A NEW SUBGENUS of the GENUS Sabei;Hes (DIPTERA: CULICIDAE) L

AUGUST1991 1 A NEW SUBGENUS OF THE GENUS sABEI;HEs (DIPTERA: CULICIDAE) l

RALPH E. HARBACH~

Walter Reed Biosystematics Unit, Department of Entomology, Walter Reed Army Institute of Research, Washington, DC 20307-5100.

ABSTRACT. A new subgenus, Peytonulus, of the genus Sabethes Robineau-Desvoidy is estab- lished for seven species previously included in the subgenus Sabethinus Lutz. The subgenus is contrasted with the other subgenera of Sabethes and the type species is illustrated.

INTRODUCTION The new subgenus is uniquely characterized by several autapomorphic features, the highly Because mosquitoes of the genus Sabethes modified larval seta l-VII and its missing pupal Robineau-Desvoidy are known to harbor and homolog being the most notable and conspicu- transmit arboviruses (Galindo et al. 1959, Mat- ous. Based on these distinctive features, the tingly et al. 1973), information on their identifi- subgenus PeytonuZus is erected for the seven cation andphylogenetic relationships isof great species listed above, and the following informa- importance. This is the third in a series of tion isprovided for its separation from the other papers that deals with taxonomic problems in- subgenera within the genus Sabethes. volving nominal taxa within this genus. The first The descriptive terminology and abbrevia- paper dealt with the transfer of a species from tions follow Harbach and Knight (1980, 1982) Sabethesto a new subgenusin WyeomyiaTheobald and Harbach and Peyton (1990a, 1990b). The (Harbach and Peyton 1990a). The second dealt illustrations are based on specimens deposited with the transfer of the subgenus Davismyia in the National Museum of Natural History, Lane and Cerqueira and its type species from Smithsonian Institution. Wyeomyia to Sabethes (Harbach and Peyton 1990b). TAXONOMIC TREATMENT On reviewing the genus Sabethes, it became apparent that the species presently included in Sabethes subgenusPeyr~nulus, the subgenussabethinus Lutz actually belong to New Subgenus two separate phyletic lines. In this paper, a new subgenus is introduced for seven of these species: Type species (Figs. l-3). Sabethinus aures- aurescens (Lutz), fabricii Lane and Cerqueira, tens Lutz, 1905. gorgasi Duret, identicus Dyar and Knab, soperi Lane and Cerqueira, undosus (Coquillett), and Sabethes (Sabethinus) in part of Lane and Cer- whitmani Lane and Cerqueira. This action leaves queira, 1942: 481,659-662,676,679-688; Lane, only intermedius Lutz, the type species, and 1953: 10551060,1082-1096; Stone et al., 1959: 93-94; Knight and Stone, 1977: 307-308; and melanonymphe Dyar within the subgenus Sabe- thinus. authors. Sabethoides (Sabethinus) in part of da Costa Lima, 1931: 56,59,60-61,62-63; and authors. lThe views of the author do not purport to reflect the Sabethinus in part of Theobald, 1907: 618; viewsof the Department of the Army or the Departmentof Theobald, 1910: 586; Howard et al., 1915: 31- Defense. 2Reprint requests: Walter Reed BiosystematicsUnit, 37; and authors. Museum Support Center, SmithsonianInstitution, Wash- Sabethes(Sabethoides) in part of Dyar, 1924: 97- ington,DC 20560. 98; and authors. 2 MEQUITO SYSTEMATICS VOL. 23, No. 1

T 1.0

T1.0 1

d Q- AUGUST 1991 3

(Facing Page) Fig. 1. Adult habitus of Sabethes(Peyfonulus) aurescens(based on specimens from the State of Sao Paulo, Brazil). Coloration is characteristic of all but one undescribed species of the subgenus. The presence of lower mesokate- pisternal setae (MkSL) distinguish species of Pqtonulus from those of Sabethinus.

(Thispage) Fig. 2. Fourth-instar larva of Subethes(Pey~onulus) aurescens (based on specimens from the State of Sao Paulo, Brazil). The modified seta l-VII is diagnostic for the subgenus. 4 hbSQUIT0 sY!3TJ34ATICS VOL..23, No. 1

Fig.3. Pupaandmouthpartsof the fourth-in&u larva of Subethes (Peyti&) nurescens(basedonspecimens from the State of Sao Paulo,Brazil). The absenceof seta l-VII is diagnosticfor the subgenus.The large, elongatemaxilla with its well developedteeth and long terminal clawlikeprocess are characteristicof all specieswithin this group. AUGUST 1991 5

Sabethoidesin part of Theobald, 1910: 584; Discussion. Peytonulusexhibits the most di- Bonne and Bonne-Wepster, 1925:22,37-41; vergent morphology of any subgenusof Sabe- Dyar, 1928: 6, 16-17,21-24; Edwards, 1932: thes. Larvae of this group possessa uniquely 80-81; and authors. specializedseta l-VII which reaches terminal developmentin the fourth instar,i.e., it doesnot Except for the adult stageof one undescribed develop in the pupal stage. This ontogenetic specieswhich lacks metallic scutalscaling and distinction represents a synapomorphy for bearscertain other stasimorphiccharacters, this Peytonulus.Nevertheless, species of this group, subgenuspossesses the general characteristics like manyother groupsof mcxsquit~ are mosaics of the genusSabethes as noted by Harbach and of primitive and derived characters.The unde- Peyton (1990b). It differs from the other sub- scribedspecies mentioned above is a good ex- genera in the diagnosticand differential fea- ample of this. It shares a plesiomorphy with tures given below. These features are illus- Wyeomyiain the retention of dull scutalscaling, trated in figures l-4 and contrasted with ho- yet bearsthe autapomorphicimmature charac- mologous features in the other subgenerain ters diagnosticof Peytonulus. Table 1. The speciesincluded in Peytonuluswere for- Adult. Adultsof the subgenusPeyfonulusare merly placed in Sabethinus.These subgenera, readily recognized by the following combina- however,have very little in commonother than tion of characters: thorax with upper proepis- overt similaritiesin the ornamentation of the ternal and lower mesokatepisternalsetae; legs adults.Based on characteragreement,Peytonu- without paddles, midleg entirely dark-scaled; Zusappears to be more closelyrelated to Sabe- most specieswith conspicuouswhite scalingon thoidesand Davkmyiu. Nevertheless,these three ventral surfaceof hindtarsomere5. subgeneradiffer rather markedly in all stagesas Larva. Maxilla large, with well developed indicatedin Table 1. Particularly noteworthy is teeth (= laciniarastrum 1) and long terminal the presence of prealar setae in the adults of clawlike process;seta 4-C in line with or lateral Davismyia.Prealar setaeare plesiomorphicfor to l-C, normally lateral; seta 8-T anterior or an- Sabethesand represent a clear caseof symple- teroventral to pleuralsetal group;seta 1-I mesa1 siomorphywith Wyeomyia. to 2-1; seta 10-I in line with or mesa1to 13-1, AlthoughPeytonulus seems to showaffinities usually mesal; seta 9-11,111anterior or anter- with Davismyiaand Sabethoides,the morpho- oventral to seta 7; seta 12-IV,V lateral to seta logical data have not been thoroughly com- 13; seta l-VII uniquely developed into a stout pared againstall speciesof the subgenusSabe- spine-, hook- or clawlike processborne on a thes. This subgenusis poorly known at present prominent tubercle, apparentlynot innervated, and appearsto be a rather heterogeneousas- without pupal homolog(Fig. 4); combwith rela- semblageof species. Thus, the relationships tively few large spines,without comb plate; se- suggestedhere are tentative and await further tae la,2a-S not duplicated;auxiliary seta 4b-X analysisof the entire genus. absent. Pupa. Seta 5-111much smaller than S-IV-VI; ACKNOWLEDGMENTS seta 9-III-VI often spinelike;seta l-VII absent; seta 8-VII ventral, small;paddle normal, with- The author is grateful to R.C. Wilkerson, out differentiated membranousdorsal area at R.A. Ward,E.L. Peyton, V.L.Mallampalli, B.A. base. Harrison and J.I. Glick for commentingon the Etymology. The subgenericname is a Lati- manuscript;Taina Litwak isespecially acknowl- nized diminutive patronym to recognize E.L. edged for masterfully preparing the illustra- Peyton for his specialknowledge of sabethine tions;and J.E. Pecor is thanked for typing parts mosquitoesand his many contributionsto the of the manuscript. field of mosquitosystematics. The three-letter abbreviationPey. is recommendedfor this subgenus. Table 1. Comparisonof diagnosticand differential charactersfor the five subgeneraof Sabethes.

Stage Character Peytonulus Sabethes Sabethoides Sabethinus Davkmyia

Adult Prealar setae No No No No Yes

Upper proepisternal Yes No, except Yes Yes Yes setae batesi

Lower mesokat- Yes Yes/no, Yes No Yes episternal setae variable Leg paddles No Yes, but not No No No all species Midtarsus largely No No, but some Yes No No, but females white-scaledon specieswith with somewhite ventral side white scalingon scalingon ventral side both sides

Larva Seta 4-C lateral Yes, rarely Yes/no, usually No, mesa1 Yes, occasionally Yes to 1-c in line with 1-C mesa1to 1-C to 1-c in line with 1-C Seta 8-T anterior Yes No, dorsalor No, dorsalor Yes/no No, dorsal or anteroventral anterodorsal anterodorsal anteroventral or to 9-T to 9-T to 9-T to 9-T ventral to 9-T Seta 1-I mesa1 Yes Yes Yes No, lateral No, lateral to 2-I to 2-I to 2-I Seta 10-I in line Yes No, in line with Yes No, in line with No, far with or mesa1to or lateral to or lateral to lateral to 13-I 13-1,usually mesa1 13-1,usually lateral 13-I Seta 9-11,111 Yes No, anterior or Yes Yes/no, sometimes Yes anterior or antero- anterolateral to 7 anterolateral on ventral to seta 7, segmentIII usuallyanteroventral AUGUST 1991 7 8 d&3!SQUlTO sYYl’EMATIcS Vot 23, No. 1

Fig. 4. Lateral view of abdominalsegment VII of the prepupalstage of Sabethes(Peyro~dus) undosus showingthe neural connectionsbetween larvaland developingpupal setae. Note that seta1 is not innervatedin the larvaand absent in the pupa. Setae 12 and 13 are innervatedin the larvabut their structuralhomologs are absentin the pupa. Illustration is basedon specimennumber TR 700-1A (from Trinidad) in the NMNH.

REFERENCES CITED eau-Desvoidy (Diptera, Culicidae). Inset. Inscit. Menst. 12:97-100. Bonne, C. and J. Bonne-Wepster. 1925. Mos- Dyar, H.G. 1928. The mosquitoesof the Ameri- quitoes of Surinam, a study on neotropical cas. Part I. Carnegie Inst. Wash. Publ. No. mosquitoes.Meded. K. Inst. Amst. 21 (Trop. 387. 616 pp. Hyg. No. 13). 558 pp., 31 pl. Edwards,F.W. 1932. Genera Insectorum. Dip- da Costa Lima, A. 1931. Sobre asespecies dos tera. Fam. Culicidae. Fascicle194. Desmet- generossabethese Sabethoides(Diptera: Cu- Verteneuil, Brussels,Belgium. 258 pp. licidae). Mem. Inst. Oswald0Cruz Rio De J. Galindo,P., E. de Rodanicheand CM. Johnson. 25:51-64,3 pl. 1959. St. LouisencephalitisinPanamaI. Iso- Dyar, H.G. 1924. A note on Sabethes Robin- lation of the virus from forest mosquitoes AUGUST1991 9 and human blood. Am. J. Trop. Med. Hyg. Lane, J. 1953. Neotropical Culicidae. Vols. 1 8:557-560. and 2. University of Sao Paulo, Sao Paulo, Harbach, R.E. and K.L. Knight. 1980. Taxono- Brazil. 1,112 pp. mists’ glossaryof mosquitoanatomy. Plexus Lane, J. and N.L. Cerqueira. 1942. OSsabetin- Publishing,Inc., Marlton, NJ. xi + 415 pp. eos da America (Diptera, Culicidae). Arq. Harbach, R.E. and K.L. Knight. 1982. Correc- Zool. Estado Sao Paulo 3:473-849. tions and additionsto Taxonomistsglossary’ Lutz, A. 1905. Novasespecies de mosquitosdo of mosquito anatomy. Mosq. Syst. (1981) Brasil. Imprensa Med. 13( 18):347-350. 13:201-217. Mattingly,P.F., R.W. Crosskeyand KG-V. Smith. Harbach, R.E. and E.L. Peyton. 1990a. A new 1973. 19. Summaryof arthropodvectors,pp. subgenusin Wyeomyia(Diptera: Culicidae), 497-532. In: K.G.V. Smith (ed.). Insectsand with the reclassificationand redescriptionof other arthropodsof medicalimportance. Brit- the type species,Sabethes femandezyepezi. ish Museum (Natural History), London. Mosq. Syst.22:15-23. Stone,A., K.L. Knight andH. Starcke. 1959. A Harbach, R-E. and E.L. Peyton. 1990b. Trans- synoptic catalog of the mosquitoes of the fer of the subgenusDavismyia from Wyeo- world (Diptera, Culicidae). Thomas Say myia to Sabethesand descriptionof the type Found. 6:358 pp. species,Mamyia petrocchiae(Diptera: Cu- Theobald, F.V. 1907. A monographof the Cu- licidae). Mosq. Syst.22:149-159. licidae or mosquitoes. Vol. 4. British Mu- Howard, L.O., H.G. Dyar and F. Knab. 1915. seum (Natural History), London. xix + 639 The mosquitoesof North and Central Amer- PP.716 Pl. ica and the West Indies. Vol. 3. Systematic Theobald, F.V. 1910. A monographof the Cu- description(in two parts). Part 1. Carnegie licidae or mosquitoes. Vol. 5. British Mu- Inst. Wash. Publ. No. 159. 523 pp. seum (Natural History), London. xv + 646 Knight, K.L. and A. Stone. 1977. A catalogof pp., 6 pl. the mosquitoesof the world (Diptera: Cu- licidae). 2nd edition. Thomas Say Found. 6:xi + 611 pp.