; designation by Stone and Knight 1956:

280|. RESURRECTED FROM SYNONYMY with evansi (Brethes 1926).

1940. Anopheles (Nyssorhynchus) strodei var. ramosi Unti 1940b:489-491. TYPE: Syntypes

larvae, Lorena (Sao Paulo), Brazil, Mar 1940 [NE]. NEW SYNONYMY.

1941. Anopheles (Nyssorhynchus) strodei var. arthuri Unti 1941:9. TYPE: Syntypes eggs, Vale

do Rio Paraiba [do Sul] (Sao Paulo), Brazil, elev. ca 500 m [NE]. NEW SYNONYMY.

1941. Anopheles (Nyssorhynchus) strodei var. artigasi Unti 1941:9. TYPE: Syntypes eggs, Vale

do Rio Paraiba [do Sul] (Sao Paulo), Brazil, elev. ca 500m [NE]. TRANSFERRED

SYNONYMY.

1941. Anopheles (Nyssorhynchus) strodei var. albertoi Unti 1941:9-10. TYPE: Syntypes eggs,

Vale do Rio Paraiba [do Sul] (Sao Paulo), Brazil, elev. ca 500 m [NE]. NEW SYNONY-

MY.

1941. Anopheles (Nyssorhynchus) strodei var. lloydi Unti 1941:10. TYPE: Holotype egg, Pana-

ma, locality not specified [NE]. NEW SYNONYMY.

Anopheles (Nyssorhynchus) strodei of Davis (1928:549-550); Shannon and Del Ponte (1928:45,

46, 54-55); Hill (1930:711-713); Shannon (1931:10; 1933:120); Curry (1932:566-568; 1934:

647); Edwards (1932:45); Root (1932:779); Senevet (1934:52-54; 1948a:278; 1948c:437);

Komp (1936a:161; 1941a:88, 93, 96, 97; 1942:39, 40, 43, 71-73, 80, 120-122, 132, 158-159);

Galvao, Lane and Correa (1937:41-42); Galvao and Lane (1937a:77, 78; 1937b:275-282; 1937c:

26; 1937d:65-69; 1937e:214, 221-223); Correa (1938:104-109; 1943:126); Galvao (1938a:52,

56-57; 1940:443-447; 1943:143-144, 149, 150; 1950:38-48); Galvao and Amaral (1938:13);

Rozeboom (193 8a:102-103; 1938b:290; 1941:102; 1942a:240-241); Senevet and Abonnenc

(1938:489-494); Galvao and Barretto (1939a:113-114); Barretto (1940:159-164); Lane (1939:

27; 1944:263-266; 1953:271-273); Gabaldon, Lopez and Ochoa Palacios (1940:34-35); Unti

(1941:3-18); Correa and Ramos (1942a:37-46; 1943:35, 36; 1944b:131; 1944c:11); Fonseca

and Fonseca (1942:94); Komp (1942:43, 71-72, 81, 120-122, 132, 158-159); Simmons and Ait-

ken (1942:39, 46, 53, 62, 96-97); Fonseca and Unti (1943:43, 45, 47-49); Kumm, Bustamante

and Herrera (1943:373-375); Unti and Ramos (1943:27-28); Causey, Deane and Deane (1944:2,

5, 6; 1946:26); Deane, L. P., Causey and Deane (1946:6, 8, 9, 12; 1948:904-906); Deane, M. P.,

Causey and Deane (1946:42, 45); Floch and Abonnenc (1946b:3-5; 1947:7; 1951:55-57); Ga-

baldon and Cova Garcia (1946b: 118-120; 1952:184, 196, 201); Amaral and Penido (1947: 168);

Arnett (1947:198; 1950:103, 106, 110, 113); Bates and de Zulueta (1949: 132); Levi-Castillo

(1949:9-11, 16, 28, 58, 67, 72-73, 77, 82, 86); Duret (1950a:372; 1950b:302; 1952:347-348);

Vargas and Martinez Palacios (1950:130-133; 1956:129-132); Carvalho and Rachou (1951:474-

477, 480); Martinez and Prosen (1951:38); Pinotti (1951:670); Rachou and Ferraz (1951:542-

543, 547-553); Rachou and Ricciardi (1951:425-426, 432-437); Horsfall (1955:185); Bejarano

(1956:9, 20; 1957:326, 327, 331-333, 337-339; 1959:317, 325); Schreiber and Pompeu Memor-

ia (1956: 1-3); Vargas V. (1956:29; 1957; 1958a; 1958b); Guedes, Amorim and Schreiber (1957:

247-248); Rachou, Moura Lima, Ferreira Neto and Martins (1958:417, 421, 423); Rachou

(1958: 148); Castro (1959a:175); Castro, Garcia and Bressanello (1959:549, 550); Stone, Knight

and Starcke (1959:32); Schreiber and Guedes (1959b: 128-129; 1960:356-357; 1961:658); Var-

gas (1959:377, 382); Cerqueira (1961:127); Forattini (1962:388-394); Aragao, Ferreira Neto

and Martins (1973:291); Consolim and Galvao (1973:177-179, 181); Cova Garcia and Sutil O.

et 19; 1975b:213; 1977:19, 43, 64, 84); Cova Garcia, Pulido F. and Amarista M. (1978:

157),

Anofeles (!) (Nyssorhynchus) strodei strodei of Unti (1940b:493-505).

. 124 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Anopheles strodei of Benarrochi (1931:691); Kumm (1932:4; 1941b:93-97); Galvao (1938c:133-

134); Gabaldon, Ochoa Palacios and Perez Vivas (1940:42-55); Kumm, Komp and Ruiz (1940:

388, 389, 391, 412, 419); Vargas (1941:110, 118, 120; 1950:106, 121); Unti and Ramos

(1942:94-105); Russell, Rozeboom and Stone (1943:49-50); Castellanos, Murrieta, Lassman and

Ortiz (1949:34); Senior-White (1951a:294); Renjifo and de Zulueta (1952:600-601, 610); Var-

gas and Martinez Palacios (1955:82-83, 106, 114, 121); Vargas V. (1958c:1, 3); Foote and Cook

(1959:22, 23); Schreiber and Guedes (1959a:97-98); Cova Garcia (1961:40-41, 67, 92-93, 114,

129-130, 155); Maciel (1962:471-473); Vincke and Pant (1962:2, 4, 6, 7, 10); Mattos and Xa-

vier (1965:273-274); Cova Garcia and Sutil O. (1975a:19; 1975b:213).

Nyssorhynchus (Nyssorhynchus) strodei of Lima (1928:100-102).

Anopheles (N.) strodei ramosi of Unti (1941:10).

Anopheles (Nyssorhynchus) evansi of Root (1927:477); Dyar (1928:438); Pinto (1930:153-154,

156-157; 1932:287, 293; 1939:425-427); Senevet (1931:108; 1932:252); Komp (1941b:801-

802); Martinez, Prosen and Carcavallo (1959:111-112).

Anopheles (Nyssorhynchus) evansae of Stone, Knight and Starcke (1959:32); Fauran (1961:10);

Stojanovich, Gorham and Scott (1966a:10, 19, 32; 1966b:18, 29, 41); Gorham, Stojanovich and

Scott (1967:17, 43, 54; 1973:114, 135, 144); Belkin, Schick and Heinemann (1968:10; 1971:

5); Vargas (1976:87, 88, 89).

Anopheles tarsimaculata in part (2?) of Howard, Dyar and Knab (1917:978); Petrocchi (1923: 139).

Cellia tarsimaculatus in part (?) of Neiva and Penna (1916:94); Peryassu (1921a:70); Neiva and Pin-

to (19224521; 19220:3'55-357):

Cellia albimana in part (?) of Neiva (1909:69-77).

FEMALE (fig. 7). Wing: 3.3 mm. Proboscis: 2.0 mm. Palpus: 2.0 mm. Forefe-

mur: 1.4mm. Abdomen: 2.8 mm. Highly variable in size and diagnostic characters.

Head: Integument light brown to very dark brown. Proboscis about 1.4 length of

forefemur. Scales on vertex and occiput predominantly white; occiput with white to

cream, cuneate scales extending caudolaterally onto median occiput; erect scales on

vertex very conspicuous, long, white, lanceolate to cuneate, and tapering into thread-

like stalk at base. Dorsal surface of palpal segments 2 and 3 with a few light scales

more extensive on segment 3; apices of 2 and 3 with a white band; segment 4 with a

basal and an apical brown band, and often with a narrow brown stripe on ventral sur-

face; segment 4 white on mediolateral surface; segment 5 completely white. Anten-

na: Flagellar segment 1 with elongate white scales on dorsomesal surface, and with

an incomplete ring of small scales at base. Flagellar segments 2-13 each with a basal

whorl of 8-10 setae. Thorax: Integument reddish brown to dark brown. Light

scales of scutum and scutellum silver to white. Anterior promontory area with elon-

gate scales extending dorsad onto acrostichal line. Humeral tuft scales numerous and

prominent. Prescutellar space moderately small and usually horseshoe shaped. Sp

with numerous (6-9), moderately long, light setae. Upper stp with 3-6 long setae and

a horizontal patch of about 10 obovate to lanceolate, gray to white scales. Lower

stp with 1,2 long setae and a small patch of scales similar to those on upper stp. Up-

per mep with 6-8 long, cream setae; scales absent. Legs: Light scales on coxae usual-

ly white or very light cream. Midfemur with both anteroapical spot and knee spot

fairly large and conspicuous. Foretarsal segments 2 and 3 with a cream to white

band in apical 0.25 (0.18-0.35) and 0.5 (0.25-0.80) respectively; segment 4 all brown,

occasionally with a few, lighter, apical scales; segment 5 usually one color, golden to

brown or with a few lighter apical scales, rarely with apical 0.5 lighter than basal 0.5.

Midtarsal segments | and 2 with a small, golden to white, apical band in about 0.1

and 0.1-0.4 respectively; segment 3 either all dark or with a small, golden to cream,

apical band; segment 4 all dark; segment 5 usually with a small cream band in less

than apical 0.3. Hindtarsal segment 2 with a brown band in basal 0.35-0.45 (0.30-

0.50). Wing: Lengths of wing spots variable. Light wing spots usually white, occa-

sionally very light cream on vein C; in general, light spots relatively large. Vein C

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 125

humeral light spot 2.0-4.0 (1.2-4.1) of basal dark spot; often sectoral light spot ab-

sent; subcostal light spot moderately large, 0.35 (0.25-0.50) of subcostal dark spot;

preapical light spot variable, 0.2-0.5 of preapical dark spot; apical dark spot small. R

presectoral and sectoral dark spots small, occasionally sectoral spot absent. Vein M

with preapical dark spot not extending onto M,,,.. Remainder of light spots on wing

usually small but variable. Apical light fringe spot small to moderately large, usually

divided; additional light fringe spots at apices of Ry,s5, My4., M3.g, Cu,, Cu, and A;

a light fringe spot at level of 0.5 distance from base of A. Abdomen: Sternite I with

a few, moderately short, lateral setae, and long median setae in basal 0.5, occasional-

ly with a few, inconspicuous, lanceolate, light scales. Dark caudolateral scale tufts

on segments II-VII with large cuneate scales.

MALE. Wing: 3.4 mm. Proboscis: 2.6 mm. Forefemur: 1.55 mm. Abdomen:

about 3.15 mm. Essentially as in female except for sexual characters. Size extreme-

ly variable. Head: Palpal segments 2 and 3 with a conspicuous, cream to white, dor-

sal stripe; segment 4 extensively light, white basally and apically with a dark subbasal

band; medially 4 predominantly cream to white, occasionally with a speckling of

dark scales. Antenna: About 0.75 length of proboscis. Flagellar segment | with nu-

merous, long, white, oblanceolate scales on dorsomesal surface; long mesal setae of

flagellar whorl | distinctly white; occasionally setae white on mesal margins of flagel-

lar whorls 2-5. Legs: Forefemur about 0.6 length of proboscis. Basal plantar surface

of foretarsal segment 5 with 6,7 long, spiniform setae; longest setae about 0.3 length

of segment. Claw on foreleg moderately large, curved and acuminate; submedian

tooth moderately thick, tapering to slightly recurved tip; basal tooth large, decurved

and blunt.

MALE GENITALIA (fig. 27). Segment VIII: Tergite and sternite with moderate-

ly narrow, light, obovate scales. Segment IX: Sternite long, about 0.2 length of side-

piece, subtrapezoidal. Anterior apodeme subtriangular to subtrapezoidal, moderate-

ly long, about 0.15-0.25 length of sternite. Sidepiece: Tergal surface with 4-6 long,

submedian tergomedial bristles and 1,2 long, subapicolateral bristles; bristles mesad

of tergomedial bristles moderately long. Parabasal spine moderately slender and ta-

pering, long, usually at least 2.0 length of its tubercle; apex of parabasal spine almost

reaching lateral margin of aedeagus on other side. Basal apodeme moderately long

and thick. Longer, more dorsal accessory spine about 0.5 length of sidepiece; more

ventral spine about 0.8 of longer spine. Internal spine subequal to shorter accessory

spine. Clasper: Spiniform heavy, fusiform, moderately short, shorter than seta b.

Dorsal Claspette: Long, equal to or slightly greater than 0.5 length of sidepiece. Pe-

dicel moderately broad; base usually not curved mesad. Leaflets slightly less than

0.5 length of claspette, strongly concave mesally; dorsal leaflet with a small basome-

sal projection. Ventral Claspette: Large, 0.45-0.52 length of sidepiece; apex very

wide, width at apex 0.5-0.6 length of claspette. Basal lobule large, expanded lateral-

ly and occasionally bending ventrad at base so that setae distributed over basal mar-

gin project caudad; setae along basal margin long, 2.0-3.5 width of aedeagus. Apex

without setae, more or less truncate and rugose or deeply striated, with a small, V

shaped median sulcus. Apex strongly expanded laterally into large rounded lobe; api-

colateral lobe with convex basal and lateral margins, and usually weakly concave api-

cal margin; depending on angle of observation, apex occasionally medially produced

into small, hemispherical cap with median sulcus at its vertex. Setae on lateral mar-

gins extending toward apex to base of apicolateral lobes. Preapical plate moderate,

oval to circular, weakly to moderately sclerotized and moderately well-defined, loca-

ted about 0.25 length of claspette basad of apex. Refringent structure in shape of

126 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

inverted horseshoe. Phallosome: Aedeagus about 0.5 length of sidepiece; apex

rounded, slightly longer than wide; without leaflets.

PUPA (fig. 27). Abdomen: 2.75 mm. Trumpet: 0.5 mm. Paddle: 0.8 X 0.5 mm.

Pupa very similar to that of triannulatus. Cephalothorax: Dark brown. Hairs 1-3-C

moderately long; 2-C 3 branched, subequal to 1,3-C; 1,3-C 2,3 branched. Hairs 4,5-

C 3,4 branched or forked, subequal in length. Hair 6-C 2,3 forked distally, long.

Hair 7-C 1-3 branched near base, long, one branch conspicuously longer, 1.5-2.0

length of other branch(es). Trumpet: In general, similar to that of aquasalis. Pinna

amber; long, about 3.8-4.6 (3.7-4.8) length of meatus; in lateral aspect, appearing

broad medially and tapered toward apex. Meatal cleft moderately open, basally

pointed. Metanotum: Hair 10-C single, moderately long, equal to or slightly shorter

than 11-C. Hair 11-C 3-6 branched usually near base. Hair 12-C 1-4 forked, about

1.75 length of 10-C. Abdomen: Pigmented as cephalothorax. Hair 2-I 5-7 branched

near base, moderately long; hair 3-I about 0.75 length of 2-I. Hair 4-I about 5-7

branched. Hair 5-I single or double, long. Hair 6-I single or double, very long (0.35

mm). Hair 7-I 3 branched, 0.5 or less length of 6-I. Hair 9-I single, subequal to 6-I.

Hair O-II-VII moderately large; 0-II 4-6 branched; 0-III-V 3-6 branched; 0-VI 2-4

branched; O-VII 1-4 branched. Hair 1-II,III 5-8 (3-8) and 4-7 (3-7) branched respec-

tively, strongly developed; 1-IV-VII strong, very long, about 1.6-2.0 length of seg-

ment. Hair 3-I1V 4-6 branched, large; 3-V 2-4 forked (2-5), long. Hair 5-II 3-5

branched, moderately long; 5-III,[V 5-8 and 3-5 (2-5) branched respectively, large; 5-

V-VII single or often 5-V double, long, all equal to or longer than length of segment.

Hair 6-II 1-3 forked, very long, at least 2.0 length of 7-II. Hair 7-II 3-5 branched near

base. Hairs 7-III-V, 8-III-VII branched near base, moderately developed; 7,8-III 3-5

branched; 7,8-IV 2-4 branched; 7-V 3,4 branched; 7-VI long, equal to or slightly

longer than 0.5 length of segment; 8-V-VII 2-4 branched. Hair 9-II thin, small, unpig-

mented; 9-III stout, short, less than 2.0 length of 9-II, unpigmented to strongly pig-

mented; 9-IV about 2.0 of 9-III; 9-V stout, slightly curved, 1.8-2.3 (1.7-2.3) of 9-IV;

9-VI curved, moderately long, 1.1-1.5 (1.1-1.8) of 9-V, much shorter than 9-VII, VII;

9-VIT 1.5-2.0 of 9-VI; 9-VII, VIII thin, tapering to fine point, very long, usually equal

to or longer than 0.5 length of segment. Hair 10-III 3-5 branched near base, moder-

ately long. Hair 4-VIII 2-4 forked, subequal to 9-VIII. Sum of branches of hairs 1-

III, 5-I1I and O-VI less than 18. Terminal Segments: Male genital lobe broad, heavy,

lateral margin tapering at slight angle from base toward apex, appearing thicker than

those of other species, with distinct mammilliform protuberance. Paddle: Large, ob-

ovate, very weakly emarginate at insertion of hair 1-P. External buttress 0.6 length

of paddle. External margin distad of buttress with moderately short, filamentous

spicules extending around apex and along inner margin to 0.5 from base. Hair 2-P 1,

2 forked, subequal to 1-P. |

LARVA (fig. 28). Head: 0.6 mm. Antenna: about 0.3 mm. Anal Saddle:

about 0.3 mm. Highly variable, several differences between Panamanian populations

and southeastern Brazilian populations. Head: Dark reddish brown, heavily pig-

mented. Inner and outer clypeal hairs (2,3-C) single and barbed in about apical 0.5;

hairs 2-C closely approximated, clypeal index about 3.0-4.0; 3-C about 0.6 length of

2-C. Hair 4-C 1-4 branched, small to moderately small, usually not reaching base of

2-C. Hairs 8,9-C 2-7 branched near base, 2.0 length of distance separating insertion

of 2-C and 3-C. Hairs 10,12-C 2-4 and 3-5 branched respectively, subequal to 8,9-C.

Hair 15-C 1-3 branched, long. Collar moderately thin dorsomedially, heavily pig-

mented. Antenna: Moderately to heavily pigmented. Mesal margin with moderately

thin, very sharp spicules. Hair 1-A 4-6 branched, short. Thorax: Submedian pro-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 127

thoracic group (1-3-P) not sharing a common tubercle; rarely 1,2-P sharing common

tubercle; palmate hair 1-P with 13-17 narrow, lanceolate leaflets; 2-P with a small

sclerotized tubercle, 16-23 (16-24) branched, apical branches subequal to lateral.

Hair 14-P 7-13 branched from a short to moderately long stalk. Mesothoracic hair 1-

M 31-35 branched, ovate in outline, apical branches much shorter than lateral. Meta-

thoracic palmate hair 3-T with 10-16 narrow, semitransparent leaflets, moderately

small. Hair 4-T 3-5 branched (2-5), small. Pro-, meso- and metathoracic pleural

group spines moderately developed, prothoracic spine slightly longer than meso- and

metathoracic spines. Abdomen: Hair 0-II-VI 5-8 branched (4-9), moderately long to

long. Palmate hair 1-I with 12-16 (11-16) semitransparent, spreading, narrow, point-

ed leaflets; 1-II-VII with pigmented, long, moderately narrow to moderately broad,

pointed leaflets. Hair 2-II 4-6 branched; 2-V single, long. Hair 5-I 3-5 branched,

moderately small; 5-III 8-11 branched. Hair 9-I 5-7 branched; 9-IV,V 10-13 (6-13)

and 7-13 branched respectively. Hair 11-1 3,4 branched, medium. Hairs 13-1, II, II

5-8, 7-10 and 7-11 branched respectively, small; 13-IV 4-6 branched (4-7), moderate-

ly large; 13-V 4-6 branched, very large, extending beyond caudal margin of segment.

Spiracular Lobe: Pecten with 14-17 teeth; median teeth mostly subequal; number

and placement of long teeth variable; serrations on basal half of teeth from short to

moderately long. Lateral arm of spiracular apparatus short to moderately short.

Hairs 8,9-S 4-7 and 5-7 (4-9) branched respectively, moderately developed. Anal Seg-

ment: Covered with very thin, moderately long spicules becoming longer at apex.

Saddle reddish brown with irregular lateral and apical margins. Hair 1-X long; insert-

ed on saddle near ventral margin, or rarely on ventral margin at base of indentation;

inserted about 0.3 cephalad of caudal margin. Hair 2-X relatively short, about 0.7

length of 3-X. Anal gills longer than saddle.

DISCUSSION. An. strodei can be distinguished from the other species in the Os-

waldoi Group in the female (except for anomalophyllus, and occasionally noroesten-

sis and trinkae) by the combination of (1) palpal segment 4 often with a narrow

brown stripe on ventral surface, (2) light scales on coxae usually white or very light

cream, (3) foretarsal segments 2 and 3 cream to white in apical 0.25 (0.18-0.35) and

0.5 (0.25-0.80) respectively, (4) foretarsal segment 5 usually entirely golden to

brown or with a few light apical scales, rarely with apical 0.5 lighter than basal 0.5,

(5) midtarsal segment 5 usually with a small cream band in less than apical 0.3, (6)

hindtarsal segment 2 brown in basal 0.35-0.45 (0.30-0.50), (7) light wing spots usual-

ly white, occasionally very light cream on vein C, (8) basal dark spot of vein C small,

humeral light spot 2.0-4.0 (1.2-4.1) of basal dark spot, (9) subcostal light spot of vein

C 0.25-0.50 of subcostal dark spot and (10) abdomen with dark caudolateral tuft

scales large and cuneate; in the male genitalia by (1) apex of ventral claspette very

wide, width at apex 0.5-0.6 length of claspette, without setae, more or less truncate,

rugose or deeply striated, strongly expanded laterally into large rounded lobe which

is convex on basal and lateral margins and weakly concave on apical margin, (2) setae

along basal margin of basal lobule long, 2.0-3.5 width of aedeagus, (3) setae on lateral

margins of ventral claspette extending toward apex only to base of apicolateral lobes,

(4) preapical plate moderate, oval to circular, weakly to moderately sclerotized, mod-

erately well-defined and (5) apex of aedeagus without leaflets, slightly longer than

wide; in the pupa by the combination of (1) hair 7-C with one branch conspicuously

longer, 1.5-2.0 length of other branch(es), (2) pinna long, 3.8-4.6 (3.7-4.8) length of

meatus, with basally pointed meatal cleft, (3) hair 10-C single, moderately long,

equal to or slightly shorter than 11-C, (4) hair 12-C 1-4 forked, about 1.75 length of

10-C, (5) hair 5-If 3-5 branched, moderately long, (6) sum of branches of hairs 1-III,

128 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

5-III and O-VI less than 18, (7) hair 6-II 1-3 forked, very long, at least 2.0 length of

7-II and (8) hair 9-III stout, short, less than 2.0 length of 9-II, 9-V 1.8-2.3 (1.7-2.3)

of 9-IV, 9-VII thin, tapering to fine point, very long, 1.5-2.0 of 9-VI and about 0.5

length of segment; and in the larva by the combination of (1) hairs 2,3-C single and

barbed, hairs 2-C closely approximated, clypeal index about 3.0-4.0, (2) collar mod-

erately thin dorsomedially, heavily pigmented, (3) hairs 1,2-P rarely sharing common

tubercle, palmate hair 1-P with 13-17 narrow leaflets, 2-P 16-23 branched (16-24),

(4) hair 1-M 31-35 branched, ovate in outline, apical branches much shorter than la-

teral, (5) hair 13-V very large, extending beyond caudal margin of segment and (6)

hair 1-X inserted on saddle near ventral margin, or rarely on ventral margin at base of

indentation.

An. strodei varies throughout its geographic range. Unfortunately with regard to

the immatures, only material from Colombia, Venezuela, Panama and Brazil was

available for this study. The only individually reared specimens examined came from

Brazil, Colombia and Panama. In general, the immatures from Brazil are much larger

and darker than those from Colombia, Venezuela and Panama. In larvae from Sao

Paulo, Brazil, hair 8-C is 4 or 5 branched, whereas in the Panamanian specimens it is

1-3 branched. Also, hair 9-III-V in the Brazilian specimens has fewer branches (1-6)

than it does in the Panamanian specimens. As in the immatures, the adult females

from the states of Mato Grosso and Sao Paulo, Brazil are much larger and darker than

those from Central America. Although not showing any geographical variation, fore-

tarsal segment 5 in the adults is usually of uniform color, which is unique in the Os-

waldoi Subgroup; only rarely is it 0.5 apically light, 0.5 basally dark. As evident in

the description of the adult female, the relative sizes of the wing spots are highly var-

iable; in one specimen from Panama the subcostal light spot of vein C is longer than

the subcostal dark spot, similar to that of rangeli.

Although it was not possible to examine the eggs of strodei for this study, it

should be mentioned that the eggs are extremely diverse. Five different varieties

(typicus, artigasi, arthuri, albertoi, lloydi) were described by Unti (1940b; 1941)

based on variations in the morphology of the egg. Rozeboom (1938a: 102-103, 106)

recognized 3 different types of eggs designated Types A, B and C from Panama,

about which he stated, “‘it seems unlikely that the different types of strodei eggs

were deposited by separate races, as they were all collected from the same breeding

place at the same times, and there were all kinds of gradations between the extremes.”’

Galvao (1943: 143-144) recognized 3 major types of eggs representing different

forms, including Rozeboom’s types. The great variation in the eggs and in the other

stages has lead to strodei being considered a complex of species similar to the Ano-

pheles (Ano. ) maculipennis Meigen 1818 complex of Europe or the An. (Cel. ) gambi-

ae Giles 1902 complex of Africa. Other important papers written on the eggs of stro-

dei are those of Galvao and Lane 1937b:272-275; Galvao 1938a:56-57, 1940:445;

Galvao and Barretto 1938:113-114; Kumm 1941b:94-97; and Rozeboom 1942a:240-

241.

Besides the diversity in the external morphology, there is considerable variation in

the banding patterns of the salivary chromosomes of strodei. Several workers

(Schreiber and Pompeu Memoria 1956: 1-3; Guedes, Amorim and Schreiber 1957:

247-248; Schreiber and Guedes 1959b, 1960, 1961) have studied the salivary glands

of strodei and have found many heterozygous inversions, asynaptic zones and (rarely)

translocations in the chromosomes. These authors stated they were able to locate 3-

11 “anomalous zones” per nucleus. This is in contrast with the cytogenetics of some

of the other species in the section which have been studied. An. aquasalis and albi-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 129

manus show no chromosomal variation in the populations that have been examined,

and nuneztovari has only one homozygous inversion. Schreiber and Guedes (1961)

hypothesized that the high degree of polymorphism in the chromosomes of strodei

is correlated with the great variation in the morphology of the eggs.

Whether or not strodei represents a complex of species cannot be ascertained with

any confidence with the information presently available to me. Until more material

and more data concerning the bionomics and genetics are available, I am treating

strodei as a single species.

The closest ally to strodei is rondoni. The male genitalia of strodei and rondoni

are almost identical except that in rondoni the setae on the ventral claspette do not

extend toward the apex to base of apicolateral lobes. Differences reported to distin-

guish rondoni from strodei are listed in the discussion of rondoni.

I no longer consider evansi as the valid name of the presently discussed species.

Brethes (1926a) described, in French, evansi from material that was sent to him from

Tucuman, Argentina by N.C. Davis. In another description (1926b) in Spanish, Bre-

thes included a mislabelled diagram of the male genitalia of evansi which is impossi-

ble to identify as any particular species in the Albimanus Section. Later the same

year, Root (1926b) described strodei from the state of Minas Gerais, Brazil, and

clearly illustrated the diagnostic characters of this species in all the life stages except

the egg. The following year, for some reason that is unclear to me, Root (1927:477)

included strodei as a synonym of evansi in a paper on Chagasia fajardi (Lutz 1904).

Davis (1928:549-550), who furnished Brethes with the type material of evansi, refer-

ring to Brethes’ description stated, “‘It is true that some Anophelines showing the

genitalic characteristics of strodei have been taken in northern Argentina... but by

far the greater number are of the type which Root accepts as tarsimaculatus.” Davis

then pointed out the errors Brethes made in labelling his diagram, and said that it is

“barely possible” that Brethes was working with both tarsimaculatus and strodei, and

that drawings of the male genitalia represent a composite. In either case I agree with

Davis that Fig. 2C of Brethes (1926b), which appears to be the spiculose apex of a

ventral claspette, does not look like anything seen in strodei. Also, the ninth sternite

in fig. 2A appears to be too broad to be strodei. Lima (1928:100-102) wrote that

Davis had told him that what Brethes described was not strodei, but that since he

(Davis) had not seen the type he was not sure of the validity of evansi; also Davis

again stated that possibly evansi was the same as tarsimaculatus. Shannon and Del

Ponte (1928:54) synonymized evansi with tarsimaculatus, giving as evidence the fig-

ure of the ventral claspette. According to Bejarano (1957:336), Del Ponte, in his un-

published 1941 thesis on the revision of the Argentine species of Anopheles, stated

that possibly specimens classified by Brethes as evansi could be oswaldoi. Casal (in

Belkin, Schick and Heinemann 1968:10) designated the lectotype male genitalia slide

of evansi which had the following Brethes label, ‘‘Cellia evansi Brethes=tarsimaculata

auct. (nec Goeld), Tucuman, N. C. Davis ded., III.1926, Pr. micr. An. M.,”’ in Del

Ponte’s hand “384,” “DP14,” ““DP14=?noroestensis G. y L., 1938 [signature in ink],

Tipo de evansi.’’ From this label it is apparent that Del Ponte looked at the male

genitalia of evansi some time after 1937 and tentatively synonymized noroestensis

with evansi, in which case evansi would become the valid name for noroestensis. Ga-

baldon (1949:765), writing about the vector status of West Indian and South Ameri-

can anophelines, also considered noroestensis and clarki as synonyms of evansi. How-

ever, not having examined the type specimen of evansi, I am not proposing at this

time any nomenclatorial changes regarding the biological species noroestensis. I am

only considering strodei as the valid name of the species described here, since it has

130 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

been extensively used in the past in referring to this taxon, and because there is sub-

stantial evidence that the lectotype of evansi does not represent this species. For the

present, I am considering evansi as anomen dubium.

As in the adult females of some of the other species in the Albimanus Section, mu-

tants or anomalous specimens of strodei have been described with extra black rings

on hindtarsal segments 3, 4 and 5, segments 3 and 4, and segments 4 and 5 (Rachou

and Ferraz 1951:542-543). |

BIONOMICS. The immatures of strodei occur predominantly in ground pools. In

addition, it has been collected from animal tracks, ponds, lakes, swamps, stream mar-

gins, potholes, marshy depressions, ditches, seepage areas and rock holes. It is found

only in fresh water, usually in the full sun or partial shade but occasionally in deep

shade. It occurs over a wide range of elevations, from near but usually not on the

coast to altitudes of 1600 m (Unti 1941). It is commonly found in mountainous

areas, plains and plateaus in the interior. The immatures usually are associated with

abundant vegetation such as grasses, algae and Utricularia sp. Galvao (1940:443-447)

stated that the immatures can tolerate large fluctuations in temperature (17.5-33.0°

C), pH (5.9-7.2) and amounts of organic material. Unti (1941) has reported collec-

ting the larvae in water as cold as 2°C. He also found strodei in water poor or rich in

organic material, varying in nitrate concentration from 0.008% to 0.064%, and in wa-

ter with sometimes high concentrations of bicarbonates or sulfates. On one occasion,

he discovered immatures in water with a pH of 4.0 and a high sulfate residue from a

nearby paper plant. The larvae have been collected with 3 other species of Nysso-

rhynchus, allopha, argyritarsis and rangeli. They have been collected with 32 other

species of mosquitoes: Aedes (Och.) crinifer, Ae. (Och.) dupreei group sp B of Heine-

mann and Belkin 1979:79, Ae. (Och.) sp, Anopheles (Ano.) ?apicimacula, An. (Ano.)

eiseni, An. (Ano.) neomaculipalpis, An. (Ano.) sp, Culex (Cux.) coronator group, Cx.

(Cux.) declarator group, Cx. (Cux.) mollis, Cx. (Cux.) nigripalpus, Cx. (Cux.) thriam-

bus, Cx. (Cux.) spp 72 and 85 of Heinemann and Belkin 1978a:184, Cx. (Mel.) bahi-

ense, Cx. (Mel.) bastagarius, Cx. (Mel.) chrysonotum, Cx. (Mel.) conspirator, Cx.

(Mel.) dunni, Cx. (Mel.) dureti, Cx. (Mel.) educator, Cx. (Mel.) ?egcymon, Cx. (Mel.)

erraticus, Cx. (Mel.) ?inadmirabilis, Cx. (Mel.) ?iolambdis, Cx. (Mel.) pilosus, Cx.

(Mel.) sp, Psorophora (Gra.) cingulata group, Uranotaenia ( Ura.) coatzacoalcos, Ur.

(Ura. ) lowii, Ur. (Ura.) pulcherrima, Ur. (Ura.) sp.

The adult females of strodei are exophilic, feeding predominantly on mammals

other than humans. They only rarely enter houses to feed; in Brazil, Deane, Causey

and Deane (1948:905) collected 1895 adults of strodei of which only one was from

inside a house. In host preference studies using a human and a horse as bait in Alaga-

dico in 1942, Deane, Causey and Deane (1948:906) collected 17 females on the hu-

man and 23 on the horse during the crepuscular hours. Kumm, Komp and Ruiz

(1940:388) in Costa Rica and Rozeboom (1938b:290, 293) in Panama occasionally

found strodei inside houses, but usually strodei showed a preference for animals and

fed outside. The only exception was reported by Correa (1938), who found a very

high density of strodei at the Fazenda Santa Alice in the state of Sao Paulo, Brazil,

and collected 165 females from inside a house which represented 95.3% of all ano-

phelines captured. In Panama (Curry 1932:566; Rozeboom 1938b:290) and Colom-

bia (Renjifo and de Zulueta 1952:601; Bates and de Zulueta 1949: 135-137), the

peak abundance of strodei is during the early part of the dry season. In Santa Catari-

na, Brazil, strodei occurs in the lowest numbers during the cold months. The adults

are usually most active around dusk, although they are reported to feed throughout

the night (Deane, Causey and Deane 1948:904-906; Barretto 1940: 160).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 131

MEDICAL IMPORTANCE. An. strodei does not seem to be an important vector

of malaria. As mentioned earlier (p 18), it has been incriminated only once as a vec-

tor of malaria, and that was by Correa (1938) at the Fazenda Santa Alice, Sao Paulo,

Brazil. In that study Correa dissected 163 females collected inside houses and found

2 of them (1.2%) naturally infected with oocysts. Galvao and Lane (1937d), Galvao

(1938c), and Fonseca and Unti(1943) have easily and successfully experimentally in-

fected strodei with Plasmodium vivax. Because of its predominantly exophilic and

zoophilic habits, strodei can be considered a possible health problem only when it

occurs at very high densities.

DISTRIBUTION (fig. 1). An. strodei is widely distributed throughout Central

America and much of South America east of the Andes. Its northernmost limit is re-

ported to be the state of Veracruz, Mexico on the Gulf of Mexico; on the Pacific side

it is reported from as far north as the Costa Rica-Nicaragua border. In South Ameri-

ca, it is found in northern and eastern Colombia, Venezuela, the Guianas, Brazil, Bo-

livia, Paraguay and northern Argentina. An. strodei is found on the eastern slope of

the Andes in Colombia, Bolivia and northern Argentina; it is presumably also found

in eastern Ecuador and Peru, although there are no records. It is not known if it oc-

curs on the Pacific slope of the Andes, although it is reported to occur in the Magda-

lena River drainage in northern Colombia. The southernmost limit east of the Andes

is in the province of Buenos Aires, Argentina. An. strodei does not occur on any of

the Caribbean islands, including Trinidad and Tobago.

Material Examined: 566 specimens: 63 males, 25 male genitalia, 146 females, 41

pupae, 291 larvae; 41 individual rearings: 21 larval, 12 pupal, 8 incomplete.

ARGENTINA (1M, 1F). Tucuman: Locality not specified, 23 Aug [21941], Alvarado, 1F; 28

Aug 1941, 1M.

BOLIVIA (6F). Santa Cruz: Loma Alta, 19 May 1944, 3092, KO 111-22, 4F. San Antonio del

Parapeti, 9 May 1944, 3151, KO 111-26, 2F.

BRAZIL. Amazonas (6F): Manaus, Jun 1931, R. Shannon, 6F. Bahia (3F,7L): Bonfim, 26

Jan 1930, N. Davis, R. Shannon, 3F. Prado, 1F [JH]. Salvador da Bahia, 1972, J. Fowler, BRS

277, 3L; same data, BRS 278, 4L. Minas Gerais (6 specimens: 2M, 1Mgen, 2F, 1L): Agua Limpa,

near Juiz de Fora, 27 Mar 1925, [F. Root] , 1M (64) and 1Mgen (750923-29, lectotype of strodei),

1F (64, paratype of strodei); same data, 1M (64), 1L (paratypes) [JH]. Lassance, 12 May 1925,

[F. Root], 1F (81) [JH]. Para (14 specimens: 1M, 7F, 3p, 3 1;3 larval ind rear): Maraba area, 7

Jun 1976, 2F (M-1); same locality, 18 Jun 1976, 1F (M-4); same locality, 21 Jun 1976, 1F (M-9);

same locality, 25 Jun 1976, 1F (M-13). Palestina, 100 km SW of Maraba, 12 Feb 1976, Moramay,

1 IpM (1001); same data except 14 Feb, 2 IpF (1002). Rondonia (1F): Guajara Mirim, May 1931,

R. Shannon, 1F. Sao Paulo (80 specimens: 14M, 10Mgen, 14F, 15P&p, 27L&l; 15 ind rear: 121,

2p, linc): Boraceia Field Station, BRA 114, 150, 151, 157. Itapetininga, 28 km W of, BRA 166,

167. Itapira, 1Mgen (No. 3, SPM-88, 89). Juquia, J. Lane, 1F (1.113). Porto Ferreira, 1M, 1Mgen.

Santos, 1F. Sao Paulo, Butantan, 1939, [Worontzow], 1M (191-3), 1M (191-9) [JH] ; same locali-

ty, 707, 3M, 3F [JH]. Sao Paulo de Piracicaba, A. Ayroza, 1Mgen (No. 4, SPM 88). Sao Paulo,

Pinheiros, 1937, A. Galvao, 5M; same locality and collector, 1940, 2M. Sao Pedro, 1F (No. 4). Lo-

cality not specified, 1Mgen (No. 5, SPM 88, 89); 1943, 1Mgen (819-27); Galvao, 1 1 (14-18) [JH].

State and locality not specified (2M, 1Mgen): J. Lane, 1M (851), 1M (854), 1Mgen (740515-33).

COLOMBIA (10 specimens: 2F, 1p, 7L&l; 1 inc ind rear). Meta: Villavicencio, Mar 1946, 2L

(425) [JH]; same locality, 6 May 1947, L. Rozeboom, CV-P 26, 1F; same locality, 9 May 1947, 1

lp, 4L [JH] ; same locality, 1947, L. Rozeboom, CV-P 32, 1F.

COSTA RICA (3F). Province not specified: Barranca, H. Kumm, 1F (84) [BM1938-696]. No

data, 2F.

PANAMA AND CANAL ZONE (402 specimens: 43M, 8Mgen, 93F, 19P&p, 239L&l; 19 ind

rear: 51, 10p, 4inc). Bocas del Toro: Punta de Pena, PA 238. Canal Zone: Chiva Chiva, 15 Aug

1944, Wood, ASM 102, 1L; same locality, 18, 20 Mar 1947, 2M, 1F; same locality, 20 Mar 1947, 2

F, [Ft.] Clayton and Corozal Army Reservations, Feb 1943, 1M, 2F. Ft. Clayton, 24 Apr 1944,

ise Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

P. Adams, 2L. Stilsons Lake [Pond] , 21 Oct 1924, D. Curry, 1Mgen. Locality not specified, Feb

1933, D. Curry, 3M [BM1933-504]. Chiriqui: El Hato del Volcan, PA 636, 637. Palo Santo, PA

646. Panama: Arraijan, 6 Feb 1940, PAX 166, 7M, 11F. Bejuco, PA 542. Juan Diaz, 4 Jan 1935,

3F; same locality and date, L. Rozeboom, 1L [JH]; same locality, 2 Dec 1936, 1F; same locality

and date, L. Rozeboom, 2L [JH] ; same locality, 7 Dec 1936, 6M, 12F; same locality and date, L.

Rozeboom, 12L [JH] ; same locality, 16 Dec 1936, 6M, 17F, 8L; same locality, 16 Dec 1936, L.

Rozeboom, 3M, 11F; same data, 1L [JH] ; same locality and collector, 29 Dec 1936, 2L [JH];

same locality, 1936, L. Rozeboom, 2M, 5F; same locality, PA 750, 838. La Chorrera, 31 May

1944, ASM 29, 7L; same locality, 4 Jul 1944, ASM 36, 1F; same locality, 26 Sep 1944, ASM 180,

2L; same locality, 6 Nov 1944, ASM 263, 1F, 1L;same locality, 6 Nov 1944, ASM 266, 4L; same

locality, 27 Nov 1944, Wood, Griffing, ASM 297, 1L; same locality and collectors, 14 Dec 1944,

ASM 337, 1F; same locality, 7 Dec 1944, K. Frick, R. Arnett, ASM 333, 1L; same locality, 13 Feb

1945, Van Doran, Turney, ASM 409, 1F; same locality, 12 Mar 1945, ASM 437, 1M, 1Mgen; same

locality, 12 Mar 1945, ASM 445, 1F. Pacora, PA 761. Tocumen and Tapia, 5M, 15F. Tocumen,

PA 548. Province not specified (1 Mgen, 181): 6 Nov 1935, L. Rozeboom, 1Mgen; 1936, L. Roze-

boom, 181 [JH].

PARAGUAY (3F). Alto Parana: Locality not specified, “‘Calle 12,” 22 Aug 1975, 3F.

VENEZUELA (20 specimens: 5Mgen, 5F, 3p, 7L&1;3 ind rear: 11, 2inc). Aragua: Maracay, 10

May 1927, F. Root, 1F [BM1929-194] ; same locality, 13 May 1927, 1 lp (10a) [JH] ; same local-

ity, 26 May 1927, 1L [JH] ; same locality, 10 Jun 1927, 1 lp, 1Mgen (63e) [JH]; same locality, 1M

gen (H36) [JH]; same locality, Jun 1929, 2Mgen. Turmero, VZ 337. State not specified: Guano-

co River, L. Wehekind, 1F [BM1930-69]. Locality not specified, 1929, 1Mgen (6) [JH] ; no data,

2F.

12. Anopheles (Nys.) rondoni (Neiva & Pinto)

Figs. 1,4, 7, 12

1922. Cellia rondoni Neiva and Pinto 1922a:322. TYPE: Lectotype large wing mounted on

slide (403) together with wing of type of cuyabensis, marked Matto Grosso, C. Pinto, re-

mainder of adult apparently lost, type locality restricted to Ladario, on Rio Paraguai (Ma-

to Grosso), Brazil [IOC; designation of lectotype and restriction of type locality by Belkin,

Schick and Heinemann 1971:6].

Anopheles (Nyssorhynchus) rondoni of Davis (1928:539, 541, 543, 545, 546, 554-561; 1933:278-

279); Dyar (1928:436); Shannon and Del Ponte (1928:42, 46, 57, 60); Edwards (1932:45);

Root (1932:779); Galvao and Lane (1937e:214); Galvao and Amaral (1938:13); Lane (1939:

26-27; 1944:264, 265; 1949:403-404; 1953:273); Pinto (1939:345); Galvao (1940:447-448;

1943:143, 149, 150); Vargas (1940a:199); Fonseca and Fonseca (1942:95); Cerqueira (1943:

19); Correa and Ramos (1943:135-137; 1944b:132); Deane, L. M., Causey and Deane (1946:7,

9, 12; 1948:906); Deane, M. P., Causey and Deane (1946:42, 45); Senevet (1948a:278); Leeson

and Buxton (1949:25 1-253); Levi-Castillo (1949:9, 10, 11, 16, 28, 58, 67, 76, 82, 86); Duret

(1950a:372; 1950b:302; 1952:347); Galvao (1950:44); Rachou and Ferraz (1951:542, 547-

553); Rachou and Ricciardi (1951 :424-426, 432-437); Gabaldon and Cova Garcia (1952:178,

196, 201); Horsfall (1955:124, 170, 171, 182); Bejarano (1956:9, 10; 1957:326, 328, 331, 336-

337; 1959:316, 325); Rozeboom and Kitzmiller (1958:244); Castro, Garcia and Bressanello

(1959:549); Stone, Knight and Starcke (1959:34); Vargas (1959:370); Cerqueira (1961:127);

Forattini (1962:421, 427); Rozeboom (1963:112); Belkin, Schick and Heinemann (1971:6);

Consolim and Galvao (1973:177, 178, 181); Cova Garcia, Pulido F., Amarista M. (1978:157).

Anopheles rondoni of Muehlens, Petrocchi and Zuccarini (1925:251,253,254, 257,264,265); Davis

and Shannon (1928:448-456); Pinto (1930:154); Russell, Rozeboom and Stone (1943:37, 41,

49); Foote and Cook (1959:16, 26, 29, 32); Maciel (1962:477); Mattos and Xavier (1965:273);

Gorham, Stojanovich and Scott (1967:19, 43, 60; 1973:115, 136, 138, 150).

Cellia rondoni of Neiva and Pinto (1922b:356-357); Bonne (1924:133, 134); Brethes (1926b:313).

Nyssorhynchus (Nyssorhynchus) rondoni of Lima (1928:102).

Anopheles (Anopheles) rondoni of Bonne and Bonne-Wepster (1925:522).

Anopheles (Nyssorhynchus) tarsimaculatus var. rondoni of Christophers (1924:40, 90).

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 133

Cellia tarsimaculata in part of Bonne (1924:133-137).

Cellia albimana in part (?) of Neiva (1909:69-77).

FEMALE (fig. 7). Wing: 3.6 mm. Proboscis: 2.05 mm. Palpus: 2.1 mm. Forefe-

mur: 1.6 mm. Abdomen: about 3.0 mm. Head: Integument dark brown. Interocu-

lar space 3,4 ommatidial diameters. Proboscis about 1.3 length of forefemur. Scales

light on vertex extending caudally onto median occiput, slightly darker on occiput,

dark brown laterad on postgenae. Erect scales on vertex similar to those of oswaldoi.

Palpal segments 1, 2, 3 and 5 as in group description; segment 4 with a basal and an

apical brown band, ventral surface with a row of brown scales, mediolateral scales

white usually with dark scales interspersed. Antenna: Flagellar segment 1 with nu-

merous, unusually long (longer than segment 2), narrow, white, oblanceolate to seti-

form scales along dorsomesal surface and a few, moderately long, white, lanceolate

scales on apicolateral surface. Flagellar segments 2-13 each with basal whorl of 8-10

long, silver setae. Thorax: Integument dark brown to black, extensively pruinose.

Anterior promontory scales extending a short distance dorsad onto acrostichal line.

Prescutellar space large, subtriangular, very dark brown to black. Bare space at cau-

dal end of posterior fossa small, distinct and rectangular. Upper stp with 2-4 long se-

tae, and a large patch of about 15 cream, obovate scales. Lower stp with 1-3 setae,

and 8-10 scales similar to those on upper stp, scales more numerous and extensive

than those of other species in group. Upper mep with about 5 long setae; scales ap-

parently absent. Legs: Light scales on coxae white to pale cream. Foretarsal seg-

ments 2 and 3 with a white band in apical 0.2-0.3 and 0.3-0.5 respectively; segment

4 all dark. Fore- and midtarsal segment 5 golden in apical 0.33-0.50. Midtarsal seg-

ments 2 and 3 with a small, cream to white, apical band. Hindtarsal segment | with

a distinct, white, apical ring; segment 2 with a brown band in basal 0.65-0.85; seg-

ment 3 with brown band in basal 0.20-0.35; segment 5 white in about apical 0.5.

Wing: Light wing spots cream. Vein C predominantly dark; basal dark spot small,

less than 0.3-0.5 of humeral light spot; subbasal, presectoral and sectoral dark spots

fused into a very long, dark spot, occasionally with a few, interspersed, light scales;

preapical light spot moderately small, 0.2-0.3 of preapical dark spot; apical dark spot

relatively large. R presectoral dark spot moderately long, longer than sectoral dark

spot. Rs-R»,3 subcostal dark spot moderately large, divided or not divided by a small

light spot. R, preapical dark spot large, about 0.5 length of vein. M sectoral dark

spot distinct, small to moderately long; preapical dark spot of vein M extending un-

broken onto M,,,, rarely with a few light scales; base of M,,. dark at furcation. Re-

mainder of dark spots small to moderate. Apical light fringe spot small, with or with-

out dividing dark scales; remainder of light fringe spots as in section description but

small and usually inconspicuous. Abdomen: Tergite I with a few, long, lanceolate to

setiform, cream scales along caudal border. Sternite I with a few setae. Light scales

on tergites IJ-VII extensive, long and narrow. Caudolateral and ventral scale tufts

with moderately long, broad scales.

MALE. Wing: 3.3 mm. Proboscis: 2.9 mm. Forefemur: 1.85 mm. Abdomen:

about 2.8mm. Essentially as in female except for sexual characters. Description

based primarily on 2 specimens from Jujuy, Argentina. Head: Palpal segments 2 and

3 with a light dorsal stripe; segment 4 predominantly white, occasionally with a

small, dark, apical band, ventrally dark scaled. Antenna: About 0.75 length of pro-

boscis. Flagellar segment 1 with long oblanceolate scales on dorsomesal margin.

Legs: Forefemur about 0.55 length of proboscis. Spiniform setae of basal plantar

surface of foretarsal segment 5 moderately long. Claw on foreleg long, acuminate;

submedian tooth tapered to a slightly recurved point; basal tooth not visible on any

134 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

specimens examined.

MALE GENITALIA (fig. 12). Segment VIII: As that of strodei. Segment IX:

Sternite moderately long, subtrapezoidal. Anterior apodeme a moderately short

band. Sidepiece: Tergal surface with 4,5 long, submedian tergomedial bristles; bris-

tles mesad of tergomedial bristles moderately long. Parabasal spine and its tubercle

long, apex of spine extending to or beyond lateral margin of aedeagus on other side;

tubercle about 0.5 length of spine. Basal apodeme moderately long, about 0.15-0.20

length of sidepiece. Longer, more dorsal accessory spine at least 0.5 length of side-

piece; more ventral spine about 0.75 of longer spine. Internal spine subequal to

shorter accessory spine, apically retrorse. Clasper: As that of strodei. Dorsal Clasp-

ette: About 0.5 length of sidepiece. Pedicel moderately narrow; base curved mesad.

Leaflets broad, subequal in length to pedicel; dorsal leaflet with moderately devel-

oped basomesal projection. Ventral Claspette: Similar to that of strodei. Large,

slightly less than 0.5 length of sidepiece; apex broad, width at apex about 0.7 length

of claspette. Basal lobule as that of strodei, with setae along basal margin long, about

2.0 width of aedeagus. Apex more or less truncate and rugose, with a small, U

shaped median sulcus. Apex strongly expanded laterally, as that of strodei, into

large, rounded lobe; apicolateral lobe appears slightly heavier than that of strodei.

Setae usually on basal lobule only, not extending toward apex to base of apicolateral

lobe. Preapical plate moderate, circular to oval, very weakly sclerotized and ill-de-

fined, located about 0.25 length of claspette basad of apex. Refringent structure in-

conspicuous, in shape of inverted horseshoe. Phallosome: As that of strodei.

PUPA, LARVA. No specimens available.

DISCUSSION. This species can be distinguished from other species in the Oswal-

doi Subgroup in the female by (1) palpal segment 4 with a ventral row of brown

scales, (2) prescutellar space large, subtriangular, very dark brown to black, (3) fore-

tarsal sement 3 light in apical 0.3-0.5, (4) hindtarsal segment 2 with a large dark band

in basal 0.65-0.85, (5) hindtarsal segment 3 with a dark band in basal 0.20-0.35, (6)

vein C predominantly dark, subbasal, presectoral and sectoral dark spots fused into a

very long, dark spot, occasionally with a few, interspersed, light scales, and apical

dark spot relatively large, (7) vein R,with large preapical dark spot and (8) the preapi-

cal dark spot of vein M extending uninterrupted onto M,,,; and in the male genitalia

by the combination of (1) ventral claspette similar to strodei, except that setae usu-

ally on basal lobule only, not extending toward apex to base of apicolateral lobe, (2)

apex of ventral claspette strongly expanded laterally into large rounded apicolateral

lobe which appears slightly heavier than in strodei and (3) preapical plate moderate,

circular to oval, very weakly sclerotized and ill-defined.

An. rondoni is the sister species of strodei. The male genitalia are almost identical

with the exception of the characters mentioned above. Although I did not have the

opportunity to study the larva of rondoni, several authors (Davis 1933:278-279; Cor-

rea and Ramos 1943: Galvao 1940:444-445) report that it is similar to that of stro-

dei, differing only in that rondoni has (1) fewer branches on hair 1-P (11-14), (2)

fewer branches on the palmate hairs (20-24) and (3) fewer teeth on the pecten.

Rozeboom (1963:112) believed that rondoni was a mutant melanic form of stro-

dei, just as bisignatus and trisignatus are melanic mutants of albimanus, cuyabensis is

of triannulatus, and guarauno, delta and deltaorinoquensis are of aquasalis. Davis

(1933) stated that rondoni could not be a variety of tarsimaculatus or strodei be-

cause it ‘““breeds true’? when isolated. However, Rozeboom (1963:112) demonstrated

that bisignatus (a mutant of albimanus) also breeds true when isolated from other

“normal’’ individuals of albimanus. 1 do not discount the possibility of Rozeboom’s

Faran: Albimanus Section of Anopheles (Nyssorhynchus ) 135

hypothesis being correct; however, at present I do not believe that rondoni is a mu-

tant form of strodei. An. rondoni has been collected in considerable numbers (Davis

and Shannon 1928) in numerous different localities. It has a fairly large distribu-

tion, occurring at times sympatrically with strodei. Besides the extra hindtarsal dark

band and large prescutellar space, the wing spots of rondoni are very different from

strodei and are unique within the Oswaldoi Subgroup. Furthermore, mutants of

strodei that exhibit extra hindtarsal bands do not possess the other characters which

distinguish rondoni. Whether or not hybridization occurs between strodei and ron-

doni has not been investigated. Until more is known about the biology of rondoni I

am considering it a separate species, albeit with a high degree of affinity toward stro-

dei. An. rondoni may be a relatively new species recently diverging from strodei.

From its present distribution it would seem that rondoni evolved in the region of

northern Argentina, Bolivia and southern Brazil.

BIONOMICS. Most of the discussion presented is from the work of Davis and

Shannon (1928) on the bionomics of rondoni in and around the city of Ledesma, in

the villages of Canitas and Calilegua, Jujuy, Argentina from March 1926 until May

1927),

Davis and Shannon first discovered the immatures of rondoni along the margin of

a reservoir in Ledesma. They soon found that a shallow ditch adjacent to a reedy

swamp behind the reservoir was a highly favored aquatic habitat of rondoni. From

that ditch 100 larvae were collected from which 75 adults emerged: 72 were rondoni,

2 were An. (Ano.) pseudopunctipennis and one was ‘“‘tarsimaculatus.’’ Later Davis

and Shannon encountered the immatures of rondoni in clear water such as in ditches,

drying pools at the edge of swamps, a puddle in a road and in heavy growths of grass

in a flooded meadow; the aquatic habitats were sometimes with considerable detri-

tus, but not with algae. The type specimens were collected on the right margin of

the Paraguay River, and later material was collected on the left margin of the S. Lou-

renco River. An. rondoni is most abundant in the province of Jujuy, Argentina and

in Rincao, Sao Paulo, Brazil in March, during the late summer. In Jujuy, it is present

all year, although occasionally in low numbers.

The adults are usually active early in the evening. On one occasion 120 females

were collected on a horse between 1730 and 1800 h, flight commencing while it was

still daylight (Davis and Shannon 1928). The females are commonly found inside

houses (Davis and Shannon 1928; Pinto 1939:402, 404; Shannon and Del Ponte

1927:60). Of 1266 anophelines collected in houses in the area of Ledesma, 118 or

9.3% were rondoni. Davis and Shannon concluded that rondoni is not as domestic as

pseudopunctipennis, “but still is found in houses in appreciable numbers.”

MEDICAL IMPORTANCE. An. rondoni has never been incriminated as a vector

of malaria. It has never been found naturally infected with Plasmodium sp. During

the malaria season in Jujuy, Davis and Shannon (1928) dissected 88 individuals and

all were negative. In 3 different experiments, Davis and Shannon unsuccessfully at-

tempted to infect rondoni with Plasmodium falciparum, P. vivax and P. malariae.

Shannon and Del Ponte (1927:60) reported that Davis was able to infect rondoni ex-

perimentally; however, I have not found any other reference to that work.

DISTRIBUTION (fig. 1). An. rondoni occurs in southern and southeastern South

America. In Brazil it has been reported from the states of Acre, Goias, Mato Grosso,

Parana, Sao Paulo, Santa Catarina and Rio Grande do Sul. In Bolivia it is known in

the departments of Santa Cruz, Beni and Tarija (Gabaldon and Cova Garcia 1952:

196; Cerqueira 1943:19). The southern distributional limit is in northern Argentina

where it has been reported in the provinces of Chaco, Corrientes, Formosa, Misiones,

136 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Salta, Santa Fe and Tucuman (Garcia and Ronderos 1962:161).

Material Examined: 69 specimens: 12 males, 10 male genitalia, 47 females.

ARGENTINA (62 specimens: 12M, 7Mgen, 43F). Jujuy: Calilegua, 5 May 1926, R. Shannon,

24F. Ledesma, 23 Mar 1926, R. Shannon, lot 18, 4Mgen (5, 6, 7, 8); same locality, 1926, N. Davis,

R. Shannon, 3F (Bac Ent nota 18); same locality, 30 Mar 1927, R. Shannon, 1Mgen; same locality,

31 Mar 1927, E. Del Ponte, R. Shannon, 1F; same locality, Mar 1927, R. Shannon, 1M (187), 1M

gen (37); same locality, Mar 1927, N. Davis, 9M, 7F [JH] ; same locality and collector, 1927, 1M;

same locality and collector, 21 Mar, 1M; same locality and collector, 1Mgen. Locality not speci-

fied, Nov [?1938], W. Paterson, 2F [BM]. Salta: Oran, 15 Mar 1961, R. Salta, 2F [AMNH]. Pro-

vince and locality not specified: No data, 4F.

BOLIVIA (3F). Beni: Guayaramerin, 15 Apr 1947,S. Blatmas, 1F; same locality, 5 Jul 1947,

LF (47-12258). San Ignacio, 17 Sep 1943, Torres Munoz, 3053B, [KO] 116-9, 1F.

PARAGUAY (1F). Olimpo: P[uerto] Esperanza, Rio Paraguay, 30 Dec 1919, U. Cornell Ex-

pedition, 1F [CU].

COUNTRY NOT SPECIFIED (3Mgen). Locality not specified, 3Mgen [JH].

13. Anopheles (Nys.) benarrochi Gabaldon, Cova Garcia & Lopez

: Figs, |) 47, 29, 30

1941. Anopheles (Nyssorhynchus) benarrochi Gabaldon, Cova Garcia and Lopez 1941:3-22.

TYPE: Syntypes male, female, pupa, larva and egg, La Ceiba (Trujillo), Venezuela

[MDM].

Anopheles (Nyssorhynchus) benarrochi of Galvao (1943:146); Vargas (1943:59; 1959:377, 383);

Causey, Deane and Deane (1944:2, 5; 1946:26); Lane (1944:263; 1949:402; 1953:277-278);

Deane, L. M., Causey and Deane (1946:6, 8, 9, 12; 1948:900-901); Deane, M. P., Causey and

Deane (1946:42, 45); Gabaldon and Cova Garcia (1946b:91; 1952:178, 189); Coher (1948:88-

89); Senevet (1948a:278; 1948c:436); Bates and de Zulueta (1949:135); Levi-Castillo (1949:10,

15, 28, 58, 67, 72, 76, 82, 86); Stone, Knight and Starcke (1959:32); Cerqueira (1961:123);

Forattini (1962:418); Belkin, Schick and Heinemann (1965:72); Morales-Ayala (1971:139); Co-

va Garcia and Sutil O. (1976:32; 1977:23, 44, 64, 89).

Anopheles benarrochi of Russell, Rozeboom and Stone (1943:37, 41, 44); Renjifo and de Zulueta

(1952:600-601); Foote and Cook (1959:24, 26, 29); Cova Garcia (1961:41-42, 68, 93-94, 114,

130-131, 155); Mattos and Xavier (1965:271); Stojanovich, Gorham and Scott (1966b: 20, 31,

40); Gorham, Stojanovich and Scott (1967:16, 47, 51; 1973:113, 139, 142); Elliott (1972:

757); Cova Garcia and Sutil O. (1975a:20; 1975b:213).

FEMALE (fig. 7). Wing: 3.3 mm. Proboscis: 2.05 mm. Palpus: 2.1mm. Forefe-

mur: 1.4mm. Abdomen: 2.7 mm. Very similar to that of aquasalis. Head: Integu-

ment light brown to reddish brown. Interocular space about 3,4 ommatidial diame-

ters. Proboscis about 1.45 length of forefemur. Palpal segments 1-3 as in group de-

scription; white apical band of segment 3 moderately large; segment 4 with a basal

and an apical brown band of subequal size, mediolaterally white, usually with single

row of brown scales on ventral surface; segment 5 with a small, brown, basal ring ex-

tending from apex of 4. Antenna: Flagellar segment 1 with numerous white scales;

distally with elongate oblanceolate scales on dorsal and dorsomesal surfaces, scales

shorter on dorsolateral surface; basally with moderately short, obovate or cuneate

scales forming an incomplete band on dorsomesal, dorsal and lateral surfaces. Flagel-

lar segments 2-13 each with basal whorl of 8-10 tan to silver setae. Thorax: Integu-

ment light brown to reddish brown. Anterior promontory scales extending short dis-

tance onto acrostichal line. Humeral scale tuft dark brown below, usually not exten-

sive. Prescutellar space small, horseshoe shaped. Upper stp with 3,4 long setae, and

a row of moderately small, cream, obovate scales. Lower stp with one long seta, and

a small diagonal patch of scales. Upper mep with about 3,4 long setae; scales absent.

Legs: Light scales on coxae usually white. Midfemur with both anteroapical spot

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 137

and knee spot moderate size, cream or white and conspicuous. Foretarsal segments

2 and 3 with a white band in apical 0.28 (0.2-0.4) and 0.65 (0.5-0.8) respectively;

segment 4 occasionally with a light apical band; segment 5 with a cream to golden |

band in apical 0.25-0.50. Midtarsal segment 2 with a moderately long, cream to

white band in apical 0.15-0.20; segment 3 with a small, cream, apical band; segment

4 all dark or with a few, golden, apical scales; segment 5 cream in about apical 0.5.

Hindtarsal segment 2 with a brown band in about basal 0.40-0.55 (0.36-0.55). Wing:

Light wing spots cream on anterior veins, white on posterior veins. Often dark wing

spots on more posterior veins indistinct. Vein C humeral light spot about 2.5 or

more of basal dark spot; subcostal light spot 0.4 (0.3-0.5) of subcostal dark spot; pre-

apical light spot 0.35 (0.30-0.45) of preapical dark spot; apical dark spot small. R

presectoral and sectoral dark spots moderately small. R3 dark spots often light

brown and inconspicuous. M mostly white, sectoral dark spot present or absent;

with a small preapical dark spot not extending onto M,,.. Mj. apical dark spot, Cu

subcostal dark spot and A subbasal dark spot often indistinct. Apical light fringe

spot large, unbroken or broken by a few, small, dark scales; remainder of light fringe

spots as in section description, except for 2 large, fairly conspicuous spots between

base of wing and vein A. Abdomen: Tergite I with a few, cream, obovate scales me-

dially on caudal border. Sternite I with a few scattered setae, no scales visible. Re-

mainder of abdomen as in section description.

MALE. Wing: 3.4 mm. Proboscis: 2.7 mm. Forefemur: 1.6 mm. Abdomen:

about 2.9mm. Essentially as in female except for sexual characters. Head: Palpal

segments 2 and 3 with a light dorsal stripe; segment 4 almost all cream to white, with

or without a small, dark, basal band, and occasionally with a dark apical band, ventral

surface cream to white with a few, scattered, dark scales. Antenna: About 0.75

length of proboscis. Flagellar segment 1 with long, thin, white, oblanceolate scales.

Legs: Forefemur about 0.6 length of proboscis. Foretarsal segment 5 with about 7

short to moderately long, spiniform setae on basal plantar surface. Claw on foreleg

large, curved and acuminate; submedian tooth thin, slightly recurved at apex; basal

tooth short, strongly decurved.

MALE GENITALIA (fig. 29). Segment VIII: Sternite and tergite with moderate-

ly long, light, obovate scales. Segment IX: Sternite long, about 0.2 length of side-

piece, subtrapezoidal. Anterior apodeme a short band across anterior margin. Side-

piece: Tergal surface usually with 4,5 long, submedian tergomedial bristles and 1,2

long subapicolateral bristles; bristles mesad of tergomedial bristles moderately short.

Combined length of parabasal spine and its tubercle long, apex of spine extending be-

yond lateral margin of aedeagus on other side and occasionally to tubercle of oppo-

site parabasal spine; tubercle about 0.5 length of spine. Basal apodeme moderately

long, about 0.15 length of sidepiece and thick. Longer, more dorsal accessory spine

about 0.5 length of sidepiece; more ventral spine 0.8-0.9 of longer spine. Internal

spine subequal to shorter accessory spine. Clasper: Spiniform fusiform, occasionally

hooked apically, moderate, subequal in length to seta b. Dorsal Claspette: Slightly

longer than 0.5 length of sidepiece. Leaflets broad and long, about 0.5 length of clas-

pette; dorsal leaflet with a large basomesal projection. Ventral Claspette: Small,

about 0.33 length of sidepiece; apex wide, width at apex about 0.5 length of clas-

pette. Basal lobule small, narrow, curving mesad; setae along basal margin short,

about equal to or slightly longer than width of aedeagus. Apex moderately expanded

laterally, apicolateral margin sharply angled and moderately pointed. Depending on

angle of observation, apical margin either appearing truncate or with a variously de-

veloped median sulcus; apical margin unsclerotized and distinctly rugose. Setae on

138 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

lateral margins extending toward apex to or nearly to base of apicolateral lobes. Pre-

apical plate small, circular, heavily sclerotized. Refringent structure rooflike, with

arms projecting basolaterally. Phallosome: Aedeagus 1.20-1.45 length of ventral

claspette; apex rounded, about as broad as long; without leaflets.

PUPA (fig. 29). Abdomen: about 2.65 mm. Trumpet: 0.5 mm. Paddle: 0.8 X

0.6 mm. Cephalothorax: Hairs 1-3-C moderately developed; 1-C 2,3 branched (1-3);

2,3-C 1-3 branched. Hair 4-C 3,4 branched, subequal to 5-C. Hair 5-C 3-5 branched

(2-5). Hair 6-C 2,3 forked (1-3). Hair 7-C 3,4 branched (2-4) near base; 6,7-C sub-

equal, moderately short. Trumpet: Pinna moderately pigmented; long, about 4.5-

5.1 (4.4-5.5) length of meatus; in lateral aspect, not appearing to taper toward apex.

Meatal cleft moderately pointed. Metanotum: Hair 10-C single or occasionally dou-

ble, moderately long, subequal to 11-C and more than 0.5 length of 12-C. Hair 11-C

3-5 branched. Hair 12-C 1-3 branched in basal 0.3, moderately long. Abdomen:

Hair 2-I 4-6 branched (4-7) about 0.3-0.5 from base, often dendritic, slightly longer

than 3-I; 3-I thin, long. Hair 4-I 4-9 branched, moderately large. Hair 5-I 1-3

branched, long. Hair 6-I single or double, about 1.5-2.0 length of 7-I. Hair 7-I 3-5

branched (3-6). Hair 9-I single, slightly longer than 7-I. Hair O-II-VII moderately de-

veloped; O0-II 3-6 branched; 0-III 4-8 branched (4-9); 0-IV,V 4-7 branched; 0-VI 4,5

branched; 0-VII 3-5 branched. Hair 1-II,[]I 6-10 and 5-9 branched respectively,

strongly developed. Hair 3-II inserted very near to or on caudal margin of segment

adjacent to 1-II; 3-III 1-3 branched (1-4); 3-IV 4-6 branched (4-7); 3-V 3-5 forked,

moderately developed. Hair 5-III,I[V 7,8 and 3-5 branched respectively, median

branches longer than lateral; 5-V-VII long, subequal to or slightly shorter than length

of segment; 5-V occasionally double. Hair 6-II 2,3 branched (1-3) near base, 1.2-1.7

length of 7-II; 6-III 3-5 forked (2-5). Hair 7-II 46 branched; 7-III-V moderate; 7-III,

IV 3-6 branched (3-8); 7-V 2-4 branched (1-4); 7-VI, VII single or occasionally dou-

ble, long. Hair 8-III-VII moderately developed; 8-III,VII 3-6 branched (2-8); 8-IV,VI

2-5 forked; 8-V 2,3 forked. Hair 9-II-VIII in general short and stout; 9-II minute, un-

pigmented; 9-III very small, about 2.0 or less length of 9-II; 9-IV heavy, small, about

2.0 of 9-III, pigmented; 9-V heavy, usually less than 2.0 of 9-IV; 9-VI slightly longer

than 9-V; 9-VII, VIII heavy, acuminate, about 0.33 length of segment. Hair 10-III 2-

5 branched, moderately developed. Hair 4-VIII 4-6 forked (3-8), subequal to 9-VIII.

Terminal Segments: Male genital lobe long and broad; apex with distinct mammilli-

form protuberance. Paddle: Large and caudally broad. Emarginate at insertion of

hair 1-P. External buttress about 0.66 length of paddle. External margin with short,

fine, filamentous spicules extending around apex and becoming indistinct along inner

margin. Hair 1-P moderately short; 2-P subequal to 1-P.

LARVA (fig. 30). Head: 0.6 mm. Antenna: 0.3 mm. Anal Saddle: 0.3 mm. In

general large. Head: Heavily pigmented. Median tooth of mental plate moderately

large, less than combined width of 2 adjacent teeth from one side. Hairs 2-C single

and barbed in about apical 0.5, widely spaced, clypeal index about 1.4; hair 3-C plu-

mose, subequal to 2-C, with moderately long, conspicuous branches in about apical

0.5. Hair 4-C 2-4 branched (1-4), small, occasionally reaching base of 2-C. Hairs 8,9-

C weakly plumose, short; 8-C 4-7 forked, slightly longer than distance between in-

sertion of hairs 5-C and 6-C; 9-C 6-10 forked (5-10), longer than 8-C. Hairs 10,12-C

3,4 and 46 branched respectively, moderately developed. Collar moderately narrow

dorsomedially and heavily pigmented. Antenna: Moderately pigmented. Mesal mar-

gin with stout, medium length spicules increasingly shorter from middle toward base

and apex; spicules usually absent in basal and apical 0.2. Hair 1-A 5-9 branched,

long, at least 2.0 width of antenna at point of insertion, inserted about 0.35-0.40

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 139

from base of antenna, branches long. Hairs 2,3-A wide and long, about 0.3 length of

antenna; mesal margins with conspicuous, featherlike, serrated fringe. Thorax: Mod-

erately to strongly pigmented. Submedian prothoracic group (1-3-P) not sharing a

common tubercle; palmate hair 1-P with 12-16 long, moderately broad, pointed leaf-

lets; 2-P relatively few branched (11-16), arising from a very small tubercle, lateral

branches equal to apical. Hair 14-P 6-9 branched from a moderately short stalk, la-

teral branches shorter than median. Mesothoracic hair 1-M 27-30 branched (23-30);

2-M 1-3 forked apically, moderately long. Metathoracic hair 2-T moderately long.

Palmate hair 3-T with 9-12 (8-12) semitransparent, broad, moderately long, pointed

leaflets. Hair 4-T 3-5 branched, moderately small. Prothoracic pleural group spine

large, longer than spines of meso- and metathoracic group. Abdomen: Many hairs

more extensively branched than those of strodei. Hair O-II,III,VI,VII 3-5 branched

(2-5); 0-IV,V 4-6 branched (4-7). Palmate hair 1-I with 8-13 (8-14) semitransparent,

moderately long, broad, pointed leaflets; 1-I]-VII with heavily pigmented, moderate-

ly broad, spreading leaflets, often apically truncate. Hair 2-II 4-6 branched (3-6); 2-

V 3,4 branched, moderately large. Hair 5-I 3-5 branched, moderately developed; 5-

III 9-13 branched (8-13), moderately large. Hair 9-I 5,6 branched (5-7); 9-IV 7-10

branched; 9-V 8-12 branched (8-13). Hair 11-I 3,4 branched, moderately large. Hair

13-1, 11,1 5-7 (4-8), 7-12 (6-12) and 7-14 branched respectively, small; 13-[V 10-13

branched (6-13), small; 13-V 4-6 branched (4-7), large. Spiracular Lobe: Pecten

with 11-15 teeth; most of median teeth subequal and of moderate length; basal serra-

tions on ventral surface of teeth fairly conspicuous. Lateral arm of spiracular appara-

tus moderately long and thick. Hairs 8,9-S 2-5 branched. Anal Segment: Covered

with moderately strong spicules. Hair 1-X moderately short, as long as or slightly

longer than saddle; inserted on saddle, on or very near ventral margin. Anal gills

about 0.35 mm, 1.00-1.25 length of saddle.

DISCUSSION. The adult female of benarrochi is very similar to that of aquasalis

and, to a lesser extent, to that of strodei. In order to correctly identify these 3 spe-

cies, the characters given below must be correlated and compared with those given in

the Discussion sections of the latter 2 species. It must be emphasized that the num-

ber of specimens of benarrochi that were available for study was not large, and the

majority were from the same general locality. For these reasons great care must be

taken in examining material from localities other than those studied here. An. benar-

rochi can be distinguished from the other species in the Oswaldoi Subgroup in the fe-

male (except for occasionally aquasalis) by the combination of (1) palpal segment 4

usually with a row of brown scales on ventral surface, (2) light scales on coxae usual-

ly white, (3) foretarsal segment 3 with a white band in apical 0.5-0.8, (4) hindtarsal

segment 2 brown in basal 0.40-0.55 (0.36-0.55), (5) light wing spots cream on anter-

_ jor veins and white on posterior veins, (6) vein C with humeral light spot about 2.5

or more of basal dark spot, (7) vein R with presectoral and sectoral dark spots mod-

erately small, (8) vein M mostly white, sectoral dark spot present or absent, (9) api-

cal light fringe spot large, unbroken or broken by a few dark scales and (10) dark

spots on veins R3, M,,. and M3,4, Cu subcostal dark spot, and A subbasal dark spot

often indistinct; in the male genitalia by the combination of (1) ventral claspette

small, apex wide, width at apex about 0.5 length of claspette, (2) apex of ventral

claspette rugose, moderately expanded laterally with apicolateral margin sharply an-

gled and moderately pointed, appearing truncate or with a median sulcus, (3) setae

on ventral claspette extending toward apex to or nearly to base of apicolateral lobe,

(4) preapical plate small, circular, heavily sclerotized and (5) basal lobule narrow,

curving mesad, with setae along basal margin about equal to or slightly longer than

140 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

width of aedeagus; in the pupa by the combination of (1) hairs 6,7-C subequal and

moderately short, (2) pinna long, about 4.5-5.1 length of meatus, not appearing to

taper toward apex in lateral aspect, (3) hair 10-C moderately long, subequal to 11-C

and more than 0.5 length of 12-C, (4) hair 2-I 4-6 branched (4-7), long, slightly long-

er than 3-I, (5) hair 9-I slightly longer than 7-I, (6) hair 6-II 1.2-1.7 length of 7-II and

(7) hair 9-III very small, about 2.0 or less length of 9-II, 9-V heavy, usually less than

2.0 of 9-IV, 9-VII, VIII heavy, short, about 0.33 length of segment; and in the larva

by the combination of (1) hair 2-C single and barbed, 3-C plumose with moderately

long branches in about apical 0.5, hairs 2-C widely spaced, clypeal index about 1.4,

(2) hair 4-C small, (3) collar moderately narrow dorsomedially and heavily pigment-

ed, (4) hair 1-A of antenna 5-9 branched, long, at least 2.0 width of antenna at point

of insertion, (5) hairs 1,2-P not sharing a common sclerotized tubercle, (6) hair 13-I1V

10-13 branched (6-13), small, (7) lateral arm of spiracular apparatus moderately long

and thick and (8) hair 1-X moderately short, as long as or slightly longer than saddle,

inserted on saddle, on or very near ventral margin.

An. benarrochi is the most highly derived species in the Oswaldoi Subgroup. The

derived characters include in the male genitalia the small distinct ventral claspette

with its narrow, mesally curved basal lobules; in the pupa the position of hair 3-II ad-

jacent to 1-II; and in the larva the large antennal hair 1-A, the small numerously

branched hair 13-IV and the short hair 1-X.

The closest allies of benarrochi are rondoni and strodei, with which it shares in the

male genitalia the nonsetaceous, laterally expanded apicolateral lobes and the rugose

apical margin of the ventral claspette, and in the larva the narrow collar and hair 1-X

inserted on the saddle. It should be stressed that the latter 2 characters in the larva

may be ancestral and may not be indicative of close phylogenetic relationship.

BIONOMICS. Very little is known of the natural history of benarrochi. The

immatures have been reported from stagnant ground pools, abandoned wells and

small streams, exposed to the full sun or partial shade, and in water containing a

large amount of organic material (Deane, Causey and Deane 1948:900-901; Cerque-

ira 1961:123). An. benarrochi is not found in the lower Amazon basin, and nowhere

is it encountered in great abundance (Deane, Causey and Deane 1948). Deane, Cau-

sey and Deane (1948) collected the larvae in association with triannulatus, An. (Nys.)

albitarsis and An. (Ano.) peryassui.

The females rarely enter houses and feed primarily on animals. Of a total of 545

adults of benarrochi collected by Deane, Causey and Deane (1948) only 46 or 8.4%

were from inside houses. Elliott (1972:757) reports that the peak hours of biting for

the females in Peru is between 1800-2000 h and 0400-0600 h, which correlates with

the observation of Deane, Causey and Deane (1948) that benarrochi is crepuscular.

MEDICAL IMPORTANCE. An. benarrochi has never been implicated as a vector

of malaria. Deane, Causey and Deane (1948) dissected 31 blood-fed females and

found none infected with Plasmodium sp.

DISTRIBUTION (fig. 1). An. benarrochi is limited primarily to the Orinoco basin

and the eastern versant of the Andes including the llanos plateau region of Colombia,

parts of the upper Amazon in Brazil (Rondonia, Acre, Amazonas) and Loreto, Peru.

Material Examined: 293 specimens: 60 males, 33 male genitalia, 85 females, 64

pupae, 51 larvae; 6 progeny rearings.

COLOMBIA (286 specimens: 57M, 30Mgen, 85F, 64p, SOL; 6 progeny rearings). Meta: Villavi-

cencio, 1947, L. Rozeboom, CV-P 34 (progeny rearing), 8M, 7Mgen, 9F, 15p, 9L; same data, CV-P

35 (progeny rearing), 6M, 3Mgen, 8F, 15p, 9L; same data, CV-P 36, 1F; same data, CV-P 37 (pro-

geny rearing), 1OM,4Mgen, 12F, 18p, 13L; same data, CV-P 40 (progeny rearing), 1F, 1p, 5L; same

data, CV-P 42 (progeny rearing), 19M, 4Mgen, 24F, 14L; same data, [CV-] P 42 (progeny rearing),

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 141

1Mgen, 10F [JH]; same data, CV-P 43 (progeny rearing), 9M, 3Mgen, 10F, 15p; same data, [CV-]P

43 (progeny rearing), 5M, 5Mgen, 10F [JH]; same data, 3Mgen [JH].

PERV (7 specimens: 3M, 3Mgen, 1L). Loreto: Iquitos, Apr 1931, R. Shannon, 3M, 3Mgen;

same locality, A. Cornejod, 1L.

1922.

1923.

1925,

1927,

1928.

1941.

TRIANNULATUS SUBGROUP

14. Anopheles (Nys.) triannulatus (Neiva & Pinto)

Piss, 147 7, 34) 32

Cellia triannulata Neiva and Pinto 1922b:356-357. TYPE: Holotype female, Fazenda Sao

Joao, right bank of Rio Cuiaba (Mato Grosso), Brazil [LU; not in IOC] (Belkin, Schick

and Heinemann 1971:6).

Cellia cuyabensis Neiva and Pinto 1923:235-236. TYPE: Lectotype small wing mounted

on slide (403) together with wing of type of rondoni, marked Matto Grosso, C. Pinto, re-

mainder of adult apparently lost, Fazenda Sao Joao, right bank of Rio Cuiaba (Mato

Grosso), Brazil [IOC; designation of Belkin, Schick and Heinemann 1971:7]. Synonymy

with triannulatus by Galvao and Barretto (1939b:150-154).

Anopheles bachmanni Petrocchi 1925:71-75. TYPE: Syntypes males and females, Corri-

entes, Entre Rios, Formosa, Argentina [NE; according to O. H. Casal in Belkin, Schick and

Heinemann 1968:10]. NEWSYNONYMY.

Anopheles (Nyssorhynchus) davisi Paterson and Shannon 1927:1277-1280. TYPE: Holo-

type male (200) with associated genitalia on slide, Tres Pozos, Embarcacion (Salta), Argen-

tina, 20 Apr 1927, Paterson, Shannon and Shannon [USNM]. TRANSFERRED SYN-

ONYMY.

Anopheles (Nyssorhynchus) perezi Shannon and Del Ponte 1928:56-57. TYPE: Syntypes

females, [La Finca] Santa Barbara, Departamento de la Capital (Tucuman), Argentina, 7

Apr 1927, Shannon and Del Ponte [NE; according to O. H. Casal in Belkin, Schick and

Heinemann 1968:10|. TRANSFERRED SYNONYMY.

Anopheles (Nyssorhynchus) triannulatus var. chagasi Galvao 1941:92-95. TYPE: Syn-

types females and eggs, type locality restricted to Manaus (Belkin, Schick and Heinemann

1971:7), originally described from Chaves (Para), and Itacoatiara and Manaus (Amazonas),

Brazil [LU]. Synonymy with triannulatus by Lane (1949:404).

Anopheles (Nyssorhynchus) triannulatus of Dyar (1928:437); Edwards (1932:45); Galvao and Lane

(1937e:214; 1941:10-18); Pinto (1939:406-425); Lane (1939:29-30); Galvao and Barretto

(1939b: 150-156); Galvao (1940:437-443; 1943:144, 149, 150, 151); Komp (1941a:93, 97;

1941b:792; 1942:37, 43, 69-71, 80, 118-120, 132, 156-158; 1956:39); Correa and Ramos

(1942a:42-43; 1944b:132); Fonseca and Fonseca (1942:94, 95, 97, 99); Rozeboom (1942a:

241-242; 1963:112); Simmons and Aitken (1942:39, 45, 53, 61, 97-99); Bruce, Knigin, Yolles

and Graham (1943:441-442); Cerqueira (1943:19-20); Fonseca and Unti (1943:49-50); Galvis

(1943:89-91); Unti and Ramos (1943:28); Causey, Deane and Deane (1944:2, 4, 5; 1946:26);

Lane (1944: 262-268; 1949:404); Deane, L. M., Causey and Deane (1946:7, 9, 10; 1948:901-

904); Deane, M. P., Causey and Deane (1946:38, 39, 42, 45); Floch and Abonnenc (1946b:3, 5;

1947:7; 1951:52-55); Gabaldon and Cova Garcia (1946a: 19-20; 1946b: 121-124); Amaral and

Penido (1947:168); Arnett (1947:198-199; 1950:106, 110, 112); Senevet (1948a:278; 1948c:

434-439); Vargas (1948:157-158); Bates and de Zulueta (1949:134, 137, 139); Levi-Castillo

(1949:9, 10, 11, 13, 16, 28, 67, 73, 77, 82, 86); Duret (1950a:474-475; 1950b:302; 1952:348);

Carvalho and Rachou (1951:475-476, 480); Rachou and Ferraz (1951:542, 547-553); Rachou

and Ricciardi (1951:424-426, 432-437); Horsfall (1955:185); Bejarano (1956:9, 10, 20, 21;

1957:327-328, 339-342; 1959:319, 325); Rozeboom and Kitzmiller (1958:244); Vargas V.

(1958b); Castro, Garcia and Bressanello (1959:550); Store, Knight and Starcke (1959:34); Faur-

an (1961:11-12); Forattini (1962:311, 400-405); Belkin, Schick and Heinemann (1968:10;

1971:6, 7); Consolim and Galvao (1973:177); Cova Garcia, Pulido F. and Amarista M. (1978:

157);

142 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Anopheles triannulatus of Rozeboom (1941:101-102); Vargas (1940a:199-200; 1941:114); Rus-

sell, Rozeboom and Stone (1943:26, 30, 37, 41, 50); Floch and Abonnenc (1944a:1-7; 1946a:

2); Renjifo and de Zulueta (1952:600-601); Vargas V. (1958c); Foote and Cook (1959: 20, 23);

Maciel (1962:475-476); Vincke and Pant (1962:2-4, 6, 7, 10); Mattos and Xavier (1965:274-

275); Stojanovich, Gorham and Scott (1966a:13, 18, 35; 1966b:22, 28, 44); Gorham, Stojano-

vich and Scott (1967:15, 45, 46, 60; 1973:111, 151).

Anopheles (Nyssorhynchus) triannulatus triannulatus of Galvao and Lane (1941:11-17); Vargas

(1941:114, 118; 1959:376, 382); Galvao (1943:149, 150); Lane (1953:274-276); Vargas V.

(1956:28, 33; 1957; 1958b); Bejarano (1957:339-342; 1959:305, 321); Cerqueira (1961:127-

128); Garcia and Ronderos (1962:137-139, 161-162); Morales-Ayala (1971:139).

Anopheles triannulatus triannulatus of Vargas (1943:58).

Anopheles (Anopheles) triannulata of Bonne and Bonne-Wepster (1925:520-521).

Cellia triannulata of Bonne (1924:133, 134).

Anopheles (Nyssorhynchus) tarsimaculatus var. triannulatus of Christophers (1924:40-41, 91).

Anopheles (Nyssorhynchus) bachmanni of Root (1926b:684-696, 714-715); Davis (1928:562);

Dyar (1928:427, 430, 435); Shannon and Del Ponte (1928:55); Hill (1930:712; 1934:425-429);

Shannon and Davis (1930:488); Shannon (1931:10, 22; 1933:118, 126-132); Curry (1932:566-

572; 1934:646); Edwards (1932:45); Antunes and Lane (1933:91-97); Townsend (1933a:101);

Senevet (1934:47-49; 1937:361-362); Komp (1936a:161); Galvao and Lane (1937a:77, 78;

1937e:214, 223); Galvao, Lane and Correa (1937:41); Galvao and Amaral (1938:13); Roze-

boom (1938a:101-102; 1938b:290-300); Kumm and Ruiz (1939:438); Pinto (1939:406-425);

Kumm, Komp and Ruiz (1940:389, 391, 410, 419); Komp (1941a:95); Floch and Abonnenc

(1942a:2; 1942b:2); Cova Garcia and Sutil O. (1976:31; 1977:22, 46, 62, 88).

Anopheles bachmanni of Petrocchi (1923:139); Muehlens, Petrocchi and Zuccarini (1925:265);

Hill (1928:355); Benarrochi (1931:691-693); Davis (1931 :46-47, 49-50); Kumm (1932:4); Pin-

to (1932:287); Rozeboom (1935:521-528); Kumm and Novis (1938:503); Clark, Komp and

Jobbins (1940:62); Gabaldon, Lopez and Ochoa Palacios (1940:38); Gabaldon, Ochoa Palacios

and Perez Vivas (1940:42-55); Cova Garcia and Sutil O. (1975a:22; 1975b:211).

Anopheles (Nyssorhynchus) triannulatus bachmanni of Garcia and Ronderos (1962:137, 138, 139,

162); Morales-Ayala (1971:139).

Anopheles triannulatus bachmani (') of Cova Garcia (1961:39-40, 68, 91-92, 113, 128-129, 154).

Anopheles buchmanni (!) of Clark (1934:642).

Nyssorhynchus (Nyssorhynchus) bachmanni of Lima (1928:98-100).

Nyssorhynchus bachmanni of Townsend (1934:494).

Anopheles (Nyssorhynchus) davisi of Shannon and Del Ponte (1928:43-45); Pinto (1939:406-425);

Castro (1959a:174); Castro, Garcia and Bressanello (1959:549).

Anopheles davisi of Renjifo and de Zulueta (1952:600-601).

Anopheles (Nyssorhynchus) triannulatus davisi of Galvao and Lane (1941:11-17); Galvao (1943:

144, 149, 151); van der Kuyp (1949a:67-68); Rey and Renjifo (1950:534, 537); Lane (1953:

276-277); Vargas V. (1956:28); Bejarano (1956:9, 20, 21; 1957:339-342; 1959:305, 320); Var-

gas (1959:382).

Anopheles triannulatus davisi of Vargas (1943:58).

Anopheles (Nyssorhynchus) cuyabensis of Dyar (1928:437); Edwards (1932:45); Galvao and Lane

(1937e:214); Pinto (1939:410, 412-414).

Nyssorhynchus (Nyssorhynchus) cuyabensis of Lima (1928:103).

Anopheles (Nyssorhynchus) tarsimaculatus var. cuyabensis of Christophers (1924:40, 90).

Anopheles (Anopheles) cuyabensis of Bonne and Bonne-Wepster (1925:521).

Anopheles (Nyssorhynchus) perezi of Edwards (1932:45).

Anopheles tarsimaculatus in part (?) of Edwards (1922:75).

Nyssorhynchus tarsimaculatus in part of Townsend (1933b:1, 7-12).

FEMALE (fig. 7). Wing: 2.6 mm. Proboscis: 1.8 mm. Palpus: 1.65 mm. Fore-

femur: 1.15 mm. Abdomen: 2.2 mm. A small, often predominantly dark species.

Head: Integument brown to dark brown. Interocular distance about 3,4 ommatidial

diameters. Proboscis about 1.55 length of forefemur. Dorsal surface of palpal seg-

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 143

ment 2 with or without a few light scales; dorsal surface of segment 3 with few to

several, light scales forming a longitudinal stripe from about 0.5 distad of base to near

apex; palpal segments 2 and 3 with a small to moderately wide, white, apical band;

segment 4 variable but always with a basal and an apical, dark band, ventral surface

dark scaled, and mediolateral surface from almost completely dark with few light

scales to predominantly white; segment 5 white with or without a few dark scales at

base extending from apex of 4. Antenna: Base of flagellar segment 1 dorsolaterally

with small, white, obovate scales; apical 0.5 on dorsomesal surface with several long,

narrow, lanceolate scales. Thorax: Integument reddish brown to very dark brown or

maroon. Light scales of scutum moderately long and narrow. Upper stp with 3-5

long setae, and a patch of silver, cuneate to obovate scales in a horizontal arc. Lower

stp with 1,2 long setae, and a patch of scales similar to those of upper stp in a diagon-

al row. Upper mep with 2-6 setae closely approximated dorsocaudally; scales absent.

Anterior mep with a conspicuous, moderately large patch of silver to white, cuneate

scales oriented horizontally at same level as scales on upper stp. Legs: Light coxal

scales usually white. Foretarsal segments 2 and 3 with a pale cream to white band in

apical 0.20 (0.15-0.35) and 0.6 (0.35-0.75) respectively; segment 4 with a pale cream

to white band in apical 0.55 (0.45-0.65); segment 5 dark, occasionally with a few,

golden, apical scales. Midtarsal segments 1 and 2 with a cream apical band; segments

3-5 completely dark or with a few, light, apical scales. Hindtarsal segment 2 with a

dark band in basal 0.5 (0.4-0.7). Wing: Dark wing spots predominantly large. Light

Wing spots usually cream to yellowish at least on veins C and R. Vein C often almost

completely dark, light spots occasionally obsolete; basal dark spot usually reaching

humeral crossvein; humeral light spot small, almost always shorter (0.5-1.3) than bas-

al dark spot; subbasal dark spot longer than subbasal light spot; subcostal light spot

about 0.15 (0.1-0.2) of subcostal dark spot; preapical light spot about 0.25 (0.1-0.4)

of preapical dark spot; apical dark spot moderately large and conspicuous. R presec-

toral dark spot usually large. R, preapical dark spot large, usually at least 0.5 length

of vein. R,3 about 0.5 of R,. M sectoral dark spot variable, usually very large, of-

ten 0.7 of vein. M3,4 preapical dark spot and vein A subcostal dark spot large. Light

fringe spots as in section description. Abdomen: Sternite I with a few, moderately

long to long setae along lateral margin, no scales visible. Remainder of abdomen asin

section description except tergites II-VII usually not as densely and extensively cov-

ered with silver to cream scales as in other species in section, and light scales concen-

trated toward midline of tergites.

MALE. Wing: 2.75 mm. Proboscis: 2.3 mm. Forefemur: 1.30 mm. Abdomen:

about 2.6 mm. Essentially as in female except for sexual characters. Head: Palpal

segments 2 and 3 with or without light dorsal stripe; segment 2 usually with only a

few light scales; segment 4 with a basal and an apical, white band, and a subbasal and

a subapical, moderately large to large, dark band, mediolaterally with or without

cream to white scales which are less extensive than those of the Oswaldoi Subgroup.

Antenna: About 0.8 length of proboscis. Flagellar segment 1 with long, white, ob-

lanceolate scales. Legs: Forefemur 0.6 length of proboscis. Basal plantar surface of

foretarsal segment 5 with moderately long, spiniform setae; setae conspicuously long-

er apically, about 0.3 length of segment 5. Claw on foreleg large, curved, acuminate;

submedian tooth about 0.2 length of claw, with conspicuously recurved apex; basal

tooth moderately short and decurved.

MALE GENITALIA (fig. 31). Segment VIII: Tergite and sternite with moderate-

ly broad, light scales. Tergite with light scales and interspersed darker scales. Seg-

ment IX: Sternite long, about 0.2 length of sidepiece, subtrapezoidal. Anterior apo-

144 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

deme large, about 0.33 length of sternite, subtriangular. Sidepiece: Moderately nar-

row, conical. Tergal surface with 3-5 long, submedian tergomedial bristles, basal bris-

tles shorter than apical; with very few, short to moderate bristles (occasionally ab-

sent) mesad of tergomedial bristles and one or 2 long subapicolateral bristles. Para-

basal spine long, its tubercle about 0.5 or less length of spine, and apex of spine of-

ten extending beyond lateral margin of aedeagus on other side. Basal apodeme thick,

moderately long, about 0.20-0.25 length of sidepiece. Accessory spines usually not

appearing blunt but sharply hooked apically; longer, more dorsal spine about 0.5

length of sidepiece; more ventral spine 0.70-0.85 of longer spine. Internal spine sub-

equal to shorter accessory spine, apically retrorse. Clasper: Spiniform thin, short,

equal to or slightly shorter than seta b. Dorsal Claspette: Pedicel moderately heavy,

indented subbasally; base rounded, mesally hooked. Leaflets very broad, strongly

curved, less than 0.5 length of claspette; dorsal leaflet with base produced into pro-

minent, mesally rounded or triangular basomesal projection. Ventral Claspette:

Without setae. Long, about 0.5 length of sidepiece; apex wide, width at apex 0.6-0.7

length of claspette. Basal lobule moderately large. Lateral margins of claspette tap-

ering slightly medially toward apex. Apex truncate and rugose. Apicolateral margins

produced into large, striated, auriculate, laterally projecting lobes. Preapical plate

near apex, small, oval, heavily sclerotized, often appearing as 2 small, heavily sclero-

tized, adjacent, circular dots embedded in a small, moderately sclerotized, oval plate.

Transparent, inverted U shaped, membranous band mesally bordering vertex and lat-

eral margins of basal lobules. Basal lobules interconnected by a weakly sclerotized,

speckled membrane; membrane conspicuously more sclerotized at vertex of mesal

cleft than basally. Phallosome: Aedeagus narrow, long, subequal to length of ventral

claspette; apex rounded, about 1.5 as long as wide; without leaflets.

PUPA (fig. 31). Abdomen: 2.13 mm. Trumpet: 0.4 mm. Paddle: 0.65 X 0.44

mm. Small and dark. Branches of hairs relatively long. Cephalothorax: Brown to

brownish black. Hairs 1-3-C 2,3 branched (1-4); 2-C slightly shorter than 1 ,3-C.

Hairs 4,5-C 2-4 branched (2-5); 5-C about 1.5 length of 4-C. Hairs 6,7-C 1-3

branched; 6-C forked about 0.5 from base, shorter than 7-C; 7-C with one branch

very long (0.3 mm), about 1.5-2.0 length of other branch(es). Trumpet: Pale yellow

to yellow. Pinna very long, about 4.0-6.5 (3.4-8.0) length of meatus, narrow; in lat-

eral aspect, not appearing to taper toward apex. Meatal cleft long, open and basally

rounded. Metanotum: Hair 10-C single, short, much shorter than 11-C and 0.5 or

less length of 12-C. Hair 11-C 3 branched near base. Hair 12-C 2,3 branched (2-4).

Abdomen: Brown to brownish black, darker dorsomedially. Float hair (1-I) with 7-

15 major branches, secondarily very dendritic. Hair 2-I 3-6 branched near base, mod-

erately long; 3-I single, subequal to 2-I. Hair 4-I 4-6 branched (4-7), moderately

large, subequal to 2-I. Hair 5-I single and long. Hair 6-I single, very long (0.35 mm),

about 2.0 length of 7-I. Hair 7-I 3-6 branched (1-6). Hair 9-I single, subequal to 6-1.

Hair O-II-VII few branched (2-5), moderately small. Hair 1-II,III strongly developed;

1-II 8-11 branched (8-12); 1-III 4-8 branched, median branches longer than lateral; 1-

IV-VII very long, 1.75-2.00 length of segment; 1-V,VI longer than 1-IV,VII. Hair 3-

IV 3-6 branched (2-6), moderately long; 3-V 2-4 forked. Hair 5-III 5-8 branched (5-

9), lateral branches shorter than median; 5-IV double or triple, long; 5-V-VII single,

subequal to length of segment. Hair 6-II single, very long, about 2.0 length of 7-II;

6-III 1-3 forked, long. Hair 7-II 3-6 branched; 7-IJI,I[V 4 or fewer branched, moder-

ately small; 7-V 1-3 branched, about 1.5 length of 8-V; 7-VI,VII single, long. Hair 8-

IfI-VII 1-3 branched, small. Hair 9-II,III thin, sharp, small, unpigmented; 9-III slight-

ly longer than 9-II; 9-IV moderately small, 2.0 or more length of 9-II,III, pigmented;

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 145

9-V 2.7-6.0 (2.3-6.0) length of 9-IV; 9-V-VIII slender, curved, acuminate, very long;

9-V,VI slightly shorter than 9-VII, VIII; 9-VII 1.1-1.4 (0.90-1.42) length of 9-VI; 9-V-

VIII about 0.5 or more length of segment. Hair 10-III 2,3 forked, moderately long.

Hair 4-VIII 2-4 forked, subequal to 9-VIII. Terminal Segments: Male genital lobe

broad, with a moderately large mammilliform protuberance. Paddle: Large, weakly

oblique, truncate; midrib fairly distinct, extending almost to apex. External buttress

about 0.68 length of paddle. External margin with moderately short, filamentous

spicules extending around apex and along inner margin to 0.66 from base; spicules

very short on inner margin. Hair 1-P moderately large. Hair 2-P less than or equal to

1-P.

LARVA (fig. 32). Head: 0.5 mm. Antenna: 0.25 mm. Anal Saddle: 0.27 mm.

In general small and dark. Head: Usually very dark and strongly pigmented. Medi-

an tooth of mental plate moderately narrow, less than combined widths of 2 adja-

cent teeth from one side. Inner and outer clypeal hairs (2,3-C) single, and simple or

barbed in about apical 0.5; 3-C shorter than 2-C; hairs 2-C widely spaced, clypeal in-

dex about 1.3. Hair 4-C 2-4 branched, small, not reaching base of 2-C, inserted

slightly laterad of an imaginary median line separating 2-C and 3-C. Hairs 8,9-C 3-5

branched (2-6), long, about 2.0 distance separating insertion of 5-C and 6-C. Hair 10-

C 2-4 branched, large. Hairs 12,13-C 4-7 branched (3-7), large. Hair 15-C 3-5 forked.

Collar narrow dorsomedially, heavily pigmented. Antenna: Darkly pigmented. Mes-

al margin with strong, stout spicules; ventral surface with stouter, shorter spicules.

Hair 1-A 4-6 branched (3-6), short. Hairs 2,3-A large, about 0.4 length of antenna.

Thorax: Moderately to heavily pigmented at least medially; grayish brown to almost

black. Submedian prothoracic group (1-3-P) with 1,2-P not sharing a common tuber-

cle; palmate hair 1-P with 15-20 (13-20) very narrow to narrow, lanceolate leaflets

(rarely filiform); 2-P 16-23 branched, arising from small pigmented tubercle, about 3.0

length of 1-P. Hair 11-P double, branched 0.25 from base. Hair 14-P 11-14 branched

(9-15) from an elongated, flattened shaft, lateral branches shorter than median. Mes-

othoracic hair 1-M 26-40 branched. Metathoracic palmate hair 3-T with 9-14 large,

narrow, semitransparent leaflets. Hair 4-T 3-5 branched, moderately long. Pro-,

meso- and metathoracic pleural group spines large; mesothoracic spine larger than

pro- and metathoracic spines. Abdomen: Hairs usually proportionally longer than

those of other species in group. Hair O-II-VI 6-11 branched (5-12). Palmate hair 1-I

with 14-17 (12-18) semitransparent, narrow, pointed leaflets; 1-II-VII with about 20-

30 pigmented, narrow, pointed leaflets. Hair 2-II 6-9 branched (5-10), large; 2-V sin-

gle or occasionally double, very long. Hair 5-I 4-6 branched (3-6), moderately devel-

oped; 5-III 7-11 branched. Hair 9-I 6-9 branched (5-10), branches long; 9-I[V,V 7-12

branched (6-12). Hair 11-I 5-7 branched (3-7), large. Hair 13-1 3 branched (2-4),

very large; 13-II 7-11 branched, moderately large; 13-III,[V 4-6 branched (3-7), large;

13-V triple, very large. Sclerotized tergal plate VII about 2.0 size of tergal plate VI.

Spiracular Lobe: Pecten with 14-18 teeth; many of median teeth subequal; serra-

tions conspicuous in basal half of teeth. Hairs 8,9-S 4-7 branched, large; 8-S smaller

than 9-S. Lateral arm of spiracular apparatus long, directed laterally, truncate at lat-

eral apex and reaching spiracular openings. Anal Segment: Covered with moderately

long spicules. Saddle reddish brown. Hair 1-X long; inserted on saddle near ventral

margin, about 0.3 cephalad of caudal margin. Anal gills slightly longer than saddle.

DISCUSSION. The monotypic Triannulatus Subgroup can be distinguished from

the Oswaldoi Subgroup in the female by (1) anterior mesepimeron with a conspicu-

ous patch of silver to white, cuneate scales, (2) palpal segment 4 with mediolateral

surface having a reduced number of light scales, (3) foretarsal segment 4 with a light

146 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

band in apical 0.55 (0.45-0.65), (4) foretarsal segment 5 dark, occasionally with a

few, apical, golden scales, (5) hindtarsal segment 2 brown in basal 0.5 (0.4-0.7), (6)

vein C with large dark spots, light spots occasionally obsolete, humeral light

spot small, 0.5-1.3 length of basal dark spot, apical dark spot moderately large and

conspicuous and (7) vein M with sectoral dark spot usually very large, often 0.7

length of vein; in the male genitalia by (1) ventral claspette without setae, apicolater-

al margins produced into 2 large, striated, auriculate, laterally projecting lobes, (2)

preapical plate small, oval, heavily sclerotized and (3) aedeagus long, subequal to

length of ventral claspette, with apex about 1.5 as long as wide; in the pupa by the

combination of (1) pinna narrow, very long, about 4.0-6.5 (3.4-8.0) length of meatus,

not appearing to taper toward apex in lateral aspect, (2) hair 7-C with one branch

about 1.5-2.0 length of other branch(es), (3) hair 10-C single, short, 0.5 or less length

of 12-C, (4) hair 3-I subequal to 2-I, (5) hair 6-I,II very long, about 2.0 length of hair

7 on same segment, (6) hair 9-III slightly longer than 9-II, unpigmented, 9-IV 2.0 or

more length of 9-III, 9-V 2.3 or more length of 9-IV, 9-V,VI slightly shorter than 9-

VII,VIUI and about 0.5 or more length of segment and (7) paddle large, weakly ob-

lique, truncate; and in the larva by the combination of (1) hairs 2-C single, and sim-

ple or barbed, widely spaced, clypeal index about 1.3, (2) collar narrow dorsomedial-

ly, heavily pigmented, (3) palmate hair 1-P with 15-20 (13-20) very narrow to nar-

row, lanceolate leaflets, 2-P 16-23 branched, arising from a small pigmented tubercle,

about 3.0 length of 1-P, (4) hair 14-P 11-14 branched (9-15) from an elongated, flat-

tened shaft, (5) hair 11-1 5-7 branched, large, and 13-I 2-4 branched, very large, (6)

hair 13-III,[V 4-6 branched (3-7), large, (7) tergal plate of segment VII large, about

2.0 size of tergal plate VI and (8) lateral arm of spiracular apparatus long, directed

laterally. |

Although the chaetotaxy of the larva and pupa varies, morphologically triannula-

tus is fairly uniform throughout its range. In the larva, there is considerable variation

in the number and width of the leaflets of hair 1-P and in the length of the lateral

arms of the spiracular apparatus which varies from long to very long. In the pupa,

hair 1-III usually has 3 branches in specimens from Brazil and 4,5 branches in those

from Panama. On an average, pupae examined from Brazil had one fewer branch per

hair than pupae from Panama. In the female, the sectoral dark spot of vein M usual-

ly is smaller in Panamanian specimens than in the Brazilian specimens, although this

is a highly variable character.

An. triannulatus possesses several derived characters including in the female (1)

presence of a patch of light scales on the anterior mesepimeron, (2) apical light band

of foretarsal segment 4 and (3) the unique wing spotting; in the male genitalia (1)

shape of the apex of the ventral claspette and (2) shape of the aedeagus; and in the

larva (1) elongated, flattened shaft of hair 14-P and (2) long lateral arms of the spira-

cular apparatus. These derived characters which distinguish triannulatus from the

Oswaldoi Subgroup necessitates, I believe, placement of triannulatus in a monotypic

subgroup.

In addition, triannulatus retains several ancestral characters which it shares with

the Albimanus Group and with some species of the Argyritarsis Section, such as, in

the adult, the small amount of white on palpal segment 4 and the predominantly

dark foretarsal segment 5; in the male genitalia, the absence of setae on the ventral

claspette; and in the larva, the number and width of the branches of hair 1-P and the

large, sparingly branched hair 13-I, IIL IV.

Within the Oswaldoi Subgroup, triannulatus shows a greater affinity to the Strodei

Complex than to the Oswaldoi Complex. This species shares many synapomorphic

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 147

characters with strodei and rondoni such as, in the male genitalia, the apicolateral ex-

pansion of the ventral claspette; in the pupa, the very long pinna of the trumpet and

the very long, slender hair 9-VII, VIII; and in the larva the relatively narrow collar

and hairs 1-3-P not sharing a common tubercle.

The taxonomy of triannulatus has been very confusing in part due to the numer-

ous mutant forms which have been described as separate nominal species. In fact,

the description of triannulatus by Neiva and Pinto (1922b) was based on an anomal-

ous specimen that had extra black basal bands on the second and third hindtarsal seg-

ments. In 1923, Neiva and Pinto described another mutant form, cuyabensis, distin-

guished from triannulatus by having only one extra basal black ring on the fourth

hindtarsal segment. Petrocchi (1925) described the more typical form she called

bachmanni, clearly illustrating its male genitalia. Later, 2 other nominal species were

described, davisi Paterson and Shannon 1927 and perezi Shannon and Del Ponte

1928, both said to be very closely related to bachmanni. Lima (1928:103), after ex-

amining a cotype of cuyabensis, stated that this species was extremely similar to

bachmanni. Dyar (1928:435) the same year listed davisi and perezi as synonyms of

bachmanni. In 1930, Pinto synonymized bachmanni with cuyabensis, stating, “Os

estudos de Davis reforcam o nosso ponto de vista de que a pequena mancha negra

existente na base do IV articulo do tarso posterior de A. cuyabensis e uma variacao.

Caso este fato venha a se confirmar o A. bachmanni caira forcosamente na sinonimia

de A. cuyabensis que foi descrito dois anos antes.’’ Galvao and Barretto (1939: 150-

154) synonymized cuyabensis, bachmanni, davisi and perezi with triannulatus based

on Pinto’s (1939:406-425) observations and on their own findings. These investiga-

tors studied material from Itapira, Brazil and discovered specimens of cuyabensis to-

gether with bachmanni. Pinto (1939) concurred with Galvao and Barretto after com-

paring the holotype of triannulatus with a female identified by Shannon as bach-

manni (no. 2899, Resistencia) and with topotypic material of cuyabensis. Pinto sta-

ted that all 3 were the same species, triannulatus being the senior synonym. He also

considered the holotypes of triannulatus and cuyabensis as atypical forms similar to

those described in other species in the section. Pinto studied the larvae, males and fe-

males of bachmanni from other localities in Brazil and found the male genitalia to be

the same as those of the topotypic material of triannulatus and cuyabensis. In the

same publication (Pinto 1939), he also listed perezi as asynonym of triannulatus.

Later, Galvao and Lane (1941:11-14) examined abundant material of triannulatus

and stated that they had observed all forms corresponding to the descriptions of

bachmanni, cuyabensis and triannulatus. However, Galvao and Lane recognized tri-

annulatus as being composed of 2 subspecies, t. triannulatus and t. davisi. The prin-

cipal characters reportedly used to distinguish ¢. triannulatus from t. davisi were (1)

pattern of the exochorion in the egg, (2) larval hair 1-P filamentous and with more

branches (20-30) than in davisi, (3) female smaller than that of davisi and (4) adult

wing with sectoral dark spot of Cu larger and reaching the furcation (Paterson and

Shannon 1927; Galvao and Lane 1941; Rozeboom 1942a:241-242; Deane, Causey

and Deane 1948:901-904). Some investigators believe that the differences between

triannulatus and davisi are so striking that davisi merits species status (Deane, Causey

and Deane 1948). I have examined adults from very near the type locality of bach-

manni, from the type locality of davisi and from Salobra and other localities in Mato

Grosso, Brazil where Galvao reportedly collected ¢. triannulatus. Though there is

some variation in the size of wing spots, I could not see anything in the adults which

would lead me to consider triannulatus as being composed of 2 subspecies. Likewise

in the larva, although the number and width of the branches of hair 1-P and the

148 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

length of the lateral arms of the spiracular apparatus varied, the variation was usually

intrapopulational and present as a continuum. It is possible, however, that I did not

ever see a true example of ¢. triannulatus, since it is considered to be much less com-

mon than ¢. davisi. In that case, my consideration of triannulatus as being composed

of a single variable taxon may be incorrect.

-Galvao (1941) described another variety of triannulatus, chagasi, from Chaves

(Para), and Itacoatiara and Manaus (Amazonas), Brazil. An. chagasi was later synon-

ymized with triannulatus by Lane (1949:404).

An. triannulatus, like several of the other species in the section, has been reported

to be highly polymorphic in the egg stage. Galvao (1940:441-442) recognized 4

types of triannulatus eggs: 2 types were of ¢. triannulatus and 2 of t. davisi; this is

discussed in detail in Rozeboom (1942a:241-242).

BIONOMICS. The immatures of triannulatus are most commonly collected in per-

manent ponds, lakes, canals, slow streams or river margins, ditches, and swamps, ei-

ther exposed to the full sun or partially shaded. Occasionally triannulatus is collec-

ted in semipermanent and temporary rock holes, small ground pools, and animal

tracks. The immatures are found in clear, fresh water, never brackish, and are usual-

ly associated with Pistia sp. Other commonly associated aquatic plants are Eichhorn-

ia sp, Azolla sp, Utricularia sp, Jussiaea sp, Elodea sp and Salvinia sp. The immatures

have been collected in association with 7 other species in the Albimanus Section: al-

bimanus, oswaldoi, ininii, nuneztovari, rangeli, strodei and on one occasion aquasalis.

Other mosquitoes collected with triannulatus are Anopheles (Ano.) apicimacula, An.

(Ano.) neomaculipalpus, Uranotaenia ( Ura.) apicalis, Ur. (Ura.) leucoptera, Ur. (Ura.)

lowii, Ur. (Ura.) pulcherrima, Culex (Cux.) coronator group, Cx. (Mel. ) bastagarius,

Cx. (Mel. ) dunni, Cx. (Mel.) educator, Cx. (Mel.) egcymon, Cx. (Mel.) elevator, Cx.

(Mel. ) ensiformis, Cx. (Mel.) erraticus, Cx. (Mel.) ocossa, Cx. (Mel.) panocossa, Cx.

(Mel.) zeteki group, Aedeomyia (Ady.) squamipennis, Mansonia (Man.) dyari and Ma.

(Man.) leberi. An. triannulatus is most abundant in Colombia during the dry season

(Bates and de Zulueta 1949:137) and is common in Panama from April to Septem-

ber, “from the end of the dry season well into the rainy season” (Arnett 1947: 198-

[99),

The adult females are primarily exophilic and zoophilic. They are rarely found in-

side houses. In French Guiana, Floch and Abonnenc (1944a: 1-7) collected 2102

anophelines in houses, of which 6 were triannulatus. Deane, Causey and Deane

(1948:901-904) reported that of 15,583 specimens of triannulatus collected, only

714 or 4.6% were taken in houses. They stated that triannulatus feeds outside, al-

though it will feed readily on man. Rozeboom (1938b: 294-297) in Panama found

that only 0.5% of the anophelines in houses were triannulatus, and they preferred to

feed on pigs rather than humans. He collected triannulatus in numbers around sta-

bles and pig sties. The only contrary finding was by Gabaldon (1949:765), who re-

ported a large number of triannulatus inside a house in the Rio Apure region of Ven-

ezuela. Hill (1934) collected triannulatus in the area around Maracay, Venezuela:in

stables, houses, dairy farms, and a farm and an orphanage where malaria epidemics

were a yearly occurrence. He performed precipitin tests on 262 blood engorged fe-

males and found 14 had fed on man, 173 on horses and 38 on cattle; 37 did not re-

act to the test. He concluded that triannulatus “definitely perfers animal blood.”

However, several investigators (Rozeboom 1935:527; Deane, Causey and Deane

1948:901-904; Floch and Abonnenc 1947:7) have stated that triannulatus is a vi-

cious biter, feeding readily on man outside even during the day although most active-

ly at dusk.

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 149

MEDICAL IMPORTANCE. Throughout most of its range, triannulatus does not

seem to be important as a vector of malaria. Only once triannulatus was found to

have a natural oocyst infection and that was in Cojedes, Venezuela (Gabaldon and

Cova Garcia 1946a:19-20). Benarrochi (1931) incriminated triannulatus as a possi-

ble vector of malaria at a boy’s school near Maracay, Venezuela, as it was the most

common species present during a marlaria epidemic. Hill (1934:428) stated that

with a high density “‘it is probable that this species can act as a malaria transmit-

toe

Several investigators (Godoy and Pinto 1923; Bonne and Bonne-Wepster 1925;

Rozeboom 1935; Floch and Abonnenc 1944a; Fonseca and Unti 1943:50) have ex-

perimentally infected triannulatus with Plasmodium vivax and P. falciparum. \n

comparing the susceptibility of triannulatus to infection with P. vivax and P. falcipar-

um with that of albimanus, Rozeboom (1935:524-525) found a larger percentage of

the specimens of triannulatus refractory to infection.

Because of the feeding behavior and low susceptibility to plasmodial infection, tri-

annulatus is probably not a serious health threat, except when present in very large

numbers.

DISTRIBUTION (fig. 1). An. triannulatus is widely distributed throughout South

America east of the Andes as far south as the Argentine provinces of Chaco, Corrien-

tes, Formosa, Jujuy, Salta, Santa Fe, Tucuman and Misiones. East of the Andes, it

occurs throughout Brazil, Paraguay, most of Bolivia, the Guianas, Colombia, Venez-

uela, Ecuador and Peru. On the Pacific side of the Andes, triannulatus extends as far

south as Tumbes, Peru. It is found in Central America as far north as central Nicara-

gua.

Material Examined: 2354 specimens: 545 males, 84 male genitalia, 1058 females,

301 pupae, 366 larvae; 262 individual rearings: 93 larval, 156 pupal, 13 incomplete;

3 progeny rearings.

ARGENTINA (52 specimens: 12M, 6Mgen, 32F, 1 lp). Chaco: Resistencia, 22 Feb 1927, R.

Shannon, 7M, 1Mgen, 5F; same data except 23 Feb, 1M, 4Mgen; same data except 25 Feb, 1Mgen;

same locality, Feb 1927, N. Davis, 2M, 1F [JH]; same locality, L. Rozeboom, 10M, 5F. Jujuy:

Ledesma, Mar 1923, N. Davis, 1F. Salta: Embarcacion, 20 May 1927, R. Shannon, 2M, 5F. Pro-

vince not specified: Salta Alto, Sierra, Apr 1960, 8F. Santa Barbara, Finca, 8 Apr 1927, E. Del

Ponte, 4F; same locality and date, R. Shannon, 2F. Santa Maria, 19 Nov 1962, R. Salta, 1F [AMN

H]. Locality not specified, N. Davis, 1 lp [JH].

BOLIVIA (63F). Beni: Guayaramerin, 19 Apr 1947, S. Blatman, 31F; same locality, 13 May

1947, 8F; same locality, 8 Jun 1947, 7F (47-12258); same locality, 23 Jun 1947, 12F (47-12258).

San Ignacio, 17 Sep 1943, 3052, KO 117-27, 1F; same locality, 18 Sep 1943, Torres Munoz, 3055-

A, KO 117-25, 1F. Trinidad, Aug 1946, E. Soracho, 3F.

BRAZIL. Amazonas (1F): Manaus, Jun 1931, R. Shannon, 1F. Bahia (7F): Bonfim, 26 Jan

1930, N. Davis, R. Shannon, 1F; same locality and collectors, 1F. Itapira, Mar 1931, 1F [JH]. R.

Barboza, 21 Apr 1930, J. Serafim, 1F. Locality not specified, H. Kumm, 3F [BM1931-302]. Mato

Grosso (3M, 2Mgen, 10F): Maracaju, Dec 1937, 2M, 1Mgen; same locality, 1937, 1M, 1Mgen, 1F.

Salobra, 1936, J. Lane, 6F. Westborder, May 1931, R. Shannon, 3F. Para (597 specimens: 71M,

IMgen, 346F, 127p, 52L&1; 127 ind rear: 401, 87p). Altamira, 150 km W of, 5 Nov 1974, J. Rei-

nert et al., coll. 77, 1 pM, 2 pF; same data except 163 km W of Altamira and 6 Nov, coll. 85, 3 lpM,

6 IpF, 6 pM, 1 pF; same data except 164 km W of Altamira, coll. 98, 2F; same data except 158 km

W of Altamira and 9 Nov, coll. 111, 1M, 3 lpM, 2lpF, 5 pM, 5 pF. Bacuri, Gleba 36, Lote 05, 23

Oct 1974, J. Reinert et al., coll. 31, 2 pF; same data except Lote 02, coll. 33, 1F, 5 lpM, 12 IpF, 16

pM, 13 pF; same data except 26 Oct, coll. 50, 1M, 2F, 3 IpM, 4 lpF, 3 pM, 1 pF; same data except

Gleba 29, Lote 03 and 25 Oct, coll. 44, 1 lpM, 1 lpF; same data except Lote 09 and 28 Oct, coll.

60, 2 pM; same data except Gleba 34, Lote 02, coll. 62, 1M, 1F, 4 pM, 5 pF; same data except Gle-

ba 38, Lote 02 and 29 Oct, coll. 64, 1 pF. Belem, BRA 42, BRB 9. Boa Vista, 12 Aug, C. Town-

150 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

send, 3F. Fordlandia, Jun 1931, R. Shannon, 1M, 1Mgen, 49F. Itupiranga, 17 Feb 197[6], Mora-

may, 1 pM (1050); same locality and collector, 1 Jun 1976, 2 pM, 1 pF (1200). Maraba area, 4

May 1976, 10F (M-6); same locality, 6 May 1976, 1F (M-8); same locality, 18 May 1976, D. Ro-

berts, 14F (R-1); same data except 19 May, 15F (R-2); same locality, 20 May 1976, 20F (R-3);

same locality, 22 May 1976, D. Roberts, 26F (R-5); same data except 23 May, 3F (R-3); same data

except 8 Jun, 6F (M-2); same data except 9 Jun, 4F (M-3); same data except 10 Jun, 5F (M-4);

same locality, 14 Jun 1976, 1F (M-6); same locality, 15 Jun 1976, D. Roberts, 6F (M-7); same data

except 16 Jun, 1F (M-8); same data except 24 Jun, 3F (A-1); same locality, 29 Jun 1976, 11F (M-

1); same locality, 30 Jun 1976, 7F (M-2); same locality, 1 Jul 1976, 6F (M-3); same locality, 2 Jul

1976, 8F (M-4); same locality, 3 Jul 1976, 10F (M-5); same locality, 6 Jul 1976, 2F (M-7); same

locality, 7 Jul 1976, 3F (M-8); same locality, 8 Jul 1976, 4F (M-9); same locality, 9 Jul 1976, 1F

(M-8); same locality, 12 Jul 1976, 7F (M-12); same locality, 17 Aug 1976, 1F (C-7); same locality,

9 Sep 1976, 1F (A-7), 1F (N-1); same locality, 8 Oct 1976, 3M (1262); same locality, 11 Oct 1976,

IF (1263); same locality, 19 Oct 1976, 6F (1); same locality, 21 Oct 1976, SF (1); same locality,

28 Oct 1976, 2F (1); same locality, 8 Nov 1976, 5F (1); same locality, 9 Nov 1976, 5F (2); same

locality, 10 Nov 1976, 1F (1), 1F (2); same locality, 12 Nov 1976, 2F (2); same locality, 19 Nov

1976, 1F (2); same locality, 1976, D. Roberts, 2M, 17F. Maraba area, Fonte Lenta, 1176, Kiu 13-

M-A, 1 pM, 2 pF. Palestina, 100 km SW of Maraba, 18 Aug 1976, Moramay, 1 pM (1045). Trini-

dad, Botanical Gardens, Oct 1924, J. Petroca, 1M. Pernambuco (1M, 8F): Recife, Sep 1929, 8F;

same locality, Oct 1929, R. Shannon, 1M. Rio Grande do Norte (11 specimens: 1M, 1Mgen, 8F, 1

L): Natal, Cajupiranga, 16 Nov 1943, MacCreary, Bricker, 3F; same locality, 19 Jan 1944, 1F;

same locality, 20 Jan 1944, 1M, 1Mgen; same locality, 24 Mar 1945, det. H. Dalmat, 1F [CU]. Na-

tal, Rio Cajupiranguinha, 21 Apr 1945, det. H. Dalmat, 1F [CU]. Natal, Lagoa Segui, 23 Apr 1945,

det. H. Dalmat, 1F [CU]. Rio Cajupiranga, on road to San Jose, 6 Jan 1945, H. Dalmat, 1L [CU].

San Jose, 13 Oct 1944, H. Dalmat, 1F [CU]. Pernambuco (1M): Afogados [de Ingazeira] , 1944,

H. Dalmat, 1M. Rondonia (14F): Porto Velho, Rio Madeira, May 1931, R. Shannon, 14F. Sao

Paulo (1M, 1Mgen): Juquia, J. Lane, 1M, 1Mgen. Sergipe (1F): Estancia, Dec 1929, R. Shannon,

1F. State not specified (2Mgen, 2F): Parahyba, 20 Mar 1930, 2Mgen; same locality, Mar 1930, R.

Shannon, 1F. Piraja, BRR 4, 1F.

COLOMBIA (270 specimens: 91M, 20Mgen, 63F, 45p, 51L&l; 8 larval ind rear; 3 progeny rear-

ings). Amazonas: Leticia, 25 Jul 1975, J. Kitzmiller, COZ 74, 2M, 2F. Antioquia: Arquia, L.

Dunn, 2F. La Pintada, COK 45A, 2M. Murindo, L. Dunn, 2F. Atlantico: Barranquilla, Nov-Dec

1924, L. Dunn, 1M, 9F; same locality, 24 Nov [1943], E. Palacio, COT 33, 1F. Calamar, 1966, J.

Bequaert, 1M [MCZ]. Boyaca: Lago de Palagua, COM 601, 602, 603. Puerto Boyaca, COM 604,

650, 651. Cordoba: Monteria, COB 12, COL 421, 427. Magdalena: Aracataca, Mar 1913, J. Eg-

bert, 1F. Meta: Villavicencio, 2 Jun 1942, W. Komp, KO 117-18, 19M, 4F; same locality and col-

lector, ?1942, KO 120A-23, 1M; same locality, 1 Jun 1944, COK 38, 4F; same locality, 15 Jun

1944, COK 44A, 4F; same locality, 1944, M. Bates, CV 90, 1F; same locality, 6 May 1947, L.

Rozeboom, CV-P 25 (progeny rearing), 16M, 3Mgen, 14F, 12p, 14L; same locality and collector,

1947, CV-P 31 (progeny rearing), 6M, 1Mgen, 6F, 13p, 13L; same data, CV-P 39 (progeny rearing),

12M, 5F, 12p, 13L; same locality, 9 May 1947, 1L. Department not specified: Cauca Valley, Jul

1943, W. Earle, KO 117-17, 1M. Locality not specified, COR 538, 1M, 1F; COR 508, 1F.

COSTA RICA (14 specimens: 6M, 1Mgen, SF, 2L). Province not specified: Barranca, H. Kumm,

No. 450, KO H-18-35, 3M, 1F; same locality and collector, No. 450, 2F [BM], 2M, 2L; same locali-

ty, 1M, 1Mgen, 2F.

ECUADOR (61 specimens: 12M, 1Mgen, 11F, 24p, 13 1; 23 ind rear: 13 1, 10p). Napo: Limon-

cocha, 4 Jun 1977, Y. Huang, coll. 48, 2 pF, 1 IpM; same locality and collector, 5 Jun 1977, coll.

49,2 pM, 1 pF, 6 lpM, 1 IpF, 1p. Tena, 1.5 km S of, 13 May 1977, Y. Huang, coll. 26, 1Mgen, 1

pM, 3 pF, 1 IpM, 3 IpF; same locality and collector except 12.5 km SE of Tena, 24 May 1977, coll.

38, 1 lpF. Pastaza: Puyo, 5-9 May 1977, Y. Huang, coll. 28, 1 pM.

FRENCH GUIANA (15 specimens: 5M, 2Mgen, 3F, 2p, 3 1; 2 larval ind rear). Guyane: Cay-

enne, 1943, H. Floch, ?3M, 2F; same locality, FG 122. Lac du Rorota, 15 Oct 1939, FGA 7-224,

IF.

GUYANA (55 specimens: 18M, 5Mgen, 28F, 1p, 3L; 1 p ind rear). East Berbice-Corentyne:

New Amsterdam, Jul 1936, 1Mgen. East Demerara-West Coast Berbice: Georgetown, 28 Sep 1945,

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 151

KO 117-21, 1F; same locality, 1 Oct 1945, KO 117-23, 1F; same data, KO 117-24, 1F; same local-

ity, GUY 40; same locality, W. Beveridge, 1M, 1F [BM193 1-210]. Hyde Park, 30 Jul 1941, BGR

3, 3M, 3Mgen. MacKenzie, Sep 1945, KO 117-26, 4F. District not specified: Coast, Sep 1949,

BGR 9, 2M, ?15F. Locality not specified, 1940, G. Bevier, KO 117-11, 8M, 1F; same date and col-

lector, 2M, 2F; 1942, G. Bevier, KO 117-4, 2F; no data, 3L (1B).

PANAMA AND CANAL ZONE (907 specimens: 291M, 37Mgen, 283F, 89P&p, 207L&I; 88 ind

rear: 281, 55p, Sinc). Bocas del Toro: Almirante, 1932, W. Komp, 1M, 1Mgen. Canal Zone: Bar-

ro Colorado Island, 20 Feb 1942, KO 111-14, 2F; same locality and date, KO 117-8, 1F. Cano

Saddle, 7 May 1923, R. Shannon, 6F. Empire, 16 Jun 1944, ASM 43-2, 7M, 4Mgen, aL \Cuy. 2.

Empire, Rio Mandinga, 24 Aug 1944, Wood, Van Doran, ASM 131-5, 3L [CU], 2L. Fort Clayton,

23 Apr 1944, 3F. Gamboa, 5 Apr 1939, W. Komp, 1M, 2F; same locality, 10 Jun 1943, Col. Elton,

OF; same locality, 10 Jun 1943, W. Komp, 2F; same locality, 22 Jun 1944, Stearns, Middlekauff,

ASM 12-1, 1L [CU]; same locality, 28 Sep 1944, Van Doran, Turney, ASM 194, 2L; same data, |

L [CU] ; same locality, 6 Jan 1947, PAX 239, 4M, 4F; same locality, 28 Jan 1947, 5M, 5F; same

locality, 18 Apr 1947, H. Brooks, 3F, 3 1; same locality and collector, 22 May 1947, 1M, 4F; same

locality and date, 1M; same locality, PA 912A. Gatun Lake, Apr 1935, PAX 47, 38M; same locali-

ty, 23 May 1939, 2M; same locality, 8 Aug 1940, W. Komp, 1Mgen. Juan Mina, Rio Chagres, 23

May 1939, 3M; same locality, PA 1122, 1124, 1125, 1160, 1161. La Pita, 11 Feb 1922, J. Shrop-

shire, 1M, 2F. Mandinga Bridge, 22 Jan 1942, 3M, 4F. Mendoza, Gatun Lake, 18 May 1939, 1M.

Pedro Miguel, 27 Jan 1947, 1M, 5F. Rio Chagres at Las Guacas, 23 Feb 1940, PAX 174, 14M, 14

F; Rio Chagres, below Las Guacas, 31 Mar 1944, E. Pierce, KO 117-14, 4M, 2F. Rio Chagres, 12

Feb 1942, 1M, 1F; same locality, 12 Feb 1943, KO 117-2, 20M, 19F; same locality, 31 Mar 1944,

3 1; same locality, 5 Jan 1947, 2M, 3F; probably same locality, KO 117-16, 2M, 2F. Rio Chilibre,

PA 27. Rio Gatun, 10 Apr 1945, KO 117-22, 6F. Rio Gatuncillo, PA 1136. Rio Mandinga, Aug

1937, W. Komp, KO H-18-31, 11M. Rio Paja, 5 Jan 1934, 1M, 21; same locality, 1 1. Locality not

specified, Feb 1933, D. Curry, 2M, 1F [BM1933-504] ; 26 Feb-22 Mar 1944, H. Hoogstraal, 1M.

Cocle: La Vieja, 7 Nov 1936, L. Rozeboom, 3L [JH]. Colon: Colon, 24 Jun 1921, W. Chidester

thru J. Zetek, 1F. Santa Rosa, 30 Aug 1935, L. Rozeboom, 2L. Panama: Juan Mina, 6 Aug 1943,

G. Fairchild, KO 117-3, 13F; same locality and collector, 5 Oct 1943, KO 117-6, 6F; same locality

and collector, KO 117-5, 4F; same locality, PA 12,31. La Chorrera, 31 May 1944, ASM 29, IL

[CU], 1L; same locality, 19 Sep 1944, Van Doran, ASM 168, 1L; same locality, 27 Nov 1944,

Wood, Griffing, ASM 298-1, 5L [CU]; same data, ASM 299-2, 1L [CU] ; same locality, 7 Dec 1944,

K. Frick, R. Arnett, ASM 333, 3L; same locality, 8, 9 Jan 1945, Wood, Van Doran, ASM

370-2, 1F; same locality, 13 Jan 1945, K. Frick, ASM 377, 1L. Las Guacas, Dec 1934, 11M, 7M

gen, 3F; same locality, 5 Feb 1935, PAX 42, 81M, 67F; same locality, 4 Apr 1935, 15M, 4Mgen;

same locality, 14 Aug 1936, L. Rozeboom, 2L; same locality and collector, 6 Feb 1937, 1F; same

locality and collector, 19 Mar 1937, 1F; same locality and date, 2M, 1F [AMNH] ; same locality,

L. Rozeboom, 24 May 1937, 1M, 1Mgen; same locality, 6 Aug 1937, 1L [JH]; same locality, Aug

1937, 1M, 1Mgen. Pacora, La Joya, Feb 1945, KO 117-20, 2F. Pedregal, PA 1163. Pequeni, PA

942. Rio Abajo, 7 Jul 1944, K. Frick, R. Arnett, ASM 59, 1L. Tocumen, PA 557, 559, 1066,

1144, 1145, 1148, 1162. Province not specified: Rio Pescado, 16 Jul 1936, L. Rozeboom, 1L.

Locality not specified, 17 Dec 1934, 1L; 14 Aug 1936, L. Rozeboom, 4L; 1937, 3M, 8F; 1937,

621001-7, 8, 1M, 1F.

PARAGUAY (2M, 2Mgen, 7F). Capital: Asuncion, 1925, F. Soper, 1Mgen; same data, 1M

[JH]. Central: Asuncion, 12 km on rd. to Villa Rica, 10 Oct 1942, KO 111-1, 3F. La Cordillera:

s bes Fiebrig, 1M, 1Mgen, 2F. Department and locality not specified: 1922, Fiebrig, 2

F [BM].

PERU (86F). Loreto: Iquitos, Mar-Apr 1931, R. Shannon, 68F; same locality, 22 Jul 1975, J.

Kitzmiller, PERZ 33, 2F. San Antonio, 17 Aug 1965, J. Hitchcock, Jr., 2F; same locality and col-

lector, 18 Aug 1965, 3F; same locality and collector, 21 Aug 1965, 1F; same locality and collector,

23 Aug 1965, 4F; same locality and collector, 25 Aug 1965, 6F.

SURINAME (27 specimens: 3M, 1Mgen, 1F, 4p, 18L&l; 4 ind rear: 2p, 2inc). Paramaribo:

Paramaribo, J. Bonne-Wepster, 1F. Suriname: Ma Retraite, SUR 206. District not specified:

Kropie Kreek, 25 Sep 1974, 16L. Nord, Kabelstation, 26 Jun 1946, E. Kuyp, 1M, 1Mgen, 2 lp.

152 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

VENEZUELA (132 specimens: 27M, 3Mgen, 79F, 8p, 1SL&1l; 8 ind rear: 21, 1p, 5inc). Aragua:

Maracay, 20 Aug 1926, M. Nunez Tovar, 9F; same locality and collector, 3 Sep 1926, 16F; same

locality and collector, 9 Oct 1926, 11F; same locality and collector, 10 Oct 1926, 9F; same locali-

ty, 11 Sep 1926, 1F; same locality, 12 Mar 1927, 1F (No. 9); same locality, 10 May 1927, 1F

[JH] ; same locality, 12 May 1927, F. Root, 1F [BM1929-194] ; same locality and collector, 24

May 1927, 2M, 2F [BM1929-194]; same locality, 17 May 1927, 3L (17) [JH] ; same locality, 31

May 1927, 1M (No. 45); same locality, 7 Jun 1927, 3 Ip (Nos. 54a, b, d) [JH] ; same locality, 8 Jun

1927, F. Root, 1M [BM1929-194] ; same locality, 18 Jun 1927, 1 lp (77a) [JH] ; same locality, 21

Jun 1927, 1 lp (83a) [JH]; same locality, 1L. Carabobo: Mariara, VZ 68. Valencia, 10 Aug 1927,

M. Nunez Tovar, 1M, 1F. Cojedes: Rd. to San Carlos, 10 Dec 1941, H. Hanson, KO 111-6, 5F.

Distrito Federal: Caracas, 19 Apr 1938, A. Gabaldon, KO H-18-32, 7F; same data, KO H-18-34,

18M, 4F. Guarico: Calabozo, 22-31 May 1941, VZK 48, 1M, 3F. Chaguaramas, 4 Jul 1945, E.

Winton, 1M. Monagas: Caripito, KO 117-12, 1M, 1F; same locality, 5 Sep 1936, KO 117-15, 1M,

2F. State not specified: Cap El Callai, CDC, 2L. Locality not specified, 4 Mar 1937, Anduze, 2M

gen [CU] ; 1949, 1Mgen, 2L; no data, 8L, 2F.

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FIGURES

Distribution of albimanus, triannulatus, strodei, rondoni and benarrochi

Distribution of oswaldoi, noroestensis, aquasalis, ininii and anomalophyllus

Distribution of galvaoi, rangeli, trinkae and nuneztovari

Hypothesized phylogenetic tree of Albimanus Section

Anopheles (Nyssorhynchus) albimanus: male and female heads and claws; male

foretarsus 5; female thorax, legs; wing and abdomen

6. Wings of oswaldoi, galvaoi, noroestensis, aquasalis, ininii, anomalophyllus and

rangeli

7. Wings of trinkae, nuneztovari, strodei, rondoni, benarrochi and triannulatus

8. Anopheles (Nyssorhynchus) albimanus: male genitalia and pupa

9. Anopheles (Nyssorhynchus) albimanus: larva

10. Anopheles (Nyssorhynchus) oswaldoi: male genitalia and pupa

11. Anopheles (Nyssorhynchus) oswaldoi: larva

12. Male genitalia of rondoni and galvaoi

13. Anopheles (Nyssorhynchus) noroestensis: male genitalia and pupa

14. Anopheles (Nyssorhynchus) noroestensis: larva

15. Anopheles (Nyssorhynchus) aquasalis: male genitalia and pupa

16. Anopheles (Nyssorhynchus) aquasalis: larva

17. Anopheles (Nyssorhynchus) ininii: male genitalia and pupa

18. Anopheles (Nyssorhynchus) ininii: larva

19. Anopheles (Nyssorhynchus) anomalophyllus: male genitalia and pupa

20. Anopheles (Nyssorhynchus) anomalophyllus: larva

21. Anopheles (Nyssorhynchus) rangeli: male genitalia and pupa

22. Anopheles (Nyssorhynchus) rangeli: larva

23. Anopheles (Nyssorhynchus) trinkae: male genitalia and pupa

24. Anopheles (Nyssorhynchus) trinkae: larva

25. Anopheles (Nyssorhynchus) nuneztovari: male genitalia and pupa

26. Anopheles (Nyssorhynchus) nuneztovari: larva

27. Anopheles (Nyssorhynchus) strodei: male genitalia and pupa

28. Anopheles (Nyssorhynchus) strodei: larva

29. Anopheles (Nyssorhynchus) benarrochi: male genitalia and pupa

30. Anopheles (Nyssorhynchus) benarrochi: larva

31. Anopheles (Nyssorhynchus) triannulatus: male genitalia and pupa

32. Anopheles (Nyssorhynchus) triannulatus: larva

WR WN

Albimanus Group joe

qrope

Triannulatus Subgroup

: triannulatus

strodei.

Equator ve aa ae

strodei complex — beskrsoeh

NORTH & SOUTH AMERICA

KILOMETERS

1000 1500 2000

500 1000

STATUTE MILES

§ MOSQUITOES OF MIDDLE AMERICA UCLA ~ ZOOLOGY - ENT ~ 1971

NYSSORHYNCHUS

anomalophyllus — ss :

4 °

oestensi

Le? SX

LOS 7” nor 8

}

{

(

{

NORTH & SOUTH AMERICA

KILOMETERS

2000

1000

STATUTE MILES

MOSQUITOES OF MIDDLE AMERICA UCLA ~ ZOOLOGY - ENT - 1971

a Wy .

( MEL,

Wig SM)

i, a YP iy

nuneztovari

NORTH & SOUTH AMERICA

KILOMETERS

1500 2000 2500 3000

1500 5 2000

1000

STATUTE MILES

MOSQUITOES OF MIDDLE AMERICA

‘UOI]IOS SNueWI|\yY 94} JO

9014] oljoueBojAud peziseyjodAy fy ‘Bi

dnoig

lOp[|EMSO

dnoibqns dnoig

ne ies @) SNURWI|\/

dnoi6qns

snjejnuueiy

x9|dwos

lopjemMSO ere

ey” Ny WS o a Ay ” w se oO A 2 5 oo yo

2 M4 “bg S 6? ig \ ; ¢) R & N4 N) m)

\\ as . & “ : 0 \ \ \) \ « J

oe oN y? : ” o ; o Or of

\) oe ws a AN

basal humeral subbasal presectoral sectoral subcostal preapical apical FORE MID HIND

2 o I eee < oe See Ch Ge << Ki =<

19. ee eee RE EE

hee > SS

ee A Y)

‘ iy

~ RY Vy iN

| Wy iy nny aa ~ <r 2 ms

ES Ta TR Sage NWA i

oy 13 A RN

ZZ 14) NN ts

_ NYSSORHYNCHUS i

| in

yin

SES

SS QoSSOo fi

SSS

WSs

SSS

YZ pg ly;

DORSAL VENTRAL

Se ae y

IN 4

Kes Yh

tal \

: a

, Wy

\ ym

\ N a

im \\

YIN a \

Sas

SELLE LEIIZOO

NYSSORHYNCHUS Fig. o.

5 ie

ST CZZZ SS

oswaldoi

ar mn mM ih AT

wy) NNN AUN AN ae \\\\\\ AN

ii ial NN i m i i HAAN iM \

galvaoi

noroestensis

Mi Hh , Ni iy i i j\ RS nai TA AY. . Nh

iii aa: Ih: aii m ie zi SN

aquasalis

Hi | a es RAN 2

mi)

anomalophyllus

i iy

iii iis

ny Hh i

TN \

HS a

i r i we

ht am ih rR oT iN

btn n

La nam ohld di A

rangeli

Re i NNN Kh it Coe a .% =~ ey N\\\\ Se

NV IY XI. «AY

A AA I

trinkae

Wha

mill) Hi Wi Wil ih iii Tw Nui

KN li

a iii

: NV...

ne RR ie " we NN ay wi AN

1 s\\\

HAVANA Ma TORT

Ni ANTE

Hutt iii

nt i

Ih ne

7 vot : a ‘\

nt AL i)

nuneztovari

strodei

rondoni

benarrochi

triannulatus

Tip,

rnin

]

=3Af

i Hii TARP,

Fig.8

NYSSORHYNCHUS

(OIG)

fused ventral lobes

ventral aspect

imanus

alb

Lf \0

NYSSORHYNCHUS

Fig. 10

NYSSORH YNCHUS

—tergomedial bristle

a oswaldoi

% median sulcus

— subapical bristle

' \ppreapical plate

refringent structure pat 3

- membranous area

= membrano e

——mesal cleft

fused ventral lobes

ventral aspect

WE a

‘ N 2 a

‘\I} | Se

(\ i Ww la\ {~~ dorsal claspette

AN parabasal spine ‘aS L of apex

\\ \ f of) :

anterior apodeme —==_.

_ NYSSORHYNCHUS

oswaldoi

Fig. 12

fused ventral lobes

ventral aspect

NYSSORHY NCHUS

\ QR \

SNS \'

SSN AW \\

\ ‘ \

X \

SS \ \

SL oe \

= \

SA ee

fused ventral lobes

ventral aspect

NYSSORHYNCHUS

fused ventral lobes

ventral aspect

Fig. 14

NYSSORHYNCHUS| | -

noroestensis

SWS S <

PSEC =

SSS

ae =

fice aaa

Fig. 15

NYSSORHYNCHUS

fused ventrai lobes

ventral aspect

QU NN | )

x “asin,

\ Z BA as

| = 4 7 3 y

Ne f 3 z —/

Jaa \ F

oe \\/ 4

. \,

IN | Lip

aquasalis

ig. 16 \

NYSSORHYNCHUS

Fig. 17

NYSSORHYNCHUS

—_

——

—-

fo \

= ——

S ‘

fused ventral lobes

ventral aspect

anomalophyllus

NYSSORHY NCHUS

STV

Fig. 21

NYSSORHYNCHUS

fused ventral lobes

ventral aspect

NYSSORHY NCHUS

rangeli nk

Fig. 23

NYSSORHYNCHUS

poe a

: = >

yy.

* \

| |

gh Y

AS far vi

‘Vit

\\ \ 1

—— subapical seta \ \

) i \

VA Wa ey)

SW Ys mesal cleft

yr \

\ }

oN \Y\

< basal lobule

aedeagus

fused ventral lobes

ventral aspect

accessory spine

sternite IX

oD s eee ;

Y La one ——= = anterior apodeme

Fig. 24

NYSSORHY NCHUS

trinkae

: Fig. 25

NYSSORHYNCHUS

fused ventral lobes

ventral aspect

ae” &

SOTERA

INNS

nuneztovari

_ NYSSORHYNCHUS

Fig. 27

NYSSORHY NCHUS

strodei

fused ventral lobes

ventral aspect

Fig. 29

NYSSORHYNCHUS

ig. 30

NYSSORHYNCHUS

benarrochi

ONE ig 31 \y

NYSSORHY NCHUS

triannulatus

===

= sl

= =

fused ventral lobes

ventral aspect

‘\ \

\\, ®

ey:

Faran: Albimanus Section of Anopheles (Nyssorhynchus) 211

TABLE OF DISTRIBUTIONS

DISTRIBUTION

7. anomalophyllus

op)

“=

™

2. oswaldoi

3. galvaoi

4, novroestensis

5. aquasalis

8, vangeli

9, trinkae

0. nuneztovari

3. benarrochi

4, triannulatus

ba)

NY)

As)

ie)

™

6, tninit

2. rondoni

™s 1 NS | ™S | om

ee ss eee ae

Dee eee a

renee 1c aa ee Od ae

a ee ie

[GU ATEMAS 6054 pve in MD os ed | als DL helene

Se es eee eee

| amiiaeameer) 0d seat MO aia TE SI Hee be fe

a eee ea ee ee

ee eee | ee

| cosrarica —-s_s§$§s§»§ «(@/e| | |e! je | || jo le

we (0 | @ @ || ee

PANAMA coms (QO | el | | | |e | le

at ele | io ||| ee | lo

en ee eed | le

sown | 1011 | leet 11 le

pant je ie TT iele

ane Ol || et le TI le

commas «cartes (@IQ]_| | | | @_ OO | |e

ee e ee

venezuera commen (WO! | @ | (@ lee Ce

Orinoco = | /@h| Ld| ld] | ie leo | le

BAHAMAS Ae ee ee

GREATER ANTILLES ee

LESSER ANTILLES °°" *"4 eo | | ie] Dee ai a

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212 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

CONSPECTUS OF TAXONOMIC CHANGES

Resurrected From Synonymy

strodei Root 1926, from synonymy with evansi (Brethes 1926), to specific rank, p

129-130.

New Synonymies

albertoi Unti 1941, from variety of evansi (Brethes 1926), to synonymy with strodei

Kool? 20, P1235, (6:

arthuri Unti 1941, from variety of evansi (Brethes 1926), to synonymy with strodei

Roegpi926, p.123, 128.

bachmanni Petrocchi 1925, from subspecies of triannulatus (Neiva and Pinto 1922),

to synonymy with triannulatus, p 141, 147.

deltaorinoquensis Cova Garcia, Pulido F. and Amarista M. 1978, to synonymy with

aquasalis Curry 1932, p 75, 83-84.

lloydi Unti 1941, from variety of evansi (Brethes 1926), to synonymy with strodei

Root 1926, p i23,,128.

ramosi Unti 1940, from variety of evansi (Brethes 1926), to synonymy with strodei

Kooe 1976,.p. 123.

Transferred Synonymies

artigasi Unti 1941, from synonymy with arthuri Unti 1941, to synonymy with stro-

der Root 1926") 123,128.

davisi Paterson and Shannon 1927, from synonymy with bachmanni Petrocchi 1925,

to synonymy with triannulatus (Neiva and Pinto 1922), p 141, 147-148.

metcalfi Galvao and Lane 1937, from synonymy with oswaldoi (Peryassu 1922), to

synonymy with noroestensis Galvao and Lane 1937, p 67, 73.

perezi Shannon and Del Ponte 1928, from synonymy with bachmanni Petrocchi

1925, to synonymy with triannulatus (Neiva and Pinto 1922), p 141, 147.

Nomina Dubia

evansi (Brethes 1926), to nomen dubium, p 129-130.

gorgasi Dyar and Knab 1907, from synonymy with albimanus Wiedemann 1820, to

nomen dubium, p 14.

Faran: Albimanus Section of Anopheles (Nyssorhynchus)

213

INDEX TO SCIENTIFIC NAMES

Invalid taxa are in italics; valid taxa treated in detail by present paper are in bold-

face. Major page references are in boldface; page reference to key is shown by suffix

**k’’; figure numbers are in italics.

albertoi (Anopheles), 8, 123, 128

albimanus (Anopheles), 1, 2,3, 4, 5, 6, 7, 8, 10,

fl ASA BOR he 18.20 2h OD. 25:

26, 27k, 29k, 32k, 35-49, 51, 82, 84, 85,

118, 128, 134, 149, figs. 1, 4,5, 8 9

albimanus of authors, 6, 56, 68, 77, 113, 124,

133

Albimanus Group, 2, 9, 11, 13, 14, 27k, 29k,

32k, 35, 40, 51, 146, 148

Albimanus Group of authors, 5, 8

Albimanus Section, 1, 2, 5, 6, 7, 8, 9, 11, 12,

13, 14-16, 18-26, 41, 83, 98

Albimanus Series, 5, 8

Albimanus Subseries, 8

albipes (Anopheles), 4, 6, 7, 8, 35, 68, 77

albitarsis (Anopheles), 6, 7, 74, 140

allopha (Anopheles), 120, 130

amazonensis (Culex), 94

angustivittatus (Aedes), 42

anomalophyllus (Anopheles), 2, 4, 8, 11, 12,

135 MO cA ieud ?. 30K, 31, 34k, SO. 52.54.

205 125 99799..127, fies. 2, 4,.6,.19,.20

Anopheles, 8

Anopheles (Anopheles) sp, 94, 120, 130

apicalis (Uranotaenia), 43, 148

apicimacula (Anopheles), 42, 85, 130, 148

aquacaelestis (Anopheles), 7, 55, 61

aquacoelestis (Anopheles), 56, 61

aquasalis (Anopheles), 2, 3, 7, 8, 10, 13, 14, 16,

D7, AS 2G 8 32k 33k, 41.42, 52. 54.

57, 58, 59, 60, 61, 62, 66, 72, 75-89, 91, 93,

99; 90/97, 100.101, 102,415, 118.120,

122, 126, 128, 134, 136, 139, 148, figs. 2,

4, 6, 15, 16

argyritarsis (Anopheles), 6, 43, 104, 118, 130

Argyritarsis Group, 8

Argyritarsis Section, 1, 2,6, 8, 10, 11, 14-15,

41, 93, 146

Argyritarsis Series, 8

argyrotarsis (Anopheles), 6, 8

arthuri (Anopheles), 8, 123, 128

artigasi (Anopheles), 8, 123, 128

atratus (Culex), 42, 85

ayrozai (Anopheles), 67, 68

bachmanni (Anopheles), 2, 6, 7, 141, 142, 147

bahiense (Culex), 130

bastagarius (Culex), 42, 94, 111, 118, 130, 148

bathana (Chagasia), 85

benarrochi (Anopheles), 2, 3, 8, 12, 13, 16, 17,

24, 26, 206, 29k, o7%, 33k, 51, 52,53, 66,

72, 81, 122, 136-141, figs. 1, 4, 7, 29, 30

bigotii (Anopheles), 8

bisignatus (Anopheles), 35, 42, 46, 84, 134

braziliensis (Anopheles), 94, 120

cancer (Deinocerites), 43

Cellia, 6, 8

chagasi (Anopheles), 8, 141, 148

Chagasia sp, 42

chrysonotum (Culex), 42, 111, 130

cingulata group (Psorophora), 42, 85, 130

clarki (Anopheles), 67, 68, 74, 129

coatzacoalcos (Uranotaenia), 130

confinnis group (Psorophora), 42, 104

conspirator (Culex), 42, 130

Corethrella spp, 43

corniger (Culex), 104

coronator group (Culex), 42, 61, 74, 85, 94,

104, 118, 130, 148

crinifer (Aedes), 130

crucians (Anopheles), 42

cubensis (Anopheles), 6, 35

Culex (Culex) sp 72, 130

Culex (Culex) sp 85, 130

Culex (Culex) sp, 74

cuyabensis (Anopheles), 4, 6, 7, 132, 134, 141,

142, 147 |

darlingi (Anopheles), 7, 10, 41, 82, 94, 118,120

davisi (Anopheles), 2, 4, 8, 141, 142, 147-148

declarator group (Culex), 42, 85, 104, 130

delta (Anopheles), 75, 83, 134

deltaorinoquensis (Anopheles), 75, 83-84, 134

Dixella sp, 43

dunhami (Anopheles), 4, 112, 117, 118, 121

dunni (Culex), 42, 130, 148

dunni group (Culex), 94, 118

dupreei group sp B (Aedes), 130

214

dureti (Culex), 130

dyari (Mansonia), 43, 148

educator (Culex), 42, 130, 148

egcymon (Culex), 130, 148

eiseni (Anopheles), 130

elevator (Culex), 148

emilianus (Anopheles), 4, 7, 75, 76, 82-83, 87

ensiformis (Culex), 118, 148

erraticus (Culex), 42, 130, 148

evansi (nomen dubium), 2, 7,68, 124, 129-130

galvaoi (Anopheles), 2,3,4, 8, 16, 17, 18, 27,

31k, 32, 54, 60, 64-67, 69, 72, 82, 93, figs.

3, 4,6, 12

gambiae complex (Anopheles), 128

geometrica (Uranotaenia), 42, 61, 94, 118

goeldii (Anopheles), 4, 7,66, 106, 111, 112,

113, 107,118,171

gorgasi (nomen dubium), 1-2, 4,6, 7, 14, 113

_ grabhamii (Anopheles), 42

guarauno (Anopheles), 75, 83, 134

guarujaensis (Anopheles), 75, 83

habilitator (Culex), 43

howardii (Psorophora), 43

idottus group (Culex), 85

inadmirabilis (?) (Culex), 130

inhibitator (Culex), 43

ininii (Anopheles), 2, 4, 8, 10, 11, 12, 13, 15,

16, 17,48,.20, 26, 27k, 30k, 31k, 34k, 50,

51, 52, 53, 54, 55, 60, 61, 89-94, 118, 120,

148, figs. 2, 4, 6, 17, 18

iolambdis (Culex), 42, 130

jamaicensis (Psorophora), 42

janitor (Culex), 43

Kerteszia, 1, 8

konderi (Anopheles), 8, 55, 56, 61

Laverania, 7-8

leberi (Mansonia), 148

leucoptera (Uranotaenia), 148

lloydi (Anopheles), 8, 123, 128

lowii (Uranotaenia), 42, 61, 74, 85, 104, 130,

148

maculipennis complex (Anopheles), 128

madininensis (Culex), 43, 85

magnus (Deinocerites), 85

maracayensis (?) (Culex), 43

mattogrossensis (?) (Anopheles), 120

Melanoconion spp, 42, 74, 94, 130

metcalfi (Anopheles), 2, 4, 7, 18, 61, 62, 67,

68, 73, 74, 77

mollis (Culex), 94, 130

Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980

Myzomyia, 8

Myzorhynchella, 1, 8

neomaculipalpus (Anopheles), 42,61, 130, 148

nigripalpus (Culex), 42, 85, 130

noroestensis (Anopheles), 2,4, 7, 8, 10, 12, 16,

18, 28k, 30k, 31k, 34k, 52, 53, 54, 60, 61,

62, 65, 66, 67-75, 82, 93, 98, 103, 104, 127,

129, fies. 2,4, 6, 135, [4

nuneztovari (Anopheles), 2, 8, 10, 12, 13, 15,

1S, t7, 28k, 50K; 32k, 33k. 41,50; 51, 352,

53, 54, 55, 60, 61, 66, 82, 91, 93, 94, 103,

107, 109, 110, 111, 112-122, 129, 148, figs.

5, 4, 7,25, 26

nuneztovari of authors, 106,111, 117,118

Nyssorhynchus, 1, 5,6, 8, 10, 11, 14, 15, 73

Ochlerotatus sp (Aedes), 130

ocossa (Culex), 43, 148

oswaldoi (Anopheles), 2,3, 7, 8, 10, 12, 13, 14,

16, 18, 28k, 31K, 32k, 33k; 41,52, 54, 55-

63, 65,66, 69, 70, /1, 72, 73, 80, 82,83.

89, 93, 94, 99, 104, 118, 120, 133, 148, figs.

2,4, 6, 10,11

oswaldoi of authors, 6

Oswaldoi Complex, 2, 10, 11, 12, 13, 15, 16,

29k, 30k, 53-55, 98, 102, 122, 146

Oswaldoi Group, 2, 11, 13, 14, 15, 27k, 29k,

32k, 40, 41, 50-52, 93, 116, 127

Oswaldoi Series, 8

Oswaldoi Subgroup, 2,9, 10, 13, 15, 16, 27k,

29%, 35k. 31, 50,31, 52-53,.55, 60, 50,31

98, 103, 109, 134, 139, 140, 145, 146

Oswaldoi Subseries, 8

panocossa (Culex), 148

perezi (Anopheles), 141, 142, 147

peryassui (Anopheles), 140

pharoensis (Anopheles), 8

pilosus (Culex), 42, 111, 130

pseudes (Deinocerites), 43

pseudomaculipes (Anopheles), 73

pseudopunctipennis (Anopheles), 42, 85, 135

pulcherrima (Uranotaenia), 42, 94, 130, 148

pulcherrimus (Anopheles), 8

punctimacula (Anopheles), 42, 61, 85, 104

pygmaea (Psorophora), 43 |

quadrimaculatus (Anopheles), 73

rachoui (?) (Culex), 94, 118

ramosi (Anopheles), 123, 124

rangeli (Anopheles), 2,3, 8, 10, 12, 16, 17, 18,

26k: 30k, 32k, 52, 54, $5, 60, 61,66, 72; 93,

99-106, 109, 110, 111, 114, 116, 117, 119,

Faran: Albimanus Section of Anopheles (Nyssorhynchus)

122, 128, 130, 148, figs. 3, 4, 6, 21, 22

rangeli of authors, 111

rondoni (Anopheles), 2, 3, 4, 6, 7, 8, 10, 13,

1S, 16, 17 ¥6, 20; 27k 29k. 25,32, 50:32,

53, 67, 122, 129, 132-136, 140, 147, figs. 1,

4,7, d2

rondoni of authors, 42

Rondoni Complex, 8

Rondoni Series, 8

sanctielii (Anopheles), 1, 8, 14

scapularis (Aedes), 42

socialis (Uranotaenia), 42

sollicitans (Aedes), 42

squamipennis (Aedeomyia), 43, 148

squamosus (Anopheles), 8

strodei (Anopheles), 2,4, 6, 8, 12, 13, 14, 15,

16, 17, 18, 28k, 29k, 31k; 34k, 50; $1.52.

235 Day 1o, 8h, 90, 96; 90s 1045 FID, T2U,

122, 123-132, 134, 135, 139, 140, 147,

148, figs. 1, 4, 7, 27, 28

Strodei Complex, 2, 10, 11, 13, 15, 16, 29k,

53, 54, 35,98, 122-123, 146

taeniorhynchus (Aedes), 42, 85, 86

tarsimaculatus.( Anopheles), 5-7, 35, 56, 60,

61, 66,/67,68. 73, 75, 76, 77, 82; 83. 86,

113, 124,229,133, 134, 135, 142

Tarsimaculatus Complex, 5, 8

Tarsimaculatus Series, 5, 8

thriambus (Culex), 130

titillans (Mansonia), 43

tortilis (Aedes), 42, 85

triannulatus (Anopheles), 2, 3, 6, 7, 8, 10, 11,

V2, 45, Pete, £6, 17,19, 20, 26, 27k, 29k,

I1K, 23h, et, 42, $0, 51, 52,53, 61, 85, 94,

104,111, 118, 120, 126, 134, 140, 141-

132, fas 34 7, 31, 22

Triannulatus Complex, 8

Triannulatus Series, 8

Triannulatus Subgroup, 2, 9, 10, 13, 15, 27k,

29k, 33k, 50, 51, 53, 141, 145-146

Triannulatus Subseries, 8, 53

trinkae (Anopheles), 2,3, 10, 12, 13, 15, 16,

V7,:13, 20,006, 20m, 32k: 33k. $0,'51,:52°

54, 55, 103, 104, 106-112, 116, 117, 118,

121, .127, figs..3, 4, 7, 23, 24

trisignatus (Anopheles), 35, 42, 46, 84, 134

Uranotaenia (Uranotaenia) sp, 130

zeteki group (Culex), 94, 118, 148

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