Assessment of the Taxonomic Status of Some Species Included in the Shannoni Complex, With the Description of a New Species of Psathyromyia (Diptera: Psychodidae: Phlebotominae)

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Assessment of the Taxonomic Status of Some Species Included inthe Shannoni Complex, with the Description of a New Species ofPsathyromyia (Diptera: Psychodidae: Phlebotominae)Author(s): P. B. Sábio , A. J. Andrade , and E.A.B. GalatiSource: Journal of Medical Entomology, 51(2):331-341. 2014.Published By: Entomological Society of AmericaURL: http://www.bioone.org/doi/full/10.1603/ME13153

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MORPHOLOGY, SYSTEMATICS, EVOLUTION

Assessment of the Taxonomic Status of Some Species Included in theShannoni Complex, With the Description of a New Species ofPsathyromyia (Diptera: Psychodidae: Phlebotominae)

P. B. SABIO, A. J. ANDRADE, AND E.A.B. GALATI1

Departamento de Epidemiologia, Faculdade de Saude Publica, Universidade de Sao Paulo, Avenida Doutor Arnaldo 715,01246-904, Sao Paulo, SP, Brasil

J. Med. Entomol. 51(2): 331Ð341 (2014); DOI: http://dx.doi.org/10.1603/ME13153

ABSTRACT The Shannoni complex, comprising sand ßies belonging to the subgenus Psathyromyia(Psathyromyia) Barretto, consists of several species, of which the females have banana-shapedspermathecae and the males have digitiform parameres on the terminalia. Among these species,Psathyromyia shannoni (Dyar) and two taxa considered as its junior synonyms (Phlebotomus limaiFonseca and Phlebotomus bigeniculatus Floch & Abonnenc), Psathyromyia pestanai (Barretto &Coutinho), and a new species are the focus of this study. On the basis of morphological andmorphometric characters, Psathyromyia bigeniculata (Floch and Abonnenc) stat. rev.; comb. n. andPsathyromyia limai (Fonseca) stat. rev.; comb. n. are resurrected from the synonymy of Pa. shannoni.Pa. pestanai is proposed as a new junior synonym of Pa. limai. Psathyromyia ribeirensis sp. n., occurringin the middle and upper Ribeira Valley in the state of Sao Paulo, Brazil is described.Pa. limai, describedfrom the Serra da Cantareira (Sao Paulo municipality) and also found in the lower Ribeira Valley andon the adjacent coastal plain, is morphologically very close to the new species.

KEY WORDS Phlebotominae, sand ßy, Shannoni series, species complex, Brazil

The Shannoni series (Psychodidae, Phlebotominae,Phlebotomini, Psychodopygina) belongs to the genusPsathyromyia Barretto, 1962, which consists of 16 spe-cies (Galati 1995, 2013). In this series, Psathyromyiaabonnenci (Floch & Chassignet, 1947), Psathyromyiacuzquena (Martins et al., 1975), Psathyromyia pestanai(Barretto & Coutinho, 1941) and Psathyromyia shan-noni (Dyar, 1929) (syn. Phlebotomus bigeniculatusFloch & Abonnenc, 1941, Phlebotomus limai Fonseca,1935, Phlebotomus microcephalus Barretto & Duret,1953, and Phlebotomus pifanoiOrtiz, 1972) present thegreatest morphological similarity, mainly among thefemales, whereas the males contribute most decisivelyin species identiÞcation. Because of the morphologicalsimilarity of these four valid species and their synon-ymies, they were grouped as the Shannoni complex.Ph. limai and Ph. bigeniculatus were considered ju-

nior synonyms of Pa. shannoni by Barretto (1946), inview of the fact that their males were morphologicallysimilar and, although some differences were observedbetween the females, they were considered as possiblegeographical or mounting technique variations. Ph.microcephalus and Ph. pifanoiwere proposed as juniorsynonyms of Pa. shannoni without any explanation,respectively by Forattini (1971, 1973) and Martins etal. (1978).

Pa. shannoni presents the widest geographical dis-tribution in the Americas, from the United States toArgentina, occurring in almost all the countries ofCentral and South America (Young and Duncan1994). In Brazil, it has been found in every state exceptthe state of Sergipe (Galati 2013).

The holotype and two paratypes of Ph. limai arefemales from Serra da Cantareira, Greater Sao PauloMetropolitan region, Sao Paulo state, Brazil (Fonseca1935). Presumptive males of this species were de-scribed based on specimens from Presidente Prudenteand Andradina municipalities, northwestern SaoPaulo state by Barretto and Coutinho (1940). Ph. bi-geniculatus were described on the basis of male andfemale character from Cayenne, French Guyana byFloch and Abonnenc (1941).

The type-locality of Pa. pestanai (Barretto andCoutinho, 1941) and Ph. limai (Fonseca 1935) is thesame. The association between sexes of Pa. pestanaiwas established through females also captured in theGreater Sao Paulo region. These females did lay eggsunder laboratory conditions resulting in males of thisspecies (Coutinho and Barretto 1941).

During studies undertaken in the SpeleologicalProvince of Ribeira Valley, a specimen of Psathyro-myia of the Shannoni series resembling to Pa. pestanaiwas collected (Galati et al. 2010). The males of the twospecies can be distinguished from each other by mor-1 Corresponding author, e-mail: [email protected].

0022-2585/14/0331Ð0341$04.00/0 � 2014 Entomological Society of America

phological characters of the terminalia, but the fe-males are virtually indistinguishable morphologically.Pa. shannoni has been recorded on the coastal plain

of the Ribeira Valley (Gomes and Galati 1989); how-ever, Domingos et al. (1998), investigating the sand ßyfauna of Pedro de Toledo municipality situated in alow area also in this Valley, found both sexes of Pa.pestanai and suggested that the specimens found byGomes and Galati (1989) had been mistakenly iden-tiÞed as Pa. shannoni,whereas in fact they belonged toPa. pestanai. Domingos et al. (1998) also commentedthat although Gomes and Galati (1989) mentioned thepresence of both sexes in their study, only females hadin fact been collected and the close similarity of thefemales of the two species could explain the mistake.

Thus, in view of 1) the Þnding of a new species veryclose toPa.pestanai,2)thesimilaritybetweenthefemaleof Pa. pestanai and that of Ph. limai, and 3) the similarityof the specimens proposed as males of Ph. limai withthose of Ph. bigeniculatus, the current study sought toinvestigate the taxonomic status of these taxa in theShannoni series. The other two taxa, considered synon-ymous with Pa. shannoni, will be addressed in a laterpublication.

Materials and Methods

Specimens of type-series of Ph. limai, Pa. pestanai,and Pa. shannoni were examined, as well as and twoother nontype specimens ofPh. bigeniculatus, from thetype-locality. Their geographical distributions andoriginal descriptions were also taken into consider-ation. In addition, 73 males and 67 females on slidesdeposited in the following collections were examined:Colecao de Referencia da Faculdade de Saude Publica(FSPÐUSP) and Colecao Entomologica do Labo-ratorio de Entomologia em Saude Publica (FSPÐLESPÐPhlebotominae), both belonging to the Univer-sidade de Sao Paulo; Colecao da Secao de Parasitologiado Instituto Butantan (IBu); Colecao Entomologicado Instituto Oswaldo Cruz (CEIOC); Colecao deReferencia Nacional e Internacional do Centro dePesquisas Rene Rachou (CRNIFÐCPqRR); and Col-lection of the Smithsonian InstitutionÕs National Mu-seum of Natural History (NMNH).

Field workers provided some specimens in severalAtlantic rainforest areas of Sao Paulo State: Cananeiamunicipality (24� 56�14.27� S and 47� 58�0.50� W, 58.3 mabove sea level [a.s.l.]) representing the coastal areaof the Ribeira Valley; Parque Estadual da Cantareira(23� 26�50.07� S and 46� 37� 59.01� W, 845 m a.s.l.)type-locality of Pa. pestanai and Ph. limai; and ParqueEstadual de Intervales (24� 16�17.18� S and 48�24�53.34� W, 811 m a.s.l.) to clarify questions about theoccurrence of Pa. pestanai and related species in theseregions. The specimens were collected using Centers forDisease Control (CDC) light traps (Sudia and Cham-berlain 1962) and black and white modiÞed Shannontraps (Galati et al. 2001). Engorged females were indi-vidually transferred to vials to obtain eggs and the prog-eny inaccordancewith themethoddescribedbyKillick-Kendrick and Killick-Kendrick (1991).

All the specimens, including the type-series, wereanalyzed morphologically, ignoring the previous iden-tiÞcation given on the label of each slide. Then themorphometric measurements (�m) of each specimenwere taken and variance analyses were performed.The multiple comparisons of the morphometric char-acters were undertaken using the Gabriel test (Rohlfand Sokal 1981; F-test; P � 0.05; degrees of freedom[df]: between groups [B] and within groups [W])using the SPSS PASW program, version 17 (PASW2004, SPSS Inc., Chicago, IL).The analyses were basedon the following characters: length and width of thehead and eyes; interocular distance; length of clypeus;labrumÐepipharynx; ßagellomere I, II, and III; andpalpomere I, II, III, IV, and V, length and width of wingand length of some alar veins (alpha, beta, gamma,delta, pi, and R5); in the male terminalia: length andwidth of gonocoxite and lateral lobe, length of gono-style, dorsal and ventral margin of paramere, Þlamentgenital, pump and piston; and in the female terminalia:length and width of the spermathecae, individual andcommon sperm ducts.

Besides the morphological characteristics of themales and females, the distribution of the thoracicpigmentation of the taxa was compared and classiÞedas: intense (brown), low intensity (straw), or absent(off-white) (Caillard et al. 1986, Andersen 2010). Thethoracic sclerites are those deÞned by Galati (2003)and Cumming and Wood (2009).

The geographical distribution of the species wasdiscussed based on material obtained in the entomo-logical collections. The terminology of the charactersfollows Galati (2003) and the abbreviations of thegeneric names follow Marcondes (2007).

ResultsShannoni Complex

Diagnosis. Both sexes. Flagellomeres (F): ascoidswith long posterior spur, which in the ßagellomeresFIIÐFXII is close or goes beyond the basis of its re-spective article and presence of simple setae on FIÐFXIV. Cervical sclerites with three spiniform sensillae.VentroÐcervical sensillae absent. Male. Gonostylewith four well developed spines and absence of thepreapical seta, paramere digitiform, and gonocoxitewithout tuft of setae. Female. Cibarium with fourhorizontal teeth well-developed, arch complete, andsclerotized area wide; lacinia of the maxilla with theexternal teeth distributed in a single row. Commonsperm duct long and individual sperm ducts shorterthan the former; spermathecae banana-shaped.

Psathyromyia shannoni (Dyar, 1929)(Fig. 1A and B)

Ph. shannoniDyar, 1929: 121.Type-series: three males:Canal Zone (Panama), May 1923, R. C. Shannon(NMNH); Barretto (1946) 11: redescribed the maleand described the female from different Brazilianareas, French Guyana, United States; Fairchild andHertig (1950): 524: described the female and rede-

332 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 51, no. 2

scribed the male on the basis of specimens from thetype-locality.Sergentomyia shannoni Barretto (1955): 186 (cat.).Lutzomyia shannoniTheodor (1965): 189 (cat.); Lewis

et al. (1977): 325 (cat.); Ibanez-Bernal (2005): 202(identiÞcation key).Lutzomyia (Psathyromyia) shannoni Barretto (1962): 99

(cat.); Martins et al. (1978): 109 (cat.); Young andDuncan (1994): 349 (cat. and identiÞcation key).Lutzomyia (Trichopygomyia) shannoni Forattini

(1971): 102 (cat.); (1973): 294 (cat., description;identiÞcation key).

Pa. shannoniArtemiev (1991): 73 (cat.); Galati (2003):43, 67 (cat. and identiÞcation key).

Material Examined. PANAMA, Canal Zone�Þvemales and one female: NMNH (type-series n.41594A,n.41594B, and n.41594C); FSPÐUSP (n.10803 male andn.10802 female); FSPÐLESPÐPhlebotominae (slideswithout number). COLOMBIA, North of Santander(Gramalote)�one female: FSPÐLESPÐPhlebotomi-nae. COSTA RICA, San JoseÐthree males and onefemale: FSPÐLESPÐPhlebotominae. VENEZUELA, E.Tachira�one male: CEIOC (n.1159).

Fig. 1. Male and femaleÕs terminalia of species ofPsathyromyiagenus. (A and B) Male terminalia and female spermathecaeof Pa. shannoni (Canal Zone, Panama; Gonostylus: as: apical spine; is: internal spine; ues: upper external spine; les: lowerexternal spine; Paramere: vm: ventral margin; dm: dorsal margin); (C and D) Male terminalia (holotype of Pa. pestanai, SaoPaulo state, Brazil) and female spermathecae (Sao Paulo, Brasil) of Pa limai;E and F) Male terminalia (Cayenne, FrenchGuiana) and female spermathecae (MatoGrosso do Sul state, Brazil) of Pa. bigeniculata (e: the other gonostylus; scalebar: 100 �m).

March 2014 SABIO ET AL.: SHANNONI COMPLEX 333

Diagnosis. Both sexes. Thoracic coloration: prono-tum and paratergite straw, prescutum, scutum, andscutellum brown, pleurae off-white. Male terminalia(Fig. 1A). Paramere: the bristles on the dorsal marginextending from its apex to the level of the most basalbristle of the ventral angle. Gonostyle with the upperexternal spine implanted equidistant between the api-cal and the lower external spine; the lower externalspine is implanted at the same level of the internalspine. Gonocoxite: ca. 1.1 times longer than the laterallobe. Female terminalia (Fig. 1B). Apex of the com-mon sperm duct not extending beyond the apex of thegenital fork; common sperm duct ca. 2.0 times longerthan the individual sperm duct and 1.9 times the sper-matheca; individual sperm duct is �0.9 longer than thespermatheca.

Psathyromyia limai (Fonseca, 1935), stat. rev.;comb. n.

(Fig. 1C and D)

Ph. limai Fonseca, 1935: 61. Type-series: female holo-type and two female paratypes: Serra da Cantareira,Greater Sao Paulo Metropolitan region, Sao Paulostate, Brazil, August 1935, L. T. Travassos Filho coll.(IBu); Coutinho (1940): 333: redescribed a femalefrom Vila Queiroz (currently Queiroz municipal-ity), Sao Paulo state, Brazil; Barretto and Coutinho(1940): 127: described a male from Presidente Pru-dente and Andradina municipalities, Sao Paulostate, Brazil; Coutinho and Barretto (1941): 74: as-sociated males with females from a laboratory cul-ture originating from wild-caught females that werecaptured in the surroundings of Sao Paulo city andin the Piedade municipality, Sao Paulo state, Brazil;Rozeboom (1944): 274: identiÞed Ph. limai in somelocalities in the United States; Barretto (1946): 11:proposed as junior synonym of Ph. shannoni. Forat-tini (1971): 102; (1973): 294; Martins et al. (1978):109; Young and Duncan (1994): 349; Galati (2003):43: accepted this synonymy.Ph. pestanai Barretto & Coutinho, 1941: 144. Type-

series: male holotype, Serra da Cantareira (SaoPaulo state, Brazil). October 1940, M. P. Barrettoand J. O. Coutinho coll. (FSPÐUSP); Coutinho andBarretto (1941): 74: description of female; Barretto(1942): 77 (biological cycle); Fairchild, (1955): 195(cat.). syn. n.Sergentomyia pestanai Barretto (1955): 179 (cat.,

comb.).Lutzomyia pestanai Theodor (1965): 189 (cat.).Lutzomyia (Psathyromyia) pestanai Barretto (1962):

99; Martins et al. (1978): 107 (cat.); Young andDuncan (1994): 348 (cat., tax.).Lutzomyia (Trichopygomyia) pestanai Forattini

(1971): 102; (1973): 294 (cat., tax.).Psathyromyia (Psathyromyia) pestanai Galati (2003):

43 (cat., tax., comb.).

Material Examined. BRAZIL, Sao Paulo StateÐ7males: FSPÐUSP (holotype of Pa. pestanai n. E-1418);FSPÐLESPÐPhlebotominae. 21 females: IBU (holo-

type of Pa. limai n. 2,271); FSPÐUSP (n. E-1,419);FSPÐLESPÐPhlebotominae. Cananeia municipality�1male and 13 females: FSPÐUSP (n. E-6027 female);FSPÐLESPÐPhlebotominae.Diagnosis. Both sexes. Thoracic coloration: prono-

tum, paratergite, prescutum, scutum, and scutellumbrown, areas of alar insertion and the halteres andpleurae off-white. Male terminalia (Fig. 1C). Gonos-tyle with the upper external spine implanted equidis-tant between the apical and the lower external spine,which is implanted at the same level as the internalspine. Paramere: dorsal margin with a curvature in itsapical third and bristles covering an area extendingfrom its apex to the level of the most apical bristles ofventral angle. Female terminalia (Fig. 1D).Commonsperm duct ca. 1.7 times longer than the individualsperm ducts and 2.4 times longer than the sper-matheca; individual sperm duct ca. 1.4 times longerthan the spermatheca.

Psathyromyia bigeniculata (Floch and Abonnenc,1941), stat. rev.; comb. n.

(Fig. 1E and F)

Ph. bigeniculatus Floch & Abonnenc, 1941: 3. Type-series: one male (n. 209), one female (n. 143), andthree other males and two females, Cayenne,French Guyana, June 1941, H. Floch and E. Abon-nenc coll. (Institute Pasteur ÐFrench Guyana); Bar-retto (1946): 11: Ph. bigeniculatus is similar to Ph.limai and proposed Ph. bigeniculatus as a juniorsynonym of Ph. shannoni; Barretto (1955): 186 (syn.Sergentomyia shannoni); Barretto (1962): 99; Mar-tins et al. (1978): 109; Young and Duncan (1994):349 (syn. Lutzomyia (Psathyromyia) shannoni); Fo-rattini (1971): 102; (1973): 294 (syn. Lutzomyia(Trichopygomyia) shannoni), and Galati (2003): 43syn. Psathyromyia (Psathyromyia) shannoni ac-cepted this synonymy.Phlebotomus limai Barretto and Coutinho (1940): 127

(description of male).

Material Examined.FRENCH GUIANA, CayenneÐtwo males and two females: FSPÐLESPÐPhlebotomi-nae. ARGENTINA, Misiones (Puerto Iguazu)�threemales and one female: CRNIFÐCPqRR (n. N.E.:3591/09, n. N.E.:3586/09, n. N.E.:3651/11, and n. N.E.:3659/11 female). BRAZIL: state of Amapa, OiapoquemunicipalityÐone female: FSPÐUSP (n. E-409); stateof Amazonas, Manaus municipality�six males and onefemale: FSPÐLESPÐPhlebotominae; state of Bahia, Sal-vador municipality�one male: FSPÐUSP (n. E-2340);state of Mato Grosso do SulÐ29 males and 9 females:FSPÐLESPÐPhlebotominae; stateofParaÐ11males and7 females: FSPÐLESPÐPhlebotominae; state of SaoPaulo: Andradina municipality�two males: FSPÐUSP(n. E-1424 and n. E-1425); Angatuba municipalityÐone female: FSPÐLESPÐPhlebotominae; DouradoÐone male: FSPÐUSP (n. E-6031); Presidente Epitaciomunicipality�one female: FSPÐLESPÐPhlebotomi-nae; Presidente Prudente municipality�two females:FSPÐLESPÐPhlebotominae.


Diagnosis. Both sexes. Thoracic coloration: prono-tum, paratergite, prescutum, scutum, and scutellumbrown, upper anepisternum straw and the other pleu-ral sclerites off-white. Male terminalia (Fig. 1E).Gonostyle with the upper external spine implantedequidistant between the apical and lower externalspine. This lower external spine is slightly more apicalthan the internal spine. Paramere with bristles of thedorsal margin extending from its apex to the level ofthe most apical bristle of the ventral angle. Femaleterminalia (Fig. 1 F).Common sperm duct smooth,extending beyond the apex of the genital fork and itslength ca. 2.1 times that of the spermatheca, and 2.5times that of the individual sperm duct.

Psathyromyia ribeirensis sp. n. Sabio,Andrade & Galati

(Figs. 2Ð5)

Diagnosis. Both sexes. Thoracic coloration: Prono-tum, paratergite, scutum, and scutellum brown andpleurae off-white. Male. Terminalia: gonostyle withthe upper external spine implanted equidistant be-tween the apical and lower external spines, and lowerexternal spines is implanted at the same level as theinternal spine; paramere digitiform with the bristles ofthe dorsal margin covering an area extending from the

apex of the paramere to the level of the apical bristlesin the ventral angle. Female. Terminalia with sper-mathecae banana-shaped.Description of the Male (HOLOTYPE). The mea-

surement result of each structure of the holotype andparatypes are given in micrometer, the holotype beingshown outside the parentheses and the paratypes inparenthesis. The conversion of the micrometer read-ing was in objective 5X � 196 �m, 10X � 100 �m, and40X � 26 �m. Head (Fig. 2A) 430 (430) in length, 440(420) in width; clypeus 127 (125) in length; eyes 286(278) in length; interocular distance 122 (127).Cibarium without teeth (Fig. 3A). LabrumÐepiphar-ynx (LE) 320 (330). Antenna (Fig. 3BÐD): ßagellom-ere length: FI 420 (450), FII 190 (180), and FIII 180.Holotype with FIVÐFVIII detached from the head andother parts of the two individual specimens examinedwere lost. Ascoids with a relatively long posterior spur,which almost reaches the basis of their article (ex-cepting in FI), and the apex of anterior spur in FIÐFIIIis close to the papilla level; in FI the ascoids areimplanted at different levels, the external more basalthan the internal (Fig. 3B), FIÐFIII with papilla im-planted in the preapical region (Fig. 3D). Presence ofsimple setae on FIÐFVIII. Palpi (Fig. 3EÐH): palpom-ere length: I 49 (52), II 166 (169), III 172 (182), IV 94(70), and V 307. Palpal formula: 1Ð4Ð2Ð3Ð5; palpom-

Fig. 2. Heads of Pa. ribeirensis sp. n. Sabio, Andrade & Galati. (A) Male holotype and (B) Female paratype. (Scale bar:100 �m).


ere II without NewsteadÕs spines; palpomere III withNewsteadÕs spines situated in the apical half (Fig. 3 F).Labial suture forming a fork (Fig. 2A).Cervix. VentroÐcervical sensillae absent. Cervical

sclerites bearing three spiniform sensillae.Thorax. Thorax 750 (830) in length, mesonotum 670

(730) in length. Pronotum, paratergite, and mesono-tum brown, pleura off-white. Two to three proepi-meral setae; 8Ð9 upper anepisternal setae. Setae on theanterior margin of the katepisternum absent. Wing(Fig. 5A): 2,732 (2,752) in length, 772 (792) in width;veins R5 1,604 (1,584); � 693 (673); � 356 (356); � 277(277); � 198 (198); � 139 (178). Legs (anterior, me-dian, posterior): coxae 455 (436), 436 (416), 436 (416).The other segments of the leg, femur, tibia and tar-someres, were lost. Abdomen.

Abdomen 2,218 (1,881) in length; absence of thetergal papillae on all tergites. Terminalia (Fig. 5C).Gonocoxite 430 (450) in length, 90 (110) in width,without persistent setae. Gonostylus: 220 (210) inlength, without preapical setae and with four well-developed spines, one apical, two external (upperexternal spine and lower external spine) and one in-ternal spine; the upper external spine is implanted ata level equidistant between the apical spine and thelower external spine; this lower external spine is slightlymore apical than the internal spine. Paramere: dorsalmargin 268 (229) and ventral 330 (351) in length.Paramere simple with rectangular base and the apicalhalf digitiform, tapering gradually toward the apex; itsdorsalmarginwithapicalhalf coveredbyspiniformsetae

pointing toward the base of terminalia. Aedeagus scle-rotizedandconiform.Lateral lobe370(380) in length,29(36) inwidthandroundedat theapex.Genitalpump177(182) in length; piston 140 (146) in length; genital Þla-ments with bevel shaped apex, 630 (570) in length, 3.6times the length of the pump (Fig. 5D).Female. Head (Fig. 2B) 450 (450Ð420) in length,

440 (440Ð400) in width; clypeus 130 (140Ð130) inlength; eyes 296 (268Ð255) in length; Interocular dis-tance 130 (127Ð127). LabrumÐepipharynx (Fig. 4B)(LE): 390 (360Ð330) and with 32Ð36 teeth on apicalregion (Fig. 4C). Lacinia of maxilla with seven exter-nal teeth and 23 internal teeth (Fig. 4D). Cibariumwith four posterior teeth well developed and 23 an-terior teeth distributed on two transverse rows; scle-rotized area short and triangular; sclerotized archcomplete (Fig. 4A). Antenna (Fig. 4EÐI): ßagellomerelength FI 310 (290Ð300), FII 130 (140Ð130), FIII 130(130Ð120), FXIII 65 (65Ð62), and FXIV 62 (52Ð55).Ascoidswith relatively longposterior spur, inFIIÐFXIItheir apex is close to the basis of the article; the an-terior spur is long and reaches almost to the base of thenext ßagellomere; in FI, both external and internalascoids are implanted at the same level; Antennalformula FIÐFXII 2, FXIIIÐ FXIV 0; FIÐIII with preapi-cal papilla; FXI without papilla. Presence of simplesetae on FIÐFXIV. Palpi (Fig. 4JÐM): palpomerelength: I 49 (55Ð55), II 198 (182Ð185), III 182 (190Ð166), IV 86 (86Ð78), and V 234 (265Ð247). Palpalformula: 1Ð4Ð(2Ð3)Ð5 (n� 3); palpomere II (Fig. 4J)without NewsteadÕs spines; palpomere III with New-

Fig. 3. Male holotype of Pa. ribeirensis sp. n. Sabio, Andrade & Galati. (A) Cibarium; (B) ßagellomere I; (C) ßagellomere II;(D) ßagellomere III; (E) palpomeres I and II; (F) palpomere III; (G) palpomere IV; and (H) palpomere V. (Scale bar: 100 �m).


steadÕs spines situated on the apical half (Fig. 4K).Labial suture forming a fork (Fig. 1B).

Cervix: VentroÐcervical sensillae absent. Scleritecervical with three spiniform sensillae.

Thorax: 800 (780) in length, mesonotum 740 (690)in length. Pronotum, paratergite, and mesonotumbrown, pleura off-white. Two to three proepimeralsetae; 8Ð9 upper anepisternal setae. Setae absent onthe anterior region of the katepisternum. Wing (Fig.5B): 2,871 (2,475) in length, 812 (653) in width; veinsR5 1,742 (1,485); � 812 (673); � 356 (317); � 277(198Ð198; n� 3); � 257 (218); � 198 (119Ð198; n� 3).Legs (n � 2) anterior, median, posterior: coxae: 416(416), 396 (396), 376 (396). Femur (n� 1): 1,010; 970;1,069.Tibiae(n�1):1,426;1,742;2,039.TarsomereI(n�1): 891; 1109; posterior missing. Sum of tarsomeres II III IV V (n � 1): 871; 990; posterior missing.

Abdomen: (n � 1): 2,079 in length. Spermathecaebanana-shaped (Fig. 5E): 73 (57) in length and 16 (16)in width; common sperm duct 83 in length and 8 inwidth; individual sperm duct 60 in length and 5.2 inwidth. The spermathecae and the sperm ducts aremembranous and with smooth walls.Type-Material.HOLOTYPE male and PARATYPES

(one male and three females), Parque Estadual deIntervales, Ribeirao Bonito municipality, state of SaoPaulo, Brazil, located between the Ribeira andParanapanema Valleys, in southwest of the state of SaoPaulo (24� 16�17.18� S and 48� 24�53.34� W, 811 m a.s.l).HOLOTYPE and PARATYPE males: forested areaaround of the Gruta Colorida (Pedra do Fogo region),27/28-IX-2002 (CDC trap); two PARATYPE females:in forest in the Tower trail, Morro do Espia, 27-X-2001(human bait in Shannon trap). Collector: EAB Galati;

Fig. 4. Female paratype ofPa. ribeirensis sp. n. Sabio, Andrade & Galati. (A) Cibarium. (B) labrumÐepipharynx; (C) apicalregion of labrumÐepipharynx; (D) apical region of lacinia of the maxilla; (E) ßagellomere I; (F) ßagellomere II; (G)ßagellomere III; (H) ßagellomere XII; (I) ßagellomere XIII and XIV; (J) palpomeres I and II; (K) palpomere III; (L)palpomere IV; and (M) palpomere V. (Scale bar: 100 �m).


one PARATYPE female: in forested area of the Sede,04-IV-2012, automatic light trap, type CDC. Collec-tors: P.B. Sabio & E.A.B. Galati. Material deposited inthe “Colecao de Referencia da Faculdade de SaudePublica (FSPÐUSP).” The association between maleand female was proposed based on the genital andextragenital characteristics, coloration patterns, andalso on the fact that they were the unique represen-tatives of the Shannoni series in the captures made inthe Parque Estadual de Intervales.Etymology. The species name alludes to the region

Ribeira Valley where the type-locality is included instate of Sao Paulo, Brazil.

Taxonomic Discussion

One of three males that comprise the type-series ofPa. shannoniwas designated as a lectotype (n. 41594A)by Dampf in 1936 (data not published) and one of theparalectotypes has been identiÞed by the currentstudy as Pa. abonnenci (n.41594B). Pa. shannonimain-tains its valid status, being morphologically distinctfrom Pa. limai, Pa. bigeniculata, and Pa. abonnenci.

Floch and Abonnenc (1941) described Pa. bigenicu-lata and distinguished it from Pa. shannoni by theposition of the spines on the gonostyle. The currentstudy corroborates this difference and identiÞes othersigniÞcant and distinct male morphometric characters

Fig. 5. Pa. ribeirensis sp. n. Sabio, Andrade & Galati. (A) Wing of the male holotype (scale bar: 200 �m); (B) wing ofthe female paratype (scale bar: 200 �m); (C) terminalia of the male holotype (scale bar: 100 �m); (D) genital Þlaments ofthe male holotype (scale bar: 100 �m); (E) spermathecae of the female paratype. (Scale bar: 100 �m).


(P� 0.05; df: B � 1; W � 4) between Pa. bigeniculata(n � 2) and Pa. shannoni (n � 4) which are respec-tively, the length of: ßagellomere FI (280 �m and 336�m; F� 9), the ratio of FI and clypeus(2.7 and 3.5 �m;F� 36) and the ratio of the gonocoxite and lateral lobe(1.2:1.0 and 1.1:1.0; F � 13). No signiÞcant morpho-metric difference was found between females of Pa.shannoni and Pa. bigeniculata.

The spermathecae of the Shannoni complex are mor-phologically very similar and the distinction among thespecies can be made by means of thoracic coloration.

The male associated with Pa. limai (Fonseca, 1935)by Barretto and Coutinho (1940) presents thoraciccoloration distinct from that of the female holotype ofthis species and also that of both sexes of Pa. shannoni(Dyar 1929), but is similar to that of Pa. bigeniculata(Floch and Abonnenc, 1941). Therefore, the maleassociated with Pa. limai by Barretto and Coutinho(1940) actually differs from Pa. shannoni and resemblesthat of Pa. bigeniculata; the synonymization of Pa. limaiandPa.bigeniculata inPa. shannoniproposedbyBarretto(1946) on the basis of the males is not justiÞed.

Several specimens deposited in collections andidentiÞed asPa. shannoni are in realityPa. bigeniculata.Thus, based on the material examined, part of thedistribution of Pa. shannoni cited for Brazil by Martinset al. (1978) corresponds to that of Pa. bigeniculata.

In the description of the female of Pa. pestanai,Coutinho and Barretto (1941) reported that Pa. limaiwould be the closest species, differing only in relationto the spermathecae. However, they did not mentionwhat that difference would be. The spermathecaeillustrated by Fonseca (1935) presents smooth walls,whereas in the allotype of Pa. pestanai some verticalstriations are present on the walls. In the analysis ofother specimens of Pa. pestanai from the type-locality(Serra da Cantareira), this character varies; there arespecimens with smooth or striated walls.

However, the thoracic coloration of both sexes ofPa. pestanai and other female characters are similar tothose of Pa. limai and both species have the sametype-locality. So, in view of these evidences and pri-ority rule of the (ICZN) International Code of Zoo-logical Nomenclature (1999), it is proposed that Pa.limai (comb. n.) be resurrected from the synonymy ofPa. shannoni and that Pa. pestanai become a juniorsynonym of Pa. limai (new synonymy).

A male obtained from laying eggs of females col-lected in Cananeia municipality, Sao Paulo State, on 10March 2013, allowed us conÞrm that Pa. limai (� Pa.pestanai) occurs in the low Ribeira Valley, consistentwith the point of view of Domingos et al. (1998). Thespecimens collected in the upper part of Ribeira Val-ley (Parque Estadual de Intervales) differ morpho-logically from other species of the Shannoni series,representing a new species, that is described here.

Psathyromyia ribeirensis sp. n

Based on the classiÞcation proposed by Galati(1995, 2003), the new species presents characters con-sistent with the genus and subgenus Psathyromyia and

Shannoni series. Its male and female morphologicalcharacteristics are very close to those of the otherspecies considered as belonging to the Shannoni com-plex. Except for Pa. limai, this new species can bedistinguished from all other species of the Shannonicomplex, known by both sexes, by the distribution ofthoracic coloration (notum and pleurae) as presentedin the identiÞcation keys.

ThemaleofPa. ribeirensis sp.n. canbedistinguishedfrom the Pa. limaimale by a straight apical third dorsalmargin of the paramere compared with a curvature inthis region in the latter species. Further, Pa. ribeirensissp. n. (n� 2) and Pa. limai (n� 6) presented severalmorphometric statistically signiÞcant differences (P�0.005; df: B � 1; W � 6), respectively: wing length(2,742 and 2,478 �m; F � 41); beta (356 and 241 �m;F� 24), delta (277 and 231 �m; F� 15), gamma (198and 313 �m; F� 63), and R5 (1,594 and 1,488 �m; F�11); terminalia length of gonocoxite (440 and 397 �m;F � 32), gonostyle (215 and 187 �m; F � 19), dorsalmargin of the paramere (260 and 229 �m; F� 7), andthe lateral lobes (375 and 323 �m; F� 41). However,the females of these two species are indistinguishable.

The males of Pa. ribeirensis sp. n. also can be dis-tinguished from the male Pa. shannoni by the exten-sion of the area of implantation of bristles on the dorsalmargin of the paramere; in Pa. shannoni this area ex-tends from the apex of paramere to the level ofthe most basal bristles of the ventral angle, while in thenew species this area runs from the apex of paramereto a little before bristles of the ventral margin. Themales of these two species also differ in the lengths ofthe followingmorphometriccharacteristics(P�0.05;df:B � 1; W � 4): Pa. ribeirensis sp. n. (n � 2) and Pa.shannoni (n� 4), respectively: labrumÐepipharynx (325and261�m;F�85),ßagellomereI(435and319�m;F�74) and ßagellomere II (185 and 144 �m; F� 76), gono-coxite (440 and 349 �m; F� 88), gonostyle (215 and 177�m;F�100),dorsalmarginoftheparamere(260and196�m;F�87)andtheventralmarginof thesamestructure(309 and 240 �m; F� 8), lateral lobe (375 and 311 �m;F�34),pump(179and123 �m;F�57), andpiston(143and 99 �m; F � 57). SigniÞcant morphometric differ-ences were not observed between the females of thesespecies.Pa. abonnenci (Floch and Chassignet, 1947) can be

distinguished from Pa. ribeirensis sp. n. by the bristleson dorsal margin of paramere, which are longer thanthose ofPa. ribeirensis sp. n. and implanted only on theapical third while in the new species they occur on theapical half in the new species. Morphometrically, Pa.ribeirensis sp. n. (n � 2) and Pa. abonnenci (n � 9)differ(P� 0.05; df: B � 1; W � 9), by the length of thefollowing structures, respectively: ßagellomere I (435and 282 �m; F� 66), ßagellomere II (185 and 128 �m;F � 40), gonocoxite (440 and 353 �m; F � 111),gonostyle (215 and 163 �m; F� 37), dorsal margin ofthe paramere (260 and 198 �m;F� 68), ventral marginof paramere (309 and 256 �m; F � 13), lateral lobelength (375 and 273 �m; F� 34), pump (179 and 139�m; F� 37), and piston (143 and 114 �m; F� 19). For


the females of these two species no signiÞcant mor-phometric differences was observed.

In the males of Pa. bigeniculata the bristles on dorsalmargin of paramere are longer than correspondingbristles on the paramere of the new species. Morpho-metrically, statistically signiÞcant differences (P �0.05; df: B � 1; W � 2) between Pa. ribeirensis sp. n.(n� 2) and Pa. bigeniculata (n� 2) were observed inthe lengths of the following structures, respectively:labrumÐepipharynx (325 and 220 �m; F � 88), ßag-ellomere I (435 and 280 �m; F� 74), ßagellomere IV(185 and 130 �m; F � 121), gonocoxite (440 and 355�m; F� 22), dorsal margin of the paramere (260 and183 �m;F� 139), and lateral lobe(375 and 285 �m;F�32). For the females, no signiÞcant statistically differ-ences were observed.Pa. cuzquena (Martins et al., 1975) differs from Pa.ribeirensis sp. n. in that the upper external spine on thegonostyle is implanted very close to the apical spine,whereas in the new species it is implanted equidistantbetween the apical and lower external spines. Mor-phometric comparison of these two species (n � 2both species) reveals signiÞcant differences in thelengths of the following structures, respectively (P�0.05; df: B � 1; W � 2): labrumÐepipharynx (325 and248 �m; F � 235), ßagellomere FI (435 and 331 �m;F� 26), gonocoxite (215 and 269 �m; F� 198), dorsalmargin paramere (260 and 170 �m; F � 300), laterallobe (375 and 221 �m; F� 951), genital Þlament (600and 317 �m; F� 73), and piston (143 and 119 �m; F�85). The female of Pa. cuzquena has not been de-scribed.

Identification Key

Males

1.Gonostyle with the upper external spine im-planted closer the apical spine than to the ex-ternal lower spine . . . . . . . . Pa. cuzquena

1�.Gonostyle with upper external spine implantedequidistant between the apical spine and thelower external spine . . . . . . . . . . . . . . 2

2.Thorax: upper anepisternum brown or strawcolored . . . . . . . . . . . . . . . . . . . . . . . . 3

2�.Thorax: upper anepisternum off-white . . . . . . 43(2).Thorax: pronotum, paratergite, anepisternum,

and postnotum brown. Paramere with the bris-tlesonthedorsalmargindistributedonlyontheapical third . . . . . . . . . . . . Pa. abonnenci

3�.Thorax: pronotum and paratergite brown,anepisternum and postnotum straw.Paramere with the bristles on the dorsal margindistributed at least on the apical half. . . . . . .. . . . . . . . . . . . . . . . . . . . Pa. bigeniculata

4(3).Thorax: pronotum and paratergite straw.Paramere with the bristles on the dorsal mar-gin distributed from apex to the basal level ofthe implantation of bristles of the ventral an-gle. . . . . . . . . . . . . . . . . . . . Pa. shannoni

4�.Thorax: pronotum and paratergite brown.Paramere with the bristles of the dorsal mar-

gindistributed fromits apex to theapical levelof the implantation of bristles of the ventralangle . . . . . . . . . . . . . . . . . . . . . . . 5

5(4).Paramere with curvature between the apicaland median region of the dorsal margin . . . .. . . . . . . . . . . . . . . . . . . . . . . . Pa. limai

5�.Paramere straight along the dorsal margin . . . .. . . . . . . . . . Psathyromyia ribeirensis sp. n.

Females

1.Thorax: upper anepisternum brown . . . . . . 21�.Thorax: upper anepisternum off-white . . . . 3

2(1).Thorax: pronotum, paratergite, upper anepis-ternum, and postnotum brown . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . Pa. abonnenci

2�.Thorax: pronotum and paratergite brown, up-per anepisternum and postnotum straw. . . . .

. . . . . . . . . . . . . . . . . . . Pa. bigeniculata3(1).Thorax: pronotum and paratergite brown. . . . .

. . . . Pa. limai, Psathyromyia ribeirensis sp. n.

3�.Thorax: pronotum and paratergite straw . . . .. . . . . . . . . . . . . . . . . . . . . Pa. shannoni

Acknowledgments

We thank the curators of the following institutions: FSPÐUSP, IOC, IBu, and CPqRR who collaborated by loaningspecimen material. We thank Ph.D. Veracilda Ribeiro AlvesInstituto Nacional de Pesquisa da Amazonia [INPA], whoprovided information regardingPa. shannonideposited at theNHNM. We also thank the staff of the Sistema de Autorizacaoe Informacao em Biodiversidade (Sisbio) for the authoriza-tions granted and the Secretaria do Meio Ambiente for grant-ing a license to the Instituto Florestal (COTEC) to collectsand ßies in Parque Estadual da Cantareira and Parque Es-tadual de Intervales. We are grateful to CoordenacaodeAperfeicoamento de Pessoal de Nõvel Superior (CAPES) fora scholarship to support themasterÕsdegree researchofP.B.S.and to Fundacao de Amparo a Pesquisa do Estado de SaoPaulo (FAPESP) (2000/06811-0; 2010-15802/7) for researchproject support of E.A.B.G. and scholarship support of thepostdoctoral fellowship of A.J.A.

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Received 5 August 2013; accepted 8 January 2014.


Documents

Assessment of the Taxonomic Status of Some Species Included in the Shannoni Complex, With the Description of a New Species of Psathyromyia (Diptera: Psychodidae: Phlebotominae)