An illustrated guide for characters and terminology used in descriptions of Phlebotominae (Diptera, Psychodidae)

Abstract

Phlebotomine (Diptera, Psychodidae, Phlebotominae) taxonomy has been studied extensively, primarily due to the role of these flies as vectors of various parasites, including species of Leishmania, Bartonella and arboviruses that cause diseases in humans and other vertebrates. We present some topics discussed at a round-table on phlebotomine taxonomy held at the Ninth International Symposium on Phlebotomine Sandflies (ISOPS IX) in Reims, France, in June 2016. To date, approximately one thousand phlebotomine species have been described worldwide, although in varying languages and mostly without standardization of characters and terminology. In the interest of standardization, we list the characters that should minimally be considered in the description of new phlebotomine taxa as well as annotated illustrations of several characters. For these characters, multiple illustrations are provided to show some of the variations. The preferred terms for all pertinent characters are listed as well as their synonyms in English, Portuguese, and French. Finally, we offer an updated list of abbreviations to be used for generic and subgeneric names.

Figure 1.

Dorsal view of the head and its appendages of a phlebotomine female: cl – clypeus; fI – 1st flagellomere; fII – 2nd flagellomere; hy – hypopharynx; ml – maxillary lacinia; lb – labium; le – labrum-epipharynx; md – mandible; pe – pedicel; pha – pharynx; p1 – 1st palpal segment; p2 – 2nd palpal segment; p3 – 3rd palpal segment; p4 – 4th palpal segment; p5 – 5th palpal segment; sc – scape; most frequently used measurements: eL – eye length; eW – eye width; clL – clypeus length; fIL – 1st flagellomere length; heL – head length; hW – head width; ioD – interocular distance; pL – palpus length – Nyssomyia intermedia.

Figure 2.

(A–G) Dorsal view of the head of Bruchomyiinae and Phlebotominae with the distribution of the setae on some sclerites, the relation of clypeus to eyes and aspects of some structures (A) Bruchomyiinae (Bruchomyia sp.). (B) Phlebotominae: Warileya phlebotomanica; (C) Brumptomyia brumpti; (D) Sergentomyia (Sergentomyia) minuta; (E) Pintomyia (Pifanomyia) maranonensis; (F) Lutzomyia (Helcocyrtomyia) tejadai; (G) Lu. (Helcocyrtomyia) blancasi; (H) Phlebotomus (Euphlebotomus) barguesae. ey – eye; fcls – frontoclypeal suture; ge – gena; ias – interantennal suture; ios – interocular suture; oc – occiput; ve – vertex.

Figure 3.

Palpus of Phlebotominae. (A) 1st–5th palpal segments of phlebotomines: Newstead’s sensilla (Ns) dispersed on p3 and present from 2nd to 5th segment – Psathyromyia naftalekatzi. (B) Newstead’s sensilla concentrated on basal part of the segment and only one simple seta (ss) – Micropygomyia echinathopharynx. (C) Several simple setae on p3 – Warileya phlebotomanica.

Figure 4.

Aspects on the presence and distribution of ascoids, papillae, and simple setae on flagellomeres of Phlebotominae: (A–C) apical flagellomeres (fXI–fXIV): (A) Evandromyia (Barrettomyia) tupynambai (♂); (B) Warileya rotundipennis (♀); (C) Trichophoromyia ubiquitalis (♀). (D1–D4) First flagellomere (fI) of phlebotomine: (D1) Warileya rotundipennis (♀); (D2) Psychodopygus squamiventris (♀); (D3) Evandromyia aroucki (♂); (D4) Sergentomyia dubia (♂). (E1–E4) 2nd and 3rd flagellomeres of phlebotomines: (E1) Warileya rotundipennis (♀); (E2) Psathyromyia shannoni (♀); (E3) Sergentomyia dubia (♀); (E4) Sergentomyia dubia (♂). ap – Apical papilla; as – ascoid; bp – basal papilla; ias – internal ascoid; mp – median papilla; p – papilla; pap – preapical papilla; pasp – preascoidal papilla; sp – spiniform papilla; ss – simple seta.

Figure 5.

Mouth parts of Phlebotominae. (A, B) Apical region of the labrum-epipharynx of phlebotomine females: (A) Micropygomyia vexator; (B) Lutzomyia longipalpis. (C–E) Apical region of the hypopharynx of phlebotomine females: (C) Lutzomyia longipalpis; (D) Sciopemyia sordellii; (E) Micropygomyia cayennensis. (F–K) Maxillary lacinia of phlebotomine females: (F) Warileya phlebotomanica; (G) Warileya rotundipennis; (H) Lutzomyia cruciata; (I) Nyssomyia intermedia; (J) Micropygomyia quinquefer; (K) Micropygomyia longipennis. et – External teeth of maxillary lacinia; it – internal teeth of maxillary lacinia.

Figure 6.

Labium of phlebotomine females in ventral view. (A) Warileya phlebotomanica; (B) Lutzomyia amarali; (C) Sergentomyia minuta; (D) Idiophlebotomus padillarum; (E) Chinius eunicegalatiae. lbI – Labellum I; lbII – labellum II; lbs – labial suture; prem – prementum.

Figure 7.

Cibarium and pharynx of phlebotomine females. (A) Micropygomyia atroclavata; (B) Bichromomyia flaviscutellata. antt – Anterior teeth; cic – cibarial chamber; f – fold; ltt – lateral teeth; pha – pharynx; pht – pharyngeal teeth; psbr – posterior bridge; ptt – posterior teeth; pp – posterior protuberance; latsc – lateral sclerite; scarc – sclerotized arch; scare – sclerotized area.

Figure 8.

Cibarium of phlebotomine females. (A) Edentomyia piauiensis; (B) Brumptomyia sp; (C) Micropygomyia pilosa; (D) Micropygomyia cayennensis; (E) Evandromyia walkeri; (F) Sciopemyia sordellii; (G) Lutzomyia (Helcocyrtomyia) kirigetiensis; (H) Lutzomyia longipalpis; (I) Trichophoromyia auraensis; (J) Psathyromyia lutziana.

Figure 9.

Cibarium of phlebotomine females. (A) Australophlebotomus notteghemae; (B) Idiophlebotomus padillarum; (C) Sergentomyia (Vattieromyia) sclerosiphon; (D) Phlebotomus (Euphlebotomus) mascomai; (E) Phlebotomus (Madaphlebotomus) vaomalalae; (F) Sergentomyia bailyi; (G) Sergentomyia (Sergentomyia) phadangensis; (H) Sergentomyia hivernus; (I) Phlebotomus (Transphlebotomus) anatolicus; (J) Chinius eunicegalatiae; (K) Chinius samarensis; (L) Sergentomyia (Parrotomyia) babu; (M) Parvidens heishi; (N) Phlebotomus (Madaphlebotomus) vincenti; (O) Sergentomyia (Vattieromyia) namo.

Figure 10.

Sclerites of cervix and thorax of Phlebotominae. anp – anepimeron; apn – antepronotum; as – anterior spiracle; at – anatergite; csc – cervical sclerite with a pair of sensilla; hal – halter; kep – katepimeron; kes – katepisternum; kt – katatergitum; las – lower anepisternum; mnt – metanotum; mscx – mesocoxa; msf – mesofurca; mtcx – metacoxa; mtf – metafurca; mtp – metepimeron; mts – metepisternum; pem – proepimeron; pes – proepisternum; pnt – postnotum; ppn – postpronotum; pps – protuberance of the prosternum; prcx – procoxa; prf – profurca; ps – posterior spiracle; psct – prescutum; pt – paratergite; sclt – scutellum; sct – scutum; uas – upper anepisternum; vcsc – ventrocervical sclerite; win – wing. Deanemyia samueli.

Figure 11.

Setae on the thorax of phlebotomines: anps – anepimeral setae; dcs – dorsocentral setae; lass – lower anepisternal setae; mnts – metanotal setae; mtps – metepimeral setae; mtss – metepsiternal setae; pems – proepimeral setae; pscts – prescutal setae; psus – postsutural setae; sas – supralar setae; sctls – scutelar setae; uass – upper anepisternal setae; vcs – ventrocervical sensilla. Brumptomyia pintoi.

Figure 12.

Thorax in lateral view of Phlebotominae with indication of characters in plesiomorphic state: arrow – indicating the long suture separating the katepimeron and metepisternum; pas – postalar seta. Oligodontomyia toroensis.

Figure 13.

Metafurca of Phlebotominae: (A–C) Lateral view: (A) completely separate vertical arms and atrophied horizontal arms: Warileya phlebotomanica; (B) completely separate vertical arms and short horizontal arms: Warileya nigrosaculla; (C) completely separate vertical arms and long horizontal arms: Chinius samarensis. (D–H) Frontal view: (D) completely separate vertical arms and atrophied horizontal arms: Wa. phlebotomanica; (E) completely separate vertical and short horizontal arms: Chinius eunicegalatiae; (F) united long vertical and horizontal arms: Sergentomyia minuta; (G) united long vertical and short horizontal arms: Bichromomyia flaviscutellata; (H) united short vertical and long horizontal arms: Brumptomyia brumpti.

Figure 14.

Wing of Phlebotominae. (A) Main indices; (B) wing with of fusion of R2 and R3: Chinius eunicegalatiae.

Figure 15.

Abdominal tergites of Phlebotominae, showing arrangements of the deciduous bristles and tergal papillae and aspects of the tergal papillae and “trumpets glands”. (A, B) 2nd–5th male tergites with the arrangement of the deciduous bristles. (A) Two transverse bands: Warileya phlebotomanica. (B) Randomly: Nyssomyia intermedia. (C, D) Distribution and aspects of the tergal papillae on 4th tergite; (C) restricted to the central area: Lutzomyia longipalpis; (D) dispersed over the surface of the tergite, among the deciduous bristle scars: Pintomyia fischeri. (E–G) Aspects of the papillae on the 6th tergite: (E) papillae without hair and without clear demarcation of their borders; (F) papillae with hair: Brumptomyia cardosoi; (G) papillae without hair and with clear demarcation of their borders: Evandromyia walkeri. (H) “Trumpet glands” of the fourth abdominal tergite: Chinius samarensis.

Figure 16.

Abdomen and genitalia of Phlebotomine female. (A) abdomen in lateral view; (B) genitalia in lateral view; (C) 8th segment in ventral view; (D) 9th segment, 10th segment and cerci in ventral view. Nyssomyia neivai. gon – gonopod; st – sternite; stem – fork stem; terg – tergite.

Figure 17.

Some structures of the abdomen and the genitalia of phlebotomine females. (A) 3rd–6th abdominal segments showing the pleural setae: Lutzomyia (Tricholateralis) sherlocki. (B, C) 10th sternite showing non-deciduous setae in the median region: (B) Micropygomyia vexator, (C) Sergentomyia minuta. (D) 9th segment showing a sclerotized protuberance on the tergite: Migonemyia (Migonemyia) rabelloi, (E) spicules in 9th and 10th tergites: Lutzomyia (Tricholateralis) cruciata. (F) 9th tergite with short bristles: Sciopemyia sordellii.

Figure 18.

Spermathecae and genital fork aspects of Phlebotominae. (A) Brumptomyia brumpti; (B) Migonemyia (Blancasmyia) gorbitzi; (C) Viannamyia tuberculata; (D) Phlebotomus (Phlebotomus) papatasi; (E) Lutzomyia (Lutzomyia) longipalpis; (F) Dampfomyia (Coromyia) vespertilionis; (G) Dampfomyia (Dampfomyia) anthophora; (H) Micropygomyia (Coquillettimyia) vexator; (I) Evandromyia (Evandromyia) saulensis; (J) Ev. (Eva.) infraspinosa.

Figure 19.

Spermathecae and genital fork aspects of Phlebotominae. (A) Evandromyia (Barrettomyia) tupynambai; (B) Psathyromyia (Psathyromyia) lanei; (C) Pa. (Psa.) shannoni; (D) Pa. (Forattiniella) aragaoi; (E) Pa. (For.) lutziana; (F) Psychodopygus panamensis; (G) Ps. chagasi; (H) Trichophoromyia auraensis; (I) Bichromomyia flaviscutellata; (J) Martinsmyia alphabetica.

Figure 20.

Spermathecae and genital fork aspects of Phlebotominae. (A) Australophlebotomus notteghemae; (B) Chinius eunicegalatiae; (C) Chinius samarensis; (D) Phlebotomus (Madaphlebotomus) vincenti; (E) Ph. (Paraphlebotomus) sergenti; (F) Idiophlebotomus padillarum; (G) Parvidens heishi; (H) Ph. (Euphlebotomus) barguesae; (I) Ph. (Par.) chabaudi; (J) Phlebotomus (Larroussius) major; (K) Spelaeomyia moucheti; (L) Sergentomyia (Rondanomyia) goodmani comorensis; (M) Se. hivernus; (N) Ph. (Transphlebotomus) economidesi; (O) Se. (Vattieromyia) namo; (P) Se. (Sergentomyia) phadangensis; (Q) Sergentomyia (Ron.) goodmani; (R) Ph. (Par.) mireillae.

Figure 21.

Lateral view of male genitalia of Phlebotominae. (A) Nyssomyia neivai; (B) Warileya nigrosaccula.

Figure 22.

(A–C) Ventral view of the epandrium and of epandrial lobes and cerci of Phlebotominae. (A) Bruchomyiinae (Bruchomyia sp); (B, C) Phlebotominae: (B) Warileya nigrosaccula; (C) Psychodopygus chagasi. (D–I) Lateral view of epandrial lobe and cercus of Phlebotominae: (D) Hertigia hertigi; (E) Sciopemyia sordellii; (F) Psychodopygus chagasi; (G) Trichopygomyia longispina; (H) Evandromyia (Aldamyia) walkeri; (I) Evandromyia (Evandromyia) infraspinosa.

Figure 23.

(A–C) Lateral view of gonocoxites of Phlebotominae. (A) Brumptomyia brumpti; (B) Phlebotomus papatasi; (C) Migonemyia (Blancasmyia) gorbitzi; (D) Trichophoromyia auraensis; (E) Micropygomyia (Micropygomyia) pilosa; (F) Psychodopygus chagasi; (G) Lutzomyia (Tricholateralis) carvalhoi; (H) Lutzomyia (Lutzomyia) almerioi; (I) Pintomyia (Pifanomyia) verrucarum; (J) Phlebotomus hindustanicus; (K) Sergentomyia dentata; (L) Phlebotomus mireillae.

Figure 24.

Lateral view of gonostyles of Phlebotominae. (A) Micropygomyia chiapanensis; (B) Brumptomyia cardosoi; (C) Sergentomyia minuta; (D) Phlebotomus papatasi; (E) Edentomyia piauiensis; (F) Oligodontomyia toroensis; (G) Deanemyia samueli; (H) Micropygomyia pilosa; (I) Pintomyia (Pifanomyia) sauroida; (J) Migonemyia (Blancasmyia) gorbitzi; (K) Pintomyia (Pifanomyia) serrana; (L) Evandromyia (Evandromyia) correalimai; (M) Pressatia triacantha; (N) – Evandromyia saulensis; (O) Chinius samarensis; (P) Parvidens heishi; (Q) Sergentomyia – Sergentomyia (Sergentomyia) dentate; (R) Phlebotomus (Transphlebotomus) economidesi; (S) Idiophlebotomus padillarum; (T) Phlebotomus (Paraphlebotomus) andrejevi; (U) Phlebotomus (Legeromyia) multihamatus.

Figure 25.

Lateral view of gonostyles of Phlebotominae. (A) Viannamyia tuberculata; (B) Psathyromyia lanei; (C) Bichromomyia flaviscutellata; (D) Trichophoromyia auraensis; (E) Martinsmyia alphabetica; (F) Psychodopygus panamensis; (G) Psychodopygus bispinosus; (H) Psychodopygus geniculatus; (I) Psychodopygus chagasi.

Figure 26.

Lateral view of paramere and parameral sheath of Phlebotominae. (A) Migonemyia (Blancasmyia) gorbitzi; (B) Evandromyia (Aldamyia) walkeri; (C) Lutzomyia (Helcocyrtomyia) guderiani; (D) Psathyromyia (Psathyromyia) lanei; (E) Psychodopygus panamensis; (F) Psychodopygus chagasi; (G) Trichopygomyia longispina; (H) Trichopygomyia dasypodogeton; (I) Viannamyia tuberculata; (J) Pressatia triacantha;(K) Lutzomyia (Lutzomyia) longipalpis; (L) Lutzomyia (Lutzomyia) dispar; (M) Sergentomyia (Sergentomyia) dentate; (N) Phlebotomus (Legeromyia) multihamatus; (O) Idiophlebotomus padillarum; (P) Phlebotomus (Euphlebotomus) barguesae; (Q) Parvidens heishi; (R) Phlebotomus (Paraphlebotomus) andrejevi.

Figure 27.

(A–R) Lateral view of terminal region of aedeagal ducts of Phlebotominae. (S) Genital pump, aedeagal ducts and hypandrial apodemes (abdominal rods). (A) Blunt apex: Lu. longipalpis; (B) striated duct with blunt apex: Ev. brachyphalla; (C) apex beveled: Pa. shannoni; (D) curved and toothed apex: Ma. gasparviannai; (E) curved and beveled apex: Mi. longipennis; (F) duct curved in its preapical region and apex provided with appendix: Vi. tuberculata; (G) lozenge apex: Pa. runoides; (H) apex with barbs: Ny. yuilli pajoti; (I–J) bifurcated apex: (I) Ny. whitmani; (J) Nyssomyia anduzei, (K) ladle-shaped apex: Ny. intermedia, (L) spoon shaped or knife to eat fish shaped: Ny. neivai; (M) clavate apex: Ev. walkeri; (N) duct with curved preapical region and blunt apex: Pa. aragaoi; (O) strongly sclerotized bifurcated apex: Ev. lenti; (P) irregular side: Se. anka; (Q) enlarged at the top: Se. sclerosiphon; (R) with apical inflated portion: Id. padillarum. (S) Terminal region of aedeagal ducts with blunt apex: Id. nicolegerae.

Figure 28.

Parameral sheath of Phlebotominae. (A) With a transparent inferior top: Phlebotomus perfiliewi; (B) rounded with a knob at the top: Idiophlebotomus nicolegerae; (C) wide and short: Chinius eunicegalatiae; (D) with transparent top: Phlebotomus (Euphlebotomus) barguesae; (E) pointed: Phlebotomus (Paraphlebotomus) chabaudi; (F) rounded at the top: Sergentomyia (Vattieromyia) anka; (G) drumstick-like: Phlebotomus (Larroussius) major; (H) finger-like: Phlebotomus (Madaphlebotomus) vaomalalae; (I) with hooked top: Phlebotomus (Paraphlebotomus) mongolensis.