Issue
Parasite
Volume 24, 2017
Special Issue - ISOPS 9 - International Symposium on Phlebotomine Sandflies
Article Number 26
Number of page(s) 35
DOI https://doi.org/10.1051/parasite/2017027
Published online 21 July 2017

© E.A.B. Galati et al., published by EDP Sciences, 2017

Licence Creative Commons
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Introduction

Phlebotomine (Diptera, Psychodidae, Phlebotominae) taxonomy has been studied extensively, primarily due to the role of these flies as vectors of various agents, including species of Leishmania, Bartonella and arboviruses that cause infections in humans and other vertebrates. To date, approximately one thousand phlebotomine species have been described worldwide, although in varying languages and mostly without standardization of characters and terminology. During the Ninth International Symposium on Phlebotomine Sandflies (ISOPS IX) held in Reims from June 28 to July 1, 2016 [4], a round-table on Systematics was co-chaired by EAB Galati, P Lawyer, N Léger, and J Depaquit. We report in this paper the results of discussions on the following topics: the use of the informal term sandflies (or sand flies); methods for permanent preservation and mounting of phlebotomine specimens and appropriate places for deposition of type specimens; a recommendation regarding terms used in morphological descriptions of sand flies and their synonyms, illustrated by several drawings with captions; the characters to be used in taxonomic discussion and to be drawn when a new species is described, and some comments about the systematics of phlebotomines and integrative taxonomy.

On the use of the common name sand flies

Throughout history, there have been discussions on the use of the popular term sandflies (as one word) or sand flies (two words). Beyond the problem of its true form, this common name has also been applied to other dipterans (e.g. Ceratopogonidae and Simuliidae). Furthermore, some feel that the name does not seem appropriate to the habitat of Neotropical phlebotomine species, whereas others argue that the name was originally given to reflect the color rather than the habitat of these flies. Some colleagues suggested using the term “phlebotomine flies” to replace “sand flies” or “sandflies”; others suggested using “phlebotomine sand flies.” The discussion ended without achieving a consensus.

On type specimens

In the past and even more recently, phlebotomine type specimens have been deposited in institutions or schools other than museums [1, 9, 10]. It is very important to deposit at least the holotype in a collection under the supervision of a curator; however, political or administrative changes in these institutions can affect the specimens’ preservation and availability under conditions suitable for examination. Consistent with Articles 16C and 72F of the fourth edition of the International Code of Zoological Nomenclature (ICZN), we strongly recommend that authors of species descriptions deposit type and voucher specimens in one or more institutions that maintain a research collection with proper facilities for preserving them and making them accessible for study. It is critical for every institution in which material is deposited to ensure that all name-bearing types are clearly marked and recognizable as such. Moreover, the repository must be capable of taking all necessary steps to preserve these specimens, make them accessible for study, publish lists of name-bearing types in their possession or custody and, in so far as possible, communicate upon request information concerning name-bearing types [6]. In addition to depositing type and voucher specimens, we recommend registration with Zoobank (http://www.zoobank.org) for all nomenclatural acts (published usages of scientific names of animals, which affect their nomenclature or the typification of a nominal taxon), publications of original taxonomic descriptions, and references to authors and designation of type specimens.

Methods of preserving phlebotomine specimens

Many type specimens of phlebotomine species and other specimens in many reference collections were mounted in non-permanent media such as Hoyer’s Medium or Berlese and have long since deteriorated so that many of the diagnostic characters of the specimens are neither visible nor distinguishable, thereby rendering the specimen useless. It was proposed that a protocol using a permanent medium such as Canada balsam be adopted and standardized, in so far as possible, for preservations of all specimens that are to be deposited in museums for future reference.

Suggested guidelines for the description of new phlebotomine species

Many pertinent characters of Phlebotominae that are currently used for their identification and classification have been described by authors using distinct terminology. This practice has been criticized by other dipterists because, in addition to hindering studies of homology within Phlebotominae, it also makes comparisons among supra-specific taxa difficult. This is particularly problematic for characters of the male terminalia. To this end, we suggest that terminology used in phlebotomine taxonomy be more closely aligned with that used in general dipterology [2, 7]. In order to standardize the descriptions and re-descriptions of phlebotomine species, a list of characters (all body parts and their appendages and sensorial structures, such as spines, setae, papillae, sensilla, sutures, etc.) deserving of comment and illustration is proposed (Table 1). Furthermore, in order to contribute to the standardization of terminology for structures that have been frequently used in phlebotomine taxonomic studies, a list is provided (Table 2) including suggested terms in English and synonyms of these terms that have been used in the past by various authors, in French and Portuguese. Some characters have been infrequently used in published descriptions and may thus be poorly known; for this reason and for the purpose of illustrating the morphological diversity of some characters, drawings have been provided with the corresponding terminology in each legend (Figures 128). A drawing is preferable to a photo, but sometimes for structures such as the pharyngeal armature, a photo can supplement or replace a drawing. We posit that the list of characters suggested permits a detailed description of the species and provides elements for species distinction and phylogenetic studies.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail Figure 14.

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

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail Figure 21.

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

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

thumbnail 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.

Table 1.

Suggestion of the characters and the respective terminology used for the description of a new phlebotomine fly species. L = length; W = width; M = male; F = female; X = include the information.

Table 2.

Suggested terminology for main characters for the description of a new phlebotomine fly species and the most common synonyms; French and Portuguese translations.

Lastly, an update to the previous proposal for abbreviating the names of genera and subgenera [8] is presented in Table 3.

Table 3.

Proposed abbreviations for genera and subgenera of Phlebotominae.

Other taxonomic approaches

It is becoming more straightforward to distinguish phlebotomine taxa using modern techniques. The use of statistical approaches, such as models based on discriminant or multivariate analyses used in morphometric studies, may also contribute to the identification of intra- and inter-specific differences.

Concerning the identification of species using methods other than traditional dichotomous keys, the development of cybertaxonomy may facilitate this task for non-specialists or when many specimens need to be identified. The limited number of characters used by this tool may restrict its application in the case of a diverse fauna on a large scale.

One of the significant challenges facing phlebotomine taxonomists is the adoption of an integrative taxonomic approach, with an increase in attention to identifying characters to be used for accurate and efficient delimitation of species. Well-known, common adult morphological and morphometric characters are important; however, others such as behavioral, biochemical, ecological, and molecular data need to be considered, as well as morphological or developmental characters related to eggs, larvae, and pupae. It is highly recommended that markers used in molecular analysis for the delimitation of taxa be standardized and gene sequences should be deposited in free-access databases to permit the analysis of species or populations, especially for widespread taxa. This information is also valuable for phylogenetic studies.

The taxonomy of American phlebotomines has recently been updated regarding the number of described species/subspecies (502: 17 fossil and 485 extant). Additionally, it was commented that there has been an increase of 16.5% in the number of species described since Galati’s classification [3] and of 22.5% since that of Young and Duncan [11], though, in this latter case, only for the groups that these authors included in Lutzomyia. It was also commented that some taxa had been resurrected, some other species had been included as junior synonyms and two artificial taxa had been excluded from the species list. It is important to note that Galati’s classification has been updated annually and is available on the website: www.fsp.usp.br/~egalati [5].

Finally, the need to strengthen the training of taxonomist groups around the world was emphasized in view of the fact that taxonomy is the basis of the eco-epidemiological studies of vector-borne diseases. In recent years, with the advances of new technologies, particularly the molecular approach, a low level of interest on the part of young researchers in classical taxonomy has been observed that could lead to a significant loss of knowledge because expertise in this field depends on the strength of training and mentorship over successive generations.

Acknowledgments

The authors are grateful to the two reviewers, José Dilermando Andrade Filho and Gregory Curler, for their work on this manuscript.

References

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  2. Cumming JM, Wood DM. 2009. Adult morphology and terminology, in Manual of Central American Diptera, vol. 1, Brown BV, Borkent A, Cumming JM, Wood DM, Woodley NE, Zumbado MA, Editors. National Research Council of Ottawa: Canada. p. 9–502. (In the text)
  3. Galati EAB. 2003. Classificação de Phlebotominae, in Flebotomíneos do Brasil. Rangel EF, Lainson R, Editors. Ed. Fiocruz: Rio de Janeiro. p. 23–51. (In the text)
  4. Galati EAB. 2016. Phlebotominae (Diptera, Psychodidae): Classificação, morfologia, terminologia e identificação de Adultos, Apostila – Disciplina PSP5127-1 Bioecologia e Identificação de Phlebotominae. Public Heath School. University of São Paulo, www.fsp.usp.br/~egalati. (In the text)
  5. Depaquit J, Pesson B, Augot D, Hamilton JG, Lawyer P, Léger N. 2016. Proceedings of the IX International Symposium on Phlebotomine Sandflies (ISOPS IX), Reims, France, June 28th–July 1st, 2016. Parasite, 23, E1. [CrossRef] [EDP Sciences] [PubMed] (In the text)
  6. International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, 4th edn., London, UK: The International Trust for Zoological Nomenclature. (In the text)
  7. McAlpine JF. 1981. Morphology and terminology – adults, in Manual of Nearctic Diptera, vol. 1, McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM, Editors. Research Branch Agriculture: Canada, Monograph No. 27, Newstead R 1912. Notes, 9–23. (In the text)
  8. Marcondes CB. 2007. A proposal of generic and subgeneric abbreviations for Phlebotomine sandflies (Diptera: Psychodidae: Phlebotominae) of the World. Entomological News, 118, 351–356. [CrossRef] (In the text)
  9. Rahola N, Depaquit J, Makanga BK, Paupy C. 2013. Phlebotomus (Legeromyia) multihamatus subg. nov., sp. nov. from Gabon (Diptera: Psychodidae). Memórias do Instituto Oswaldo Cruz, 108(7), 845–849. [CrossRef] (In the text)
  10. Ubeda Ontiveros JM, Morillas Marquez F. 1983. Designacion del holotipo de Phlebotomus fortunatarum Ubeda Ontiveros y col., 1982 (Diptera, Phlebotomidae). Revista Iberica de Parasitologia, 43, 307–308. (In the text)
  11. Young DG, Duncan MA. 1994. Guide to the identification and geographic distribution of Lutzomyia sand flies in Mexico, the West Indies, Central and South America (Diptera: Psychodidae). Memoirs of the American Entomology Institute, 54, 1–881. (In the text)

Cite this article as: Galati EAB, Galvis-Ovallos F, Lawyer P, Léger N & Depaquit J: An illustrated guide for characters and terminology used in descriptions of Phlebotominae (Diptera, Psychodidae). Parasite, 2017, 24, 26.

All Tables

Table 1.

Suggestion of the characters and the respective terminology used for the description of a new phlebotomine fly species. L = length; W = width; M = male; F = female; X = include the information.

Table 2.

Suggested terminology for main characters for the description of a new phlebotomine fly species and the most common synonyms; French and Portuguese translations.

Table 3.

Proposed abbreviations for genera and subgenera of Phlebotominae.

All Figures

thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail Figure 14.

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

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail Figure 21.

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

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text
thumbnail 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.

In the text

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